Open Collections

UBC Theses and Dissertations

UBC Theses Logo

UBC Theses and Dissertations

Systematics of Saxifraga rufidula and related species from the Columbia River gorge to southwestern British… Perkins, Walter Ethen 1978

Your browser doesn't seem to have a PDF viewer, please download the PDF to view this item.

Item Metadata

Download

Media
831-UBC_1978_A1 P47.pdf [ 7.52MB ]
Metadata
JSON: 831-1.0094491.json
JSON-LD: 831-1.0094491-ld.json
RDF/XML (Pretty): 831-1.0094491-rdf.xml
RDF/JSON: 831-1.0094491-rdf.json
Turtle: 831-1.0094491-turtle.txt
N-Triples: 831-1.0094491-rdf-ntriples.txt
Original Record: 831-1.0094491-source.json
Full Text
831-1.0094491-fulltext.txt
Citation
831-1.0094491.ris

Full Text

SYSTEMATICS OF S J ^ I F R J G A BOFIDULJ AND BELATED S P E C I E S FBCM THE COLOMBIA BIVEB GOBGE TO SOUTHSESTEBN B B I T I S H COIUUBIA  By HALTER ETHEN PERKINS B.Sc. , U n i v e r s i t y o f O k l a h o m a , 1S73 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOB THE DEGREE OF DOCTOB OF PHILOSOPHY in THE FACULTY OF GRADUATE STUDIES {Department of Botany)  Se a c c e p t t h i s t h e s i s a s c o n f o r m i n g required  t o the  standard  THE UNIVERSITY OF B B I T I S H COLUMBIA October,  1978  (8) W a l t e r E t h e n P e r k i n s , 1978  In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of B r i t i s h Columbia, I agree that the  Library shall make it freely available for reference and study.  I further agree that permission for extensive copying of this for  thesis  scholarly purposes may be granted by the Head of my Department or  by his representatives.  It  is understood that copying or publication  of this thesis for financial gain shall not be allowed without my written permission.  W. E t h e n  Department of The  Botany  University of B r i t i s h Columbia  2075 Wesbrook Place Vancouver, Canada V6T 1W5  5 O c t o b e r , 1978  Perkins  Research  S u p e r v i s o r : D r . R a t h e r i n e I . Beamish.  ABSTRACT  In  the  variation  Pacific  have  produced  Saxifraga rufidula entities  from  difficult  to  recognized  several  one  highly  the  of  species  meiosis,  and  observations  of  Polyploid  Biver  study and  of  related  of  pollinators,  treatment  and  occidentalis nomenclature S. o c c i d e n t a l i s S. o c c i d e n t a l i s *>• o c c i d e n t a l i s  as  a  have  recognize  taxa  natural  hybrid  system  experiments, observations.  with  introgressant  w i t h t h e S. i n t e g r i f o l i a of  morphological,  exist  dent a t a ,  and  S. r u f i d u l a  to  ecological  becomes becomes  v a r . , o c c i d e n t a l i s becomes S.  the  latipetiolata,  S. o c c i d e n t a l i s to  be  species  substantiate  var.  becomes  var., latipetiolata dentata  to  of  studies  many o f w h i c h a p p e a r  species. , According  var.  systematic  studies,  and  plants  o f S. r u f i d u l a . S. O c c i d e n t a l l s var.,  the  ecological  discontinuities  S. o c c i d e n t a l i s  authors  subspecif i c  breeding  and  hybridization  geographic  The  particularly  while others  artificial  complex. S u f f i c i e n t c o r r e l a t i o n s and  Some  from n u m e r i c a l  populations,  intermediates  result  are  approached  c h a r a c t e r i s t i c s a r e shown t o o c c u r , the  among  i t s relatives.  Gorge  from t h a t area  with data  observations  individuals  polypoid  widely d i s t r i b u t e d species with v a r i e t a l  present  Wats.  and  i n the r e l a t i o n s h i p s  taxoncmically.,  S. r u f i d u l a  S. o c c i d e n t a l i s of  confusion  Columbia  separate  The  hybridization  ( S m a l l ) J a m e s Macoun a n d  variable,  components. treatment  Northwest,  the S. S.  var.  rules  of  aeguidentata. latipetiolata,  S. g o r m a n i i . Occidentalis.  and  ii  Table Of Contents  ABSTRACT .  ..  ..  .* .  '/  LIST OF TABLES  i  iv  LIST OF FIGURES . .:. .......  .. .... .. .......,>. y*  AC K NOWL E EG E H ENT S .................... ,<••• • •  .... . .....  INTRODUCTION ......... .... . . . . . . . . . . . . . , , . . . . . . . / MATERIALS AND METHODS .v.w. .V. .V. »V  >v>./.  V  w.-, .-.V.-. .  .V. V.  «^ . Y  ix 1 6  Chromosomal S t u d i e s  13  H y b r i d i z a t i o n Studies ...................................  14  Numerical S t u d i e s ... .............. ...................... .  15  Breeding  17  Systems  ....... .... .... ...... •. .-.........  Habitat And P o l l i n a t i o n S t u d i e s  18  RESULTS AND DISCUSSION  19  Morphology .....».........•  19  Chromosomal Studies ......... ............................  42  H y b r i d i z a t i o n .............................. ...... .......  63  Numerical S t u d i e s .......................................  87  Breeding  114  System Observations  •• ••  i i i  B a g g i n g T e s t s •».......*.........* * . • . « . * . . . .  •.. 114  F l o r a l Observations G e o g r a p h y , H a b i t a t And P o l l i n a t i o n  118 Ecology Observations  . 121  TAXONOMY . . . . . . . . . . . . . . . . . . . . . . . . . . . . ... . . V . . . . . . . . . . . . . . . .  130  S p e c i e s D e f i n i t i o n I n T h i s Complex- •..«-. i*>-..>V»«-*v»VV»-*V»»- .' 130 Key To The S p e c i e s ......................................  131  CONCLUSIONS  145  LI TEE AT 0 EE CITED  155  APPENDIX .... . . . . . . , . i . ..... . . . . . . . . . . . • .. ... ..... • • I n s e c t S p e c i m e n s ........................................  161 161  iv  L I S T OF TABLES  Table I , L i s t  of C o l l e c t i o n  Table I I . L i s t  of  Sites  Morphological  Computer A n a l y s i s . . . v . . Table I I I . ,  Summary o f  9 Characters  Used f o r  v . . . . v . . . . . . . . . . . . . . . . 16  Chromosome  Number f o r  Natural  P o p u l a t i o n s ...................... .... ...... ............ 44 Table IV. P o l l e n F e r t i l i t y  in Artificial  Hybrids  ........76  T a b l e V. Baw P r i n c i p l e C o m p o n e n t s f o r PCA A n a l y s i s ........99 Table VI., Bagging Compatibility  Tests f o r Automatic  ..................,.  Self ing  and  Self 115  T a b l e V I I . . B a g g i n g T e s t s f o r O u t c r o s s i n g and A p o m i x i s .....116 Table V I I I .  A v e r a g e S o i l D e p t h B e n e a t h P l a n t s ............. 125  T a b l e I X . Amounts o f S o i l  M o i s t u r e Under P l a n t s ...........126  T a b l e X. Summary o f I n s e c t V i s i t o r s .....•....•.•........••128  V  L I S T OF  Figure  1 : Map o f C o l l e c t i o n S i t e s .  Figure 2 : J?» Figure  Cut-away  occidentalis 3 :  var.  F l o w e r and  Fruit  Flower  Drawings o f  o c c i den t a l i s , S. r u f i d u l a . S. i n t e q r i f o l i a  S. i n t e a r i f o l i a var.  vari  Drawings o f  S,.occidentalis  s.--, o c c i d e n t a l i s  var.  inteqrifolia,  :  Var.  claytoniifolia. and S.  inteqrifolia  l e p t o p e t a l a . ................................... 23  4 :  Sketch  of  S. o c c i d e n t a l i s Plants. Figure  .......................7  v a r . l a t i p e t i o l a t a . ••/.... .............. 20  Cut-away  dentata,  Figure  FIGURES  5 :  Artificially  Hybrid  and  Produced S . , t a f i d u l a  .Representative  ... ................. Sketch  of  x  Parental  ,•>•  Artificially  27  P r o d u c e d S. r u f i d u l a X  S. O c c i d e n t a l l s - v a r . d g n t a t a H y b r i d a n d R e p r e s e n t a t i v e P a r e n t a l P l a n t s . ......... • Figure 6 : Sketch J.  29  of A r t i f i c i a l l y  inteqrifolia  Hybrid  Produced  and  S. r u f i d u l a  Representative  x  Parental  P l a n t s . ,....-»....... * .................................... 31 Figure 7 : Sketch  of Artificially  S. o c c i d e n t a l i s Representative Figure  Representative  var.  latipetiolata.  of A r t i f i c i a l l y  S. o c c i d e n t a l i s  F i g u r e 9 : Leaf  S. r u f i d u l a  x  H y b r i d and  P a r e n t a l P l a n t s . .................... y . . . 3 3  8 : Sketch  x  var.  Produced  Produced  var.  S. o c c i d e n t a l i s  dentata  H y b r i d aud  P a r e n t a l P l a n t s . .,......,....,.,........ 35 Outlines of  occidentalis.  latipetiojata.  S. r u f i d u l a ,  S. o c c i d e n t a l  S. o c c i d e n t a l i s  a n d S. i n t e g r i f p j j - i a  var.  var.  inteqrifolia  vi  v. . .... Figure  10 :  Leaf  Outlines  S. o c c | d e n t a l i s Figure  11 : Camera  Chromosome in Figure  Figure  12 : Camera  13 :  Camera  Lucida  H y b r i d s and  14 :  Showing  Population  o f S.  15 :  Drawings  Showing  Population  o f S.  16 :  Lucida  C e l l s from  Drawings o f  rufidula  Drawings  Lucida  var.  ...........................•  Number o f  Meiotic  «,...... y .59  Showing  Cells cf  clavtcniifclia •.... .61 Variation i n  Figures i n  Artificial  .. .... v . . . . . . . . . v . . . . ..........................66  19 : P r i n c i p a l  Depicting A r t i f i c i a l 20 :  Drawings  var.  Meiotic  var.  .55  Heiotic  Re l a t i v e s .  Drawings o f  S. i n t e q r i f o l i a  Lucida  dentata.  Showing  Populations.  18 : Camera  Variation i n  S. o c c i d e n t a l i s  and S. i n t e q r i f o l i a Lucida  Cells cf  o c c i d e n t a l i s . .53  and  Hybrids.  Populations of  Showing  Drawings  Numbers f o r  Camera  Chromosome  Meiosis i n  Meiotic  var.  Number f o r S. o c c i d e n t a l i s Camera  17 :  Showing  S. o r e g a n a  Figure  Variation in  49  Drawings  Normal  Lucida  Camera  latipetiolata  Figure  rufidula  ..........................47  and S. o c c i d e n t a l i s  Chromosome  Figure  Variation i n  i n t h e C o l u m b i a R i v e r G o r g e . .............. 51  Camera  Chromosome  Figure  Tetraploid  t h e C o l u m b i a R i v e r Gorge  5. r u f i d u l a  Figure  Lucida  and  ........................40  Drawings  Number f o r One  S. r u f i d u l a  Figure  Lucida  t h e Columbia R i v e r Gorge.  Cryptic  Figure  Diploid  dentata.  Number f o r One  Chromosome in  var.  of  38  Principal  Components  Analysis  Axis  1 vs.  H y b r i d s and P a r e n t a l P o p u l a t i o n s . Components  Analysis  Axis  2 .68  1 vs. 3  vii  Depicting Figure  Artificial  21 : P r i n c i p a l  Depicting  Components  Artificial  Figure  22 ; C r o s s i n g  Figure  23 :  H y b r i d s and P a r e n t a l P o p u l a t i o n s .  H y b r i d s and P a r e n t a l  Success Chart.  S. i n t e g r i f o l j a  var.  T e t r a p l o i d S. o c c i d e n t a l i s 24 :  Scatter Plot  Between  Depicting  S. i n t e q r i f c l i a  var.  25 :  Scatter Plot  Between S. r u f i d u l a S. O c c i d e n t a l l s Figure  26 :  Introgression  c l a y ton i i fo 1 i a  Character  Introgression  claytcniif olia  Figure  from t h e Columbia  Introgression  River  Gorge and  ..................84  T o t a l S a m p l e Osed i n C l u s t e r i n g  Figure  Populations.  30 : H y p e r s p a c e  Cloud  1 v s . 3 from  Populations.  ..95  29 : P r i n c i p a l C o m p o n e n t s A n a l y s i s A x i s  Canonical  1 vs. 2 frco  P o p u l a t i o n s . ........................93  28 : P r i n c i p a l C o m p o n e n t s A n a l y s i s A x i s  Samples of N a t u r a l Figure  ..86  27 : P r i n c i p a l Components A n a l y s i s A x i s  Samples o f N a t u r a l  and  .... ...... ................82  var. ocoidentalis .  Samples o f N a t u r a l  and  . ........ ....80  P r o g r a m , UBC-CGBOUP. Figure  3  Populations.,.72  Depicting Character  Dendrogram o f  2 vs.  ....................... 74  v a r . dentata  S y m p a t r i c S. r u f i d u l a P l a n t s . Figure  Axis  Scatter P l o t Depicting Character  Between  Figure  Analysis  .70  P l o t of  Variables i n  Bepresentation  SDF  2 vs. 3 from  ........................97  Grouping  Relationships cf  A n a l y s i s Program and P o i n t  Of  S. i n t e q r i f c l i a  var.  leptop„etala . ......................................... 101 Figure  31 :  Individual  Points  in  Hyperspace  R e l a t i o n s h i p s a b o u t t h e Group Means f o r Sk var.  claytoniifclia  and  Cloud  integrifolja  S. i n t e q r i f o l i a  var.  viii  l e p t e p e t a l a . • •.. .•.......•................. .......... 104 Figure  32 :  Individual  Points i n  R e l a t i o n s h i p s about t h e Group Population Figure  33 :  dentata  var.  Hyperspace  34 :  a n d S. o c c i d e n t a l i s  Individual  R e l a t i o n s h i p s about t h e Gorge  S. i n t e q r i f o l i a Figure  35 :  Artificial Figure  36 :  •S. r u f i d u l a Figure  37 :  var. occidentalis  Fiqure  Cloud  Means f o r C o l u m b i a Vancouver  the  Points i n Group  River Island  Olympic  Columbia  Cloud  S. r u f i d u l a Mountains and  River  Gorge  with  ..........................,.«>......112  Speculative and R e l a t e d  Polyploid  Relationships i n  T a x a . ......................... 151  Distribution  Map o f  and S. o c c i d e n t a l i s  S. x S.  gormanii.  inteqrifolia  ... •........... .... ..... ............. 162  Distribution  S. l a t i p e t i o l a t a .  Hyperspace  Means f o r  I s l a n d and t h e  Hybrids.  Population.  38 :  ... .......108  Hyperspace  and  from t h e  S» o c c i d e n t a l i s m Hybrid  Group  Individual  Vancouver  S. r u f i d u l a  Cloud  . ......................... ... .......... 110  R e l a t i o n s h i p s about from  Points i n  S. i n t e q r i f c l i a  . ..106  Means f o r S. c c c i d e n t a l i s  . .................. ...... . . . . . . . . . Figure  Hybrid  latipetiolata  Points i n  R e l a t i o n s h i p s a b o u t t h e Group var.  Cloud  Means f o r H e x a p l o i d  and S. O c c i d e n t a l i s Individual  Hyperspace  Map o f  S. r u f i d u l a  and  ..... .y«. ..... . • •>•;#•• .. «... ........... .164  ix  ACKNOWLEDGEMENTS "The j o u r n e y n e e d n o t be a l o n e a t a l l moments. He c a n a n d do s p a r k o n e a n o t h e r , a n d c a r r y e a c h o t h e r o n . " Paulus To  Berensohn  many who h a v e h e l p e d  insufficient, appreciation.  but, alas,  along the  way  simple  thanks  are  t h e y a r e a l l I have t o d e m o n s t r a t e my  My w i f e , Mary Dee, knows a l l t o o w e l l t h e k i n d s o f  teamwork and e f f o r t  which  she has a p p l i e d  to  this  paper.  Her  c a p a c i t i e s as f i e l d a s s i s t a n t , data r e c o r d e r , computer o p e r a t o r , illustrator, of  f r i e n d and m o t h e r were i n v a l u a b l e i n t h e c o m p l e t i o n  t h i s l a b o r o f l o v e . My d a u g h t e r ,  j o y s and much n e e d e d d i v e r s i o n s final  phases  Rachel, provided  when  she  during  o f t h i s work. Kay B e a m i s h h a s b e e n a most  a d v i s o r and r e s p e c t e d f r i e n d t h r o u g h o u t travel,  arrived  this  study.  Careers  Beamish,  a  O.B.C. , G r a d u a t e  »75 S t i p e n d , and a U.B.C. S t u d e n t  thoughtful  the final  by my  good  provided in  Funds  for  Travel Stipend t o the  manuscripts, friend,  The f l o w e r c l o s e - u p d r a w i n g s  Joan  his statistical  reading c e r t a i n  numerous h e r b a r i u m many  Miller.  Gary  Bradfield  for  reading  were done generously  e x p e r t i s e , c o m p u t e r p r o g r a m , and t i m e  sections of this  paper.  Johann  van  o t h e r s gave v a r i o u s k i n d s o f s u p p o r t each  Seenen  assisted i n  d u t i e s and o p e r a t i o n a l a s p e c t s o f t h i s  a l c n g t h e way i t i s d i f f i c u l t t o t h a n k non-human  thanks  s u g g e s t i o n s a l o n g t h e way a n d c a r e f u l  i d e n t i f i e d t h e i n s e c t s p e c i m e n s and J o h n P i n d e r - M o s s  So  helpful  Fellowship,  a u t h o r . The members o f my a d v i s o r y c o m m i t t e e d e s e r v e  of  the  e q u i p m e n t a n d e x p e n s e s were p r o v i d e d by an N.R.C, G r a n t  (No. ,67-6901) t o D r .  their  intangible  study.  and encouragement one  here.  Of  the  c o h o r t s , t h e UBC c o m p u t e r was c f g r e a t u s e f u l n e s s f o r  many a s p e c t s o f t h i s s t u d y .  But m o s t l y  there  are  the  plants.  X  whose s p i r i t s  l i v e on where t h e w i l d t h i n g s grow.  xi  1 S o r t o f a Song  L e t t h e snake wait his  under  weed  and the w r i t i n g be o f w o r d s , s l o w to s t r i k e ,  and g u i c k ,  sharp  guiet to wait,  sleepless.  Through metaphor t o r e c o n c i l e the  p e o p l e and t h e s t o n e s . Compose.  but  (No i d e a s  i n things)  Invent!  S a x i f r a g e i s my f l o w e r the  that  splits  rocks.  -William Carlos  Williams  (1883-1963)  1  INTBODOCTION  The  genus  Saxifraga  L. i s composed o f h e r b a c e o u s , m o s t l y  p e r e n n i a l , p l a n t s which a r e found i n a r c t i c , places  i n  temperate  restricted  mostly  Saxifraga,  comes  fragere  meaning  reference likely  to  regions.  to from to  the  the  The  Northern  the  break  of  the  or  habitats  genus  rocky  i s c i r c u m b o r e a l and  Hemisphere.  The  fragment,  which  for the  and  i s  name,  probably  a  many members o c c u p y . A l e s s t o the f o l k  treatment  use  of  of  kidney  ( S p o n g b e r g 1 9 7 2 ) . E t h n o b o t a n i c a l u s e s o f Saxi,f r a g a as  and  L a t i n w o r d s , s a j s i m e a n i n g r o c k and  d e r i v a t i o n c o u l d be a t t r i b u t e d  members  genus  alpine  some stones  species  are  a l e a f e x u d a t e f o r t r e a t m e n t o f s u p e r f i c i a l wounds and b o i l s  (Hunan 1 9 7 4 ) , a s a r o o t e x t r a c t u s e d f o r t h e t r e a t m e n t o f a wide range o f d i s o r d e r s from a c h i l d ' s t e e t h i n g dysentery  pains  to  relief  of  (Watt 1 9 7 2 ) , a s a l p i n e r o c k g a r d e n p l a n t s , and as t h e  hanging basket h o u s e p l a n t ,  S. s a r m e n t o s a  Schrceder,  known  as  strawberry begonia, s t r a w b e r r y geranium, or mother-of-thousands. Although s e v e r a l early b o t a n i s t s c o n t r i b u t e d g r e a t l y t o the knowledge the  of  t h e genus  comprehensive  brought  together  ( L i n n e a u s 1753, Don 1 8 2 2 , H o o k e r  monograph the  of  earlier  Engler  and  1833),  Irmscher  (1916)  t a x o n o m i c i n f o r m a t i o n and today  c o n s t i t u t e s t h e s t a n d a r d r e f e r e n c e f o r t h i s group o f p l a n t s . In North Irmscher  America  the  section  Boraphila  of  Engler  (1916) h a s two t e m p e r a t e c e n t e r s o f s p e c i e s  and  diversity,  one i n t h e E a s t e r n A p p a l a c h i a n R e g i o n a n d t h e o t h e r i n t h e R o c k y Mountain Region of t h e P a c i f i c  Northwest  (Spongberg  a r e a s were s t r o n g l y i n f l u e n c e d  by r e c e n t P l e i s t o c e n e  1972).  Both  glaciaticns  2  and  the e v o l u t i o n a r y h i s t o r y  many  respects,  a d v a n c e and evolving single  closely  of the p l a n t s i n t h o s e  linked  with  the  history  retreat. This i s especially evident i n  species  complexes  morphological  of  those  characteristics  areas.  the  name, r u f i d u l a , was  genus  Micranthes  subsequently Macoun  1887.  whose  Persistent  Vancouver  confusion  among S a x i f r a g a r u f i d u l a  (Small)  Northwest r e l a t i v e s  Kjvali-virginiensis present  time.  Early  variability  extant  recognizing  a  Bydberg  1905,  reduced  to  (Engler  and  within  large  number  Johnson two  this  1S23).  al.  1961,  of More  or three broadly  Hitchcock  and  the type  Irmscher,  was  James  material  Columbia  and  1916)  treatments  its  in  up  dealt  entities  recently,  close  subsection  related  specific  of  the r e l a t i o n s h i p s  Macoun  closely  It  by  i n the s e c t i o n Boraphila  monographic  entity  (1905).  British  these  to  the  with  the  group  by  (Small have  and been  d e f i n e d s p e c i e s complexes with  s u b s p e c i f i c or v a r i e t a l components ( B a c i g a l u p i et  one  find.,  Hydberg  surrounded James  and  rapidly  used t o d e s c r i b e a s p e c i e s  Island,  has  glacial  the  which s e p a r a t e  f a t h e r , John, c o l l e c t e d  F i n l a y s o n on  Pacific  Small  of  t r e a t e d as a p a r t o f the genus S a x i f r a g a  (1906)  f r o m Mt.  by  first  i s , in  I n such complexes  from i t s c l o s e r e l a t i v e s a r e o f t e n d i f f i c u l t t o The  areas  Cronguist  1973,  1944, K r a u s e and  Hitchcock Beamish  1972,1973). The  channeling  permitted  a  e l e m e n t s and otherwise plants  in  zone  of the C o l u m b i a R i v e r through of  contact  between d r i e r i n t e r i o r  more m e s i c c o a s t a l s p e c i e s . The  generally  ecologically  situations  where  i t s gorge  and  flcristic  intimate contact  geographically  environmentally  has  of  isolated  intermediate  or  3  perhaps unique micro-habitats are a v a i l a b l e a p p a r e n t l y has l e d t o complicated  p a t t e r n s o f v a r i a b i l i t y and  evolution within several  groups of p l a n t s , i n c l u d i n g t h e Clay_tonia complex  (Miller  1976),  and  § 3 1 gg|?*i?-53ffl- j-dahoense- l i t t o r a l e (Henderson  the  authors  have  morphological  intermediary  S.  S.,Wats.  occidentalis  subspecies  occidentalis  Hitchcock of  and S.  transitional Hitchcock  commented  to  Cronguist  variety Gorge  variety  occidentalis  C.L.Hitchcock.  and  Saxif raga  extremely  rufidula  variety  a  variable entity  occidentalis  unexplored. Beamish  The  introgressant  the  entity  (1972)  recognized  (1975)  in  as  that  a S.  was  confirmed  the also in  latipetiolata  confirmed  by K r a u s e  was  and  considered  w i t h o u t s u b s p e c i f i c t a x a . They  Biver  i d a h o e n s i s had subspecies  characteristics  in  variety,  occidentalis  Columbia  marshallii  C.L.  Oregon. , H i t c h c o c k  new  a  rufidula  Irmsch.)  n o t e d t h a t t h e e x t e n t o f i n t e r g r a d a t i o n b e t wee n S. S.  as  (Piper)C.L.Hitchcock  r u f i dnla  as  S.  (1961),  (Small)C.L. Hitchcock  S,  taxa;  Hitchcock et a l .  ( E n g l , and  adjacent  (1973) a s a s p e c i e s and  subspecific  considered  allenii  described  two  treated  idahoensis  and  of  (1944),  (1S73)  S.  extent  B a c i g a l u p i , i n Shram* s f l o r a  dentata  and  the  and  Beamish an  group  rufidula  and r u f i d u l a .  variety  Biver  included S.  species  on  between  occidentalis  and  Columbia  Sisvrinchium  duodecaploid  t e n t a t i v e l y d i v i d e d S. o c c i d e n t a l i s i n t o  variety  polyploid  1976) .  Several  and  perfpliata  subsp.  the c l a s s i f i c a t i o n  was  b e e n t r e a t e d by  of  of  Gorge  rufidula  S.  and  largely  Krause  and  marshallii  but  they  idahoensis  has  some  ,S. o c c i d e n t a l i s . by H i t c h c o c k e t j | l .  Elvander (1961) ,  4 and  H i t c h c o c k and C r o n g u i s t  idahoensis,  rufidula.  v a r i e t i e s d e n t a t a and and  vary  (1973) o f S. o c c i d e n t a l i s  and  latipetiolata.  he  considered  a l i e n i i to lack morphological  continuously  conseguently,  He  with  treated  variety  them  as  varieties the  distinctions  occidentalis synonyms  of  and variety  occidentalis. None  of  addressed  these  the  problem  r e l a t i v e s o f S. (1961,1967)  previous of  systematic  Krause  and  Beamish  s t u d i e s on r e l a t i o n s h i p s m a i n l y  in  Elvander  Biver  G o r g e and o t h e r  (1975) b a s e d h i s s t u d y  chromatographic  evidence  R o n t a n a . He b r i e f l y d£JLt§il# and include  in  British  data  from  o f the  Gorge.  Columbia  plants  r u f i d u l a . S.  var.  populations  the  Washington.  from  Idaho  occidentalis  latipetiolatabut  of those  their  although  cytological,  largely  the  Eeamish  m a t e r i a l from  a r e a s o f O r e g o n and cn n u m e r i c a l ,  d i s c u s s e d S.  S. o c c i d e n t a l i s  Biver  (1972,1973) f o c u s e d  t h e y r e p o r t e d s e v e r a l chromosome c o u n t s Columbia  have  t h e c o n f u s i o n t h a t e x i s t s among  r u f i d u l a i n the Columbia  and  investigations  did  and and var. not  entities i n his basic  study. The  r e l a t i o n s h i p s among S.  occidentalis. S. and  occidentalis  S,,occidentalis var.  with  i n the  morphological approaches  dentata  and  into  Southwestern  paper. C o r r e l a t i o n s of  B.C.  are  hybridization  o b s e r v a t i o n s w i t h c y t o l o g i c a l s t u d i e s and  analyses  are presented  classification.  northward  present  s t u d i e s and e c o l o g i c a l  var.  var.  l a t i p e t i o l a t a i n the Columbia B i v e r Gorge  other areas extending  dealt  r u f i d u l a . S. o c c i d e n t a l i s  based  on  three  different  numerical  i n an e f f o r t t o p r o v i d e a more n a t u r a l  S i n c e S. r u f i d u l a a n d i t s  close  relatives  are  5  often S.  found  growing  Integrif olia  1916),  the  leptopetala  are  have  this  also and  taxa  some  paper.  close  at  d e n t a t a may  and  i n the and  to  group  of  here  and of  studies  Uashington  include  subsp.  S.  marshallii  l e p t p p e t a l a . The  idahpensis  Proposed comecclature of  changes the  are  (1972, 1973),  synonymous w i t h  taxa  var. in  and  var.  subsp.  species.  ferruginea Saxifraja  here as e q u i v a l e n t  p r e s e n t paper  treated  uses  to the  rufidula  and  as r e c o g n i z e d  by  ,S. o c c i d e n t a l i s  var.  ccci.dentaj.is. the  studied  Conclusions sections of t h i s  PhD  califcrnica*  r h o m b o i d e a . S,  columbiana i s t r e a t e d  in  marshallii.  S«  American  a  study  S a x i f r a g a •• o c c i d e n t a l i s  North  in to  (Elvander,  subsp. , Beamish  ties  sensu l a t o i s c u r r e n t l y  S.  and  discussed  A systematic  H i t c h c o c k f o r S. . , o c c i d e n t a l i s b u t 5 .  marshallii  study  i s b r o u g h t f o r t h aisd as  c o l u m b i a n a . S.  var.,  this  close  S. n i v a l i s ,  Eastern  var.  of  t e r m i n o l o g y of  a l l e n i i as  the  taxa  has  marshallii  reflexa,  var. of  j,ntegrjfclja  oreqana  S.  the  to species.  preparation).,  S.  integrifolia  Krause  Irmscher  sources  in  main t h r u s t  oreqana  key  S.  mentioned  isiiflfifsiia S.  potential  latipetiolata  introgress  taxa  a  and  of  claytoniifolia,  introgression  S.  University  in  idahoensis, J»  that  var.  the  dissertation,  Other  as  members  Engler  r e l a t i o n s h i p s with  Evidence  integrifolia  progress  included character  result i t i s included S.  of  inteqrifolia.  p e r i p h e r a l t o the  S.•..occidentalis  of  with  here.  Several may  Integrifcliae  varieties  hybridization presented  (sect.  sympatrically  classification appear  paper.  in  the  and  revised  Taxonomy  and  6  MATERIALS AMD METHODS  Live  plants,  pressed  specimens,  flower buds and s e v e r a l  s o i l samples were c o l l e c t e d from p o p u l a t i o n s from t h e the Columbia R i v e r e a s t to beyond  the Columbia  mouth  of  River Gorge. Some  c o l l e c t i o n s were a l s o made i n areas extending north and south i n the  lower  Columbia  River  area  and  adjacent  P o p u l a t i o n s from t h e Olympic Mountains, Island,  and Cascades i n Washington  B r i t i s h Columbia collections Fifteeen  ( F i g . 1,  Table  from  localities  came  to  populations  twenty  rosettes  Coast  Southeastern  Range.  Vancouver  and the Southern Mainland of  I)  were  also  with  from  a  two  sampled. sympatric  large  number  Many taxa.  of  the  sampled were t r a n s p l a n t e d t o c c l d f r a m e s f i l l e d with  a p e a t - v e r m i c u l i t e bedding mixture which  were  University  c o l l e c t i o n s of buds i n  of  British  Columbia.  Mass  located  at  the  meiosis were sampled. Buds from i n d i v i d u a l p l a n t s were c o l l e c t e d and a l a r g e number o f p l a n t s were pressed as Specimens  are on d e p o s i t a t the u n i v e r s i t y of B r i t i s h  Specimens  studied  classification,  from  for  herbarium  the  sheets  of Oregon  Berkely  Washington (WTU).  (UC),  British  State U n i v e r s i t y  Several  hundred  (MIN),  Brigham U.S.  (ORE) , Oregon  U n i v e r s i t y of B r i t i s h Columbia at  representing  following herbaria:  (BEY), U n i v e r s i t y o f Minnesota (NA) , U n i v e r s i t y  morphological  material. Columbia, comparison,  annotation, and geographic d i s t r i b u t i o n  1,507  a t o t a l of plants  voucher  National  5,000  Herbarium  State U n i v e r s i t y  Provincial  (MS) , and U n i v e r s i t y  additional  about  Young U n i v e r s i t y  (UBC), U n i v e r s i t y Columbia  included  of  California  Museum of  (OSC) ,  (V)  w  Washington  herbarium sheets were a l s o  7  F i g u r e 1: C o l l e c t i o n s i t e s i n t h e P a c i f i c Northwest f o r the present study. One collection from Berthoud Pass, C o l o r a d o , i s n o t shown.  9  • T a b l e L. C o l l e c t i o n s f o r Study. 7  Tjaxpn S. r o f i d u l a  Collection Code Number 616 608 610 612A 613A 637 673 675A 647A 76-1 617A 669A 672 6 18A 645A NMO-8  location V i e n t o , r e s t a r e a on I n t e r s t a t e 8, 0.9 mi w. o f V i e n t o S t . P k . , Hood B i v e r Co., O r e . T r o u t d a l e , 1.3 mi s . o f S a n d y B. B r i d g e , Multnomah Co., O r e . Yeon P k . , c a . 1.5 mi a l o n g t r a i l t o M c C o r d F a l l s , M u l t n o m a h Co. Ore. , Mayer P k . , 0.8 m i n. o f M a y e r S t . P k . , Wasco C o . , Ore. T h e D a l l e s , 2. 0 m i w. o f C h y n o w e t h C r . on O l d U.S. , Hwy. 3 0 , Wasco Co., O r e . M o s i e r , 1.0mi e. c f H o s i e r on O l d C o l u m b i a 8. Hwy., Wasco C o . , Ore, L a k e c r e s c e n t , 8.5 mi e. o f F a i r h o l m on U.S. Hwy.,101, C l a l l a m Co., Wash. . Mt. P l e a s a n t , 1.9 mi e. o f t h e C l a r k - S k a m a n i a Co. b d r y . on S t , Hwy. 1 4 , C l a r k Co. , Wash. W a s h o u g a l , 1 . 0 mi w. o f C l a r k S k a m a n i a Co. b d r y . on S t . Hwy. 14, C l a r k Co., Hash. Marmot P a s s , c a . 5 mi s. o f Camp M y s t e r y , s . e . o f Marmot P a s s , J e f f e r s o n C o , , Wash. B i n g e n , 5.6 mi e . o f B i n g e n , on S t . Hwy 1 4 , K l i c k i t a t Co., Wash. B i n g e n L k . , 7.4 mi e. o f w. b d r y . , o f K l i c k a t a t Co. on S t . Hwy. 14, Wash. C o c k , 9.8 mi w, o f e. b d r y , o f S k a m a n i a Co. o n S t . Hwy. 14, Bash., S k a m a n i a Co., 3 . 5 mi e. o f C l a r k S k a m a n i a Co. b d r y . on S t . Hwy. 1 Wash, C o l l i n s , 0.9 n i e. o f C o l l i n s Depot Bd. on S t . Hwy. 14, S k a m a n i a C o . , Wash. N a n a i m o , c a , 5 m i ' s , o f Nanaimo c n W h i t e R a p i d s Rd., V . I . , B.C.  10  Table I  t  Taxon  Collection Code Number NOOS-B UCL-R 623A 624B 627A  S. o c c i d e n t a l i s var. denata  JconJMt)  629A 606 630 607  Nanoose H i l l , c a . 15 m i n. o f Nanaimo o n Nanoose H i l l , V.I., B.C. Upper C a m p b e l l L k . , 7.4 mi w. o f C a m p b e l l L a k e B r i d g e , V . I . , B.C S o o k e , s . s i d e o f Sooke B. a t Sooke P o t h o l e s P k . , V.I.,B.C.. Hill Hill, C a p i t a l D i s t r . Pk., V i c t o r i a , V . I . , B.C. Mt. F i n l a y s o n , G o l d s t r e a m P r o v . P k . , V . I . , B.C. S a d d l e M t . , c n t r a i l t o summit near westernmost Peak, C l a t s o p Co., Ore. Delena, Beaver Creek F a l l s , 2.8 mi w. o f D e l e n a , C o l u m b i a Co. Ore, T i l l a m o o k C o . , 2.0 mi w. o f e. b d r y . o f T i l l a m o o k Co. c n S t , Hwy. 6, O r e . Kalama B., 3 mi e. o f I n t e r s t a t e 5 on Kalama B. B d . , C o w l i t z Co., Wash.  S. o c c i d e n t a l i s var. latipetiolata  629C  S a d d l e M t . , on t r a i l t o summit n e a r t o p , C l a t s o p Co. , O r e .  S. o c c i d e n t a l i s var. occidentalis  605  C h e h a l i s 8. , ( h y b r i d ) , 3.3 m i w. o f L i t t e l , L e w i s Co. Wash. , Mt. B a k e r N a t l , f o r e s t . Y e l l o w A s t e r Meadows, Whatcom Co., Wash, Y a l e , c a , 1.5 mi e. a n d n. o f Y a l e below C a n . Hwy. 1, B.C. Liumchen B i d g e , s. o f S a r d i s , B » C» Cornwall Lookout, se. o f Hat C r e e k , B.C. B o o t a n i e V a l l e y , Skwaha Mt. a r e a n. o f L y t t o n , B.C.  BMR  626 666 COEN 687 S. i n t e q r i f o l i a 641 var. claytoniifolia  G i l i a m C o . , 1.7 mi e. o f w, b d r y . G i l i a m Co. on I n t e r s t a t e 80 n . , Ore.  11  Table I. (con't)  HiLSSLE  Collection Code Number 609 611 612B 613B 667 692 675B 668 669B  Location T r c u t d a l e , 1.3 mi s. o f Sandy 8. B r i d g e , Multnomah C o . , Ore. Bowena, 2,4 mi w. o f Bowena o v e r p a s s on I n t e r s t a t e 80-N,, Hasco c o . , Ore, Mayer P k . , 0.8mi n. o f Mayer St. P k . , Wasco C o . , O r e . The D a l l e s , 2.0 mi w, o f C h y n o w e t h C r . on O l d U.S. Hwy. 3 0 , Wasco Co. O r e . B i g g s , 1.25 mi e. o f j e t . w i t h U,S Hwy. 97 on U.S. Hwy. 3 0 , Sherman Co., O r e . S t e p h e n ' s P a s s , 11 mi e. o f S t e p h e n ' s P a s s S k i Lodge on U.S. Hwy. 2 , C h e l a n C o . , H a s h . M t i P l e a s a n t , 1.9 mi e. o f t h e C l a r k - S k a m a n i a Co. b d r y . on St.Hwy. 14, C l a r k C o . , l a s h . L y l e , 1.6 mi n. o f j e t , w i t h S t . Hwy. 142, K l i c k i t a t C o . , Wash. B i n g e n L k . , 7.4 mi e. o f w. b d r y . o f K l i c k a t a t C o . o n S t . Hwy. 1 4 , Wash. ., Benchmark, U.S.G.S. V 4 2 5 , 1 S 6 8 , 5.2 mi e. o f The D a l l e s b r i d g e , K l i c k a t a t C o . , Wash. B i n g e n , 5.6 mi e. o f B i n g e n , on S t . Hwy 14, K l i c k i t a t C o . , B a s h . C l a r k Co. l i n e , b d r y . w i t h S k a m a n i a Co. on St.,Hwy, 14, Sash. :  642 617B 619  5A i n t e g r i f o l i a var. i n t e g r i f c ^ i a •  671 MIMA 6 23B 624B 627B  G r i z z l y L k . , a b o u t 0.5 mi downstream from G r i z z l y L k . S i s k i y o u Co., C a l i f o r n i a Mima Mounds, w. o f L i t t l e B o c k on B d . t o Mima, T h u r s t o n C o . , Wash. S o o k e , s . s i d e o f Sooke B. a t P o t h o l e s P k . , V . I . , B.C. Mill Hill, M i l l H i l l Capital D i s t r . P k . , V i c t o r i a , V.I.,B.C. Mt. F i n l a y s o h , G o l d s t r e a m P r e v , Pk. , V . I . , B.C.  12  T a b l e Ja. Tax on  ( c o n * t)  Ccllecticn Code Number NJ30-B NOOS-B 625 617B 78-1 682  S. i n t e g r i f o l i a var. l e p t o p e t a l a  648  S*. orjejgajja  BEBH  Location Nanaimo, c a , 5 mi s, o f Nanaimo on W h i t e B a p i d s Bd. , V. I . , E. C, N a n o o s e H i l l , c a . 15 mi n. o f Nanaimo on Nanoose H i l l , V . I . , B.C. Y a l e , c a . 1.5 mi e. a n d n. o f Y a l e b e l o w Can. Hwy 1, B.C., B i n g e n , 5.6 mi e. o f B i n g e n on S t . Hwy. 14, K l i c k a t a t C o . , Wash. H a r r i s o n L k . , 1.5 mi w. o f H a r r i s o n , H o t S p r i n g s , B.C. E l k B a l l s , below C a m p b e l l L k . Dam, V . I . , B.C.  P r i n c e t o n , o f f Hwy. e. o f P r i n c e t o n , road t c g o l f c o u r s e , B. C. B e r t h o u d P a s s , B d r y . o f G r a n d Co. Colorado  13  inspected  from m a t e r i a l  Chrcmosomal  on  loan  to the  U n i v e r s i t y of  Studies  S q u a s h e s o f a n t h e r t i s s u e n e r e made material fluid. and  in  Carnoy s f  fixing  bud  3:1,absolute e t h a n o l : g l a c i a l a c e t i c  acid  Buds were t h e n s t a i n e d  warmed i n a 40  were  Washington.,  preserved  in  using  Snow*s  C o v e n f o r a b o u t 24 Hoyer*s  by  permanent  first  (1963)  bulk  hours. S l i d e  method  preparations  medium ( f i l e x o p c u l c s  and  Beneke 1952) . Pollen s t a i n a b i l i t y of l i v e , was  measured by c o u n t i n g  stained  and  1958).  Large,  Preliminary  mounted  be  t o 200  investigations (1958)  correlated  i n v e s t i g a t i o n was  and  grains of  reports with  were  pollen that  flowers  w h i c h had  been  stain  (Sass  blue  considered  grain  pollen  chromosome  discontinued.  dried  pollen grains  lactophenol-aniline  dark-staining  Sokolovskaya•s to  in  up  preserved  size  fertile. confirmed  s i z e does not  number  and  appear further  14  HYbridlzatiGjp Studies  P l a n t s used as female water-resistant artificially Only  of  a  parchment  bags  cross-pollinated  crosses  successful.  p a r e n t s were e m a s c u l a t e d , bagged  involving  under  stigmatic  watery  appearance  surface.  lactophenol-aniline  a  stigmatic receptivity.  stain  Whole  flowering  and  microscope.  blooming  plants  was e s t i m a t e d f r o m  the  were onset  and p a p i l l a t e c o n d i t i o n o f t h e  Increased blue  to  dissecting  simultaneously  Stigmatic receptivity  slight  prior  with  stainability  (Sass  anthers  1958) were  also  used  using indicated  tc  transfer  p o l l e n masses t o t h e s t i g m a s . Samples  of  hybrid  w e l l - d r a i n e d sandy beneath in  about  1-2mm  large-mouthed  were  placed  conditions  o u t c r o s s e d seed  mixture  in  o f s a n d and a 1cm  early  misted  By  April  g e r m i n a t e d and l a t e r  December  under  ambient  periodically seedlings  in  rows  layer of pea-sized gravel water.  temperature to  of  maintain  nearly  were t r a n s p l a n t e d t o s a n d y  where t h e y were a l l o w e d t o grow f o r one the  were p l a n t e d i n a  p o t s s e t i n a s h a l l o w pan o f outside  and  environment.  soil  and  every  Plantings and  light  a  moist  cross  had  p r o p a g a t i o n beds  season. In the s p r i n g  of  s e c o n d s e a s o n s e e d l i n g s w e r e t r a n s p l a n t e d i n t o p o t s . As t h e y  flowered,  samples  were  examined  for  their  morphology  b e h a v i o u r i n m e i o s i s , then p r e s e r v e d as p r e s s e d specimens.  and  15  Numerical  Studies  Forty-three combinations plants  natural  for  infestations not  of  developmental  certain  rust,  were a v o i d e d of  treatment.  Population  whorl  after  minor  choosing of  flowering  artifically  variation  infestation  i t was  growing  produced Periodic  may  or  study  be  the Such  but  a  the result of  subsequent  pesticide  s a m p l e s c o n s i s t e d o f 8-10 p l a n t s c h o s e n a t transplantations  carefully  Flower  grown  o f a mature  dehiscence  of  removed from t h e o u t e r  measurements  and  by  i n cold  leaf  were  trying  to  each  rosette  standardized  f l o w e r s a s c l o s e as p o s s i b l e t o t h e s e q u e n t i a l  anther  weevils  plants.  wherever p o s s i b l e i n t h e  well-established  and pressed.  live,  treatments.  some  f r a m e s . L e a f measurements were t a k e n plant  character  mildews, s l u g s and r o o t  of  spurious  to  from  or  plant m o r t a l i t y but o c c a s i o n a l l y a l t e r e d  responses  random  aphids,  263  and  taxonomic  morphology  amount  on  populations  numerical  only increased  individuals  characters  ( T a b l e I I ) were measured  (from  hybrids)  quantitative  by  midpoint  obtain flowers  from  e q u i v a l e n t p o s i t i o n s i n t h e i n f l o r e s c e n c e . L i n e a r v a r i a b l e s were converted  t o metric v a l u e s , counted v a r i a b l e s  using  the  square  root  matrix  was  subjected  to  transformation three  were  transformed  and t h e r e s u l t a n t data  different  available  computer  p r o g r a m s . The h i e r a r c h i c a l c l u s t e r i n g a n a l y s i s p r o g r a m w h i c h was used  i s called  UBC-CGBOUP,  available  B r i t i s h Columbia Computing Center. o f Hard  (1963) a n d t h e c o d i n g  Principal  Components  the  The a l o g a r i t h m  i s modified  analysis  at  (PCA)  University of used  i s  that  f r o m Veldman  ( 1 9 6 7 ) . ft  program c a l l e d  PBINCOMPS  16  Table I I C h a r a c t e r s used i n N u m e r i c a l X  No. 1. 3. 5. 7,  No.  Character petal In. sepal In, g l a n d wd, style ht.  9. 11. 13, 15,  p e r i a n t h wd. f i l a m e n t wd. a t b o t t o m h a i r no. a t l e a f t i p h a i r no, on p e d i c e l  17,  angle o f l o w e s t b r a c t t o stem  19, h a i r no.  at lowest  branch  2 1 . h a i r no. on p e t i o l e 23. l o w e s t b r a c t l n . / w d .  ratio  25. s c a p e d i a m . a t l o w e s t 27. p l a n t h t . 29. i n f l o r e s c e n c e l n . / w d . 31. 33. 35. 37. 39.  branch ratio  i n f l o r e s c e n c e I n . from p a r t t o top l e a f ln./wd. r a t i o hair In. petiole angle l e a f shoulder p e t i o l e ln./wd. r a t i o  widest  41. s t y l e I n . • g l a n d h t . / gynoecium I n . 43,  leaf In./petiole In.  abbreviations:  Studies  ratio  Character  2. p e t a l l n . / w d . r a t i o 4. s e p a l l n . / w d . r a t i o 6. g l a n d l n . / w d . r a t i o 8. f i l a m e n t wd. a t middle/wd. a t bottom r a t i o 10. g y n o e c i u m h t . 12. p e d i c e l I n . 14.,hair In. l e a f t i p 16.,angle of lowest i n f l o r e s c e n c e branch 18. h a i r I n . a t l o w e s t branch 20. h a i r no. a t s c a p e bottom 22. l o w e s t b r a c t I n . 24. t e e t h no. on l o w e s t branch 26. i n f l o r e s c e n c e a n g l e 28. f i l a m e n t I n . 30. p l a n t I n . / I n . t o lowest branch r a t i o 32. l e a f I n . ;  34. t e e t h no. on l e a f 36. a n g l e o f l e a f t i p 38. p e t i o l e wd. 40. I n . o f t o o t h a t l e a f tip 42. g l a n d wd. s g u a r e d * f i l a m e n t In. sguared/ ( s t y l e In. + g l a n d In.) sguared.  ht.=h e i g h t , l n . = l e n g t h , no.-number, wd.=width  17  (available  from  Department  of  Brad f i e l d . Botany)  University  was  used  c o r r e l a t i o n m a t r i x w h i c h was eigenvector second  matrix.  vs..third  groups  First  hybrid  and a n a l y s e d u s i n g a program  hyperspace  Breeding  plots  upon  first  were  standardized to  form  vs. t h i r d ,  and  produced.  Ten  also  discriminate  1970)  to  function  determine  discriminating  analysis  categorizing  variables  and  group  Systems  for  bagging f l o w e r pollinating  autogamous buds  for  p o l l i n a t i o n were c a r r i e d the  emasculated,  flowering  also  served  season  bagged f l o w e r s w i t h  f l o w e r s on t h e same p l a n t . T e s t s  for  out by;  and  2)  mature  within  unfertilized  p o l l e n from o t h e r seed  production  as p r o c e d u r a l c o n t r o l s f o r hand  pollination  s e e d s were The several  the  pollination  ovules  were  Seed  number  inflcresence.  1)  hand  asexual  e x p e r i m e n t s were c o n d u c t e d by a l l o w i n g e m a s c u l a t e d , bagged to  an  relationships.  Tests  which  operated  a  Columbia,  and i n t e r m e d i a t e g r o u p s were a s s i g n e d  Dixon  important  British  obtain  vs. second,  stepwise  (BMD07M,  functions,  then  eigenvector  including  to  of  recorded was  buds  bags.  Undehisced seed  in  subsample  a  of  and each  e s t i m a t e d where h i g h numbers o f  produced.  number o f s e e d s i n i m m a t u r e c a r p e l s plants  of  each  i n f l o r e s c e n c e were g r o u p e d  taxon.  Repeated  according to  the  was  counted  counts  from  on a  single  determinate  order  and p o s i t i o n o f t h e f l o w e r s on t h e i n f l o r e s c e n c e  branches.  18  Habitat find P o l l i n a t i o n  Observations  of  Studies  insect  behavior on Sax.jfraqa f l o w e r s were  taken over a period of 21 hours on 5 days a t  3  locations.  The  l o c a t i o n s i n c l u d e d two on Vancouver I s l a n d where S. r u f i d u l a and S. i n t e q r i f o l i a  are  sympatric and a S. o c c i d e n t a l i s p o p u l a t i o n  at Y a l e , B r i t i s h Columbia. Observations the  time  spent f o r a g i n g  of  behaviour  ( f o r nectar and/or p o l l e n ) or r e s t i n g ,  the numbers of and d i s t a n c e s between f l o w e r s and visited, in  and i f p o s s i b l e , the v i s i t o r s '  various  Sasifraga  stages flowers  of  included  anthesis.  were  preferences for flowers  Insects  collected  inflorescences  observed  visiting  and i d e n t i f i e d t c a s s i s t i n  i d e n t i f i c a t i o n of s i m i l a r uncaptured v i s i t o r s . P o l l e n was  washed  or scraped from the c o l l e c t e d i n s e c t s t o determine the amount o f g a x i f r a g a p o l l e n present. S o i l depth t o rock s u b s t r a t e depth of 30 cm)  was  (or i n r a r e cases to a oaximum  measured from beneath 15 t o 30 p l a n t s chosen  at random and r e p r e s e n t i n g p o p u l a t i o n s on Vancouver I s l a n d , Lower  Mainland  the  of B r i t i s h Columbia, and a t r a n s e c t of s t a t i o n s  along the Columbia B i v e r Gorge.  About 20 cc samples of s o i l were  taken from beneath 15 p l a n t s chosen a t random from 7 p o p u l a t i o n s |6 cn Vancouver I s l a n d , 1 locations  received  flowering  season.  homogenized,  at  repeated The  weighed  Yale,  sampling  samples and  British  dried  from in  over each  Columbia),  Three  the course of one population  were  a 60 C vacuum oven f o r 24  hours before dry weight measurements were t a k e n .  19  RBSCLTS ftMP DISCUSSIGN  Horphological include  both  taxonomic  f e a t u r e s which  floral  and  t r e a t m e n t s , /The  distinguish  vegetative  between  characteristics  created  t h e degree midpoint fruit.  of  anther  For i n s t a n c e ,  S.  proceeds  inteqrifolia  superior  ovary  dehiscence) in  using midpoint of  gland  the  latipetiolata  i n f e r i o r ovary almost  species anthesis  as  a  complexes, i s  clearly  completely  the  as  fruiting  in  live  difference to detect  where  petals  precise  ovary  anthesis. may  distort  m a t e r i a l t h a t i s s t a n d a r d i z e d by  dehiscence  in stylar  condition  as  a  precise  event  in  elongation, r e c e p t a c l e width  and  positioning create d i f f i c u l t i e s i n guantitive  Nonetheless,  as  h a s l e d t o some a m b i g u i t i e s i n p r e v i o u s  anther  anthesis, v a r i a b i l i t y  an  between  at maturity of  var,  to  which  by  conditions i s d i f f i c u l t  drying processes  Even  later  as  (judged  (Fig.2ft,B). E v i d e n t l y  i s interpreted  relationships.  and  S, o r e g a n a  t r e a t m e n t s i f an e a r l y  P r e s s i n g and  feature  anthesis  occidentalis  fruiting  cn p r e s s e d s p e c i m e n s a n d  are p e r s i s t e n t  early  from a g r e a t e r than h a l f  fruit  b e t w e e n f l o w e r i n g and  taxonomic  S.  and  in  i n most  f o r taxcnomists i s the d i s p a r i t y  of ovary i n f e r i o r i t y a t  development in  problems  taxa  p o s i t i o n o f t h e o v a r y h a s b e e n used  an i m p o r t a n t k e y c h a r a c t e r i s t i c , ft d e v e l o p m e n t a l has  the  characteristic ovary  position  which i s  measurement.  separates useful  the  major  provided  that  d e f i n e d t y d i s c r e t e e v e n t s such as  anther  20  F i g u r e 2: D r a w i n g o f S. o c c i d e n t a l i s var, latipeticlata f l o w e r (A) and f r u i t (£). F l o w e r i s a t m i d p o i n t o f a n t h e r dehiscence. S e p a l s , n e a r e s t p e t a l s , f i l a m e n t s and a p i e s h a p e d segment o f o v a r y a r e removed t o show t h e position of t h e o v a r y . N o t e t h e d i f f e r e n c e s i n o v a r y p o s i t i o n and g l a n d s t r u c t u r e between f l o w e r and f r u i t .  .21  22  d e h i s c e n c e . On interpret  herbarium  flowers  material  which  are  i t i s often  past  possible  to  anther dehiscence as being  t h o s e f r o m w h i c h p o l l e n h a s b e e n removed by p o l l i n a t o r s . The  b r e a d t h and  characteristics  in  c o m p l e x e s . ., • I n  the lower portion  dentata  (Fig.  and  many  of  the  ovary  the  are  useful  and  species  populations  of  ringlike  band  ( F i g . 3,A,B).  The  a r e hidden a t t h e base o f t h e appressed  ( F i g . 3 , C ) , S. o c c i d e n t a l i s  3,D-F)  nectar  gland  species  i n many o f t h e s e p l a n t s .  gland  covers a considerable  at  portion  anthesis  I n S. o c c i d e n t a l i s var.  l a t i pe t i e 1 a t a  - oregana-  complex  i s an o b c o n i c d i s c  which  o f t h e t o p o f t h e o v a r y and exudes  i n diffuse, glistening droplets.  Filament  shape  S. o c c i d e n t a l i s  A  S» r u f i d u l a slightly  few on  clavate  but  produces  filaments  relatives  of  from  high  filaments mountain  are  l i n e a r or  populaticrs  of  Vancouver I s l a n d and t h e Olympic Mountains have filaments., filaments  some  apparently  S. i n t e q r i f o l i a  among  c c c i d e n t a l i s ( F i g . 3 , A - C ) . I n S. r u f i d u l a  plants  the on  widely  var., dentata  clavate  occidentalis  varies  var.,  5. o c c i d e n t a l i s  subulate.  of  some  2 A ) , and members o f t h e S. i n t e g r i f o l i a  (Fig.  and  nectar  var. occidentalis i ti s a small  and f i l a m e n t s  var.  the  separating  g l a n d and i t s s e c r e t i o n s petals  of  S. r u f i d u l a  S« O c c i d e n t a l i s encircling  shape  pressed linear  -oregana  In  are  S. o c c i d e n t a l i s  usually  specimens filaments.  complex  have  at  shrinkage  scuewhat in  A l l members linear  ( F i g . 3D-F) . I t i s i n t e r e s t i n g t h a t  S. O c c i d e n t a l j s  least  var.  or  drying c f the  subulate  i n other r e l a t i v e s  s u c h a s S. m a r s h a l l i i , , w h i c h h a v e y e l l o w  green p e t a l spots the f i l a m e n t s  are broadly clavate  and may  or  have  u r e 3: D r a w i n g o f S. o c c i d e n t a l i s v a r . , o c c i d e n t a l i s ( A ) , £J3JLi£LsIa (B) # S. o c c i d e n t a l i s v a r . ,dentata ( C ) , §* i n t e q r i f o l i a var. claytoniifolia (D) , S. i n t e q r i f o l i a v a r . i n t e q r i f o l i a IE). S. i n t e q r i f o l i a var. lggtoeetala (F), flowers a t midpoint o f anther dehiscence. Nearest s e p a l s , p e t a l s , f i l a m e n t s and a p i e s h a p e d seqment o f t h e o v a r y w a l l a r e removed t o show t h e position o f t h e o v a r y . Note d i f f e r e n c e s i n n e c t a r q l a n d f e a t u r e s , ovary p o s i t i o n , filament structure and p e t a l shape i n t h e f l o w e r s . The q l a n d i s t h e s w o l l e n s t r u c t u r e i m m e d i a t e l y above t h e b a s e s o f t h e f i l a m e n t s .  25  a special function these  features  in  pollinator  as  well  pollination strategies evolutionary Petal taxa  size,  from  its  petals  clawlike  base.  parts.  with  broad  often  as  diverse  interesting  var.  study  somewhat  as  S.  anthocyanic  small  of  usually  narrowed  into  may  var.  be  nearly  and  flower  dentata  have o v a t e  intggx^fojLia  greenish-petaled  a  occidentalis  inflorescences  S.  the  by h a v i n g  rufidula  l a t i p e t j.olat a u s u a l l y  among  occidentalis  f o r m s o f S.  occidentalis  Varieties  with  r u f i d u l a has  S.  dentata 6,B)  a  have  var.  and  petals  range  forms  which  to  from large,  S.  f r o m few  convex,  Some  Columbia  inflorescences.  occidentalis  (Fig.  4,B),  occidentalis  most v a r i e t i e s o f S.  integrifolia  conic in  flower  7.ft).  In  S.  or  interrupted-conic  density  occidentalis  integrifolia  few-flowered, dense, c a p i t a t e inflorescence  obconic  var.  have  c o n g e s t e d , M o n t a n e f o r m s o f S,  broadly  #  latipetiolata(Fig.  range  or  ( F i g . 4, ft, 5 B) .  broadly  ( F i g . 5,ft) and usually  as  flat-topped  inflorescence  occidentalis  occidentalis  well  this  are  well  var.  plants  inflorescences  as  an  of  forms,  B i v e r Gorge  (Fig.  with  shape vary c o n s i d e r a b l y  in  which  diffuse-flowered  var.  provide  relatives  darkly  or  Saxifraga  contrast,  perhaps  occidentalis  bases.  white-petaled  S.  and  S a x i f r a g_a r u f i d u l a . 5.  apetalous  characters  C e r t a i n dwarfed a l p i n e  occidentalis  with  gland  Saxifraga  occidentalis  apetalous  S.  color,  2,3).  elliptic  var.  cculd  Correlations  story, ,  (Fig.  differs  as  attraction.  var.  panicles.  var, apetala  from  open  occidentalis are  usually  F l o w e r number r a n g e s  (4-42) i n S. ...-rufidula,  to  especially  per  those  26  on  Vancouver  (13-81)  Island  and  i n S. o c c i d e n t a l i s  var.  dentata  Saxifraga var.  and  occidentalis  exceeding  Olympic  Mountains,  var., occidentalis,  •• ~s. i n t e q r i f o l i a var.  clavtoniifolia  a large  and  occidentalis  clavtoniifolia  networks  of  Such  s u b s p.  leptopetala.  The  r h i z o m e s w h i c h do  not  replacement  short  rosette. fewer  brittle,  networks  taxa  form  have  deep  vertically  penetrating  features The  of  terminal  teeth  and  S. i n t e q r i f o l i a short,  stout,  var.  horizontal  althouqh  b r a n c h e s and b r a n c h l i k e  the  leaves  leaves are  qrowth  rosette  basal  bulblets  i n this  qroup  plastic  response  to  occidentalis  feature  var.  rather  transition inteqrifolia  from  more  pubescent  i n s i z e than the leaves  The l e a v e s may  shaded c o n d i t i o n s  also  indistinct  narrowed, blade  to  with from  i n w h i c h t h e y become  area. of  taxonomic  petioles and  a  demonstrate  siqnifieance.  1 a t i p e t i o l a t a , as t h e name i m p l i e s ,  i t r e s e m b l e s S. o r e q a n a  w h i c h have d i s t i n c t l y  i n most t a x a v a r y w i t h i n  usually  a r e much s m a l l e r  outer whorl of the r o s e t t e .  broad, short,  S,  a l s o be p r e s e n t i n  networks,  Seme l e a f c h a r a c t e r i s t i c s a r e  The  usually  inteqrifolia  may  and  e l c n q a t e and h a v e a q r e a t e r s u r f a c e  S.  inteqrifolia  plants. Many  a  S.  and * j§.  a p p e a r s t o be a common means o f v e q e t a t i v e  the  inteqrif clia.  number o f f l o w e r s ,  dentata  mar s h a l l i i  other  from  var.  have f i n e ,  rhizoires.  S. j a r s h a l I i i  cf  several  occidentalis  var.  latipetiolata  have  S.  to  75.  Saxifraga var.  the  ( F i q . 9D).  differs  from t h e o t h e r  evident petioles petiole  var.. c l a v t o n i i f o l i a  In  is  has that taxa  ( F i q . 9,A-C,E).  most  abrupt  and S. o c c i d e n t a l i s  in var.  27  Figure 4: Habit sketch cf artificial hybrids and representative parental plants. T e t r a p l o i d S. r u f i d u l a ( A ) , t e t r a p l o i d S. o c c i d e n t a l i s v a r . occidentalis IB), and an artificially produced S. r u f i d u l a x occidentalis var. o c c i d e n t a l i s F1 h y b r i d p l a n t (C)., M a g n i f i c a t i o n x 2/3.  28  29  Figure 5: Habit sketch of artificial hybrids and representative parental plants. Tetraploid S. o c c i d e n t a l i s v a r . , d e n t a t a {&), d i p l o i d S. r u f i d u j a (B) , and an artificially produced S. r u f i d u l a x S. o c c i d e n t a l i s var. dentata F1 hybrid plant (C). M a g n i f i c a t i o n x 2/3.  31  Figure 6: Habit sketch of artificial hybrids and representative p a r e n t a l p l a n t s . D i p l o i d S. rufidula-(fl). tetraploid S. i n t e q r i f o l i a ( B ) , and an artificially produced S. r u f i d u l a x S. i n t e g r i f o l i a F1 h y b r i d p l a n t (C) . M a g n i f i c a t i o n x 2/3. .  32  gure 7: Habit sketch of artificial hybrids and representative parental plants. N=38 S . o c c i d e n t a l i s var. 1 a t i p e t i o l a t a (A) , t e t r a p l o i d S. r u f i d u l a (B) , and an a r t i f i c i a l l y p r o d u c e d S. r u f i d u l a x S. o c c i d e n t a l i s var. l a t i p e t i o l a t a F I h y b r i d p l a n t <C). M a g n i f i c a t i o n x 2/3.  34  35  Figure 8: Habit sketch of artificial hybrids and representative parental plants., Tetraploid S. o c c i d e n t a l i s var. dentata (A), tetraplcid S. o c c i d e n t a l i s (B) , and an artificially produced 5* o c c i d e n t a l i s x S. o c c i d e n t a l i s var. dentata F1 h y b r i d p l a n t (C). M a g n i f i c a t i o n x 2/3.  36  37  dentata other  ( F i g . 10)  usually  (Fig.9).  groups  squared  and  teeth  S.  of  blade-petiole  The  rather  rufidula  transition  teeth  a c u t e whereas  Si  of S.  Columbia  character reddish S.  occidentalis  dentata var.  vary  on  the  var., from  green  a  feature  to are  somewhat make  the  (Fig.  teeth  rufidula  surfaces and  and are  r u f i d u l a p l a n t s from  introgressant  S.  the 10).  d e e p l y rounded  occidentalis  of  lower  taxa  t o be  apparently  occidentalis  l a t i p e t i o l a t a leaves  surfaces,  these  tend  Leaves  the  the a c t u a l angle c f  o r o b t u s e . However. S.  9,A-C).  tinged  occidentalis  but  var.  B i v e r Gorge a r e  (Fig.  S.  obtuse i n  occidentalis  usually right-angled the  and  rufidula  cf  d i s t i n c t rounded or  evident  blade base i s u s u a l l y r a t h e r Marginal  more g r a d u a l f o r members  S.  for  are  while  this  usually those  occidentalis  of var.  reddish.  Saxifragaoccidenta 1 is  usually  light  r e s e m b l i n g members o f t h e  green S.  on  both  integrifolia-  oreqana group. Seme m o r p h o l o g i c a l f e a t u r e s and  further attention include:  stcmate c h a r a c t e r i s t i c s , f i n e coat  sculpturing,  nectar  glands.  features, p o s s i b l e on the  the  extent  S. the  correlations  of  may  be  taxa  the  observed most and  that  genus.  its  in  in  groups data  usefulness It  leptopetala apparently in  section, seed  petals  was is  are  in  and  these  were  not  concerning  members o f S a x i f r a g a .  of c o n s i d e r a b l e  within var.  i n ether  i n cross  survey  petals, filaments,  was  with  a broader  differences  basis of l i m i t e d m a t e r i a l  subsections  observed  structural  variation  of v a r i a b i l i t y  inteqrifclia  leaf thickness  papillcsity  Although  definite  characteristics of  and  which r e q u i r e  These  definition noted  unique without  that amonq  adaxial  38  F i g u r e 9: L e a f o u t l i n e d r a w i n g s h e w i n g variability in shape and margin characteristics i n samples of diploid S. r u f i d u l a (A), tetraploid S. o c c j . q e n t a l i s var. occidentalis (B) , t e t r a p l o i d a n d h e x a p l o i d 5. r u f i d u l a i n the Columbia B i v e r Gorge ( C ) , n=38 S. o c c i d e n t a l i s var. latipetiolata ( D ) , and t e t r a p l o i d S. i n t e g r i f o l i a var. i n t e g r i f o l i a (E) l e a v e s . Each s a m p l e i s f r o m one population. Each leaf i s taken from t h e l o w e r w h o r l of l e a v e s f r o m a s e p a r a t e p l a n t . L i n e i s 5 cm long. Note differences in petiole l e n g t h and w i d t h , petiole-blade transition and dentition. increased variability of Columbia Biver Gorge polyploid S. r u f i d u l a p l a n t s and r e s e m b l a n c e t o S. o c c i d e n t a l i s i s a l s o a p p a r e n t . ,  39  40  F i g u r e 10: L e a f o u t l i n e d r a w i n g s h o w i n g v a r i a b i l i t y in shape and dentition in s a m p l e s o f d i p l o i d (A) and t e t r a p l o i d IB) S. o c c i d e n t a l i s var* d e n t a t a . Samples a r e from f o u r d i f f e r e n t p o p u l a t i o n s . Each l e a f i s taken from the lower w h o r l o f l e a v e s f r o m a s e p a r a t e p l a n t . L i n e i s 5 cm l o n g . Note t h e v a r i a t i o n i n l e a f s i z e and e x t e n t o f d e n t i t i o n .  .41  42  p a p i l l a e and t h e f i n e s t r u c t u r e o f s e e d ridges  or  projections.  differences i n f l o r a l taxcn  I t  be  of  interest  to  study  u l t r a - v i o l e t reflectance patterns f o r this  and i t s r e l a t i v e s .  especially helpful  would  coat s u r f a c e s i s without  in  Use o f t h e s e c h a r a c t e r i s t i c s may p r o v e  studies  of  Eastern  and  Western  North  American s p e c i e s - p a i r s .  Chromosomal S t u d i e s  Polyploid-aneuplcid  series  are  common f o r many g r o u p s o f  plants  (Tobgy 1 9 4 3 , L e w i s and Raven 1 9 5 8 ,  1965,  Carr  Pcdlech  1975, S u b h a s i  1975) i n c l u d i n g  1 9 6 5 , D a m b o l t 1968, E l v a n d e r  Jackson  1962,  Kyhos  Saxifraga{Dambclt  and  1975) and a r e c o n f i r m e d  in  the present study as w e l l . Diploids hexaploids  with  with  10  29  S. o c c i d e n t a l i s  cr  pairs, 28  tetraploids  pairs  have  with  been  19 p a i r s and reported  v a r . o c c i d e n t a l i s a n d S. r u f i d u l a  (Krause and  Beamish 1972,1973). I t i s i n t e r e s t i n g t h a t  in  10-paired  diploid  tetraploid  fRandhawa  and  parallel  }  and  Eeamish  19-paired 1972)  polyploid-aneuplcid  S. r u f i d u l a a n d i t s r e l a t i v e s . the  case  i n S. i n t e q r i f o l i a  has  reduced evolved  pattern  to  5. f e r r u g i n e a  in  an  the  one  The same s i t u a t i o n  may  var. ijjtejrj,folia  for  where  a  situation apparently seen also  in be  19-paired  p o p u l a t i o n s a r e common. Chromosome c o u n t s in  Table  f o r Saxifraga populations are  I I I . P o p u l a t i o n s o f S« r u f i d u l a  t h e O l y m p i c P e n i n s u l a and t h e Opper are,  summarized  from Vancouver I s l a n d ,  Willamette  River  dxainaqe  w i t h one e x c e p t i o n , composed o f d i p l o i d i n d i v i d u a l s w i t h 10  43  pairs  of  chromosomes  (see F i g . 1 4 C ) . K r a u s e and  have r e p o r t e d a h e x a p l o i d p o p u l a t i o n Vancouver  Island.  Attempts  tc  u n s u c c e s s f u l , p o s s i b l y because  (n=29)  relocate of  local  Beamish  at  Elk  (1913)  Falls  that population extinction  on were  resulting  from h y d r o e l e c t r i c water d i v e r s i o n . . Saxifraga  rufidula  G o r g e a r e more c o m p l e x Diploid individuals the  more  populations  from  the  Columbia  River  and v a r i a b l e i n t h e i r chromosome numbers.  (n=10) o c c u r i n some mixed  common t e t r a p l o i d  populations  (n=19) ( F i g . 1 1 ) . O t h e r  with  populations  had o n l y t e t r a p l o i d o r h e x a p l o i d  r e p r e s e n t a t i v e s ( F i g . 12),  the  i n d i v i d u a l s c a n n o t be r u l e d c u t  possibility  considering the S.  rufidula  of rare d i p l o i d small  plants  from  R i v e r Gorge undergo between  plants  number  an  and  counts  populations  abnormal  meiosis  hexaploid  No.  previously referred to 690001  intermediate  along the Columbia typical  plants  and  of  hybrids  their  putative  identical.  S. o c c i d e n t a l i s  i n K r a u s e and B e a m i s h  between  S.  1972)  occidentalis  and  I n d i v i d u a l s sampled  were u n i f o r m l y h e x a p l o i d  This  probably  population  allopolyploid populations  hybrid elsewhere  entity. in  represents Attempts  Fort  Lewis  and  and  S.  inteqrifolia.  (n=29, F i q . 13C,D). a  to  rare, locate  stabilized similar  t h e a r e a were u n s u c c e s s f u l , a l t h o u q h  Olympia, Washinqton  f r c m n e a r Mima, W a s h i n q t o n  (Krause  i s morphologically  t h e r e a r e r e s e m b l a n c e s t c c e r t a i n S. i n t e g r i f o l i a the  the  p o p u l a t i o n from a l o n g t h e C h e h a l i s R i v e r i n S o u t h w e s t e r n  Washington Maze  of  l e v e l s o f p o l y p l o i d y ( F i q . 13A,B).  natural hybrids are morphologically almost A  made. , Some  most  of d i f f e r e n t  Diploid, tetraploid  of  but  specimens  r e q i o n s . One  of  specimen  had an u n c e r t a i n chromosome c o u n t  of  44  Table I I I . Summary o f Chromosome Numbers. Taxa, l o c a t i o n s ^ c o l l e c t i o n no.) occidentalis Y a l e , B.C. (626) Cornwall Lookout, (COBN) Ht. B a k e r , Hash. (BAK)  No. Plants  B.C.  ata S a d d l e Mt., O r e g o n (629A) T i l l a m o o k , Oregon (630) Kalama B i v e r , Hash. (607) D e l e n a , Oregon (606) .dula Mt.Finlayson, V.I., B.C. (627A) Nanoose H i l l , V . I . , B.C. (NOOS-B) Nanaimo,V.I.,B.C. , (NMO-R) Upper C a m p b e l l L a k e , V.I.,B.C. (UCL-R) l a k e C r e s c e n t , Wash. (673) B i n g e n L a k e , Hash. , (669A) H t . P l e a s a n t , Wash. (675A) Yecn P a r k , O r e . (610) V i e n t c , Ore., (616) Troutdale,Ore. (608) B a y e r P a r k , Ore. <612A) The D a l l e s , Ore. S k a m a n i a Co., Wash. (618A)  No. Cells  M e i o t i c Chromosome I l l o_r I I + I ) *  2  6  19  3  10  19  2  5  19  3  8  10  11  44  10  2  7  20  8  23  3  20  10  4  7  10  4  31  10  3  5  10  4  16  10  2  10  29  1  3  10  6  21  5  13  5  7  4  11  4 4  16 26  20(6)  No.  ,19+2(2)  10(2) , 1 9 ( 3 ) , c a . 2 0 1 9 ( 4 ) , ca.19 c a . 2 0 * 1 , ca.27+2. 2 9 , 19+12, 19+13 ca.19,ca.16+8, c a . 2 9 , ca.20+5 c a . 2 9 (3) , c a . 26+6 19"(3) , c a . 14+8  •Numbers i n p a r e n t h e s e s i n d i c a t e t h e number o f p l a n t s w i t h t h a p a r t i c u l a r chromosome number.  45  Table I I I . (con*t) Taxa, l o c a t i o n s j c o l l e c t i o n no ) A  Ho. Plants  o c c i d e n t a l i s x i n t e q r i f o l i a pop, C h e h a l i s R., Hash. 4 (605) 2£ti£ejkiciata S a d d l e Mt., O r e . 5 (629C) integrifolia Mt.Finlayson, V.I., 3 B.C. (627) Nanaimo, V . I . , B . C , 1 (NMO-I) E l k F a l l s , V.I.,B.C. 1 (682) H a r r i s o n L a k e , B.c. 1 (78-1) Y a l e , B.C. 1 (625) Mayer P a r k , Ore. 4 (612B) G r i z z l y L a k e , Ca. 1 (671) B i n g e n , Hash. 1 (617) Mima Bounds, Hash. 1 (MIMft) SlSltcniifolia T r o u t d a l e , Ore. 5 (609) Bowena, Ore. 5 (611) Mayer P k . , O r e . 4 (612) The D a l l e s , O r e . 2 (613) B i g g s , Ore. 10 (640) C l a r k - S k a m a n i a Co. l i n e 3 Hash. (619) G i l l i a m Co., Ore. 1 (641) Cape H o r n , Hash. 1 (646) L y l e , Hash. 1 B i n g e n , Hash. 6 (617) oreaa^a Berthoud (BEE)  Pass, Colo.  No. Cells  M e i o t i c Chromosome No • j l l or II+I)*  11  28,29 (3)  23  ca.36,38j(3) ,ca.40  3  19  2  19  3  19  3  19  2  19  20  1 9 , c a . 17+5(3)  4  19  1  19  4  ca.47  16  10(4) ,19  12  10,8 + 11 (4)  27  10 (3) ,9+5  6  10  55  10  20  10  2  10  10  10  1 30  10 10 (2) ,14+10, ca.14+13, ca. 18+9, ca.23+10 ca.  36  46  n=ca.  47  hybrid  (see  Fig.  population  16D)  or  and  to  may  be r e l a t e d t o t h e C h e h a l i s  S. o r e g a n a  (Elvander,  personal  communication), The  chromosome  counts  for  S. o c c i d e n t a l i s  var.  o c c i d e n t a l i s agree with the majority of e a r l i e r counts 1961, K r a u s e and B e a m i s h sampled  in  of  n=19  Elvander  n=10  (1975)  from  where S.  the  records  Storm  individuals  marshallii  higher  numbers  of  P a c k e r (1968) Mt.,  two m i x e d d i p l o i d a n d  has  Alberta  tetraploid  Mts.,  Montana  Lake P a s s i n t h e Anaconda  Range,  Montana,  apparently  are  idahoensis  Krause  populations  Bitterroot  sufesp.  o c c i de ja t a l i s .  a l l  p o p u l a t i o n from Blakeston  p o p u l a t i o n s f r o m T r a p p e r Peak i n t h e and  for  t h e p r e s e n t s t u d y ( F i g . 14A,B,0)•  reported a diploid and  1972)  (Beamish  ( K r a u s e and  n=28  intermediate  and  between  S. o c c i d e n t a l i s  Beamish  1972)  has  var,  recorded  o r c a , 29 f o r p o p u l a t i o n s i n N o r t h e r n  B r i t i s h C o l u m b i a and t h e m o u n t a i n s o f I d a h o . Plants (Hitchcock  identified  as  Saddle  occidentalis  e t a l , 1973) w e r e c o l l e c t e d  o f t h e C o l u m b i a R i v e r Gorge River.  S.  Clatsop  County,  are  Co.,Washington  or  plants  have  20  i n c l u d i n g t h o s e from  diploid  (EP606)  chromosomes  #  the  allotetraploid clavtoniifolia  Willamette  hybridizations and  diploid  River  as  Valley  between  (n=10)  O r e g o n , and  sometimes  ( F i g . 15A,B). These t e t r a p l o i d s  from  west  and t o w a r d t h e mouth o f t h e C c l u m b i a  15C-E). P l a n t s from Columbia County (EP607),  dentata  from f o u r l o c a t i o n s  P l a n t s o f t h e C o a s t Range o f O r e g o n , Mountain,  var.  19 well  (Fig. Cowlitz  pairs as  other  probably represent  S. i n t e q r i f o l i a  S. o c c i d e n t a l i s  of  var.  j u d q e d by t h e i r m o r p h o l o g i c a l r e s e m b l a n c e and t h e  var.  dentata  proximity  as of  47  Figure  1 1 : Camera lucida drawing of pollen mother c e l l s (PMC*s). N u c l e o l a r o r g a n i z e r s a r e g r e y c i r c l e s . Drawings here a r e from four p l a n t s i n one p o p u l a t i o n a l o n g t h e Columbia R i v e r Gorge. P a r t s D and E a r e from t h e same plant. &=PMC a t d i p l o t e n e - d i a k i n e s i s ; S. r u f i d u l a - (EP610) n=20 B=PMC a t d i p l o t e n e - d i a k i n e s i s ; S. r u f i d u l a (EP610) n=19 C=PMC a t d i a k i n e s i s ; S. r j g f j j u i a (EP610) n=19 D=PMC a t d i p l o t e n e - d i a k i n e s i s ; S. r u f i d u l a (EP610) n=10 E=PMC a t m e t a p h a s e I I ; S. r u f i d u l a (EP610) n=10, each daughter c e l l . ;  C.  10JJ  4*  v«  49  F i g u r e 1 2 : Camera l u c i d a d r a w i n g s o f PMC* s . S a x i f r a g a r u f i d u l a p l a n t s from t h e T r o u t d a l e p o p u l a t i o n . A=PMC a t d i p l o t e n e - d i a k i n e s i s : S. r u f i d u l a (EPeoSMe) n=ca. 27 b i v a l e n t s + 2 u n i v a l e n t s i n t h e l o w e r l e f t B=PMC a t metaphase I ; S. r u f i d u l a (EP6 0 8-24) n=ca.20 bivalents + 1 univalent C=PMC a t d i p l o t e n e ; S. . r u f i d u l a (EP608-26) n=29  51  F i g u r e 1 3 : Camera l u c i d a d r a w i n g s o f PMC's., C r y p t i c hybrid p l a n t s f r o m T r o u t d a l e (A,B) a n d C h e h a l i s R i v e r (C,D)• a=PMC a t l a t e m e t a p h a s e I ; S, r u f i d u l a c r y p t i c h y b r i d (EP608-14) n=ca.19 b i v a l e n t s + 1 3 u n i v a l e n t s ( 2 b i v a l e n t s i n upper r i g h t p o s s i b l y multivalents?) B=PMC a t l a t e m e t a p h a s e I ; S, r j i f i d u l a c r y p t i c hybrid (EP6G8-5) n=ca.19 b i v a l e n t s • 12 u n i v a l e n t s C=PMC a t d i p l o t e n e - d i a k i n e s i s : S. o c c i d e n t a l i s v a r . , o c c i d e n t a l i s h e x a p l o i d h y b r i d (EP605-12) n=29 D=pcllen grain mitosis; m e t a p h a s e ; S. o c c i d e n t a l i s var. o c c i d e n t a l i s h e x a p l o i d h y b r i d (EP605-26) n=29  53  Figure 14: Camera l u c i d a drawings c f PMC»s. P l a n t s from Y a l e , Mt. Baker and C o r n w a l l p o p u l a t i o n s of S. . o c c i d e n t a l i s var, occidentalis and Mt. F i n l a y s o n p o p u l a t i o n of S. r u f i d u l a . Upper s c a l e i s f o r c e l l s A-C, lower scale i s f o r c e l l s D and E. A=PMC a t d i p l o t e n e - d i a k i n e s i s ; S. o c c i d e n t a l i s v a r . o c c i d e n t a l i s (EP626) n= 19 I=PMC at metaphase I; S. ^ c c i d e a t a l i s var. o c c i d e n t a l i s (BAKEB) n= 19 C=PMC a t d i p l o t e n e - d i a k i n e s i s ; S. r j j f i d a i a (EP627) n=10 D=PMC at metaphase I; S. o c c i d e n t a l i s var. o c c i d e n t a l i s lEPCOBN) n=19 ~E=PMC at~~metaphase I ; S. r u f i d u l a (EP618-17) n=19  55  Figure  15: Camera l u c i d a d r a w i n g s o f PMC*s. S. o c c i d e n t a l i s var, dentata plants from Kalama Biver, Delena, Tilamook, and Saddle Mountain. A=FMC a t d i p l o t e n e - d i a k i n e s i s ; occidentalis var. d e n t a t a t e t r a p i c i d (EP607) n=19 B=EMC a t d i p l o t e n e - d i a k i n e s i s ; S. o c c i d e n t a l i s v a r . d e n t a t a t e t r a p l o i d (EP606-24) n=20 C=PMC a t d i p l o t e n e - d i a k i n e s i s ; S. o c c i d e n t a l i s v a r . d e n t a t a d i p l o i d (EP630-25) n=10 D,E=PMC a t diakinesis and diplotene-diakinesis; S. o c c i d e n t a l i s v a r . d e n t a t a d i p l o i d (EP629A-25) n=10  57  S.  inteqrifolia  the  S.  var.  occidentalis  Biver  intergrade  var. in  S . , m a r s h a l l i i subsp. expanded  and  claytqniifoliaplants. dentata plants  features  such  aarshallii  but  petal-like  than  from  as  the  clavate  the  those  H o w e v e r , seme o f Willamette filaments to  filaments  are  o f S. m a r s h a l l i i  less  subsp.  marshallii. Plants (Hitchcock  known et  al,  Oregon C o a s t Bange nuaber  of  as  S. o c c i d e n t a l i s  1973)  from  isolated  chromosomes. C o u n t s  varied  38 o r 39  pairs  ( F i g . 16A,C).  of  oreqana  from  at.  personal 36)  1atipetiolata  peaks of the N o r t h e r n  (Chambers 1 9 7 4 ) , w e r e f o u n d t o  around S.  var.,  communication)  Hood, and  have  a  hiqh  somewhat, w i t h most b e i n g  T h i s corresponds with  Oregon  (n=ca.  38,  counts  Elvander,  from Berthoud P a s s , C o l o r a d o  (n=ca.  ( F i g . 17,A). S a x i f r a g a i n t e q r i f o l i a has  Mainland of B r i t i s h and s o u t h w a r d Gorqe  and  into  19 chromosome p a i r s on t h e Lower  Columbia, or Vancouver Washinqton.  P l a n t s from  Island the  ( F i q . 16,B,E) Columbia  Upper W i l l a m e t t e B i v e r a r e more v a r i a b l e and  cytological  studies  are  chromosomal  differences.  necessary At  least  to  detect  the p r a i r i e s s o u t h o f Tacoaa  related  t o S.  chromosomes personal  o r e g a n a and one (2n=ca.  47,51)  further  correlated  some o f t h e p l a n t s f r o m  C o l u m b i a B i v e r G o r g e have t h e 1 9 - p a i r e d c o m p l e m e n t . from  Biver  and O l y m p i a  individual  had a  the  Populations  Washington high  may  be  number  of  (my c o u n t s . F i g . 1 6 , D and E l v a n d e r ,  communication).  Saxifraga  integrifolia  have  been  repeatedly  pairs  ( F i g . 1 7 , B - F ) . One  var.  counted  as  clay, t o n j i f o l i a diploids with  1 9 - p a i r e d i n d i v i d u a l was  populations 10  chrcuoscme  r e c o v e r e d from  58  the otherwise from  near  10-paired p o p u l a t i o n at T r o u t d a l e , Oregon.  Bingen,  W a s h i n g t o n were m o s t l y  with abnormal m e i o s i s although found.  Further  cytelogical  Eastern  Columbia  unravel  the  related  integrifolia  Gorge  var.  Gorge  may  be  with  l. integrifolia I n  populations  are  S.  study.  integrifolia  against  the  var.  A  diploid  Eastern  relatives  lack  of of  such  entities  19-paired  plausible hypothesis  that 10-paired  p r o g e n i t o r s gave r i s e  20  chromosomes  of  chromosome p a i r . 20-paired be  followed  19-paired  by  followed  by  the  argues  directly  from  parental  10,19,20,28,29,38 evidence  subsequent is  loss  apparent  var.  polyploid-aneuploid  plants  frcm  to polyploid offspring  Some p a i r i n g i n s t a b i l i t y  10-paired  and  i n nature  dentata  an i n d i c a t i o n t h a t i t i s o f r e c e n t o r i g i n and  and  they  chrcioscme  9-paired  f o r the  many p o p u l a t i o n s o f t h e o t h e r t a x a . C o n t a c t 19  where  and  plants  p o p u l a t i o n o f S. o c c i d e n t a l i s  conform to the  19  Columbia  with 9  p o l y p l o i d - a n e u p l o i d s e g u e n c e b a s e d on t h e a v a i l a b l e  pairs  there.  clavtoniifolia  from l i t e r a t u r e r e p o r t s  production  more  and  l € P t o p e t a , l a has  chromosome d o u b l i n g o f a h y b r i d b e t w e e n 10 and plants.  p l a n t s and  occur  chromosomally var,  to  inteqrifolia,  close  The  complex  the  necessary  also  p o p u l a t i o n s from the  p a i r s are notably absent present  were  inteqrifolia  which  inteqrifolia  more  var.  Saxifraga  var.  leptppetala  p a i r s o f chromosomes b u t  intergrade  individuals  morphological analyses of  S. i n t e g r i f o l i a  N o r t h w a r d i n i t s r a n g e S.  River  and  10-paired  h y b r i d forms  c o m p l i c a t e d r e l a t i o n s h i p s between t h e s e  the c l o s e l y S.  River  a few  sterile  Plants  may  pattern  and  in which  is with of a one may  eventually found  in  c r o s s i n g between  d o u b l i n g o f chromosome  59  Figure  1 6 : Camera l u c i d a d r a w i n g s o f P M C s o f S. o c c i d e n t a l i s var. l a t i p e t i o l a t a a n d S. i n t e g r i f oljj, a p i a n t s. , &=PBC a t d i p l c t e n e - d i a k i n e s i s ; S. o c c i d e n t a l i s var. l ^ i i e e t i o l a t a <EP629C) n=ca. 38 B=P0C at metaphase I; S. i n ^ a E i J p J L i a var. i f i t e a r i f o l i a (EP617B) n=19 C=PMC a t e a r l y metaphase I ; S. o c c i d e n t a l i s var. latipetiolata (E0629C) n= 3 8 D=PMC at diakinesis; S.- . i n t e g r j f o ^ j a - S . o r e q a n a ? (EPMIHft) n=ca.47 E=PHC at diakinesis; S. i n t e g r i f o l i a var. i n t e q r i f o l i a (EPNMO-I) n=19 ,  M  B.  A.  60  X c.  D.  y  3 ^  \  a  10JJ  61  Figure 17; Camera lucida d r a w i n g s o f PHC*s. P l a n t s from B e r t h o u d P a s s , The D a l l e s , L y l e , B i n g e n , T r o u t d a l e , a n d Bowena p o p u l a t i o n s . The u p p e r s c a l e i s f o r c e l l s A-C, t h e l o w e r s c a l e i s f o r c e l l s D-F. A=PMC a t a n a p h a s e I ; S. o r e g a n a (BPBEfi) n=ca. 36 Upper p o l e shows 34 b o d i e s o f w h i c h a t l e a s t 2 a r e overlapping. l o w e r p o l e shows 35 b o d i e s o f w h i c h a t l e a s t 2 are overlapping, B=PMC a t d i p l o t e n e - d i a k i n e s i s ; S. i n t e g r i f o l i a v a r . c l a v t o n i i f o l i a (EP613-34) n=10 C=PHC at diakinesis; 5. / i n t e g r i f o l i a var. c l a v t o n i i f o l i a (EP668-14) n=10 D=PMC at metaphase I; S. i n t e g r i f o l i a var. c j j j j t o n i i f o l i a (EP617-21) n=10 E=PMC at diakinesis; S. i n t e g r j - f o l i a var. c l a j t o n i i f o l i a (EP6C9-310) n=10 F=PHC at late diakinesis; S. i n t e g r i f o l i a var. c l a y t o n i i f o l i a (EP611) n=10  62  63  number p r o d u c e d 2 9 - p a i r e d be  further  represent  aneuploid  hybrid  p o p u l a t i o n s , and  reductions., Probably  combinations  of  combinations i n v o l v i n g d i p l o i d  and  Although  the  i n v o l v e mostly  polyploid  two  one  processes  r e l a t e d t a x a . Moore an  apparently  (1959) and  autopolyploid  t a y l o r i . an e n d e m i c t o Brooks Peninsula  the  pairs  entities  or  i n t h i s group appear  reflexa  of  to  autopclyploidy  (Krause  and  Beamish  h e x a p l c i d s were f o u n d i n  autopolyploidy  T a y l o r and  Mulligan  occurs  (1968)  Charlotte  on V a n c o u v e r I s l a n d , B r i t i s h  Islands  in  report  derivation f o r populations Queen  may  components.  a l l o p o l y p l o i d y ,the p o s s i b i l i t y  seem t o i n d i c a t e t h a t  counts  19-paired  where d i p l o i d s , t e t r a p l o i d s , and population  n=28  c o u n t s o f 38  hexaploid  c a n n o t be e l i m i n a t e d . R e p o r t s i c S. 1973)  the  of  and  S. the  Columbia.  Mlbridization  Most S.  artificial  occidentalis  dentata,  and  intermediate However,  var. S.  the  two  multivariate  e s p e c i a l l y • S.  19-21). P o l l e n f e r t i l i t y Artificial  among  occidentalis.  hybrids closer to t h e i r parents,  hybrids  integrifolia,  between  the  F1  are  generally  analyses  and  seed  a b o u t by  set  hybridizations  per  inefficient  {Fig.  4-8).  placed  certain  than to t h e i r  diploid  S.  occidentalis (Fig.  of h y b r i d s i s summarized i n T a b l e were  successful  c o m b i n a t i o n s o f t a x a w h i c h were a t t e m p t e d in  var.  morphologically  entities  p o l y p l o i d parents integrifolia•  rufidula.  S. o c c i d e n t a l i s  parental  computer  S.  ( F i g . 22).  among  IV. a l l  Differences  attempt p o s s i b l y r e f l e c t s v a r i a t i o n s brought crossing techniques  as w e l l as s m a l l  sample  64  sizes. be  S i m i l a r v a r i a t i o n i n seed germination  the result of d i f f i c u l t i e s  improper  germination  i n assessment  conditions,  apparently i t i s possible t o  obtain  among most p l a n t s i n t h i s r e l a t e d The  artificial  F1  or F1  to  a  hybrids  fraction  absent i n nearly a l l crosses the  cross  which  var.  latipetiolata  (n=ca.  small  sample  sizes,  hybrids  from  crosses  difficulty.  survived  to  maturity  Bivalent formation  was  i s  (n=19) a n d S. o c c i d e n t a l i s v a r .  The  cells.  had  a hybrid origin involving  morphological  affinities  of  meiosis  and  formation  i n artificial  tetraploid  ploidy  var. latipetiolata  ( n = 1 9 ? . c a . 3 6 . c a . 38) was p r o b a b l y  analysis  Bivalent  p o s s i b l y i n d i c a t i n g t h a t S. o c c i d e n t a l i s  38) o f S. o c c i d e n t a l i s  S. o r e g a n a  maturity,  (n=38) where l a r g e r f i g u r e s , p o s s i b l y m u l t i v a l e n t  (15-19 p a i r s ) ,  S. r u f i d u l a .  seed  (Fig. 18). a p o s s i b l e exception  a s s o c i a t i o n s , c o u l d Jse s e e n i n seme was h i g h  (.18-.90) may  o f t h e genome and m u l t i v a l e n t s were  between S. r u f i d u l a  latipetiolata  of  group w i t h o u t  demonstrated various m e i o t i c behaviours. restricted  rates  level  suggest t h a t  the other  parent.  intervarietal  and  i n t e r s p e c i f i c c r o s s e s i n d i c a t e s t h a t t h e r e h a s been c o n s i d e r a b l e genomic  divergence  among t h e t a x a . When d i p l o i d  crossed  t o t e t r a p l o i d S. o c c i d e n t a l i s ,  10  fewer  or  the r e s u l t a n t hybrid  b i v a l e n t s . H y b r i d s b e t w e e n S. r u f i d u l a  -S. ,. o c c i d e n t a l i s . bivalents  S. r u f i d u l a i s  var.,  indicating  occidentalis that  (n=19)  has  (n=19) a n d  produced  about  t h e y h a v e o n l y a p a r t i a l chromosome  c o m p l e m e n t i n common ( F i g . 1 8 , C ) . I n c o n t r a s t , t h e c r o s s b e t w e e n pcGi den t a l i s dentata completely  var., Occident al i s  (n=20) sterile  produced  F1  and  S. o c c i d e n t a l i s  individuals  which  were  and f a i l e d t o u n d e r g o m e i o t i c d i v i s i o n .  var. almost When  9  65  S»  ruf i d u l a  usually  (n=10)  10  resultant  or  fewer  hybrids  occidentalis  an or  situations  F1  hybrid  b e t w e e n S.  inteqrifolia  resembled  S.  19  individuals pollen. §»  S.  rufidula  by  similarities  S.  hybrids  cross  a  but  between  S.  rufidula  higher  (Table I V ) .  even  pollen  Occasionally  no.  S.  should  Crosses  leptcpetala had  from  the  higher  pairing  produce  highly  sterile  occidentalis  F1  hybrids  s h a p e and  relationships  parental  3*).  var.  in  The  intermediates  polyploid  inteqrifolia  inflorescence  and  18).  i n p e t a l c h a r a c t e r i s t i c s and  hybridization  group p r o b a b l y  and  abnormalities x  biological  elucidated this  in  F i g . 6 ) . The  ( F i g . 19-21, p l a n t  bivalents  occidentalis  Although  and  anaphase  sterile  (n=19)  d e v i a t e d e n t i r e l y from e i t h e r  integrifolia  nearly  the  (Fig*  h y b r i d s w h i c h had  intermediate morphologies  6 to  (see  inteqrifolia  forced  latipetiolata  v i g o r o u s F1  S.  are  resemble  than other a r t i f i c i a l  artificial  with  bivalents  var.  produced s e v e r a l fertility  crossed  closely  f o u n d i n many f i e l d S.  is  and  size  resembled  (see  affinities  Fig. may  be  experiments, taxonomic d e c i s i o n s be  based  on  discontinuities  the  of  more  4).  in  pragmatic  morphological  and  merphcraetrie d a t a . , Since a l l possible many  aspects  of  the  c r o s s e s were n o t hybrid  Even i f a l l h y b r i d s were caution  in  literature  reports  homeclogous and  Law  bivalent  1965, or  that  r e l a t i o n s h i p s are  available  interpretation  would in  completed  for be  wheat,  Thomas 1972,  multivalent  formation  as  necessary oats,  may  yet  analysis  p a i r i n g i s under g e n e t i c c o n t r o l R a j h a t h y and  (Fig.  unclear.  considerable in  rye,  light  and  of  ethers,  (alley  1960  H o s s a i n 1977)  and  be  22),  a function  #  Riley that ef  the  66  F i g u r e 18: Camera l u c i d a d r a w i n g s o f PMC*s f r o m artificial hybrids. The u p p e r scale i s f o r c e l l s A-B, t h e l o w e r s c a l e i s f o r c e l l s C-D. A=PMC at metaphase I; S. o c c i d e n t a l ' s var. l a t i p e t i o l a t a x S. r u f i d u l a t e t r a p l o i d n=ca.18 b i v a l e n t s + 14 u n i v a l e n t s E=PMC a t l a t e m e t a p h a s e I ; S. r u f i d u l a d i p l o i d x S. i n t e q r i f o l i a v a r . i n t e q r i f o l i a n=ca.7 b i v a l e n t s + 14 univalents C=PMC a t m e t a p h a s e I ; S. r u f i d u l a tetraploid x S« o c c i d e n t a l i s v a r . o c c i d e n t a l i s n=ca.9 b i v a l e n t s • ca.19 u n i v a l e n t s D=PMC a t metaphase I : S. o c c i d e n t a l i s v a r . dentata tetraploid x S. r u f j i d u ^ a d i p l o i d n=ca. 10 b i v a l e n t s + 10 univalents  68  F i g u r e 19: P r i n c i p a l components a n a l y s i s graph of a x i s one vs. two showing a r t i f i c i a l l y produced h y b r i d i n d i v i d u a l s and r e p r e s e n t a t i v e p a r e n t a l p o p u l a t i o n s . , Symbols are as f o l l o w s : D= S. o c c i d e n t a l i s var. / d e n t a t a 1 = S. I n t e q r i f o l i a v a r . i n t e q r i f o l i a L- S. o c c i d e n t a l i s var. 1 a t i p e t i o l a t a o= S. o c c i d e n t a l ' s B= S. r u f i d u l a (from Vancouver I s l a n d and the Olympic Mountains) r= S. r u f i d u l a (from the Columbia B i v e r Gorge) 1= S. r u f i d u l a (B) x S.- i n t e q r i f o l i a (I) 2= S. r u f i d u l a (B) x S. o c c i d e n t a l i s (o) 3 = S, o c c i d e n t a l i s (o) x S. o c c i d e n t a l i s var. dentata {D) 4= S. r u f i d u l a (B) x S, o c c i d e n t a l i s v a r . dentata (D) 5= S. r u f i d u l a (r) x S. o c c i d e n t a l i s var. dentata (D) 6= S. r u f i d u l a (B) x S. o c c i d e n t a l i s v a r . 1§M BftA-ol§ia (L) :  3* occupies a point to the r i g h t of i t s placement w i t h i n the p l o t . I t i s probably an a b e r r a n t i n d i v i d u a l . , The percentage of the t o t a l v a r i a n c e accounted f o r by the f i r s t three axes i s 2 1 . 3 6 , 1 1 . 7 3 , and 7 . 8 7 percent respectively.  U)  H  Eigenvector 2 O  U>  1  5  +  "2° S3  ro-  ro  ro ro  m  ro  ~  ro  *  01 o  CD  ro  < n <-+ O  M M M  ~5  05  O rD  CD o-  0)  (Jl  O O O M  o  o  o  M  LO  Of  ro-  DO  70  Figure 20: Principal c o m p o n e n t s a n a l y s i s g r a p h o f a x i s one vs. t h r e e showing artificially produced hybrid individuals and representative parental populations. S y m b o l s a n d t o t a l v a r i a n c e f i g u r e s a r e a s i n F i g u r e 19.  .71  .24-  L  1  c_  O +-> u £  0  §.  Li  R R  I  4  r  D  r R R R  r6  1  L x l  D  1  o o D °I  o  o  D D  D D  D D DD  D  -.2-h  D  • 1 0  Eigenvector 1  D  D  D D  3  72  F i g u r e 2 1 : P r i n c i p a l components a n a l y s i s graph cf axis two vs. three showing artificially produced hybrid individuals and representative parental populations. S y a b c l s and t o t a l v a r i a n c e f i g u r e s a r e a s i n a s i n F i g u r e 19. The a c t u a l p o i n t f o r h y b r i d p l a n t 3* l i e s f u r t h e r t o t h e l o w e r r i g h t c o r n e r t h a n drawn h e r e .  !L L  2  r  L  r 2  c  1 6  D  I Ri R  1  L  r £ J 5 R L ° r~ 3 o°  1  L  1  o  D 2  3  RR  LR 0  •  R  0  0  4  K  I D  D D D D D D  D  0 _  ^Eigenvector 2  DD  74  F i g u r e 22: C r o s s i n g success c h a r t f o r t h e t a x a considered in the present s t u d y . , L i n e number and t h i c k n e s s i n d i c a t e s relative success of seed s e t . Numbers o u t s i d e the hexagons a r e t h e h a p l o i d chromosome number. Numbers a d j a c e n t t o t h e l i n e s a r e t h e a v e r a g e number o f s e e d s s e t per a t t e m p t . , N u m b e r s i n s i d e p a r e n t h e s e s a r e t o t a l seed set followed by number o f c r o s s i n g a t t e m p t s , . L e t t e r s r e p r e s e n t a s f o l l o w s : C=S, o c c i d e n t a l i s • • var, dentata. L=S. o c c i d e n t a l i s v a r , l a ^ i p e t j o l a t ^ , 0=S. o c c i d e n t a l i s var, occidentalis, a n d B=S, r u f i d u l , a . The a b s e n c e o f connecting l i n e s i n d i c a t e s t h a t those crosses were n o t attempted.  75  Crossing  19  76  Hybrid  Cross  Between:  T a b l e IVj. Pollen F e r t i l i t y .  No. Individuals  r u f . (10) o c c . (19) r u f . 119) o c c . (19) r u f . (10) dent.|19) r u f . (19) dent.(19) o c c . (19) d e n t . (19) l a t i . ( c a . 38) x r u f . (19) i n t . (19) x r u f . (10) * i n t . (19) x l e p t . ( 1 9 ) *  4 4 3 1 3 3 4 2  J  average lertiliti 9.9 19.9 37.1 1.0 3.0 31. 0 2.4 1.8  Bange (0.8-24.G) (4. 1-33. 1) (0.7-73.4) (1.0) (0.0-8. 2) (2.3-78.5) (0.0-4.6) ( 1 . 7 - 1 . ?)  * T h e s e c r o s s e s were made e a r l i e r by K.I. Beamish and are included h e r e f o r c o m p a r i s o n . V a r i e t a l names a r e a b b r e v i a t e d t o t h e f i r s t t h r e e o r f o u r l e t t e r s . Numbers i n p a r e n t h e s i s i n d i c a t e h a p l c i d number.  77  presence o f c e r t a i n genes numerous a u t h o r s can  Beamish  which  modified  1961) . O r n d u f f  argue  against  crossability  gene  combinations.  h a v e shown t h a t m e i o t i c e v e n t s  be e n v i r o n m e n t a l l y  1975,  or  and/or  {Skovsted  (1969) h a s  rigid  i n anther  1934,  Sax  presented  species  interfertility  Furthermore,  1937,  Singh  several  cases  definitions  in  plant  tissue  based  on  hybridization  studies. Naturally S.  occurring  r u f i d u l a and  I s l a n d and  hybrids  S. -, i n t e < j r i f o l i a  i n places along  tentatively  recognized  resembles the m e i o s i s , and  the  by  between  var.  intejr.if.plia  Columbia  Biver  vegetatively  years. S.  integrifolia  frames at the Judging  vigorous F1  have  and  may  hybrids  can  of  exist  abnormal  .  S.rjifidula  Columbia  comparison  individuals to  their  are,  in  nature,  for  12  usually  more  S.  inteqrifolia  var.  i n d i v i d u a l s . I n o t h e r p o p u l a t i o n s , somewhat a b e r r a n t resemblinq  S.  rufidula  in  habitat  and  years. sterile  rare  p a r e n t a l e n t i t i e s . O n l y a b o u t 30  h y b r i d s were s e e n c o m p a r e d t o a b o u t 3,000 S. r u f i d u l a considerably  and  i n the c o l d  f r o m a l a r g e p o p u l a t i o n s o u t h o f N a n a i m o , B.C.,  intermediate  but  i n n a t u r e f o r many  under c u l t i v a t i o n  British  be  which  t o s e t good s e e d  between  flourished  University  Vancouver  6),  high degree of p o l l e n s t e r i l i t y  Artifical  on  Gorge  (Fig.  Hybrid intermediates c o n s i s t e n t l y f a i l are  sympatric  t h e i r i n t e r m e d i a t e morphology  a r t i f i c i a l l y produced h y b r i d their  locally  in  putative  plants  and  integrifolia individuals  morpholoqy are  perhaps  more common. The difficult  exact to  nature  of  determine  hybrid  relationships i s  much  more  i n the Columbia B i v e r Gorge a r e a  where  78  S. r u f i d u l a a n d up t o t h r e e occur i n close populations  proximity.  of  claytoniifolia  varieties  Variability  S, r u f i d u l a  as  S. i n t e g r i f o l i a  i n levels of ploidy  well  Individuals  which a r e m o r p h o l o g i c a l l y  the other  of the parental e n t i t i e s  i r r e g u l a r and p o l l e n s t e r i l e . locations,  notably  of  var. hybrid  c l o s e t o one o r  may p r o v e t o  be  meiotically  Samples f o r c e r t a i n C o l u m b i a  the  S. i n t e g r i f o l i a  Bingen, Washington, c o n t a i n a l a r g e p r o p o r t i o n  may  within  a s S. i n t e g r i f o l i a  a l s o confound p r e c i s e determinations  origins.  Gorge  of  population  of t h e s e  River near  cryptic  hybrids. Judging in  several  herbarium m a t e r i a l , hybrid  areas.,  collection and  from  of  var.  putative  idahoensis)  x  hybrid  rare  or  numbers o f p u t a t i v e var. the  in  hybrid  occidentalis  and  Hybrid  claytoniifolia recorded Willamette short-term  plants  were  Many o f t h e p u t a t i v e as  well  Another between  S. i n t e g r i f o 1 i a  as  area  with  being high  S. o c c i d e n t a l i s  var. leptopetala i s  Mountains  of  northeastern  swarms p r o b a b l y i n v o l v i n g S. i n t e g r i f o l i a a n d S. m a r s h a l l i i s u b s p .  mass  { o r S. m a r s h a l l i i  Parental  fertility  individuals  a  v a r . l e p t o p e t a l a {or  collected.  appearance.  made  Washington near Spokane  found.  Petes P o i n t area o f t h e Wallowa  Oregon.  and  were  inadeguately  intermediate  1916  S, o c c i d e n t a l i s  p l a n t s had reduced p o l l e n  somewhat  in  S. i n t e g r i f o l i a  columbiana?) hybrids  either  Suksdorf  one s u c h a r e a o f E a s t e r n  S p a n g l e where  subsp.,  Wilhelm  swarms a r e common  marshallii  have  var. been  f o r t h e Mary's Peak a r e a o f t h e C o a s t Range and Opper r e g i o n o f w e s t e r n and c e n t r a l O r e g o n . I n t h e s e hybrid  f i t n e s s , longevity, vegetative  l a r g e areas o f possibly intermediate  areas  reproduction,  h a b i t a t may e x p l a i n t h e  79  a p p a r e n t abundance o f h y b r i d  forms  and  scarcity  of  parental  forms., Fertile common  in  polyploid several  p o p u l a t i o n s of p r o b a b l e h y b r i d  locations  Two t e t r a p l o i d p o p u l a t i o n s which  conform  which a r e near  to  i n t h e Lower C o l u m b i a R i v e r  of  S. o c c i d e n t a l i s  E n g l e r and I r m s c h e r ' s  the  type  origin are  locality  var.  area.  dentata  (1916) d e s c r i p t i o n  f o r S. o c c i d e n t a l i s  and var.  d e n t a t a , a r e m o r p h o l o g i c a l l y i n t e r m e d i a t e between d i p l o i d C o a s t Range  S. o c c i d e n t a l i s  claytoniifolia  var.  d e n t a t a a n d S. i n t e g r i f o l i a  var.  ( F i g . 2 3 ) . These resemble t h e type specimens  §. o c c i d e n t a l i s  var.  dentata  from  Elk  Rock,  of  n e a r Oswego,  Clackamas County, Oregon  ( t h e f i r s t s p e c i m e n c i t e d i n E n g l e r and  I r m s c h e r 1916, i s H e l l e r  10059) ., K r a u s e and B e a m i s h  previously but  referred  synonymized  the  C l a t s o p Co. P l a n t s  S. o c c i d e n t a l i s -  S. m a r s h a l l i i s u b s p . Introgression  t o S.  var.  (1972)  occidentalis  dentata  s e e n by  of  comparing  non-sympatrie §• E2£i&£±&  S. i n t e j r i f o l i a  S. o c c i d e n t a l i s resulted  from  S. o c c i d e n t a l i s  of  are  past  also  var.  plants  24) .  clavtoniifolia  from  Since  completely  by  the  from t h e Columbia R i v e r  o f t h e l e a k a g e o f S. i n t e g r i f o l i a  local Gorge  o r i g i n o f S. r u f i d u l a i n t h e  extinction or  and of  i s the  genes, e s p e c i a l l y  var. c l a v t o n i i f o l i a .  to  intermediacy  S. r u f i d u l a  area  and  introgressant  introgressant  between the  Gorge can  sympatric  i t i s u n c l e a r whether  hybridization  followed  S. i n t e g r i f o l i a  hybrid  (Fig.  ( F i g . 25)  S. o c c i d e n t a l i s  result  S. r of id,a l a  sites plants  with  ,marshal,lij.  c h a r a c t e r i s t i c s i n t o S. r u f i d u i a i n t h e C o l u m b i a R i v e r be  have  those  i n t o S. r u f i d u l a . The  Columbia  Biver  Gorge  may  80  Figure 23: Scatter diagram showing i n t r o g r e s s i c n of c e r t a i n characteristics between S. i n t e g r i f o l i a var. claytoniifolia ( s g u a r e s ) and t e t r a p l o i d 5. o c c i d e n t a l i s var. dentata plants ( c i r c l e s ) . Diploid S. o c c i d e n t a l i s var. dentata plants are t r i a n g l e s .  81  Gland Wcl.(mm) 50"  y  H  O  y  > .6 4~.5 9 < .39  £  A  25 +  A  C  05 CL  A  A A A  A 5 +  + 2  Petal Ln.(mm)  +  3  A  82  Figure 24: Scatter diagram showing introgression of characteristics between S. j n t e g r . j f o l i a var. clai+^iiifolia (squares) and sym pa t r i e " S. r u f i d u l a (circles) individuals. Plants in lower right are J' SJfiduia p l a n t s from a nearby p o p u l a t i o n which i s not locally sympatric with S. i n t e q r i f o l i a var. c l a j f t o n i i f o l i a ( t r i a n g l e s ) (EP608,609,610). ~  83  51  Gland Wcl.(mm)  T  • ^.5 mm © .3- .49 mm O K29mm. ;  E U  ©  i'27 c _CTJ  o  CL  A  A  A  ^  A  3 + 12  2.5 Petal Ln.(mm)  •3.7  A A A A .  84  Figure 25: Scatter diagram showing i n t e r m e d i a c y o f c e r t a i n characteristics between S. o c c i d e n t a l i s var, occidentalis <sguares, E P 6 2 6 , Y a l e , B.C.) and C o l u m b i a Biver Gorge S. r u f i d u l a plants (circles). Triangles represent presumably non-introgressant diploid S. r u f i d u l a p l a n t s f r o m V a n c o u v e r I s l a n d .  Leaf Ln.(cm) 0-2.9 3-4.5 4.6-7.6  0  O  O  O A  A A  A  + 75 125 I n f l o r e s c e n c e A n g l e (degrees)  86  have  led to  a  S.  occidentalis.  S»  Ofidula  secondary  r e s e m b l a n c e b e t w e e n S. r u f i d u l a a n d  The  variable  more  are mostly  those o f higher  have p r o b a b l y a r i s e n v i a d o u b l i n g of  introgressant  morphs  of  p l o i d y l e v e l s and t h u s  o f t h e chromosome  complements  s t e r i l e hybrid individuals. Variability  within  S. o c c i d e n t a l i s  demonstrated t o c o r r e l a t e probably  reflects  completely  a  with  hybrid  perhaps l e s s u s e f u l document  than  ploidy  level  which  T e t r a p l o i d s appear t o be  with  23).  (Fig.  higher  origin.  introgressant  claytoniifolia  a  v a r . d e n t a t a c a n be  S. i n t e g r i f o l i a  var.  S c a t t e r diagrams i n t h i s group a r e  multivariate  analyses  as  tools  i n t r o g r e s s i o n s i n c e they u t i l i z e a rather small  to  number  of c h a r a c t e r i s t i c s . Further  evidence  of  introgression  i s provided  by  P r i n c i p l e Components A n a l y s i s and S t e p w i s e D i s c r i m i n a n t programs. S.  The  programs  show  that  r u f i d u l a p l a n t s a r e more v a r i a b l e  populations Columbia  Function  t h e Columbia  Biver  than  S. r u f i d u l a  diploid  River  and  plants  also  form  intermediate  t h e S, o c c i d e n t a l i s  t h e Vancouver  or  spatial  S. i n t e q r i f o l i a  I s l a n d S. r u f i d u l a g r o u p s  (see F i g .  27,28,30-35). The l a t t e r a r e a p p a r e n t l y  not introgressant  either  var.  sympatric  allopatric  Gorge  f r o m V a n c o u v e r I s l a n d a n d t h e O l y m p i c M o u n t a i n s , The  r e l a t i o n s h i p s between groups  both  S. i n t e q r i f o l i a -  S. o c c i d e n t a l i s  var. occidentalis.  integrifolia  with or  87  Numerical  Studi.es  I d e n t i f i c a t i o n o f m a j o r g r o u p s and p o p u l a t i o n s using the h i e r a r c h i c a l  clustering  Relationships  among  were  regardless  similar  included all  or  excluded  S. i n t e q r i f o l i a  similarity usually  larger  in  absence  S.  It  plants  is  and  for  §• O c c i d e n t a l i s  var.,  S.rjifidula  S.  dentata  have i n  features  and  var.  ovary  of  of  the  common  position  with  similarity  (Crovello could  between  were few  rufidula  populations body  Gorge.  as  of  a  a  of  Diploid  group  with  counterparts.  var.  dentata  S. i . n t e q r j | g l i a var./  several  than  tc  and  ;  the  group.  characteristics  which  S. i n t e g r i f o l i a shape.  to  occidentalis  l a t i p e t i o l a t a and S.„. o c c i d e n t a l i s  inflorescence  OTU»s  g r o u p . S.  Columbia R i v e r Gorge  such  Alternatively  d i s t o r t i o n o f t h e r e l a t i o n s h i p s between groups  hybrids  although  River  together  occidentalis  Perhaps t h i s i s because of occidentalis  levels  l a t i g e t i . p l a t a show c l o s e r a f f i n i t i e s  varieties  J.  that  larger  S. o c c i d e n t a l i s  e a c h o t h e r and t o and  the e x c e p t i o n  at greater  the  Columbia  clustered  that  are  group  with  { F i g . 26)  hybrids  C o l u m b i a R i v e r Gorge  the  s i m i l a r i t y to their interesting  parental  grouped  rufidula populations  phenetic  artificial  or with a n o n p a r e n t a l  northern  OBC-CGBOUP.  of h y b r i d s . A r t i f i c i a l  one  S. o c c i d e n t a l i s  S. r u f i d u l a  whether  from the a n a l y s i s with  clustered independently  while  called  possible  i n t h e phenogram  p l a n t s group t o g e t h e r  the  into  groups  of  clustered within  separated  program  was  var.  as  gland  i t may  be a  phenetically  dissimilar  1 S 7 0 ) . More p r e c i s e measurement o f  perhaps  have  S. o c c i d e n t a l i s  increased var.  the  dentata  phenetic and  the  u r e 2 6 : Dendrogram u s i n g DBC-CGBOUP c l u s t e r i n g program. F o r t y - t h r e e s t a n d a r i z e d v a r i a b l e s from 262 individuals were u s e d . P l o t i s t h e l o g o f t h e e r r o r value a t a given c l u s t e r i n g step. Individuals are c l u s t e r e d u n d e r one l i n e b e l o w a n e r r o r v a l u e o f 15. Beference to the numbers i s given i n the a c c o m p a n y i n g k e y . V a r i e t a l names a r e a b b r e v i a t e d t o the first 3 or 4 l e t t e r s . B.C=British Columbia, 0 . N . — O l y m p i c M o u n t a i n s , C.E.G.=Columbia R i v e r G o r g e , V.I.=Vancouver I s l a n d , Hybs.=hybrids. Key  t o l i n e numbers: 1. rufidula x occidentalis, 2. 2 a t i e t i c l a t a x rjjiidula 3. r u f i d u l a x dentata 4. i n t e g r i f o l i a Oliiuia 5. r u f i d u l a x 605 q c c i d e n t a l j s ( C h e h a l i s fi.) 6. integrifclia x rufidula 7. r j a f i ^ u l a (618) ; i f i f i t a t a {629) 8. d e n t a t a (630D) 9. 2 r u f i d u l a ; i n t e g r i f o l i a x r u f i d u l a : dentata x r u f i d u l a 10. o c c i d e n t a l i s x dentata 11• rufidula x occidentalis 12. o c c i d e n t a l i s x dentata 13. rufidula x occidentalis 14. 2 r u f i d u l a x o c c i d e n t a l i s 15. rufidula x occidentalis 16. dentata x r u f i d u l a : r u f i d u l a x o c c i d e n t a l i s 17. occidentalis: rufidula 18• occidentalis 19. 3 r u f i d u l a : c l a v t o n i i f o l i a 20. EP605 ( C h e h a l i s B.) 21. 1 £te^rifs 1 i a ; le£topetaja 22. integrifolia 23. o c c i d e n t a l i s x dentata 24. rufidula 25. rufidula 26. rufidula 27. rufidula ; .integrifolia 28. rufidula 29. o c c i d e n t a l i s x dentata 30. 2 d e n t a t a " 31. ifltga£ifclla x I s J i o ^ g t a l a 32. integrifolia x leptopetala ; leptcpetala x dentata 33. r u f i d u l a x "(EP605) C h e h a l i s fi. 34. rufidula 35. dentata 36. r u f i d u l a x dentata 37. (EP605) Chehalis R, , 5a. o c c i d e n t a l i s x S. i n t e g r i f o l i a h e x a p l o i d p o p u l a t i o n £  x  1 5 0 0  —  5 0 0 —  oo  VO  90  S. ^ g u j i j u l a  and  S. o c c i d e n t a l i s  h i e r a r c h i c a l grouping analysis and  S. o c c i d e n t a l i s  groups.  o f S. o c c i d e n t a l i s -  var., l a t i p e t i o l a t a  t h e s e two g r o u p s d e s e r v e s p e c i f i c g  the  extensive  plants  in  numbers  the  claytoniifolia S.  integrifolia  Gorge  var. of  var.  clearly  leptopetala  i  f r c m S. i n t e q r i f o l i a  of  the  dentata  indicate  that  status.  S a x i f r a g a in t e,gr.j.fo jLia - v a r . i  Results  failed  to separate  inteqrifolia.  probably  because  intermediate  Columbia  River  sample.. I n  contrast,  populations  form  i n s p i t e of c l o s e  Gorge  S..inteqrifolia  a fairly distinct  of  var.  group from  contacts i n the Columbia  River  ( F i g . 26). The  population  independently  from C h e h a l i s , Washington  with  some  S. i n t e g r i f o l i a ,  origin  var.  analysis  clustered  group  and  hybrids.  Its  allohexaploid  integrifolia.  of g r o u p s i n t h e h i e r a r c h i c a l  clustering  compare o n l y r o u g h l y w i t h p r e s e n t t a x o n o m i c t r e a t m e n t s  ( K r a u s e and Beamish  1972,1973, H i t c h c o c k e t a l , 1 9 6 1 ,  and C r o n g u i s t  1973,  Elvander  latipetiolata  and  to c l u s t e r  S. O c c i d e n t a l l s  does  •S. O f i d u l a  individuals  S. o c c i d e n t a l i s involving  not  by S.  o f S. o c c i d e n t a l i s  p o p u l a t i o n from near  group  with  although  K r a u s e and  v a r . , d e n t a t a do n o t a p p e a r  entities  The a l l o h e x a p l o i d  Washington  Hitchcock  1975) . S a x i f r a g a o c c i d e n t a l i s v a r .  a s good s u b s p e c i f i c  occidentalis.  hybrid  o f mixed  leptopetala.  (n=29) s u g g e s t a n  S. o c c j , d e n t a j . j s a n d S.  The r e l a t i o n s h i p s  the  var.,  claytoniifolia,  and chromosome number  between  for  S. i n t e g r i f o l i a  S. i n t e g r i f o l i a morphology  affinities  (EP605)  i t  Beamish  occidentalis.  The  var.  Chehalis,  the  S. o c c i d e n t a l i s  is  considered  (1972)  or  within  and i s p r o b a b l y a  clustering  of  this  91  population  as  well  as  that  i n d i v i d u a l s i s s i m i l a r to the Smith that  clustering  (1969) i n V a c c i n i u m where n a t u r a l  usefulness of  a clustering  study appears l i m i t e d t c the  pattern  the  basis  of  technique  multiple  hybrid  affinities,  guestionable  and  using  high grouping error S.  they  are  1965,  H e i s e r 1976)  with  distinct  1968,  Principal  relationships  several  groups  multivariate  and  hybrids  and  (Bloom 1976)  Components  to  involve  increased the  (Fig.  27-29).  processes  probable  groups,  var.  dentata  suqqests  that  occidentalis  var.  others 1969,  (Eyles  and  Schilling  and  (PCA)  to  of  manner  contrast  correlations  than  standard  to  parent at the  within  the  parental of  a  repeated  crossovers  and  entities.  such  in  diploid level  present group  polyploidization  groups  i n d i v i d u a l s among  which i n v o l v e s  one  origins  treats  recognized taxonomic  In  ;  w i t h i n the  hybrid  by  intermediate  introgression  introgression  variability  of  surprisingly  Analysis  precise  with  backcrossing the  the  other factors including  overlaps  s t u d y of  strongly  f r o m S.  h i e r a r c h i c a l c l u s t e r i n g methods. The spatial  among  Hhitehouse  c y t o l o g i c a l d a t a i n a more  present  groups  rufidula.  Rising  that  natural  occidentalis  present study confirmed reports  Blackith  hybrid  S.  present  However, t h e  latipetiolata  morphologically  g c c i d e n t a l i s and The  technigue.  var.,  the  t o chromosome number are a l l  l e v e l b e t w e e n S.  occidentalis  clusters  c h a r a c t e r i s t i c s . Determination  correlations  this  by  parents.  in  d e f i n i t i o n of l a r g e r  hybrid  described  h y b r i d s formed  b i o l o g i c a l r e l a t i o n s h i p s , phenetic distance  and  artificial  were d i s t i n c t f r o m c l u s t e r s o f t h e i r presumed The  on  o f a number of  appears  consequently Considering  polyploid  taxa  as  92  S.  S. j j i t e ^ K i f a l i a ,  occidentalis,  latipetiolata.  and  S« . o c c i d e n t a l i s reasonable  certain  var.  S.  occidentalis  populations  dentata.  of  interaediacies  rufidula  populations  separated  w i t h n o r t h e r n d i p l o i d s g r a d i n g i n t o a mixed tetraploid,  and  hexaploid  individuals  ( F i g . 27,28) . T e t r a p l o i d  populations  showed  intermediate S.  and  represent  allohexaploid  var.,  to  diploid,  var.  tended  dentata  to group i n  S. o c c i d e n t a l i s ,  J=ati£etiolata,  and  h y b r i d s are  variability  19-21). O c c a s i o n a l s e g r e g a t e s and  of  clouds  from the C o l u m b i a B i v e r  S. . o c c i d e n t a l i s  close  two  S. i n t e g r i f o l i a  and  and  are  probably  included is clearly  (Fig.  in  PCA,  represented  19-21,  extremely  the  plant  unbalanced  their  (see F i g .  3*)  appear  monstrosities  w h i c h w o u l d n o t s u r v i v e i n n a t u r e . Most h y b r i d i n d i v i d u a l s closer S.  affinities  to  o c c i d e n t a l i s o r S.  parental  group.  solutions  Ten grouping  the  Columbia  integrifolia  The  ( T a b l e V,  artificial  raw  River  parent  principal  hybrids excluded)  p a r e n t a l and  principal  taxonomic treatments discriminant discriminating  hybrid  function  (SDF)  v a r i a b l e s and  to  show  rufidula. the  other  (eigenvector) whether or  not  analysis. defined  components for  analysis F values  than  are s i m i l a r  groups  were r e c o g n i z e d  Gorge S.  component  hybrids are i n c l u d e d i n the  and  the  p l a n t s from C h e h a l i s , Washington.,  When a r t i f i c i a l intermediacy  into  group  g r e a t e r v a r i a b l i l i t y and  hyperspaces  occidentalis  different  rufidula  assessments of the ccmplex r e l a t i o n s h i p s i n v o l v e d .  Saxifrajga  Gorge  S.  var.  by  analyses use (Fig.,  in  hierarchical and  standard  the  stepwise  30-35).  for inclusion  The  (critical  p r o b a b i l i t y 0.05), i n t h e o r d e r u s e d , were: p l a n t h e i g h t  six F  (51.9),  93  F i g u r e 27: P r i n c i p a l components q r a p h o f a x i s one v s . two showing individuals frcm s e v e r a l n a t u r a l taxa and p o p u l a t i o n s . A r t i f i c i a l h y b r i d s a r e not i n c l u d e d . Symbols are as f o l l o w s : B=diploid Vancouver I s l a n d and Olympic Mountains S. r u f j d u l a r = p o l y p l o i d C o l u m b i a B i v e r G o r g e S. uiiiLaJLa D= S. o c s i d s n t a l l s var. deflt&ta 0= S. o c c j , < J g n t a i i s v a r . sssiijgfltalis L= S. o c g i i g n t a l i s v a r . laJijaetloJaia X=hexaploid hybrid population from C h e h a l i s Biver EP6C5 1= S. i n t e g r i f o l i a var. integrifolia c= s. i a t s a t i i c i i a var. cj^itsniUeiia. M= S. i f i t e g £ i f c l i a v a r . l&eio.pe.tala. The p e r c e n t a g e o f t h e t o t a l v a r i a n c e a c c o u n t e d f o r by t h e f i r s t three axes i s 19.95, 12.17, and 6,87 percent respectively.  Eigenvector 2 o  to 33  33 S L  3 3  3  ^  3  ^VS3 3  50 33  LO  -i 33  S3 S3 S3 33  S3  S3  S3  * S 3 ^  siV  -, _S3  S3  -J -J  ~>  -J  S3.  D  m O  CD < CD  o  ,  ox oo a  O  o-f-  M  M M  o o  o  o M  o  0  H  7  D  o  o  H °  o  o o_c9  o ^  O  O O  o o  o n  n  o °o  o  ^  H  O  o  o  o o  o  O O  ^  o  n  o o o oo  n o  o  M X  o  c o  o o  D  M  M M  M  n  X  r-  O  X  o o o c oo  95  Figure 28; P r i n c i p a l c o m p o n e n t s g r a p h o f a x i s one v s . t h r e e showing individuals from several natural taxa and p o p u l a t i o n s . A r t i f i c i a l h y b r i d s a r e n o t i n c l u d e d . Symbols a r e a s i n F i g u r e 26.  96  C C  I I .3-  I  M  M M M  °I I C  C M  O  R  00  „ 1  £_  o  P  +-»  u (D 0 c CD  Lu  R  r  r  R  RR R R R R R R_ R R  r  r r r  r r  p  R  B  r r  r R K r  rR r  x  o  O  1  I  °  R r r r R r r  r  .o  r  u  D  D  r D  D  D  c  0  L  •  C  ^  D n  c ix. c oi  0 L  D  X  I  1  D-  ib  OO  R  r  X x  r  r  M  M IC  c  c  C C  I  C D D D D  L  C  D  C  L  _ D  D  D  D  -.3-  M  D C  H_  0  Eigenvector 1.  C C  DC c C CC D C  D D  .2  97  Figure 29: P r i n c i p a l components g r a p h o f a x i s two v s . t h r e e showing i n d i v i d u a l s frcm several natural taxa and p o p u l a t i o n s . A r t i f i c i a l h y b r i d s a r e not i n c l u d e d . Symbols are a s i n F i g u r e 26, P l a n t *C i s d i s p l a c e d s l i g h t l y t o the r i g h t i n t h i s representation.  98  M 1  c  M  M M  i  M  .M  c  c  c  <D  D)  Lu  r  r  x  X  0_  X 0  L  X r Oo r OO  rn  r  c  1  •+->  >  Si rX  L  r r  r D M L r L r r i r 0 ° O. r r r R Xr OR r D R" r -j-%r R Rr D L r D D r ° DL L D D , D. D D D  IC I D  o  u 0) o c*c  M  M I  L.  O  1  CC  00  M  r  c c  &  R  IB °R '  C  c  c  C  R  r  1  R  C  c c c  D  D  1  Eigenvector 2  D  O  O O  99  T a b l e V.. The raw p r i n c i p a l c o m p o n e n t Variable  No.  1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42. 43. 44.  solution.*  Jst  2nd  3rd  4 th  -0.1174 0.0941 -0.0110 0.1087 0.2609 -0.0485 -0.1232 0.1223 0. 1798 -0.0812 -0.0397 0.0768 0.0114 -0.0368 0.1485 0.1247 -0.0268 0.0922 0.1828 0.1451 -0.1307 0.1143 0.0214 0.1383 0.2367 -0.2871 0.2956 -0.0455 0.0639 0.2615 0.2743 0.1939 0.1864 -0. 1018 0.G595 0.2190 0.0748 0. 1621 -0.1733 0.2593 0.1418 -0.0074 0.0821 0.0821  0.2560 0.1533 0.1943 0.123 8 -0.0537 -0.0380 0.2034 0.0767 -0.0644 0.2692 0.0204 0.1146 0.3009 0.1845 0.2158 0.1240 0.0197 0.1000 0.1409 0,1982 0.2130 0.0221 -0.0568 0.1932 0.0714 0.0182 -0.0747 0.2652 0.1619 -0.0067 -0.0533 -0.0791 0.2205 0.2619 0.0153 -0.002 5 0.1716 -0.1472 0.2272 0.0451 0.0209 0.0292 0.1830 0.1830  0.0 362 0.0639 0.1528 -0.0 270 0.1897 -0.0228 0.0256 -0.0233 0.3628 -0.0656 0.1641 -0.3414 0.0120 0.0738 0.1051 C, 10 93 -0.0664 0.0863 0.1536 0.1312 0.0324 -0.2373 -0.1147 -0.2079 -0.1214 -0.0S38 -0.0520 0.1701 -0.1869 -0.2043 -0,1184 0.0 717 -0.1632 0.0607 -0.2388 0.2913 0.1181 -0.1351 -0.2316 -0.1651 0.1137 C.C793 -0.0541 -0.0541  0.0733 0.0 938 0.0401 0.2 052 -0.0 963 -0.1*23 0.2571 0.0258 -0.0733 0.1438 -0.2045 0.1300 -0.0150 0.0048 -0.0034 0.0663 -0.0991 -0.1326 0.1225 0.0790 -0. 1724 -0.1960 -0.0713 -0.OCC1 -0.2404 -0.1099 -0.0546 0.1720 -0.0798 -0.0916 0.1539 0.3015 -0.0470 -0.0 926 0.1640 0.0396 -0.3577 0.3517 0.0896 -0.1621 0.0500 0.2037 -0.1629 -0.1629  * These e x p r e s s e d as e i g e n v e c t o r s ; r e s u l t s o f t h e p r i n c i p l e components w i t h o u t a r t i f i c i a l h y b r i d s a r e s i m i l a r t o t h o s e with hybrids (above). **• V a r i a b l e s a r e a s c o d e d i n T a b l e I I .  100  leaf  blade angle a t p e t i o l e  p e t i o l e length/width length  classification  (Variable diploid,  27)  p o l y p l o i d , and  the  IgPjopetala  S.  from  distinguishes  leaf  discriminates  var.  latipetiolata  (Variable  10)  cases, over  length/width  r u f i d u l a from  Vancouver  of  S.  var.  2aii£§M2l3iS  Island  plants  63.7% were  of  the  claytoniifolia  and  gynoecium  grouped  and  with  S.  height  occidentalis separates  group. by SDF  Of  var.  integrifolia, var.  claytoniifolia,  Mountains  S.  rufidula  jlntegrifolia  S.  var.  integrifolia population  (Fig. (FP605)  (20%), t e t r a p l o i d or  d i p l o i d S. r u f i d j i l a  the  analysis  S. o c c i d e n t a l i s  allohexaploid  among S. o c c i d e n t a l i s (30%),  the  var.  S.  var. 39)  remaining  Olympic  S. r u f i d u l a ,  ( V a r i a b l e 32)  integrifolia  and  var.  (Variable  and  integrifolia  31,B), while i n d i v i d u a l s o f the  S. r u f i d u l a  var.  the  separates  occidentalis  ratio  occidentalis.  31) , S.  Some  were s c a t t e r e d  S.  were c o r r e c t l y c l a s s i f i e d  65%  J . occidentalis  leptopetala  and  leptopetala  65.79%  individuals.  diploid  t  the  while gland height/width  p o p u l a t i o n . Leaf length  both groups of S.  and  from  S. i n t e q r i f o l i a  s e p a r a t e s S. o c c i d e n t a l i s and  hexaploid hybrid  including  groups  inteqrifolia  var.  height  occidentalis  ( V a r i a b l e 37)  l a t i p e t i o l a t a and  Petiole  S.  plant  claytoniif olia  angle  6)  S.  S« i n t e g r i f o l i a  shoulder  other  groups.  that  var.  the  (Variable  dentata  and  (15.52), and  S. r u f i ( | u l a i n d i v i d u a l s from  i n t e g r i f o'\ja.  var.  occidentalis  total  S.  show  discriminates  hybrid  others.. Leaf  occidentalis  ratio  functions  effectively  remaining taxa and  S.  (17.81), gynoecium height  (21.09),  (12.76).  Group  all  (23.16), gland height/width  (20%)  groups  hybrid (Fig,  101  F i g u r e 30: S t e p w i s e d i s c r i m i n a n t f u n c t i o n s p l o t o f t h e f i r s t three canonical variables. The f i r s t two canonical v a r i a b l e s a r e s t r a t i f i e d on the t h i r d v a r i a b l e t o p r o d u c e a stratified three-dimensional plot which clarifies hyperspatial relationships. This representation depicts r e l a t i o n s h i p s o f g r o u p means as s p h e r e s on supporting lines. Group means s y m b o l s a r e as f o l l o w s : A= S. r u f i d u l a plants from t h e C o l u m b i a R i v e r Gorge and h y b r i d s i n t e g r i f o l i a p l a n t s from the B - S. i n t e g r i f o l i a v a r . Columbia R i v e r Gorge clajftoniifolia O S. i n t e q r i f c l i a v a r . dentata D= S. o c c i d e n t a l i s v a r . l3ti£etiolata L= S. o c c i d e n t a l i s v a r . lentopetala M = S. i n t e g r i f o l i a v a r . occidentalis 0= S. o c c i d e n t a l i s v a r . J« r u f i d u l a p l a n t s f r o m V a n c o u v e r Island and t h e Olympic Mountains I S. i n t e g r i f o l i a var. integrifolia p l a n t s frcm Vancouver I s l a n d X = H e x a p l o i d S. o c c i d e n t a l i s x S. i n t e q r i f o l i a plants from C h e h a l i s , Hash. B  =  =  103  3 2 , A ) . Of S. o c c i d e n t a l i s placed Biver  into  the  Gorge  S. i n t e q r i f o l i a  S. i n t e g r i f o l i a  intermediacies  with  group  plants  34,B) b u t p r o b a b l y  frcm  the  they  have  Island  S. i n t e q r i f o l i a  which 12%  are classified.  intermediate  integrifolia and  S. o c c i d e n t a l i s  increased  v a r . dentata  variability  in  var.  Several  differences inteqrifolia  Columbia  that  (Schueler  (12.5%).  SDF a n a l y s i s  parental  S. r u f i d u l a  probable  var.  hybrid  35,B),  and  S. o c c i d e n t a l i s  conjunction with program,  in  natural  1976). I t has been s u g g e s t e d  preferred  method  nature  f o r separation  of  Columbia Biver a  the  hexaploid  Gorge  population  var. latipetiolata  program  t r e a t m e n t which c l a s s i f i e s lsii£§ij=2JsSta,  S. r u f i d u l a from  and  results  morphology.  agree  S. r u f i d u l a .  S. o c c i d e n t a l i s  groups Chehalis,  as i s t h e u n i q u e n e s s o f  results of the hierarchical clustering SDF  1966).  o f t h e t e t r a p l o i d S. c c c i d e n t a l i s  W a s h i n g t o n i s d o c u m e n t e d by SDF a n a l y s i s the  and  r e l a t i o n s h i p s have been  means ( D a n i c k and B u r n s 1 9 7 5 , Namkoong  dentata plants,  (Fig.  (10%)  s p e c i e s g r o u p s , e s p e c i a l l y where t h e y a r e d i f f i c u l t t o  s e p a r a t e by o t h e r The  i s a  Rising  var.  ( 5 % ) . Such i n t e r m e d i a c i e s  SDF a n a l y s i s  and  (Fig.  S. i n t e q r i f o l i a  used a s e v i d e n c e f o r t h e o c c u r r e n c e o f i n t r o q r e s s i o n populations  Biver  a r t i f i c i a l hybrids  between  ( 5 % ) , S. o c c i d e n t a l i s  Columbia  demccstrate  insiqnificant  Vancouver  are  were  v a r . c l a v t o n i i f o l i a (19%)  G o r g e S. r u f i d u l a i n d i v i d u a l s i n c l u d i n g 35,B)  28%  ( F i g . 33,A).  ( F i q . 34,A)  S. i n t e q r i f o l i a  (Fiq.  qroup w i t h  var. dentata i n d i v i d u a l s ,  Taken  in  analysis  favorably  S. o c c i d e n t a11s  with  a  var.  v a r , d e n t a t a as d i s t i n c t  s p e c i e s f r o m S. o c c i d e n t a l i s . According  to  the  rules  of  botanical  nomenclature  104  Figure 31: Stepwise d i s c r i m i n a n t f u n c t i o n s p l o t s of t h e f i r s t three canonical v a r i a b l e s . The f i r s t two canonical v a r i a b l e s a r e s t r a t i f i e d on t h e t h i r d v a r i a b l e t o p r o d u c e a stratified three-dimensional plot which clarifies h y p e r s p a t i a l r e l a t i o n s h i p s . G r o u p means a r e shown a s i n the Figure 29. P a r t A shows i n d i v i d u a l p l a n t p o i n t s f o r the S. i n t e g r i f o l i a var. claytonjifglja cloud as triangular symbols and P a r t B shows s i m i l a r h y p e r s p a c e p o i n t s f o r t h e S. i n t e g r i f o l i a var. leptopetala cloud. S y m b o l s a r e a s i n F i g u r e 29.  105  106  Figure 32: S t e p w i s e d i s c r i m i n a n t f u n c t i o n s p l o t s o f t h e f i r s t three canonical v a r i a b l e s . The first two canonical v a r i a b l e s a r e s t r a t i f i e d on t h e t h i r d v a r i a b l e t o p r o d u c e a stratified three-dimensional p l o t which clarifies h y p e r s p a t i a l r e l a t i o n s h i p s . G r o u p means a r e shown as i n the Figure 29. P a r t A shows i n d i v i d u a l p l a n t p o i n t s f o r t h e S. o c c i d e n t a l i s h e x a p l o i d hybrid (Chehalis River, EP605) cloud as t r i a n g u l a r symbols and P a r t E shows s i m i l a r h y p e r s p a c e p o i n t s f o r t h e S. • , O c c i d e n t a l i s - v a r , l a t i p e t i o l a t a c l o u d . .Symbols a r e a s i n F i g u r e 29.  5 Canonical Variable  G 2  tr  108  re 33: Stepwise d i s c r i m i n a n t f u n c t i o n s p l o t s o f t h e f i r s t three canonical variables. The f i r s t two canonical v a r i a b l e s a r e s t r a t i f i e d on t h e t h i r d v a r i a b l e t o p r o d u c e a stratified three-dimensional plot which clarifies h y p e r s p a t i a l r e l a t i c n s h i p s . G r o u p means a r e shown a s i n the Figure 29. P a r t A shows i n d i v i d u a l p l a n t p o i n t s f o r t h e S. o c c i d e n t a l i s var. dentata cloud as triangular symbols and P a r t B shows s i m i l a r h y p e r s p a c e p o i n t s f o r t h e S, o c c i d e n t a l i s var. occidentalis clcud, Symbcls a r e a s i n F i g u r e 29.  Canonical Variable  6br  2  110  Figure 34: s t e p w i s e d i s c r i m i n a n t f u n c t i o n s p l o t s o f t h e f i r s t three canonical v a r i a b l e s . The first two canonical v a r i a b l e s a r e s t r a t i f i e d on t h e t h i r d v a r i a b l e t o p r o d u c e a stratified three-ditiens i c n a l plot which clarifies h y p e r s p a t i a l r e l a t i o n s h i p s . Group means a r e shown a s in the Figure 29. P a r t A shows i n d i v i d u a l p l a n t p o i n t s f o r t h e §. i n t e g r t f g l i a var. integrifolia cloud frcm the Columbia River G o r g e as t r i a n g u l a r s y m b o l s and P a r t B shows s i m i l a r h y p e r s p a c e p o i n t s f o r t h e S. i n t e g r i f o l j . a var. integrifolia c l o u d f r o m V a n c o u v e r I s l a n d and t h e B.C. M a i n l a n d . S y m b o l s a r e as i n F i g u r e 29.  i n  112  F i g u r e 35: Stepwise d i s c r i m i n a n t f u n c t i o n s p l o t s of the first three canonical v a r i a b l e s . The first two canonical v a r i a b l e s a r e s t r a t i f i e d on t h e t h i r d v a r i a b l e t o p r o d u c e a stratified three-dimensional p l o t which clarifies hyperspatial r e l a t i o n s h i p s . G r o u p means a r e shown a s i n t h e F i g u r e 29. P a r t A shows i n d i v i d u a l p l a n t points f o r the S. r u f i d u l a cloud from Vancouver Island and t h e O l y m p i c M o u n t a i n s a s t r i a n g u l a r s y m b o l s and P a r t B shows similar h y p e r s p a c e p o i n t s f o r t h e S. r u f i d u l a c l o u d f r o m the C o l u m b i a Biver Gorge and t h e S. r u f i d u l a hybrid c l o u d . Symbols a r e as i n F i g u r e 29.  Canonical Variable  2  114  S. r u f i d u l a  becomes  latipetiolata dentata  S. a e g u i d e n t a t a ,  S. o c c i d e n t a l i s  var.  becomes S. l a t i p e t i o l a t a .  S. o c c i d e n t a l i s  var.  becomes  S. g o r m a n i i .  S. o c c i d e n t a l i s  (see d e s c r i p t i o n s  B r e e d i n g System  Observations  and  S. ... o c c i d e n t a l i s  i n Taxonomy  becomes  section).  Baacfing, T e s t s  The  results  incompatibility  of  tests  are  for  presented  automatic  in  Table  a u t o m a t i c s e l f i n g t e s t s i n t h e bagged, frequently  higher  than  This probably r e s u l t e d  stigmatic  and  the  leptopetala  may  self-fertilization  have than  S, . i n t e g r i f o l i a •  var.  latipetiolata.  Elvander  no  seed  a  as  set  during coincide  S. o c c i d e n t a l i s  of  var,  var.  potential  integrifolia.  o r S. o c c i d e n t a l ' s  communication)  f o r members o f t h e S.  var.  var,  reports  that  inteqrifoljabut  similar  s e t i n bagged h a n d - s e l f e d t e s t s t o t h o s e r e p o r t e d h e r e f o r  S. i n t e g r i f o l i a cited  i s  condition.  to  S. o r e g a n a c o m p l e x i n b a g g e d , u n t r e a t e d c o n d i t i o n s seed  from  self-compatible  degree  S, i n t e j r i J a l i a  (personal  set  S. i n t e q r i f o l i a  higher  claytoniifolia.  self  condition  pollinations  o f S. r u f i d u l a , e . g .  and  especially  r e c e p t i v i t y . A l l taxa are c l e a r l y  relatives  obtained  Seed  untreated  from damage i n h a n d l i n g  d e n t a t a and S. o c c i d e n t a l i s a s w e l l  he  VI,  seed s e t i n the h a n d - s e l f e d  e m a s c u l a t i o n and d i f f i c u l t y i n t i m i n g with  selfing  here  and i t s r e l a t i v e s . D i f f e r e n c e s  are possibly  in  the  results  due t o t h e b a g g i n g method u s e d i n t h i s  115  T a b l e VIj, •, Bagging t e s t s f o r s e l f i n g . Selfed by. Hand Ro. Bruits (Plants)  Bagged Untreated  Total Seed  % Good* Seed  No. Fruits (Plants)  Total Seed  %  Good Seed  Abbreviated Taxa ruf. (dipl.)  57 (6)  1402  55.2  19(3)  990  48.  ruf. (poly.)  33 (5)  646  25. 1  4(1)  230  7.8  dent, (dipl.)  82(4)  1313  55.9  7(1)  228  0.0  dent, {tetr.)  35(3)  1312  59.1  11(1)  254  76.4  occ. (tetr.)  40(3)  1719  27.9  27(3)  lati. (n=38)  112(3)  1087  6.3  59(2)  530  5.8  int. (tetr.)  13(2)  646  13.9  5(2)  407  0.4  clay, (dipl.)  67(3)  603  16.9  20(1)  179  15. 1  lept. (tetr.)  8(1)  37  21.6  37(3)  1516  470  48.9  35.5  * "Good" s e e d d e f i n e d as l a r g e , s w o l l e n s e e d v e r s u s s m a l l , s h r i v e l e d s e e d w h i c h was c o n s i d e r e d u n f e r t i l i z e d p o t e n t i a l s e e d .  116  Table V I I . Bagging t e s t s f o r o u t c r o s s i n g Outcrossed Ml l a n d  and a p o m i x i s . Emasculated Bagged  Outcrossing C cntrcl.Ontreated  No. T o t a l % Good* F r u i t s Seed Seed (Plants)  No., T o t a l % Good* No. T o t a l % Good Fruits Seed Seed F r u i t s Seed Seed (Plants) (Plants)  abbreviated Taxa ruf. 10(2) (dipl. )  505  59.6  ruf., 20(3) (pcly.)  932  56. 1  103(5)  4054  38.9  21(4)  621  0.5  dent. 17(1) (dipl. )  206  33.5  355(5)  3304  69.4  6 (1)  149  0.0  dent. (tetr.)  17(1)  675  71.0  61 (4)  2253  63.3  occ,  2 1 ( 2 ) 1210  .6-2.5  41 (3)  2767  45.5  4 7 ( 7 ) 2554  0.2  lati. (n=38)  52(2)  39.6  6(1)  348  0.0  15(2)  676  0.0  3 0 ( 6 ) 1691  0.0  (tetr.)  707  i n t . ./ (tetr.) clay. (dipl.) lept. (tetr.)  22 (2) 8(4)  315(8)  7698  35(5)  1852  66.3  75.1  10 (2)  608  0.0  1271  74.3  41 (1)  401  0.0  32(2)  410  O.O  681  82. 5  12(1)  544  1.3  44(6) 1 175  0.3  * Gocd seed d e f i n e d i n T a b l e V I .  117  s t u d y w h i c h may have mechanical j a r r i n g  allowed  autodeposition  result  of  in  S. o c c i d e n t a l i s r e l a t i v e s , t o s e e d s e t i n t h e s e l f i n g  tests.  In  tests.  cases  the outcrossed  Ter-Avanesian  different deposited  has  reported  on r e c e p t i v e s t i g m a s r e s u l t i n a This  unsuccessful.  i a t i£ e t i o 1 a t a ,  general  large  several grains  failure  of  amounts  of  pollen  S i n g l e i n d i v i d u a l s o f S. o c c i d e n t a l i s v a r .  var.  var.,  harvesting  claytoniif olia .  leptopetala failed to set large  o f s e e d when e x p o s e d t c p o l l i n a t o r s , premature  in  may a c c o u n t f o r f a i l u r e o f some i n d i v i d u a l s  S. i n t e g r i f o l i a  S. i n t e q r i f o l i a  that  p l a n t g r o u p s , l o w numbers o f p o l l e n  s e t s e e d where e f f o r t s t o d e p o s i t  were  or  s e e d s e t was l o w e r t h a n i n s e l f i n g  (1978)  flowering  fertilization. to  relatives,  V I I ) was u s u a l l y egual  few  J3. i n t e g r i f o l i a  (Table  nearly  a  in  a  o f t h e bagged, u n t r e a t e d i n f l o r e s c e n c e s .  Seed s e t i n o u t c r o s s i n g e x p e r i m e n t s higher  as  or  perhaps  infertility  as  in  a  those  and  amounts  result  of  particular  individuals. In a l l cases seed s e t levels  (less  than  was  the  wide  individual plants offspring  which  range  and  the  differ  against i t s occurrence production.  or  at  low  background  1.0%) i n e m a s c u l a t e d , bagged i n f l o r e s c e n c e s  (Table V I I ) . Although a p o m i x i s possible,  absent  which  of fact  reguires  variability that  of  pollination  o f f s p r i n g from  hybridizations  produce  markedly from the female parent at  least  as  a  major  i s  means  of  argues seed  118  flojEili  Observations  Nectar continues  production  for  papillate,  several  days  stigmatic  flies. there  w h i c h may  Saxifraga  flowers  stigmatic  overlap  be  and  close that  i n s e c t s and stigma may  observed  may  dehiscence  and  separation  ( F i g . , 2,3),  b r i n g t h e a n t h e r s and auto-depcsition  been p r e v i o u s l y r e m o v e d by  other  bring  There i s a l s o the of  water  plants  by  from  Hagerup  (Hyde 1950,1969, Hyde and since  properties after  the  observations  i n the  demonstrate  p o l l e n was  h y d r o p h o b i c and  s t i g m a on w h i c h r a f t s o f  float  apparently  and  contact  the  as  blown Proctor  air.  However,  Saxifraga  pollen  were  stigma. Also  No  water-mediated casual  pollen  is  accumulate  in  producing a continuous water surface  a n t h e r s and  pollen  o r be  with  d r o p l e t s from a l i g h t m i s t i n g can  flowers  and  observed to l o s e i t s  present study. fresh  anther  1961,  prolonged contact  that  foraging  stigma,  Williams  of  possible  anther  (1950, 1951)  but  stigmas  that  to  or  and  possibility  f o r w i n d - p o l l i n a t i o n or repeated t e s t s f o r were c a r r i e d o u t  is  in  exists  filament  not  1973) , e s p e c i a l l y  Saxifraga  onset  p o l l e n has  a film  bees  protandrcus  that  for  self-adherent  selfing  stigmas  curvature  contact.  wetness.  present  so  a b o u t by w i n d  tests  anther  that s l i g h t external forces  into  Yeo  are  proximity  f l o a t across  and  substances  and  receptive,  loses i t s characteristic  r e c e p t i v i t y . , A l t h o u g h some s p a t i a l  style elongation  provided  apparently  characteristically  in final  a n t h e r s and  dehiscence  visual stimuli to foraging  are  between d e h i s c e d  into  anther  the  re-emitting  nectar  i s an  with  until  surface  Some U V - a b s o r b i n g and Saxifraga  begins  between  observed  to  a reduction  was  119  noted  i n t h e a d h e r e n c e o f p o l l e n g r a i n s t o e a c h o t h e r and  tapetal wall after reduction  in  prolonged  oily  anemophily f o r these animal  sympatric  e x p o s u r e t o t h e a t m o s p h e r e . Such  a d h e r e n c e may  enhance t h e chances of  open, d i s h - t y p e f l o w e r s i n t h e  separation  of f l o w e r i n g times  t a x a . H o w e v e r , i n most c a s e s  there i s a  are  p o s s i b l e . No  of  insect  testifies Whether  s u c h p o l l i n a t i o n s were r e c o r d e d  observation  are  have  from the present require  of i n d i v i d u a l polyploidy. d i p l o i d and  broad  samples.  more e x t e n s i v e and  It  is  interestinq  p o l y p l o i d taxa occur  to  number o f o v u l e s p e r The  more  sanplinq  levels  the  of  different polyploids Seasonal  within taxa Southern  those at higher  over  lowland latitudes  transplant conditions.  earlier-maturing  maturing  differences  diploids.  t  and  i s unclear  t h a t where  ranges.  with  Lewis  carpel varies considerably  p o s i t i o n s c l o s e r to the c e n t r a l later  such  or  t o some e x t e n t  topographic  or a l t i t u d e s , even under u n i f o r m  the  correlated  sympatrically,  p o p u l a t i o n s t e n d t o bloom e a r l i e r t h a n  inflorescence.  note  events.  separated  i n the season than the  a l s o appears to occur or  of  temporally  p o p u l a t i o n s w h i c h e x h i b i t two  periods  hybridizing  differences  Determination  period  individuals  described for Claytonia v i r q i n i c a  qeographic  The  of  between  during  t h e presence of hybrid  phenological  flower later  isolation  than  of  pollinations  l e v e l s of p o l y p l o i d y w i t h i n p o p u l a t i o n s a s  (1976)  to  but  to the o c c a s i o n a l occurrence there  different  an  absence  short  p h e n o l o g i c a l o v e r l a p d u r i n g which i n t e r s p e c i f i c  tend  a  limited  i s evident  of  would  the  pollen vectors.  Temporal  Suda  to  branches  within  flowers  which  occupy  produce  more  ovules  o n e s w h i c h a r e a t p r o g r e s s i v e l y more  120  terminal  positions  differences earliest, var. and  in  most c e n t r a l 5.  in  that  var.  integrifolia  numbers o f  var.  Although f e a t u r e s do  size,  s i z e with  ovules  extent  do n o t  of  carpel  or  C1978) var.  which  plants, apparently  is  has  and  observed  inteqrifolia  v a r i e t i e s o f 5.  and  that are  on  and  outcrossing  in  associated  Evidently  populations  with  less  which  gynodioecious.  evidence  for  some  S.  similar  significance  populations  of an  male-sterile  Island.  He  has  degree of compaction  inflorescence  synchrony or  var.,  inflorescences  of  s y n c h r o n y among  that  compacted  integrifslia  compacted  are  gynodioecious,  Vancouver  concluded  with  numbers  studies  f i n d i n g of  flowering  populations  inflorescences.  and  He  floral  number.  the degree of f l o w e r i n g  integrifolia.  compaction  in  with  number.  other  biological  s u p p o r t e d by t h e  seed-fertile  inflorescence  are  flower  inflorescence. Further  n o t e d a p o s i t i v e c o r r e l a t i o n between the the  the  leptopetala.  lacking,  of these observed d i f f e r e n c e s i n o v u l e / c a r p e l  observation  with  seem t o show s u c h a marked r e d u c t i o n  integrifolia  series  seem t o be c o r r e l a t e d  are  needed t o d e t e r m i n e t h e s y s t e m a t i c  S.  form a  var.  branching,  observations  p o s i t i o n i n the  Elvander  inteqrifolia  plants  integrifolia  When t h e  S. o c c i d e n t a l i s  var.  per  Some  c l a y t o n i i f o l i a a n d S. o c c i d e n t a l i s p l a n t s  detailed not  exist.  compared,  latipetiolata  o r d e r . ,• T h e s e t r e n d s  infloresence  or  are  numbers a p p e a r i n g i n S.  integrifolia  branches.  number among t a x a  r u f i d u l a . S.  frcm lowest t o higher  S.  inflorescence  flowers  occidentalis  highest  the  ovule/carpel  dentata. S.  on  are  promotes congested integrifolia the  ones  Evidence of m a l e - s t e r i l e i n d i v i d u a l s correlations  involving  degree  of  121  c o m p a c t i o n and g y n o d i o e c y i n t h e t a x a o t h e r t h a n S. is  l a c k i n g from The  integrifolia  the present study.  r e a s o n s f o r c o m p l e t e seed and p o l l e n s t e r i l i t y  individuals are  unclear,  but  one  possibility  i n some  i s that  represent v e g e t a t i v e l y vigorous segregant hybrid  they  o r mutant forms  w h i c h c l o s e l y r e s e m b l e one o r t h e o t h e r p a r e n t i n m o r p h o l o g y and habitat of  preference,  instability  recent origins. mycelial  A n o t h e r e x p l a n a t i o n may i n v o l v e t h e d e g r e e  i n h e r e n t i n a p o l y p l o i d genome w i t h In certain s t e r i l e  fragments  may  p o l l e n mother c e l l s .  be  geologically  i n d i v i d u a l s what a p p e a r t o be  seen  among t h e a b o r t i v e p o l l e n o r  Fungus-induced s t e r i l i t y  may  indeed  be. a  common f e a t u r e o f c e r t a i n p o p u l a t i o n s .  Whatever t h e e x p l a n a t i o n ,  such  i n d i v i d u a l s may compose a  vegetatively  vigorous,  sterile  large portion of certain populations Although the bagging t e s t s interesting  biological  useful  in  uncovering  related  taxonomic e n t i t i e s  Geoggaghy• H a b i t a t  and f l o r a l  in  entity  1961).  observations  reveal  and e v o l u t i o n a r y  t e n d e n c i e s , they a r e not  characteristics  t o separate the closely  involved i n t h i s study, .  And P o l l i n a t i o n  G e o g r a p h i c a l l y S« o c c i d e n t a l i s widespread  (Beamish  Ecology Observations  var.  occidental's  i s a  w h i c h i s f o u n d on m o u n t a i n t o p s a n d c l i f f s i d e s  t h e B o c k y M o u n t a i n s and C a s c a d e s f r o m s o u t h e r n A l a s k a and t h e  Yukon t o Montana and I d a h o . I t g r o w s on e x p o s e d grcund  in  rock  crevices  mossy  or  and cn r o c k y peaks i n v e r n a l l y  bare mesic  h a b i t a t s w h i c h become r a p i d l y x e r i c a s t h e s e a s o n p r o g r e s s e s . I t is  n o t found on Vancouver I s l a n d , t h e O l y m p i c  Mountains  or i n  122  the  Columbia  losland  B i v e r Gorge. S a x i f r a g a  cliffs  abundant.  and o t h e r s i t e s  rufjulula  where  early  grows commonly o n spring  runoff  I t i s f o u n d f r o m Lane C o u n t y i n Oregon n o r t h i n t o t h e  C o l u m b i a B i v e r Gorge a n d  then  northward  s i d e o f Vancouver I s l a n d . I t occurs  on  the  east  mountaintop areas i n t h e Island  as  well  as  in  introduction S. r u f i d u l a i n  Verbal  of  the  Olympic  Olympic  Mountains  lower  elevations.  at  e x t i n c t i o n caused by i n c r e a s e d  and  mountain  may  Obstruction  Pass  Area  on  on  Literature  reports  population  with the failure  o c c i d e n t a l i s o r S. n i v a l i s s i n c e  and  The  Irmscher  British  var.  specimen reported  occidental's  plant  S. y j . r a l l i e n s i s , a l t h o u g h  are  rufidula  a t Ames, I o w a , o r  (1916) i s a l s o  Ridge  Columbia h a s been  w i t h S. o c c i d e n t a l i s v a r .  the U n i v e r s i t y of Alaska, the State  BYU.  to f i n d  Hurricane  on t h e  there  §• r u f i d u l a o r S. o c c i d e n t a l i s v a r .  from  recent  (Welsh 1974) o f S. r u f i d u l a i n  A l a s k a may be t h e r e s u l t o f c o n f u s i o n  Anderson c o l l e c t i o n  on  o f the Olympic Mountains.  r e p o r t s t h a t S. r u f i d u l a o c c u r s  documented.  and  Vancouver  Local  explain  p l a n t s w h i c h have b e e n r e p o r t e d  the  and  grazing pressure  goats  Mountains  m a i n l a n d were a l s o i n v e s t i g a t e d b u t o n l y one o c c u r r e n c e  at  i s  no  i n the c o l l e c t i o n s  Welsh's  f o r southern  since  a  seen  specimens  Alaska  small  i t was  i t was n o t  of  Museum a t J u n e a u , i n t h e among  probably  specimens  by E n g l e r  S. o c c i d e n t a l i s  first  during  labeled the  as  present  study. The tc  those  h a b i t a t s o f S. o c c i d e n t a l i s  var. dentata  o f S. r u f i d u l a b u t S. o c c i d e n t a l i s  var.  are s i m i l a r dentata  i s  found west o f t h e C o l u m b i a R i v e r Gorge i n t h e C o l u m b i a R i v e r and Willamette  River  lowlands  and  higher  i n t o t h e C o a s t Range o f  123  Oregon. faces  I t i s commonly f o u n d on s h a l l o w s o i l i n open o r p a r t l y s h a d e d s e e p a g e  m o i s t u r e i s abundant. allows and  i t  obtain  propagate  Chambers sunny §*  to  I t s extensive  by  vegetative  rufidula  locality  on  while  are of  locality  the  in  heliophilic  Saddle other  nearer  Examination  a r e a s where e a r l y  rhizome  means.  system  is  which  in  S. o c c i d e n t a l i s  present  open,  study  few  sunny  areas that  S. o c c i d e n t a l i s  o c c i d e n t a l i s var.  isolated volcanic  p l u g s and  T i l l a m o o k C o u n t i e s , Oregon. a r e a s i n somewhat d e e p e r in  the  season  than  It  dentata., t h e same  the  snaller,  var.  dentata  tetraploid  vicinity. i s endemic t c a  mcuntaintop balds of C l a t s o p grows  in  open,  rather  drier s o i l .  occidentalis  s y m p a t r i c w i t h i t on S a d d l e M o u n t a i n as  resembles  at  reveal  latipetiolata  and p e r h a p s  S.  m a t e r i a l from a  var.  indicates  diploid  that  s h a d y a r e a s f r o m t h e same  r u f i d u l a plants i n the  Saxifraga  spring  apparently  closely  p l a n t s but f u r t h e r c y t o l o g i c a l s t u d i e s might o r h e x a p l o i d S.  rock  interesting  has c o l l e c t e d  plants  from  are  It  Mountain  to  plants  forms  volcanic  f o o t h o l d s on l o o s e o r c r u m b l i n g s u b s t r a t e  (personal communication)  location  of  var,  and  grassy  I t blooms  later  d e n t a t a which i s  and p r o b a b l y a t o t h e r  sites  well., The  varieties  segregated Gorge  over  they  often  populations. possible varieties  of  are  microhabitat  hybridization  inteqrifolia  much o f t h e i r  Seasonal  which  S.  are  found  are  geographically  range, but i n the Columbia growing  differences differences  in  together  in  flowering  times  apparently  isolate  Siver mixed and the  sympatric although there i s evidence that  and i n t r o g r e s s i o n  a r e common i n t h e C o l u m b i a  Eiver  124  Gorge. S a x i f r a g a i n t e g r i f o l i a grassy  soil  pockets  and  var.  ledges  f r o m t h e Upper f r a s e r V a l l e y and through Gorge S.  the and  Puget  Sound and  southward  integrifolia  as  var.  Columbia  O r e g o n and Biver  h e a d l a n d s and  as  south is  southward  S. . i n t e g r i f o l i a  often gravelly  strearabanks  as  well  central  south  to  var.  ccnderosa  Montana, of  on  ledges  Differences i n s o i l  S. / i n t e g r i f o l i a • •  soil  varieties  depth  occurs i n  of  rocky  of  southern  soil S.  integrifolia var.  to  depth  indicate  hybrids  s a m p l e d a t one  and  Columbia  Idaho west  and  a  significant  o c c i d e n t a l i s p l a n t s and  var.  (Table  larger  VIII).  sample  claytoniifolia  plants  Similarly  from  no c l e a r  the  populations,  have a d e e p e r s o i l  b e t w e e n S.  locality  and  S. depth are  average s o i l  occidentalis  depth  does  Naturally  integrifolia is  and var.  preference than  S.  and  differences i n  necessary.  r u f i d u l a and  of  Columbia  r o c k y s u b s t r a t e emerge b e t w e e n n o r t h e r n  r u f i d u l a but f u r t h e r o b s e r v a t i o n s  occurring  the  o f S. i n t e g r j l f o l j a  integrifolia  rufidula  o c c i d e n t a l i s •• may S.  western  and  outcrops  to  p r e f e r e n c e s f o r one  B i v e r G o r g e were l e s s o b v i o u s . depth  and  clavtoniifolia  or d i s t u r b e d s o i l a l o n g roadcuts  as  on  sympatric  S.  Idaho,  the  Columbia  d i f f e r e n c e between t h e S. r u f i d u l a o r S,  sympatric  in  California.  Observations  the l o c a l l y  i n open  the  f r o m n o r t h e a s t O r e g o n and  to Northern  south  Oregon.  commonly f o u n d  c l i f f s i d e s from the Washington Cascades south B i v e r Gorge and  outcrops  then e a s t i n t o the Columbia B i v e r  savannah beneath P i n u s  interior  open  C e n t r a l Vancouver I s l a n d  w e s t a s f a r as t h e e a s t e r n p o r t i o n  Gorge.  s h a l l o w and  of r o c k y  leptcpetala  rocky p l a c e s or grassy British  far  i n t e g r i f o l i a o c c u r s on  were  intermediate  125  Table V i l l i A v e r a g e s o i l d e p t h (cm) b e n e a t h p l a n t s r e p r e s e n t i n g two s y m p a t r i c t a x a f r o m l o c a l i t i e s i n t h e C o l u m b i a B i v e r G o r g e and B r i t i s h Columbia. rufidula No. . Ave. Hants Depth  IfOca 1 i t i e s Columbia  Biver  Gorge  Troutdale Mt.Pleasant C l a r k Co. L i n e Hosier Mayer P a r k The D a l l e s British  i£tejrilglia No. , Ave. Plants Depth  10 9 9 10 10 10  4.3 3.6 3.3 4.4 4.9 4.5  10 9 10 10 10 20  4.9 6.7 9.6 8.6 14.0 11.6*  15 15 15 15  6.2 3.3 3.6 3.1  15 15 15 15  10.7 6.5 6.5 5.7  Columbia  Ht.Finlayson Sooke Nanaimo** Nanoose  * A subsample o f i n t e g r i f o l i a var. integrifolia (n=20) from this locality had an a v e r a g e d e p t h o f 14.9cm. O t h e r C o l u m b i a B i v e r G o r g e p l a n t s a r e v a r . c i a y t o n i j f o l j a and B.C. p l a n t s are var. integrifolia. ** An i n t e r m e d i a t e h y b r i d s u b s a m p l e (n=15) f r o m t h i s s i t e had a n a v e r a g e s o i l d e p t h o f 5.5 cm.  126  Table  Soil  IJU  m o i s t u r e i n g. H 2 0 ( g . d r y w e i g h t ) under p l a n t s a t f i v e sympatric l o c a l i t i e s . 1  Site  Date  rufidula  integrifolia  Nanoose  March April May  1.039* .801 .514  .565 .675* .216  Nanaimo  March May  1.117 1.014*  .487 .517*  Ht.  March  .995*  .638  Sooke  March  .752*  .725  lale  April  Finlayson  * These d a t e s c l o s e s t t c h e i g h t each l o c a l i t y . ,  occidentalis .614* ~*  integrifolia .426  o f bloom f o r t h a t p l a n t t a x o n a t  127  for  the  hybrid  A few  sympatric  document S.  plants.  d i f f e r e n c e s i n w a t e r r e l a t i o n s b e t w e e n S.  inteqrifolia  (Table  IX).  b e n e a t h S. beneath  and  b e t w e e n S.  Especially  S.  integrifolia  natural  o c c i d e n t a l i s and  during  r u f i d u l a plants  a b o u t A p r i l o r May, under  l o c a t i o n s were s a m p l e d f o r s o i l  peak  contains  more  as i t  does  for  temperature  light  differences i n soil  m o i s t u r e l e v e l s may  habitat  at  selection  the  that the soil  artificial  moisture  sustained  taxa  parameters  soil  Insect  spectra  divisible  by t h e i r with  deerflies  much  their smaller^  could  in  cultivation  establishment  failed  S.  inteqrifolia  to m a i n t a i n  adequate or  rujidala.  s u m m a r i z e d i n T a b l e X* , F l o w e r s  pollen  behavior  into  likely  ground-loving  to  also attracted  and  nectar.  motile  the l a r g e r anthophilous more  the  integrifolia " Numerous  greater upon  and  Syrphids,  The  authors  have  a l l flies  to flies  more  the are  sedentary  Bombylids,  and  e f f e c t c r o s s - p o l l i n a t i o n than  cohorts.  dependence  flies  of  p r e d o m i n a n t l y by n e c t a r - l o v i n g  both  in  observed  p o s s i b l e p o l l i n a t o r d i f f e r e n c e among t h e t a x a be  similar  phase. Such  higher  duplicated  o c c a s i o n a l l y wasps a r e  collect  categories  One  are  were v i s i t e d  a l t h o u g h b e e s and and  but  occurs i n  r u f i d u l a specimens, assuming  regime  w a t e r l e v e l s f o r S.  studied  flowers  watering  S.  soil  be i m p o r t a n t f a c t o r s  seedling  rates i n transplanted  soil  than  conditions,  d i f f e r e n c e s i n water r e l a t i o n s help t o e x p l a i n mortality  times the  germination  seedlings  and  integrifolia  moisture  plants. If field  to  r u f i d u l a and  S.  flowering  moisture  as  of  S.  pollinating  suggested  that  observed  inteqrifolia agents  (Table  var. X).  b e e s i n p a r t i c u l a r show  128  Table X Summary o f m a j o r g r o u p s o f i n s e c t v i s i t o r s o b s e r v e d i n 4 p o p u l a t i o n s . Numbers i n d i c a t e t h e t o t a l number o f i n s e c t v i s i t o r s r e c o r d e d . A  No. B e e s , gasps  % Bees, Wa s£s  §jt O f i d u l a (NOOS-B) 10.5 h r s . ; 4 , 5 A p r i l  14  32.53  29  43  Sj, r u f i d u l a (NMO-B) 5.25 h r s . ; 2 1 , 2 2  10  19.0%  43  53  Total Visitors  13  54.2%  11  24  2  5.8%  32  34  April  Sj. o c c i d e n t a l i s (Yale-626) 2 h r s . ; 19 A p r i l S integrifolia (NHO-I) 4 h r s . ; 10,11 May A  No. pjpt era  129  l e s s i n t e r e s t and c o n s t a n c y i n a s p e c i e s as i t s f r e g u e n c y i n t h e immediate f l o r a 1956,  (Brittain  and  Newton  S i m p s o n and Duncan 1956, L e w i s 1 9 6 1 , F r e e  and the  declines  Anderson  1970) . C o m p e t i t i o n  more i n t e n s e  judging from the greater  concurrently  flowering species  competition  1963,1968, L e v i n  var.  integrifoliacculd  v a r i e t y and  abundance  w h i c h a r e p e r h a p s more  flowers.  S l i g h t s e l e c t i o n pressure  (WcNeilly  and  s p e c t r a between s y m p a t r i c plants,  i f further,  (Antonovics  detailed  the  development  the enlarged  nectar  u l t r a - v i o l e t absorbing petal  size,  which  nectar  papillae,  sympatric  presents  type  confirm  evolutionary  o f such p o s s i b l y f l y - a d a p t e d gland  1967,  5. r u f i d u l a  observations  s p e c u l a t i o n , may h a v e been an i m p o r t a n t in  seasonal  1968). A s h i f t i n v i s i t o r  S. i n t e g r i f o l i a a n d  more  this  influence  structures as  diffuse,  glistening,  { P e r c i v a l 1965) a s w e l l a s r e d u c e d  and  s h o w i n e s s i n S. i n t e g r i f o l i a  var.  l e p t o p e t a l a . The e v o l u t i o n o f g y n o d i o e c y i n S. i n t e g r i f o l i a a l s o be c o r r e l a t e d t o  such  of  due t o  Antovonics  F a t e r n i a n i 1969) and g r e a t e r s e l f - c o m p a t i b i l i t y w i t h i n populations  be  attractive  f o r p o l l i n a t o r s may f a v o r more p r o n o u n c e d  d i f f e r e n c e s i n f l o w e r i n g time  S. r u f i d u l a  Stephens  f o r Hymenopteran p o l l i n a t o r s i n  l a t e r f l o w e r i n g S. i n t e g r i f o l i a  to bees than S a x i f r a g a  1933,  a  pollinator  shift  may  i f selection  f a v o r s p r o p o r t i o n a t e l y mere o u t c r o s s i n g e v e n t s t o c o m p e n s a t e f o r i r r e g u l a r and u n r e l i a b l e f l y v i s i t o r s van  der P i j l  1966).  ( P e r c i v a l 1965, F a e g r i and  130  TAXQNGJ3Y  SJBfcies D e f i n i t i o n  The  In This  definition  of a s p e c i e s i n t h i s complex  must a l l o w f o r o v e r l a p s i n intermediacy  of  Complex  many  c l u s t e r s of v a r i a b i l i t y  variability  specimens.  and  Hence,  the  group  cf taxa  morphological  i t must f o c u s on t h e  which e x i s t around c e r t a i n  morphological  characteristics.  I s o l a t i n g mechanisms which r e s t r i c t  are  of secondary importance to morphological  considered  in arriving  at a  group., I f  the  workable presence  f l o w a r e u s e d as main subspecific level, variable  with  perhaps  in  this  for  the r e s u l t i n g  unmanageable  treating  trends  particular  the  taxa  species are extremely  ranges  entities  in are  a functional approach  included  at  a  l a r g e and  morphology, in  w o u l d be t o t r e a t  s p e c i e s a g g r e g a t e s i n t h e main k e y w i t h a s e p a r a t e , key t o t h e a g g r e g a t e t a x a .  flow  o f i n t e r m e d i a t e s and e v i d e n c e o f gene  criteria  c y t o l o g y , e t c , When t h e s e floras,  classification  gene  ecology, regional them a s  more p r e c i s e  131  Key  To The S p e c i e s  The  following  sufcspecific  taxa  key within  H i t c h c o c k and C r o n q u i s t to  does  5. o c c i d e n t a l i s  not  discriminate  S. inteqrifolia  or  ;  (1973) o r t h e t a x a such  as  varietal  S. o r e g a n a  peripherally  subspecies  of  not  (1972,1973).  included  simplified  in  the  integrifolia  following  related  Krause  and  and S. o r e g a n a a r e  descriptions.  A  shorter,  key i s presented i n t h e Appendix. I t i s designed f o r  identification intermediate that  Saxifraga  sensu  S. m a r s h a l l i i ,  S. r e f l e x a . o r t h e S. n i v a l i s - t e n u i s c o m p l e x • s e n s u Beamish  or  of  the  or hybrid  majority  of  specimens,  specimens w i l l  but  many  not i d e n t i f y e a s i l y  using  key. ,  A. O v a r y l e s s t h a n h a l f i n f e r i o r a t a n t h e r d e h i s c e n c e and s c a p e s ,5m o r l e s s t a l l hybrid  (nearly  entities  to  about  half  shallowly  square-dentate lead  AA);  seme  petiole  as wide; l e a f  nargins  s i n u a t e - d e n t a t e t o , more commonly, r o u n d e d o r ( i f teeth  petals  small,greenish,  in  o f S. o c c i d e n t a l i s and S. g o r m a n i i :  d i s t i n c t , u s u a l l y more t h a n 2x a s l o n g from  inferior  markedly reduced o r near absent  usually  reddish  or  2.5mm lacking  or  greater  in  alpine  see  (sometimes forms  of  S. , o c c i d e n t a l i s . C a n d S. a e g u i d e n t a t a . C C ) . B.  Nectar  qland  at  anthesis  douqhnut-shaped r i n q t h a t  a  swollen,  cylindric  almost covers t h e ovary;  f o r m i n q a f i n e network r a r e l y p r e s e n t o r  as  reddish-brown  material;  tapering  fraqments  abruptly  in  pressed  i n t o an e l e n q a t e  petiole;  a  rhizomes  few  ovary  or  small leaves usually  132  from  1/3 t o a b o u t  1/2 i n f e r i o r  a t anther  dehiscence,  •,...............>,«.•••• • v . . , . . S a x i f r a g a g o r m a n i i BB. N e c t a r g l a n d  1.  reduced t o a narrow band, r i n g i n g t h e o v a r y  w a l l ; rhizomes s h o r t , s t o u t , w i t h few b r a n c h e s , h o r i z o n t a l ; leaves  t a p e r i n g more o r l e s s g r a d u a l l y  a p e t i o l e ; ovary  C.  1/3 o r l e s s i n f e r i o r  Infloresence  clustered  usually  headlike  from  to  dehiscence.  a  tightly  panicle; filaments usually clavate to  petals  narrowed base; gland broadened  at anther  conical  sometimes n a r r o w l y o b l a n c e o l a t e ; sguare-dentate;  from the blade i n t o  teeth  usually  from  on  tapered  inconspicuous  leaf  margins  i n t o a somewhat to  a  somewhat  band.  ................... ................... S. O c c i d e n t a l i s CC., I n f l o r e s e n c e not  usually  tightly clustered  filaments  flat  into  a  oblanceolate  i n montane  sinuate-dentate  topped t o obtuse c o n i c a l , dense  l i n e a r or subulate,  headlike  panicle;  rarely s l i g h t l y clavate or  forms;  teeth  to sguarish-dentate;  broad base; gland  2.  an i n c o n s p i c o u s ,  on  leaf  margins  petals with a rather narrow band.  .... .. S. a e g n i d e n t a t a 3, ,•  Ah.  Ovary  h a l f o r more i n f e r i o r a t a n t h e r d e h i s c e n c e  superior i n f r u i t inferior,  then  i n some scapes  cases)  or,  exceeding  .5  i f less  (becoming than  meters; p e t i o l e from  a l m o s t l a c k i n g , t o e l o n g a t e and n a r r o w ; l e a f m a r g i n s or  minutely  sinuate-dentate;  denticulate p e t a l s about  to  shallowly  2.5mm  half  or  less  or  entire unevenly  (greater  in  133  S. i n t e g r i f o l i a * s e e DD and S. l a t i p e t i o l a t a . £ ) . D,  Petioles  almost  lacking,  s h o r t and b r o a d u s u a l l y l e s s  t h a n 2x a s b r o a d a s l o n g ; l e a f into  petiolar  shallcwly  region;  and  inflorescence  tapering  gradually  margins uneven u n d u l a t e d e n t a t e t o  unevenly often  blades  sinuate  with  dentate;  long,  clear  vestiture  or merely  in  pinkish  glandular h a i r s . , E, S c a p e s u s u a l l y l e s s t h a n 0.5m; soil  of  Northwest  Oregon  plants i n  Coastal  moist  Range  grassy  mountaintop  «balds". ..................................... S. l a t i f i e t i s l a t a 4 . EE.  S c a p e s o f t e n 0.5m  swampy  places  or  from  the  greater; Sierra  C a s c a d e s o f c e n t r a l H a s h i n g t o n and Rockies  plants Nevada west  in north  boggy  or  to the  throughout  the  Mountains.,  ..........•................... ............. S. o r e g a n a . CD.  Petioles elongate  n a r r o w e d and e v i d e n t ; abruptly  into  even i f r e d u c e d i n l e n g t h , leaf  petiolar  blades  tapering  distinctly  gradually  r e g i o n ; m a r g i n s v a r i o u s , commonly  nearly e n t i r e ; v e s t i t u r e i n i n f l o r e s c e n c e u s u a l l y with reddish-tipped  glands.  or  dark  134  1»  Saxifraga  gormanii  Gorman 4 0 8 1 , S.  f.  dentata  I V , 1 1 7 , 1 : 3 6 . 1916. C l a c k a m a s Co.,  {Heller  e t a l . I l l : 149.  10059,  Irmsch.  Elk  dentata  rosette-forming  gradually  Bock,  (Engl.  herb  with  to  ovate,  into a distinct  with  dentate;  Inflorescence paniculate; regular 2.5  mm  base,  S.  w i t h S.  teeth  nearest  & Irmsch.)  reddish,  5-merous;  indefinite sepals  long or  longer,  mostly  deciduous  gland  about  anther  encircling cliffsides  dehiscence; the  0.5  mm  i n v e r n a l l y wet  to  long.  open,  conical  evident.  Flowers  perfect,  or a s c e n d i n g ; p e t a l s about gradually  to  orange to  (rarely  clavate  in  marshallii  fruit.,  sparsely  claytoniifolia)  anthers  gland  2x  tetraploid  fruit;  swollen,  or  (20-81),  tapering  in  m a r s h a l l i i subsp.  or  ( i n some  var.  spreading  white,  simple,  abruptly  a  broad yellow;  intermediates  : carpels broadly  pyriform  doughnut-shaped, surrounding  u p p e r p o r t i o n o f t h e o v a r y ; o v a r y u s u a l l y l e s s t h a n 1/2 at  CL.  Hitchcock  dark,  tapering  apex  several-flowered rachis  anthesis;  Oswego,  p e t i o l e u s u a l l y mere t h a n  'ntegrifolia  f i l a m e n t s l i n e a r to s u b u l a t e  at  near  branching rhizomes. Leaves  r n s t y v i l l o u s below; margins from s u b e n t i r e  sinuate  HSI)  Pflanzenr.  l o n g as b r o a d , g l a b r o u s above, from near g l a b r o u s t o  introgressants  (8.W.  1961. ,  exstipulate; blades e l l i p t i c  as  &  B o r t h w . , U n i v . Wash. P r e s s ,  d e e p l y - g r o w i n g and  somewhat  1923.  O r e g o n , J u n e 2,1917  Engl.  Wats. v a r .  H i t c h c . , V a s e . P I . Pac.  Perennial,  23:106.  Oregon)  occidentalis  delicate,  Torreya  E l k B o c k , Multnomah Co.,  marshallii  S.  Suksdorf  remnant  (n=10,19,20)  often March  seeps or m o i s t  a to  places  the  inferior  linear  ridge  April.,  Bocky  from  Clatsop  135  County,  Oregon  east  to  Cowlitz  Co,  Washington  and s o u t h t o  L i n c o l n and M a r i o n C o u n t i e s i n Oregon.  £S£E£§§J3iative  Specimens  OREGON: C l a t s o p Co.: e x p o s e d b a r e s l o p e o f main  westernmost  summit peak. S a d d l e Mt. S t a t e P a r k , on S a d d l e M t . , c a . 3 2 2 5 * , June  1 9 7 3 , K.L.  bluffs,  Chambers  3752 • (WT0,0BC);  r o c k c r e v i c e s and  n e a r A s t o r i a , 6 May 1 9 3 3 , G.P. B a k e r s . n . ( Q B E ) ;  Co.: O t t e r C r e s t , 26 March  burned f o r e s t , g r a v e l l y  T i l l a m o o k B u r n a r e a , N. W i l s o n B i v e r  var. more  following  specimens  claytoniifolia i ntheir nearly  half  inferior  IMlJson  Creek,  qreater plant s i z e  { c a . 20 c m ) ,  o v a r i e s , a n d r e d u c e d t e e t h : OREGON:  above  about  waterfalls,  Clatskanie,  s. f o r k  of  15 May 1927, J..W. 1938,  (ORE).  s p e c i m e n i s p r o b a b l y a h y b r i d between  •S. i n t e g r i f o l i a OREGON:  mi  (OSC).  24j47 (STD) ; Lane C o . : H i l l ' s C r e e k , 3 0 0 ' , 29 May  D g t l i n a 2783 This  10  of  t e n d t o r e s e m b l e S. i n t e q r i f p l i , a  C o l u m b i a Co.: damp r o c k y b l u f f s , Clatskanie  slopes,  Hwy. , e x a c t l y 3 mi s e .  B l u e L a k e , 3 0 0 0 * , 6 J u n e 1 S 7 5 , K.L. Chambers 4065 The  Lincoln  I QBE): T i l l a m o o k  1 9 3 0 . anonymous s.n.  Co.: e x p o s e d k n o b s , r i d g e s above  3  var. claytonij.folia  Clackamas  q o r m a n i i and  ( p o l l e n i s 100% s t e r i l e ) :  Co.; n e a r M i l w a u k e e , E l k R o c k , 28 M a r c h 1 8 8 5 ,  anonymous 239 (OBE).  2. S a x i f r a g a o c c i d e n t a 1 i s S. Wats. P r o c . Am. A c a d . 23:264 ,1888. §•,saximontana E. N e l s . E r y t h e a 7:168. 1899. M i c r a n t h e s o c c i d e n t a l i s S m a l l , N. Am.  F l . 22 ( 2 ) : 1 4 4. 1905. ;  136  (J.  Macoun. N.c.  Yale  Mt.,  B.C.  May  M i c r a n t h e s s a x i m c n t a n a S m a l l , N. &  E.  1889  N e l s o n , 5917,  17,1875.  Am.  Macoun, l y t t o n , B.C.  April  16,1889)  Micranthes  Small,  A l l e n 242,  allenii  S. a l l e n i i  Fl.  N.  Am.  22 ( 2 ) : 145.  Bot.  17,  Missoula,  occidentalis  5:60.  1947.  1896)  31(1):613.  Jahresb.  W a l l o w a Co.,  1906.  31(1):613.  wallowensis  18542,  var.  Peck,  above  Oregon J u l y 4,  occidentalis  Ice  19C6.  seldom-branching  1923.  (M.F•  allenii  rosette-forming rhizome.  Leaf1.  Lake,  West. B o t . ,  Wallowa  C.L.  H i t c h c . , Vase. P i . £l a l . , 111:49.  herb  Leaves  with  simple,  p e t i o l e , usually glabrous  short,  stout,  exstipulate;  sparsely  to  distinctly  sguare dentate; longer. densely  teeth nearest  Infloresence  villous apex  ca.  (13-50),  paniculate, conical to  individuals;  rachis  p e t a l s about 2.5  mm  long,  into a  puberulent,  indefinite  5mm  white,  long  or  paniculate  to  spherical  in  or  F l o w e r s p e r f e c t , r e g u l a r , 5-merous; s e p a l s s p r e a d i n g reflexed;  blades  below; margins u s u a l l y  usually  several-flowered  clustered headlike headlike  rusty  above, or r a r e l y  Pac.  1961.  o v a t e t o obovate t a p e r i n g g r a d u a l l y o r sometimes a b r u p t l y distinct  Mts.,  1894)  N o r t h w . , U n i v . Wash. P r e s s , H i t c h c . Perennial,  (O.D.  Mont. MINN!)  var.  (Peck  (John  F l . 22(2):144.,1905.  Jahresb.  Fedde, J u s t Bot.  1905.  m i c r o c a r p a J o h n s o n , M i n n . S t . P I . S c i . 4:25.  E l r o d 98a,  more  (A.  N a t l . Park J u l y  Goat M o u n t a i n s , Sash. June 27,  S. l a t a F e d d e , J u s t  S.  19C5.  MINN!) Am.  S.  F l . 22 (2) :145.  Yancey«s, Y e l l o w s t o n e  M e r a n t h e s la£a S m a l l , N.  S.  CAN)  distinct. to souewhat  usually  tapering  137  a b r u p t l y or g r a d u a l l y t o a narrow, c l a w l i k e base; to  yellow;  definitely  filaments  usually  at  c l a v a t e ( a l t h o u g h not  least  anthers  slightly  petaloid);  ovary  w a l l ; ovary  anther  1/3  dehiscence,  clavate to  carpels  b o t t l e shaped a t a n t h e s i s ; g l a n d a b a n d - l i k e r i n g or l e s s  (sometimes almost  Stylar  beaks  recurved  orange  elongate,  encircling  1/2) in  the  inferior  fruit;  at  gland  r e m n a n t s u s u a l l y i n c o n s p i c u o u s , , (n= 10, 19, 28,29) A p r i l  to  August,  Bocky o u t c r o p s , c l i f f s i d e s  to  vernal  s t r e a m l e t s , seeps, (about  7  cm  wet  mountaintops adjacent  rock f a c e s or moist  deep), widespread  n o r t h C a s c a d e s from Scutwestern  and  i n shallow  soil  i n the Bocky Mountain r e g i o n  Washington,  A l a s k a and  areas  Idaho,  adjacent  and  Montana  and  north  to  Yukon,  ISlU§e£iaili§ S p e c i m e n s ALEEBTA: N a t l . Park, COLUMBIA: I.J.  rocky  21 J u n e among  1930,  rocks,  A n d e r s o n 2420  2419  slope  above jB.-C.  Atlin  Bertha McCalla  Hot  (V) ; L i l l o o e t ,  glacier,  Vrugtman  13  60805  Cougar V a l l e y , S e l k i r k  B u t t e r s and  E.W.D. Holway ,340  Sphinx G l a c i e r f o r e l a n d .  758  B,  Eraser  1975,  s.n.  C.C,  E.JH.  ERITISH 1914,  Andersfin  calcareous  1600», 18 J u l y Mt.  Garibaldi  1908,  Bobson  Park,  rockslide 1934,  1960,  foot  Chuanjg 75J[399 (V) ;  (UBC);  10 mi s .  Lakes  B l a c k w a l l , 14 J u l y  w a l l , A l e x a n d r i a B r i d g e , 3500», 18 A p r i l (UC,WTU); C r o n i n Mt.,  (MIN);  1916,  (MIN) : meadow,  moraines.  1965,  Aug.  Mts.,  Pass,  e.  Hay  (V,UBC);  Snowbird  Aug,  9  3618  Waterton  S p r i n g s , 2250', 5 J u l y  (V); r o c k o u t c r o p s , Manning P a r k ,  K . I . B e a m i s h . F.  Lake,  F.K, Park,  glacial  500 0*,  above r i v e r E.T.  of Smithers, 1 J u l y  of  26 on  HcCabe 1967,  G.  138  Mendel  127  (V);  wet  A s h n c l a D i s t r . , 7000',  crevices. 12 J u l y  C a t h e d r a l L a k e s , Mt. B c m f o r d ,  1951, T.M.C. T a y l o r  M a r b l e M t s . , L a k e B o o t a h n i e , 5000', -20,25 E.M. w.  Thompson 70 o f sw,  S.L. J.B.  HeIsfa, K. R i q b y 9109 s.n.  f r o m snowbank, 11 Aug. top  Goat  on  C u s t e r Co.: among r o c k s i n s e e p a g e of Alturus Lake, Stanley,  (UEC)MONTANA:  on  Goat  Flat  9100», 7 J u l y  Co.: g r a s s l a n d on w e l l 1950,  side  1974. P.  Strawberry  Grant  Mt.,  18  Co.:  Hitchcock  mi  s.w.  of  a  large  2.6  limestone  (WTO). G l a c i e r  mi n. o f  St.  Mary,  Co.:  mi a b o v e Thomas  15 June 1 9 4 1 , A.H.  rocky  8900»,1 Aug.  1/2  slope,  ne.  Holmgren of  5428 ( O S C ) . WASHINGTON: S n o h o m i s h Co. :  cliffs,  Mt.  July  at 1940,  the  head  J .  Ownby.,  2  July  1962,  dripping 1 9 3 2 , J.H.  below  Slate  o f the S l a t e Fork of the Paysatan B i v e r ,  Co.: r o c k y s l o p e , Mt. Henry's,  Co.: r o c k y o u t c r o p s ,  of  (WS^HTD) :  under  D i c k e r s o n , C a s c a d e Mts.„ 5000», 17 J u l y (WTO) ; Okanagan  1122  sumiit  1953, A. C r o n g u i . s t 7703  G» l a s o n  Peak  E.G.  Meyer  2222  Bainier Natl.  1928,  4  Canyon  W a l l o w a Co.; s t e e p e x p o s e d s l o p e s o f P e t e ' s P o i n t , 19 J u l y  2 h £ J £ S o n 8853  of  (HS) : NEVADA: E l k o Co.: i n f r e g u e n t i n  La M o i l l e Canyon,  OBEGON:  Head  1 9 4 6 , C.L.  E l v a n d e r 443  c r e v i c e s of r o c k s a l o n g s t r e a m bank,  (UC).  n=10,  b e t w e e n S t o r m L a k e P a s s and  drained s o i l ,  D. L y n c h 6284  Campgrounds,  Beaver  (HS) ; D e e r l o d g e Co.: c a .  s l o p e above t r a i l  just  outcropping,  July  1892,  K r a u s e 68  e.  Flat  anji  ( B B 1 ) . IDAHO; K o o t e n a i C o . : J u n e  (OBE);  M u h l i c j c 14925  Anaconda  J.jjl.  1969,  O d e l l P e a k , P i o n e e r Bange, 24 J u l y  a n d C.V.  1938,  Mt., 4 5 0 0 * , 14 J u n e  common, Mt. s s w .  1969* D.L.  of  (8S.PC):  (UC.WS.MIN); McComal C r e e k Q u a d r a n g l e , c a . 2 mi  corner of Thutade, S t i k i n e  Hieberq  June  1346  F.A.  28  (HS,0BI,OSC,MIN,0C);  Pierce  Park,  Indian  Warren  785  Crystal  Mt.  (WS); S k a m a n i a  Co.:  139  s h a l l o w r o c k y s o i l . S i s t e r s Bock, Columbia M a t i . F o r e s t , 7 June 1945, D.C. hillside,  near  I n q r a h m .1877 Latah  (HS,OSC) ; Spokane Co.: damp r o c k y  Creek,  1916, J . S . S u k s d o r f 8616  4000*,  se. of  (WS);  S p a n g l e , 12  Whatcom  Co.:  flay,  shallow  27 June soil  on  r o c k y s l o p e s a t t i m b e r l i n e , a t . B a k e r , 6 2 0 0 ' , 11 J u l y 1 9 2 2 , H.J..-; >  Mason  3871  hillside.  {DC).  WYOMING: G r a n d T e t o n N a t l .  Park:  moist  mossy  C a s c a d e C a n y o n , 7 5 0 0 , 19 J u n e 1933. I . W i l l i a m s  J135  f  {CSC).  3. / S a x i f r a g a 37, B e i b l .  aeguidentata  83:70,  Saxifraga rufidula  ( S m a l l ) B o s e n d . J.n E n g l . B o t . J a h r b .  1905. (Small)  James  Macoun  Ottawa  Nat.,  20:162.  1906. JiSfantbes  r j i f i d u l a S m a l l , N. Am.  F l . 22(2) : 140.  Macoun, Mt. F i n l a y s c n , V a n c o u v e r I s l a n d May Micranthes (Suksdorf  aeguidentata  S m a l l . N. Am.  1905.  17, 1887  S. r u f i d u l a f .  HY)  F l . , 22(2):145.  9 6 7 , Lower C a s c a d e s , S k a m a n i a Co., Wash.  (John  1905.  WS)  major E n g l . 8 I r m s c h , , P f l a n z e n r .  IV,117,1:39.  minor E n g l . S Irmsch.,Pflanzenr.  IV,117,1:39.  1916. S. r u f i d u l a  f.  1916. '§.* J t l i c k i t a t e n s i s {Suksdorf,  Johnson,  Klickitat  M i n n . S t u d , P I . S c i , 4: 25.  Co., Wash. A p r i l  S. o c c i d e n t a l i s Wats, s u b s p . 111.  F l . Pac. S t .  S. o c c i d e n t a l i s  rufidula  9 and May  1883  1923.  HS!)  Bacigalupi i n  Abrams,  2:366.,1944. var.  rufidula  Hitchc,  Vase  PI.  Pac.  N o r t h w . , O n i v . Wash. P r e s s , B i t c h c . E% a l , , 111:49. 1961. Perennial  rosette-forming  herb  with  short,  stout,  few  140  branching, blades  horizontal  elliptic  abruptly  to  into  rhizomes. ovate,  a  Leaves  tapering  distinct  simple,  somewhat  petiole,  somewhat  usually  shallowly  greater  than  several-flowered convex  or  perfect,  .5mm  long.  teeth  2.5  usually  broad  mm  5-merous;  long  nearest  flat-topped,  spreading  or  base, mostly  soil  w a l l ; ovary  deciduous linear  <ca.  moist,  1/3  or  in fruit;  anthers  oblanceolate  from  to  3-4  often  cm)  then  found  from  slightly  dehiscence.  inconspicuous  cliffsides  in  into  the  in  shallow vernally  washes o r r i v u l e t s f r o m t h e east  at  encircling  F e b r u a r y t o J u l y . On  o f r o c k y o u t c r o p s and  W i l l a m e t t e B i v e r a r e a n o r t h and Gorge,  g l a n d remnant  c a , 28,29) M i d  dripping seeps,  dark  c a r p e l s narrow, bottle-shaped  or l e s s i n f e r i o r at anther  s t y l a r beaks r e c u r v e d ; (n=10,19,28  Flowers  ascending;  a n t h e s i s ; gland a narrow, i n c o n s p i c o u s b a n d - l i k e r i n g  fruit.  apex  I n f l o r e s c e n c e f e w - (4-42) t o  sepals  i n some montane f o r m s ;  Fruiting  dentate  or l o n g e r , white, t a p e r i n g g r a d u a l l y to a  to yellow; f i l a m e n t s  the ovary  to rusty  sinuate  ( a s h i g h as 7 4 ) , o p e n , s p r e a d i n g ,  regular,  clavate  dentate;  above,  obtusely conical; rachis usually i n d e f i n i t e .  petals  red  sinuate  gradually  glabrous  tcmentose to r u s t y v i l l o u s below; margins deeply to  exstipulate;  Columbia  the Olympic Mountains to east  Upper Biver  central  Vancouver I s l a n d . ,  Be p r e s e n t a t i v e  Specimens  BBITISH COLOMBIA: Carter  CI.59-  Vancouver I s l a n d ,  A l b e r n i , Vancouver I s l a n d ,  (V); 24  April  1914,  r o c k y b l u f f s , C o w i c h a n L a k e , B a l d Mt. ,  aarch  1940,  I.M.  Cowan  s.n.  (V);  in  141  crevices  of  rocks,  soil  damp,  n. end  of  V a n c o u v e r I s l a n d , (n=10), 20 F e b . 1 9 7 1 , D.L. mossy  wet  rocks.  J . B. A n d e r s o n mi  n. o f  84  Hill  H i l l , Vancouver I s l a n d ,  Lk.,  12.5 mi w . o f  Hwy.,  11 A p r i l  1958. C L .  portion  28  June  1938,  L.E.  Detling  e. f a c i n g c l i f f , S a n t i a m fi., 20 1951,  JU  Cron^uist  Falls,  9 April  6828  r o c k s and r o c k y c l i f f s s.n. of  J?£ck  1452J[  (OSC);  The  Dalles  sterile-14, 274  2800»,  moss mats on  (OSC);  Multnomah 1903.  Co.:  M.».  (WTB); a l o n g Sandy B i v e r a t C o l u m b i a Hwy.,  22  March  fertile-190,  Columbia  Biver  on  Gorman junction  1926,  m o i s t s l o p e s , n e a r Multnomah F a l l s ,  cn  21501  H.E.  18  April  wet c l i f f s , 6 mi  Hwy.,(pollen  E. P e r k i n s ) , 27 March  w«  fertility,  1946, H.H.  Baker  (OSC). WASHINGTON: C l a r k Co.: on wet r o c k s b e s i d e r o a d , 1 mi  w.  of  Clark-Skamania  1 9 7 1 , D.L. Mt.  K r a u s e 2-7J  Colonel  1931,  J.H.,  Co.  l i n e , St.  B i n g e n , 300«, 26 A p r i l  14, n=10,  6583  wet  14  (OSC);  Bidge,  Klickitat  1950, L.S. Rose 50073  5500*, Co.:  (OC);  March  cliffs  1930, J . J J . Thompson 9404  r o c k y c r e s t of Constance  Thompson  Hwy.  (DEC); G r e y s H a r b o r Co.:  Bob, 3500», 12 J u l y  J e f f e r s o n Co.:  rock  Mul'ck  Creek  1935, J . j f . Thompson 17370 (BS) ; U a s c o Co.; of  Columbia  Sweethome,  n e a r E l k B o c k , 12 A p r i l  and e.  Selagjnella  ( H S ) ; M a r i o n Co.: c l i f f . S i l v e r  ( h e r b a r i u m no. 22637)  e., S t a r k e Bd.  9-7.1  April  s.n.,  e. o f  2  7  1940, M. H r i j h t  mi  Krause  G»Leary H t n . ,  30.82. (OBE) ; L i n n Co.:  1895,  Cliffs  Old  H i t c h c o c k and C. V.  (SS) ; Lane Co.: s t e e p , g r a v e l l y , wet s o i l ,  (UBC);  17 K a r c h  knolls with  Hood B i v e r a l o n g  Lake,  1-71  (n=10) , 17 H a r c h 1 9 7 1 , D.L.  (UBC,UBC) ; Hood B i v e r Co.: o p e n b a s a l t i c  Biver  Krause.  ( V ) . CBEGON: C l a c k a m a s Co.; E l k Bock  Oswego  wallacij,  Shawnigan  (PC) :  30  3 mi e. Mason  of  May of Co.;  o u t c r o p s where p r o t e c t e d i n n a r r o w c a n y o n s , n e a r summit o f  142  fit. E l i n o r ,  11 J u n e 1 9 4 0 , E.G. J e j e r J 7 8 3  wet  Cape H o r n , 10 A p r i l ,  cliffs.  (§S) ;  Skamania  Co.:  27 May 1920, 1. S u k s d o r f  10365  (WS,UC) . The show  f o l l o w i n g specimens, u s u a l l y  slight  paniculate  resemblances  inflorescences  Strathcona  1931,  B., 6 0 0 0 * , 29 J u l y 1921,  J.W. Thompson 739S (UC, These  Linn  Co.:  specimens  longer pedicels e.  slope,  3082 (UC); M.J.  Peck  facing  14638  l . P . Taylor  Angeles,  head  s.n.  5000*,  (UCJ;  16  S. m a r s h a l l i i - i n t h e i r  cliff  reflexed  along  July  Santiam  clavate  sepals: R.  OREGON:  20 mi e.  of  A. C r o n q u i s t 6828 (UC) : L a n e C o . ; s t e e p  t o p o f House  are tenatively this  taxon  M t . , 31  classified  i s doubtfully  May  1926,  here  a s S.  distinct  from  F u r t h e r work i s n e c e s s a r y on t h i s p r o b l e m . OREGON:  J o s e p h i n e Co.: d r i e d up b u t l a t e l y Selma,  26  March  moist  Feb.  1 9 7 3 , M. W i l l i a m s  4.  Saxifraga  bluffs.  Eight  Dollar  1 9 2 6 , L . F . H e n d e r s o n .5845 (QBE) ;  Douglas Co.: r o c k y h i l l s i d e , t h i n s o i l ,  Elvander.  ridge,  (OSC).  specimens  rufidula.  near  Co.:  OSC).  and o f t e n  Marion Co.: c l i f f ,  h o w e l l i i Greene b u t  Mt.  COLUMBIA:  m o i s t , Mt. 0»Leary, 4 800*, 28 J u n e 1 9 3 8 . L.E. P e t l i n g  These  S«  Ht.  approach  Sweethome, 7 A p r i l 1951. n.  and c l u s t e r e d  t o S. o c c i d e n t a l i s : B R I T I S H  Co.: m o i s t r o c k y banks,  filaments,  filaments  10481 ( V ) . WASHINGTON: J e f f e r s o n  Dosewallips  Clallam  clavate  areas,  P a r k , Mt. B o o s t e r Comb, V a n c o u v e r I s l a n d , J u l y 1937,  N.C. S t e w a r t of  in  o f higher mountain  10  mi  Beston  Rd.,  24  s . n . (OBE) .  latipetiolata  (C.L. H i t c h c o c k )  Perkins  and  143  S. o c c i d e n t a l i s v a r .  l a t i p e t i o l a t a C.L. H i t c h c . .  Pac,  Northw.,  1961.  (fl.H.Gorman 3 5 6 1 , S a d d l e Mt., C l a t s o p  20,  1915.  Univ.  Hash.  Press,  rhizome.,  Leaves  above, s p a r s e l y  long  gradually  a  to  puberulent margins  many-flowered  to  paniculate to  sepals  ovary  anthers  a t anther 1/2  linear  early  or  mountaintop northern  broad,  ciliate  widely  ovate,  ciliate,  faintly  to  region,  sericeous  below;  Infloresence  Flowers  perfect,  Tillamook  base,  linear;  grading  usually somewhat  regular,  persistent carpels obconic  sell in  becoming  s t y l a r beak r e f l e x e d ; g l a n d  remnant  the of  moist,  Counties,  the top c f the  dehiscence,  anther  fruit.  higher grassy  the  or  tissue;  at  into  into  stylar  inferior  encircling  "balds"  broad  a f l a t t e n e d disc covering  Fruiting  Shallow  a  filaments  dehiscence, more  fie p r e s e n t a t j v e  tapering  petiolar  rusty  erose.  indistinct.  gland  ridge  July.  e x s t i p u l a t e blades  ( o c c a s i o n a l l y corymbiform) head; c e n t r a l  yellow;  superior i n f r u i t . a  seldom  r e f l e x e d ; p e t a l s a b o u t 2.5 mm l o n g o r l o n g e r ,  umbonate o b c o n i c ; ovary  stout,  below,  undulate  tapering gradually  fruit;  Co., O r e g o n , J u n e  (46-231), congested, c o n i c p a n i c l e , t o a  evident  5-merous; white,  villous short,  dentate  rachis  H i t c h c . e t a l . , 111:49.  herb w i t h s h o r t ,  simple,  above and b e l o w t o  congested,  PI.  WTU!)  Perennial, rosette-forming branching  Vase.  (n=ca. 38) L a t e  May t o  volcanic  plugs  areas  C l a t s o p and  of  and  Oregon.  Specimens  OBEGGN: C l a t s o p Co.: r o c k y  slopes.  Saddle  Mt.,2800-3300*,  144  20  June  1915. a . l .  S a d d l e Mt., fir-spruce  28  June  forest,  S a d d l e Mt., 2 2 0 0 ' , The resemble  Gorman 3 5 6 1 I S O T Y P E  following  1952, J . T .  very  19 J u n e  April  (0C); Douglas  as  7906 ( O R E ) .  S . i n t eg r i f o l i a  i n l e a f shape, broad s h o r t leaf  shade,  surface:  but  petioles,  OREGON:  Pclk  p l a c e s , Monmouth, 20 May 1893 . W.J. S p i l 1 m a n 78  <WS); M a r i o n C o . : g r a v e l l y 5  s.n.  1932, L.E. B e t l i n g  are classified  S. l a t i p e t i o l a t a  wet  Howell  slopes.  r o c k c r e v i c e s nw. e x p o s u r e , m o d e r a t e  and w h i t e p u b e s c e n c e c n t h e u p p e r Co.:  (WS); m o i s t o p e n  soil,  common, S a l e m , B r o o k s  Pasture,  1 9 1 9 , fl.W. Gorman 4J4JJ (WS) ; WASHINGTON: T h u r s t c n C o . :  Rock P r a i r i e ,  12 May 1934, I.£. Q$i§ JL§93 (IS) .  cojsayjsioNS The  present s t u d i e s i n d i c a t e that  S» o c c i d e n t a l i s which H i t c h c o c k list  evolutionary  e t a l . (1961) a n d H i t c h c o c k  trends  occur  ranges.  among  some  .polyploid i n d i v i d u a l s c e r t a i n combinations each  of  these,  t a xa c a n  be  involving  in  character  certain  l e s s importance  of characters are  to  or  of  tetraploid  cf  areas, esjpecially i n  tp  from  the  the  S. i n t e q r i f o l i a  and  species  g r c u p a p p e a r s t o be  hexaploid  of  diploid  S.. a e g u i den t a t a and  i s confirmed  without  species  i n  a s a c y t o l o g i c a l l y and  agreement  with  (Small and Bydberg  1916, K r a u s e and Beamish Columbia  Biver  Gorge  hybridization  several  1 9 0 5 , Hacoun 1S73).  area  m o r p h o l o g i c a l l y b u t t h e r e s e m b l a n c e may  complex  maintain  a 11 p p p i y g 1 o i d y  and  have a r i s e n d i r e c t l y  treatments  E n g l e r and I r m s c h e r  S. o c c i d e n t a l i s result  variable  taxonomic  individuals  separated  taxa.  aeguidentata  morphologically  are  inter<jr.adation  sufficient  hybridization  members  with other  Saxifraga  1906,  Jhejsr  --.tlsej-taxenoiiic -^dif f 4euitiej§^®sn.a----these -  p l a n t s f r o m t h e same l o c a t i o n  previous  distinct  but i t i s p o s s i b l e that nearly i d e n t i c a l  tetraploid  and  ;  J.anx-  various  taxa  .  and-- c y ^ p l p g ^ i c a j . l y  Althpush  (1973)  dentata,  1  taxa  and p o p u l a t i o n s , t h e ^|.^t4]|.qtjQBS;- b.ased-,on-  attributed  hybridization  and C r o n q u i s t  rufidula,  complex. A u t o p o l y p l o i d e v o l u t i o n i n t h i s  and  The v a r i e t a l  S. o c c i d e n t a l i s - -  within  9S030fifei£§llX#  much o f t h e i r  does  namely  c f the  are n o t merely e n t i t i e s which r e p r e s e n t  I S * PJipJ^SJ-SaAA X# over  reassessment  s p e c i e s complex i s necessary.  f o r S. o c c i d e n t a l i s .  latipetiplata,  a  and p o l y p l o i d i z a t i o n  Some  resemble be  the  probably  146  i n v o l v i n g S. i n t e g r i f o l i a in  var.  t h a t a r e a . There i s e v i d e n c e  S. i n t e g r i f o l i a occurring Gorge.  var.  between  populations  tetraploid  the r e s u l t o f  t h a t some i n t r o g r e s s i o n  claytoniifolia  i n sympatric  The  claytoniifcliaor i t s relatives  and  along  ancient  hybridizations  S. o c c i d e n t a l i s  resemblance  with  and  and  polyploidy  var.  S.  perhaps  j.ntegrifpjLj,a  leptopeta•la.  Even  is  g e o g r a p h i c a l l y i s o l a t e d and g e n e t i c a l l y d i s t i n c t  with  hybrids  between  crosses  Numerical  group.  other  or  Artificial  more  distantly  tend  to  Biver  Gorge to  Vancouver I s l a n d  siiilar  cytolcgically Oregon  and  the  S. a e g u i d e n t a t a  and  unknown northwest  pollen  similarities related  distinguish S.  Olympic and  a  aeguidentata  S. o c c i d e n t a l ' s  h y b r i d i n d i v i d u a l s t h a n do S.  between  southwestern  also  similarities  relationship and  cytological  presumably  Columbia  closer  polyploid  greater  treatments  p o p u l a t i o n s show  p l a n t s from  features.  S. a e g u i d e n t a t a and S. o c c i d e n t a l i s a r e  involving  S. a e g u i d e n t a t a  that  morphological  s e e d s t e r i l e a n d show no  entities.  Biver  S. a e g u i d e n t a t a  characteristic  than  Columbia  sympatric  i t appears  several  and  the  i s t h e r e s u l t o f p a s t c o n t a c t and i n t r o g r e s s i o n  S. p c c i d e n t a l i s  presently  S. a e g u i d e n t a t a i s  S. o c c i d e n t a l i s genome i s a l s o p r o b a b l y  p r o g e n i t o r s s i m i l a r t o S. i n t e g r i f o l i a if  between  and  aeguidentata  Mountains.  The  the morphologically  entity,  S. h o w e l l i i .  C a l i f o r n i a deserves  of  further  study. S a x i f r a g a o c c i d e n t a l i s i s d e f i n e d as taxon its  which range.  S. n i v a l i s  shows e v i d e n c e Detailed a n d S.  a  variable,  montane  of h y b r i d i z a t i o n i n s e v e r a l areas of  examination  of  i t s relationships  t e n u i s as w e l l a s w i t h S. m a r s h a l l i i  with subsp.  147  Mishallii using  and  subsp.  artificial  S.  r e f l e x a . S.  S.  aeguidentata.  idahoensis  is  h y b r i d i z a t i o n s among d i p l o i d  nj.vaJJLs, and  subspecies  S. g o r m a n j i  the present study. , A major problem the  strong  plants  seasonal  which  are  a l t i t u d e s and  necessary.  of  treated  S.  o c c i d e n t a l i s and  a  distinguishable  Clark  R o c k , O r e g o n and  to  w i t h such  be  studies  to  flowering  regimes  S.  here  as  a  a e g u i d e n t a t a . f o r m e r l y S.  m o r p h o l o g y and  County,  Washington.  specimens  that  S. o c c i d e n t a l i s  S.  aeguidentata  where  b u t i s i r r e g u l a r and S.  gormanii  S.  i n t e g r i f o l i a and  var. . and  (n=20,19).  and c o n s i s t e n t l y  shows  pollen closer S.  latipetiolata.  fail  Some  specimens  frcm  rufidula•,  with  closely  Tetraploids  and  extending  resemble  group  Artificial  tetraploid  S.  germanii  t o undergo m e i o t i c d i v i s i o n s  fertility phenetic  latipetiolata.  tend toward  with  hybrids  involving  S.  ajSIiajaii  m e i o s i s i n t h e F1 h y b r i d d o e s  t h a n t o S.  var.  species  Oregon  site  anther t i s s u e , i n c o n t r a s t t o crosses  and  diverse  type l o c a t i o n i s Elk  populations i n the numerical s t u d i e s .  between t e t r a p l o i d  plants  frcm  The  diploid  in  in  these  a geographical distribution i n  tetraploid  sterile  would  occidentalis  separate  nearby  are  diploid  w o u l d be a u s e f u l a d d i t i o n  Range and L o w e r W i l l a m e t t e R i v e r o f  to  occidentalis,  s e p a r a t i o n i n f l o w e r i n g t i m e s among  adapted  work  latitudes.  is  north  S.  S. .ffiargfoaAlii-  S a x i f r a g a g o r m a n i i S u k s d o r f , f o r m e r l y S.  the Coast  Further  is  low.,  affinities formerly  ocjej, d e n t a l i s  As  a  occur group,  to v a r i e t i e s S.  gccidentaljs  o r S.  aeguidentata  S. m a r s h a l l j i s u b s p .  marshallii  f u r t h e r s t u d i e s , e s p e c i a l l y i n the Willamette River area  needed t o c l a r i f y t h i s  problem.  of  are  148  Evidence Hitchcock) S.  is  presented  P e r k i n s and  occidentalis.,  similarities  Elvander  to  a  S.  is  and  a  Therefore  most  contact  where with  cytclogically  S.  of  in  S. S.  related  distinctive  aeguidentata  integrifolia  T h e s e can  or  the  be  t o S.  in  several  areas,  and  S,  or  oregana.  previous  S.  contact  between  i d a h o e n s i s and  S.  of  more  in  meiosis  However  f r o m S.  oregana  features.  close  syapatric var.  morphologically  which var,  Hybrid may  swarms involve  the or  S.  result  of  mixed  diploid  h e x a p l o i d S.  aeguidentata  as of  marshallii  ?ar, , leptopetala. parental  entity  or e x h i b i t reduced f e r t i l i t y  n o t uncommon, e s p e c i a l l y i n t h e C o l u m b i a H i v e r Gorge a r e a populations  and  integrifolia  likely  C r y p t i c n a t u r a l h y b r i d s w h i c h r e s e m b l e one abnormal  confirm  species.  occidentalis  integrifolia  of  S. i n t e g r i f o l i a  jntegrajglia are  undergo  relatives  intermediate habitats i n  comes  seme  (n=3 8) .  integrifolia  morphological  recognized  p a r e n t a l e n t i t i e s but o t h e r s  and  S.  by s t u d i e s o f p o l l e n f e r t i l i t y .  aeguidentata  subsp,  analysis a l l  Californian  occur  from  several  S.oregana  geographically isolated  intermediates  claytoniifolia.  occur  of  i t i s t r e a t e d here as a s e p a r a t e  areas  has  numerical  from s t u d i e s  more c l o s e l y  number  Sterile  species  S. i n t e q r i f o l i a - c o m p l e x .  a f f i n i t i e s with  and  e c o l o g i c a l l y and  has  the  (CL,  ( E l v a n d e r , p e r s o n a l c o m m u n i c a t i o n , 1978)  that i t i s probably it  and  S. r h o m b o i d e a ,  inteqrifolia  and  <n=ca.38), and  hybrid origin  latipetj.olata  latipetiolata  S. o r e g a n a  Other s y s t e m a t i c evidence group*  S.  i s a l s o a separate  Saxifraga  Morphological, cytogical indicate  that  and t e t r a p l o i d  p l a n t s , but a l s o i n  are  among  or t e t r a p l o i d  and  ether  and  areas  149  for  o t h e r t a x a s u c h as S,  is  difficult  to  tell  inteqrifolia  whether these  the Columbia a i v e r Gorge, are siblinq  entities  interspecific m o r p h o l o g y and One is S«  apparently  an  inteqrifolia.  possibility a  The  closely  most  more  resemble  between  appears  to  the  inteqrifolia  treated  here as b e l o n g i n g  most l i k e l y b e e n one  region  needs  t o S,  of  several  be  to  and  of  classify  Its  this i t with  Mima  s p e c i e s ) . T h i s c y c l e has  forms.  rise In  S.  mixed p l o i d y l e v e l s , through  close  and  Ft.  e v o l u t i o n i n the group independent  ( i n t h e a b s e n c e o f an a v a i l a b l e  give  population  apparent  the  The  further investigation. I t i s  probably  parental  occurred  in  (n=29)  occidentalis  f o l l o w e d by chromosome d o u b l i n g and  to  of  occidentalis.  reduction  introgressant  (EP605),  treatment  p l a n t s from  aneuploid  areas  S.  entity.  general pattern of p o l y p l o i d  hybridizations  parent  that i t i s a r e l i c t u a l  widespread  Washington,  The  two  f u r t h e r document i t s u n i g u e b a c k g r o u n d .  Lewis,  have  of c r o s s e s between  reasonable  c a n n o t be e x c l u d e d  r e l a t i o n s h i p t o S.  exhibit  in  preferences.,  population  once  several  individuals, especially  p l o i d y l e v e l s or segregates  intermediate  o c c i d e n t a l i s and  of  It  p o p u l a t i o n west of C h e h a l i s , Washington  problematical §•  which  habitat  claytoniifolia.  the r e s u l t  with d i f f e r e n t  crosses  var.  to  apparently higher  aeguidentata  higher  levels  the combination  diploid  coupled  itself in  levels  populations of  may  gametes  f r o m t h e same p o p u l a t i o n  but  followed  d o u b l i n g i s more l i k e l y . A s p e c u l a t i v e  by  c h a r t of the  chromosome relationships  involving  S.  with  and which  polyploidy  of r a r e unreduced  hybridization  with  nine-paired  repeated  ploidy  has  other  aegui.deBta,|a  and  taxa,  its  150  allies  i s presented  limited a r t i f i c i a l genetic  control  i n F i g u r e 36. The c o m b i n e d d i f f i c u l t i e s o f  hybrid combinations, of  synapsis,  parental entities limits  and  possible differences i n  the p o s s i b i l i t y  of extinct  t h e p r e c i s i o n o f any such s p e c u l a t i o n .  Bagging t e s t s demonstrate that t h e p l a n t s i n v e s t i g a t e d a r e most  likely  evidence  facultative  sexual  i n favor of apomictic  earlier  flowering  seed  members  of  S. S£gJdentajLj.s, S, a e g u i d e n t a t a . of  setting  relatively pollinators var. §*  more  autogamous  higher  numbers  than  e  observations floral  S.  may  of  intearifolia  bagging  a n d S, g o r m a n j i ) and  may  capable  Dipteran  (S. i n t e g r i f o l i a  var.  claytoniifolia. These  be c o r r e l a t e d w i t h t h e g e n e r a l d i f f e r e n c e s i n between  t h e two  gynodioecy  in  var., i n t e g r i f o l i a .  tests  {  f r e q u e n t e d by  versus  relatives  S. i n t e g r i f o l i a  group  are  be  Hymenopteran  somewhat  v a r . l e p t o p e t a l a . a n d S. l a t i p e t i o l a t a ) .  ia  morphology  occurrence  The  the o c c i d e n t a l i s  seed of  There i s l i t t l e  production.  the integrifolia  istegrifolia,  JB£ ggJf93-  outcrossers.  and  pollination  groups  as  well  as the  certain  populations  of  Although  the  of  studies  show  results  interesting  d i f f e r e n c e s between t h e major s p e c i e s complexes s t u d i e d ,  within  t h e m o r p h o l o g i c a l l y s i m i l a r S. Q c c ' d e n t a l i s - a e g u i d e n t a t a  species  group, they a r e l e s s u s e f u l a s a taxonomic The apparently evidence  post-glacial  v a r i e d . Bandhawa for glacial  r e f u g i a l areas in  contrast  glaciated  history and  of  tool.  t h e group  Beamish  (1972)  r e f u g i a i n northwestern  apparently contain r e l i c t u a l tc  widespread  areas,  Bandhawa  i s complex and  North  diploid  reviewed America.  the The  populations  polyploid c o l o n i z e r s of surrounding and  Beamish  (1972)  used  the  151  Figure 36: Speculative polyploid formations within the S. o c c i d e n t a l i s and S. a e g u i d e n t a t a ( t h e synonym- r u f i d u l a i s used i n t h e f i g u r e ) lineage with e m p h a s i s on t h e Columbia River Gorge r e l a t i o n s h i p s . Several other constructions are also logically p o s s i b l e . Numbers i n parenthesis are not supported i n the l i t e r a t u r e . J o i n i n g lines indicate possible allopolyploid r e l a t i o n s h i p s but in certain instances autopolyploidy i s p r o b a b l e {CF. S. r e f l e x a ) . F o r most o t h e r s , a l l o p o l y p l o i d o r i g i n s are p o s s i b l e but l e s s l i k e l y .  152  ntegrifolia ca 5 5  oregana latipetiolata 38 38  SPECULATIVE RELATDNSHIPS  occidentalis 29->28  integrifolia (20)-> 19  rufidula 29^28  leptopetala. (20) ->19 / occidentalis gormanii 19 "20-> 19  rufidula 10  (integrifolia) 10?  P r o t o integrifolia (10)  rufidula . (20)^19  Proto occidentalis (10)  153  d i s t r i b u t i o n o f 10 and an  example  of  19 p a i r e d  p o p u l a t i o n s o f S. f e r r u g i n e a  s u c h a p a t t e r n . The  narrow s o u t h e r n and  d i s t r i b u t i o n o f d i p l o i d s and  widespread  within  some  S. o c c i d e n t a l i s  and  ranges  members  c o n f o r m w e l l t o t h e p a t t e r n e x h i b i t e d by diploid  S. a e g u i d e n t a t a p o p u l a t i o n s  glaciated  areas  have  Gorge.  and  presumably  interior  Similar  processes  are  complex also  S. l a t i p e t i o l a t a . Columbia  River  resulting  floristic  formerly Island,  i n areas south c f the contacts  between  elements i n the Columbia and  i n the h i s t o r y  Zones o f complex G o r g e may  Although  Vancouver  from  hybridization  evident  integrifolia  recclonized  o f t h e O l y m p i c M o u n t a i n s and  g l a c i a l boundary  S.  western  polyploids  S. f e r r u g i n e a .  p o l y p l o i d s and i n t r o g r e s s a n t s a r e a b u n d a n t  coastal  of  of  River  polyploidization o f S. g o r m a r i i  hybrid a c t i v i t y  p r o v i d e t h e raw  such  and  as  the  m a t e r i a l s i n t h e form  o f more v a r i a b l e gene p o o l s upon w h i c h s e l e c t i o n c a n o p e r a t e f u t u r e g l a c i a l or i n t e r g l a c i a l  epochs  (Stebbins  in  1571).  H e x a p l o i d and o c t o p l o i d p o p u l a t i o n s i n r e s t r i c t e d  areas  may  r e p r e s e n t the remnants o f a once  more w i d e s p r e a d d i s t r i b u t i o n  such  result  restriction  sympatry  may  between  be  two  the  tetraploid  S. o r e g a n a  the l a t t e r  from  category  r e l a t i o n s h i p t o S. A its  or  tetraplcid-diploid  S. c a l i f o r n i c a , with  The  absence  t h e C o a s t Range o f O r e g o n a p p e a r s t c be i n with  respect  to  its  probable  parental  jLatjpetiolata.  t h o r o u g h e x a m i n a t i o n i s n e c e s s a r y o f S, i n t e g r i f o l i a  relatives  or  of narrow ecogeographic  p r o g e n i t o r s w h i c h h a v e s i n c e become l o c a l l y e x t i n c t . of  as  including  S. .,  oregana.  S.  rhomboidea,  t h e l a t t e r two p e r h a p s h a v i n g r a t h e r c l o s e r  S. o c c i d e n t a l i s and S.  and and ties  m a r s h a l l i i r e s p e c t i v e l y . There i s a  154  s t r o n g need f o r a s y s t e m a t i c s t u d y o f t h e r e l a t i o n s h i p s disjunct  Western  (Spongberg  1972)  and  Eastern  such  as  §•  oregana-S. p e n s y l v a n i c a .  S.  r e f l e x a - S.  where  schedules  and  those  are i n c o m p l e t e field  season  S.  for  many  conflicts  careliniana  taxa,  with  especially  institutional  which are l e s s a c c e s s i b l e . A t t e n t i o n  and  a r e a s o f i n t r o g r e s s i o n i n t h e Opper W i l l a m e t t e  should  f u r t h e r s y s t e m a t i c i n v e s t i g a t i o n o f h y b r i d swarms Biver  area,  Mountains d f n o r t h e a s t e r n Oregon, the Spokane a r e a  o f e a s t e r n W a s h i n g t o n , and C a l i f o r n i a and  pairs  virginiensis,  mags[bap. 1 j i - S .  given  Wallowa  species  S. f e r r u g j n e a - S . m i c b a u x i i .  be  the  tc  the  American  S. o c c i d e n t a l i s - 5 .  m i c r a n t h i d i f d j i a and  . Chromosome c o u n t s those  North  between  southern  the h i g h e r mountain areas  Oregon..  of  northern  155  LITERATURE C££ED  A l e x o p o u l o s , C , J . , and E.S.Beneke. 1952* Laboratory manual for introductory mycology. Burgess P u b l i s h i n g C o . , M i n n e a p o l i s , Minn. Antonovics,J. 1968. populations. V. 23:219-238.  Evolution in closely adjacent plant Evolution of self-fertility. Heredity  B a c i a g a l u p i , B . 1944. J n L. Abrams. I l l u s t r a t e d flora of the P a c i f i c States. Vol, I I . Stanford University Press, Stanford, California, B e a m i s h , K . I . 1961. S t u d i e s c f m e i o s i s i n t h e g e n u s t h e P a c i f i c N o r t h w e s t . Can. J . B o t , 3 9 : 5 6 7 - 5 8 0 .  Saxifragaof  Beamish, K.I. 1967. A P a c i f i c C o a s t S a x i f r a g e w i t h 10 p a i r s o f chromosomes: m e i o s i s , d e v e l o p m e n t o f t h e f e m a l e gamete, and s e e d p r o d u c t i o n . Can, J . B o t . 45:1797-1801. B l o o m , ».L. 1976., M u l t i v a r i a t e a n a l y s i s o f t h e i n t r o g r e s s i v e r e p l a c e m e n t o f C l a r k i a n i t ens by CjLajckja s p e c i c s a p e l vantha (Onagraceae). Evcluticn~30:4*12-4 247" Brittain, H.H, And D.E. Newton. 1933. A s t u d y i n t h e r e l a t i v e c o n s t a n c y o f h i v e b e e s and w i l d b e e s i n p o l l e n gathering. Can. J . B e s . 9:334-349., Carr, G.R. Calycadenia  1975, Chromosome e v o l u t i o n and reduction in p a u c i f l c r a ( A s t e r a c e a e ) . E v o l u t i o n 29:681-699,  C h a m b e r s , K. 1974. N o t e s on Madrono 2 2 : 2 7 8 - 2 7 9 .  the  f l o r a o f C l a t s o p C o u n t y , Oregon.  C r c v e l l e , T . J , 1970. A n a l y s i s o f c h a r a c t e r v a r i a t i o n i n and s y s t e m a t i c s . A n n u a l Bev, E c o l , S y s t , 1:55-98, Damiolt, J, 1968, Zur c y t o t a x o n o m i c der g a t t u n g I I I . B e r l i n D e u t . B o t . Ges. 84:43-52. Dambolt, J., and D. Podlech. u n t e r s u c h u n g e n an S a x i f r a g a - S i p p e n Engl, e t I r m s c h . , B e r l i n deut. Bot.  ecology  Saxifraga  L,  1965. Zytotaxonomische der grex Exarato-moschatae Ges. 7 7 : 3 3 2 - 3 3 9 .  Danick, B.P. And B.V. Burns. 1975. M u l t i v a r i a t e a n a l y s i s o f h y b r i d p o p u l a t i o n s . N a t u r . Can. 102:835-843.  156  D i x o n , 8 . J . 1970. B.M.D. b i o m e d i c a l computer programs. Health Sciences Computing F a c i l i t i e s , Department o f P r e v e n t i v e Medicine, U n i v e r s i t y o f C a l i f o r n i a a t Los Angeles. Don, D. 1822. A monograph Soc. 13:341-452.  o f t h e genus S a x i f r a g a .  Trans.  Linn.  llvander, P.E. 1975. B i o s y s t e m a t i c s t u d i e s o f a new s p e c i e s o f S a x i f r a g a and i t s r e l a t i v e s , MSc. T h e s i s . University of Washington, S e a t t l e . llvander, P.E. 1978. S y s t e m a t i c r e l a t i o n s h i p s within the S. i n t e g r i f o l i a c o m p l e x (Saxif ragaceae). Abstract I n Bot. S o c i e t y o f A m e r i c a , M i s c . S e r i e s P u b l . 156:82. Engler, A., a n d E. I r m s c h e r . 1916. Das P f l a n z e n r e i c h . Ed. A. E n g l e r . W i l h e l a Engelmann, L e i p z i g I V , 117, I . E y l e s , A. C. , a n d 8. E. B l a c k i t h . 196 5. S t u d i e s on h y b r i d i z a t i o n in Scolopostethus Fieber {Heteropteraj lygaeidae). Evolution 19:465-479. F a e g r i , K,, a n d L. v a n d e r P i j l , 1966. The p o l l i n a t i o n e c o l o g y . Pergamcn P r e s s , O x f o r d . Free, J . B . 1963. The E c o l . 32:119-131.  flower constancy  pollination.  of  o f h o n e y b e e s . J . Anim.  F r e e , J . B . 1968. D a n d e l i o n a s a c o m p e t i t o r bee v i s i t s . J . A p p l . E c o l . 5:169-178. H a g e r u p , 0. 1950. R a i n B i o l . M e d d e l . 18:1-19.  principles  to  fruit  trees  for  K g l . Danske V i d . S e l s k .  H a g e r u p , 0. 1 9 5 1 . P o l l i n a t i o n i n t h e F a r o e s — i n spite of rain and p o v e r t y o f i n s e c t s . K g l . Danske V i d . S e l s k . B i o l . M e d d e l . 18:3-48. Henderson, D.M. 1976, Northwestern blue-eyed B r i t t o n i a 28:149-176.  A biosystematic study of Facific grasses tsisyrinchium. Iridaceae).  Hitchcock, C.L., A. C r o n q u i s t , M, Ownbey, and J , a . Thompson. 1961. V a s c u l a r p l a n t s o f t h e P a c i f i c N o r t h w e s t . U n i v e r s i t y o f Washington P r e s s , S e a t t l e . H i t c h c o c k , C.L. , a n d A. C r o n q u i s t . 1973. F l o r a o f t h e Northwest. U n i v e r s i t y o f Washinqton P r e s s , S e a t t l e . Hooker,J. 1:249.  1833. S a x i f r a g a i n t e q r i f o l i a .  Facific  F l o r a B o r e a l ! Americana  157  H o s s a i n , M.G. 1977. The s i g n i f i c a n c e o f chromosome association in an a d v a n c e d p o p u l a t i o n o f t e t r a p l o i d r y e . C a n . J . G e n e t . C y t o l . 18:601-652. Hunan, C. 1974. A b a r e f o o t d o c t o r ' s m a n u a l ( t r a n s l . ..of C h i n e s e ) . U.S. Dep. o f H e a l t h , E d u c a t i o n and W e l f a r e , P u b l i c Health Service, Nat. Inst, of Health. H y d e , H.A. 1950. S t u d i e s i n a t m o s p h e r i c pollen. IV.,Pollen d e p o s i t i o n i n G r e a t B r i t a i n . , N e w P h y t o l . 49:398-420. Hyde, H.A. 1969. A e r o p a l y n o l o g y Phytol.,68:579-590.  i n Britain--an  outline.  New  Hyde, H.A., a n d D.A. W i l l i a m s . 1961. A t m o s p h e r i c p o l l e n a n d spores as causes o f a l l e r g i c d i s e a s e : h a y - f e v e r , asthma, and t h e a e r o s p o r a . Advancement S c i . 526-533. Jackson, B.C. 1962. I n t e r s p e c i f i c h y b r i d i z a t i o n i n H a p l o p a p p u s and i t s b e a r i n g on chromosome e v o l u t i o n i n the Blepharodon s e c t i o n . Amer. J . B o t , 49: 119-132. Johnson, A.M. 1923. A r e v i s i o n o f t h e N o r t h A m e r i c a n the section Boraphila Engler o f the genus ( T o u r n e f . } L . M i n n . S t u d . B i o l . S c i . 4: 1-109.  species of Saxifraga  Krause, D.L., a n d K . I . B e a m i s h . 1 9 7 2 . Taxonomy o f S a x i f r a g a o c c i d e n t a l i s and S. m a r s h a l l i i . C a n . J . B o t . 50:2131-2 141. Krause, D.L., and K . I . B e a m i s h . 1973. N o t e s o n S a x i f r a g a o c c i d e n t a l ' s and c l o s e l y r e l a t e d s p e c i e s i n B r i t i s h C o l u m b i a . S y e s i s 6:105-113. :  Kyhos, D. 196 5. The i n d e p e n d e n t a n e u p l o i d o r i g i n o f two s p e c i e s c f C h a e n a c t i s ( C o m p o s i t a e ) f r o m a common a n c e s t o r . E v o l u t i o n 19:26-43. ~ Levin, D.A., and W.W. A n d e r s o n . , 1970. C o m p e t i t i o n f o r p o l l i n a t o r s between s i m u l t a n e o u s l y f l o w e r i n g s p e c i e s . Amer, N a t u r . , 104: 455-467. Lewis, H.; 1 9 6 1 . E x p e r i m e n t a l s y m p a t r i c p o p u l a t i o n s c f C l a r k ' a . Amer. N a t u r . 95:155-168. L e w i s , H., a n d P. Baven, E v o l u t i o n 12:319-336.  1958. B a p i d  evolution  in  Clarkia.  Lewis, W.H.,and Y. Suda. 1 9 7 6 . D i p l o i d s and p o l y p l o i d s f r o m a s i n g l e species population: temporal a d a p t a t i o n s . , J . Hered. 67:391-393.  158  L i n n a e u s , C. 1753, S p e c i e s  plantarum. F i r s t ed. Stockholm.  Macoun, James 1906., C o n t r i b u t i o n s Ottawa Nat. 20:162-171.  t o Canadian botany.  XVIII.  M c N e i l l y , T., and J . A n t o n o v i c s . 1967. E v o l u t i o n in closely a d j a c e n t p l a n t p o p u l a t i o n s . I V . B a r r i e r s t o gene f l o w . B e r e d . 23:205-218. Miller, J.M. 1976. Variation i n populations of p e r f o l i a t e ( P o r t u l a c a c e a e ) . S y s t . B o t . 1:20-34.  Claytonia  M o o r e , fi.J. 1959. I n C a l d e r , J . A . , and D.B.C. S a v i l e , S t u d i e s i n Saxifragaceae, I I . Saxifraga sect. Trachyphyllum i n North A m e r i c a . B r i t t o n i a 11:228-249.. Namkoong, G. 1966. Statistical B i o m e t r i c s 22:488-502.  analysis  of  Crnduff, B. 1969. Beproductive s y s t e m a t i c s . Taxcn 18:121-133.,  biology  in  P a c k e r , J.G.  introgression. relation  1968. IOPB chromosome number r e p o r t s . T a x o n  Paterniani, E. 1969. Selection f o r reproductive between two p o p u l a t i o n s o f m a i z e , Z e a mays L. 23: 534-547. P e r c i v a l , M.S.  1965. F l o r a l b i o l o g y . Pergamon P r e s s ,  Proctor, M., and P. C o l l i n s , London.  Yeo.. 1973.  The p o l l i n a t i o n  to  17:287.  isolation Evolution Oxford. of flowers,  B a j h a t h y , T., a n d H. Thomas. 1972. G e n e t i c c o n t r o l o f chromosome p a i r i n g i n h e x a p l o i d o a t s . N a t u r e (London) 2 3 9 : 2 1 7 - 2 1 9 . Bandhawa, A.S., and K . I . B e a m i s h . 1972. The distribution SaxjLfraga ferruginea and the problem of r e f u g i a n o r t h w e s t e r n N o r t h A m e r i c a . Can. J . B o t . 5 0 : 7 9 - 8 7 . Biley, B. 1960, 15:407-429.  D i p l o i d i z a t i o n of  polyploid  wheat.  of in  Hered.  Biley, B., a n d C, N, Law, 1965, G e n e t i c v a r i a t i o n i n chromosome p a i r i n g . A d v a n c e s i n G e n e t . 13:57-114. B i s i n g , J.D. 1968. A m u l t i v a r i a t e a s s e s s m e n t o f interbreeding b e t w e e n t h e c h i c k a d e e s P a r u s a t r i c a p i l l u s and £, c a r o l i n e n s i . S y s t . Z o o l . 14: 131-132. ~  159  Sass, J . E . 1958, Botanical S t a t e C o l l e g e P r e s s , Ames,  microtechnique,  S c h i l l i n g , E.E,, a n d C B . Heiser. numerical taxonomic study of Taxon 2 5 : 4 5 1 - 4 6 2 .  3 r d e d . The  1976. Re-examination of a Solanum s p e c i e s and h y b r i d s .  S c b u e l e r , F.8., and J.D. Rising. 1976. P h e n e t i c n a t u r a l h y b r i d i z a t i o n . S y s t . Z o o l . 25:283-289. Sax, K. 1937. E f f e c t o f v a r i a t i o n c e l l d i v i s i o n i n Trandescantia.  Iowa  evidence  of  i n t e m p e r a t u r e on n u c l e a r a n d Amer. J . B o t . 2 4 : 2 1 8 - 2 2 5 .  S i n g h , I . S . 1975. E f f e c t s de l a t e m p e r a t u r e e t du n i v e a u x f l o r a l sur l a f e r t i l i t e ' p o l l i n i q u e chez l a p e t u n i a . Ann. Amelior. P l a n t . 25:365-37 0. S i m p s o n , D.H., and E.N. Duncan. 1956. C o t t o n i n s e c t s . Agron. J . 48:305-308,  p o l l e n d i s p e r s a l by  S k c v s t e d , A. 1934, C y t c l o g i c a l s t u d i e s i n t h e t r i b e Dansk. B o t . A r k . 8: 1-50. Small, J . K . , and P.A. F l o r a 22:81-158.  Saxifrageae,  R y d b e r g . 1905. S a x i f r a g a c e a e . I n N. Amer.  S m i t h , D.H, 1969, A taximetric study of Vacciniu n n o r t h e a s t e r n O n t a r i o . Can. J . B o t . , 4 7 : 1 7 4 7 - 1 7 5 9 .  in  Snow, R. 1963. A l c o h o l i c h y d r o c h l o r i c a c i d - c a r m i n e a s a s t a i n for chromosomes i n s g u a s h p r e p a r a t i o n s . Stain Techncl. 23:9-13. Sckclovskaya, A.P. 1958. On t h e c o r r e l a t i o n b e t w e e n t h e number o f chromosomes a n d t h e s i z e c f p o l l e n g r a i n s i n t h e Arctic species o f R a n u n c u l a c e a e and S a x i f r a g a c e a e . ( I n R u s s i a n ) . Bot. Zhur. 43:1146-1155. Spongberg, S.A. 1972. T h e genera o f Saxifragaceae i n the S o u t h e a s t e r n O n i t e d S t a t e s . J . A r n o l d Arboretum 53:409-453. Stebbins, G.L. 1971. Chromosomal Edward A r n o l d L t d . , I c n d o n , S t e p h e n s , S.G. segregating  evolution i n higher plants.  1956, The composition o f an c o t t o n p o p u l a t i o n . Amer. N a t u r .  open pollinated 90:25-39.  Subhasi, U. 1975, I n t e r s p e c i f i c h y b r i d i z a t i o n and a n e u p l o i d y the genus N i c o t i a n a . C y t o l o g i a 40:735-741.  in  160  T a y l o r , B.L., and G. A. M u l l i g a n . 1 9 6 8 . F l o r a o f t h e Queen C h a r l o t t e I s l a n d s , P t . 2. C y t o l o g i c a l a s p e c t s o f t h e v a s c u l a r plants. Plant Research I n s t i t u t e , C e n t r a l Experimental Farm, Ottawa, Ont. T e r - A v a n e s i a n , E.y. 1978. S i g i n i f i c a n c e o f p o l l e n f e r t i l i z a t i o n . B u l l . T o r . B o t . C l u b 105:2-8.  amount f o r  Tobgy, H.A. 1943.„ A c y t o l o g i c a l s t u d y o f C r e p i s f u l i q i n o s a . £• n e g l e c t a . and t h e i r F1 h y b r i d , and i t s b e a r i n q on phyloqenetic reduction i n chromosome number. J . Genet, 45:67-111. V e l d m a n , D.J. 1967. FORTRAN proqramminq s c i e n c e s . H o l t , R i n e h a r t , and W i l s o n .  f o r the  behavioral  Ward, J . H . 1963. H i e r a r c h i c a l q r o u p i n q t o o p t i m i z e f u n c t i o n . J . A m e r . . S t a t i s t i c a l A s s n . 58:236-244.  objective  W a t t , G. 1972. A d i c t i o n a r y o f t h e e c o n o m i c p r o d u c t s o f VI. P t . 11. R e p r i n t . P e r i o d i c a l Experts, Vivek S h a n d a n a , Delhi.„  India. Vihar,  W e l s h , S.L. 1974. A n d e r s e n ' s f l o r a o f A l a s k a a n d a d j a c e n t parts o f C a n a d a . B r i q h a m Young U n i v e r s i t y P r e s s , P r o v o , O t a h . W h i t e h o u s e , B.N.H. 1969. Ah a p p l i c a t i o n o f c a n o n i c a l p l a n t b r e e d i n g , G e n e t i c a A g r a r i a 23:61-96,  analysis to  161  APPENDIX  Insect  Specimens  List of insects captured while visiting Saxifraga flowers: P = w i t h g a x i f r a g a p o l l e n on body o r l e g s ; (No.)-number of v i s i t o r s c o l l e c t e d .  HYMENTOPTEBA: V e s p i d a e P ( 2 ) : P o l i s t e s fuscatus (1) ; Formicidae (1); Apidae: A | i s m e l i f e r a P ( 3 ) : Bombidae: • Bprobus spp. P (2) ; D i p r i o n i d a e ( 1 ) ; A n d r e n i d a e P f 1 1 ) .  COLEOPTEBA:  Elateridae (1).  DIPTEBA: Anthomyidae (3): Scatophaga spp., P ( 6 ) ; Tachinidae P(8); Bombyliidae: Bombvlius major P ( 2 ) ; Empidae P(7);Calliphoridae ( 1 ) ; Muscidae: Musea- d p m e s t i c a ( 1 ) ; Ceratopogonidae ( 2 ) ; M y c e t o p h i l i d a e P<2); Chironomidae {H); Syrphidae ( 3 ) : Metasyrphus s p p . (5) ; D a s v s v r o h u s sp. (1); S p h a e r o p h o r i a s p . (1) ; A g r c m y z i d a e (1) .  162  F i g u r e 3 7 : Map o f d i s t r i b u t i o n s o f S. o c c i d e n t a l i s ( i n O r e g c n , Washington, and s o u t h w e s t e r n B r i t i s h C o l u m b i a o n l y ) and S* g c r m a n i i . The single sguare i s the hexaploid intermediate population <S. o c c i d e n t a l i s x S. i n t e g r i f o l i a ) . ,  164  F i g u r e 38: Hap o f the distribution of S, a e g u i d e n t a t a (the synonym, S. r u f i d u l a is used in the figure) and S. l a t i p e t i o l a t a . D o t t e d l i n e i n d i c a t e s t h e w e s t e r n l i m i t of S. o r e q a n a and m o r p h o l o g i c a l l y r e l a t e d ! i n t .  166 A B B R E V I E T E D KEY  A. O v a r y l e s s t h a n h a l f i n f e r i o r sinuate-dentate  to evenly  at anthesis;  leaf margins  shallowly  dentate  B. N e c t a r g l a n d a d o u g h n u t - s h a p e d r i n g a t r h i z o m e s , when p r e s e n t , a f i n e , r e d d i s h  anthesis;  brown network  S_; ' g o r m a n i i BB. N e c t a r g l a n d r e d u c e d t o a n a r r o w b a n d ; r h i z o m e s stout,  horizontal C. I n f l o r e s e n c e filaments  flat-topped  subulate  CC. I n f l o r e s e n c e  filaments  unevenly sinuate-dentate  usually  to narrowly c o n i c a l  a  panicle,  S_. o c c i d e n t a l i s  at anthesis;  leaf,margins e n t i r e t o  ( d i s t a n t l y s e r r a t e i n S_. o r e g a n a )  D. P e t i o l e s a l m o s t l a c k i n g , g r a d i n g infloresence  with  long,  into blades;  vestiture  clear or pinkish-glandular  E. :Scapes u s u a l l y 0.5m  hairs  or. .less-. .^S.  EE.  obtuse-conical,  S_. a e g u i d e n t a t a  clavate  AA. O v a r y h a l f o r m o r e i n f e r i o r  or  not flat-topped,  clustered, headlike  in  short,  Scapes u s u a l l y more t h a n  latipetiolata 0.5m S_. o r e g a n a  DD. P e t i o l e s e l o n g a t e , b r o a d e n i n g m o r e o r l e s s into blades; glands  vestiture i n infloresence with  abruptly  reddish  S_. i n t e g r i f o l i a  

Cite

Citation Scheme:

        

Citations by CSL (citeproc-js)

Usage Statistics

Share

Embed

Customize your widget with the following options, then copy and paste the code below into the HTML of your page to embed this item in your website.
                        
                            <div id="ubcOpenCollectionsWidgetDisplay">
                            <script id="ubcOpenCollectionsWidget"
                            src="{[{embed.src}]}"
                            data-item="{[{embed.item}]}"
                            data-collection="{[{embed.collection}]}"
                            data-metadata="{[{embed.showMetadata}]}"
                            data-width="{[{embed.width}]}"
                            async >
                            </script>
                            </div>
                        
                    
IIIF logo Our image viewer uses the IIIF 2.0 standard. To load this item in other compatible viewers, use this url:
http://iiif.library.ubc.ca/presentation/dsp.831.1-0094491/manifest

Comment

Related Items