Science, Faculty of
Botany, Department of
DSpace
UBCV
Perkins, Walter Ethen
Doctor of Philosophy - PhD
University of British Columbia
In the Pacific Northwest, hybridization and polypoid variation have produced confusion in the relationships among Saxifraga rufidula (Small) James Macoun and its relatives. The entities from the Columbia River Gorge are particularly difficult to separate taxonomically. Some authors have recognized several species from that area while others recognize one highly variable, widely distributed species with varietal components. The present study approached the systematic treatment of S. rufidula and related subspecific taxa of S. occidentalis Wats. with data from numerical studies, studies of meiosis, observations of artificial and natural hybrid individuals and populations, breeding system experiments, observations of pollinators, and ecological observations. Polyploid intermediates and plants with introgressant characteristics are shown to occur, many of which appear to be the result of hybridization with the S. integrifolia species complex. Sufficient correlations of morphological, ecological and geographic discontinuities exist to substantiate the treatment of S. rufidula. S. occidentalls var. latipetiolata, S. occidentalis var. dentata, and S. occidentalis var. occidentalis as a species. According to the rules of nomenclature S. rufidula becomes S. aeguidentata. S. occidentalis var. latipetiolata becomes S. latipetiolata, S. occidentalis var. dentata becomes S. gormanii. and S. occidentalis var. occidentalis becomes S. occidentalis.
SYSTEMATICS OF SJ^IFRJGA BOFIDULJ AND BELATED SPECIES FBCM THE COLOMBIA BIVEB GOBGE TO SOUTHSESTEBN BBITISH COIUUBIA By HALTER ETHEN PERKINS B.Sc. , U n i v e r s i t y o f Oklahoma, 1S73 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOB THE DEGREE OF DOCTOB OF PHILOSOPHY i n 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 as conforming t o the r e q u i r e d s t a n d a r d THE UNIVERSITY OF BBITISH COLUMBIA Octo b e r , 1978 (8) W a l t e r Ethen 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 British 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 thesis for 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 P e r k i n s Department of Botany The University of British Columbia 2075 Wesbrook Place Vancouver, Canada V6T 1W5 5 O c t o b e r , 1978 R e s e a r c h S u p e r v i s o r : D r . R a t h e r i n e I . Beamish. ABSTRACT I n t h e P a c i f i c Northwest, h y b r i d i z a t i o n and p o l y p o i d v a r i a t i o n have produced c o n f u s i o n i n the r e l a t i o n s h i p s among S a x i f r a g a r u f i d u l a (Small)James Macoun and i t s r e l a t i v e s . The e n t i t i e s from t h e Columbia B i v e r Gorge a r e p a r t i c u l a r l y d i f f i c u l t t o s e p a r a t e t a x o n c m i c a l l y . , Some a u t h o r s have r e c o g n i z e d s e v e r a l s p e c i e s from t h a t area w h i l e o t h e r s r e c o g n i z e one h i g h l y v a r i a b l e , w i d e l y d i s t r i b u t e d s p e c i e s w i t h v a r i e t a l components. The p r e s e n t s t u d y approached t h e s y s t e m a t i c t r e a t m e n t of S. r u f i d u l a and r e l a t e d s u b s p e c i f i c t a x a o f S. o c c i d e n t a l i s Wats. w i t h d a t a from n u m e r i c a l s t u d i e s , s t u d i e s of m e i o s i s , o b s e r v a t i o n s o f a r t i f i c i a l and n a t u r a l h y b r i d i n d i v i d u a l s and p o p u l a t i o n s , b r e e d i n g system e x p e r i m e n t s , o b s e r v a t i o n s of p o l l i n a t o r s , and e c o l o g i c a l o b s e r v a t i o n s . P o l y p l o i d i n t e r m e d i a t e s and p l a n t s w i t h i n t r o g r e s s a n t c h a r a c t e r i s t i c s a re shown to o c c u r , many o f which appear to be the r e s u l t o f h y b r i d i z a t i o n w i t h the S. i n t e g r i f o l i a s p e c i e s complex. S u f f i c i e n t c o r r e l a t i o n s o f m o r p h o l o g i c a l , e c o l o g i c a l and g e o g r a p h i c d i s c o n t i n u i t i e s e x i s t t o s u b s t a n t i a t e t h e tr e a t m e n t 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. 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 v a r . , dent a t a , and 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 as a s p e c i e s . , A c c o r d i n g t o the r u l e s o f nomenclature S. r u f i d u l a becomes 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 v a r . , l a t i p e t i o l a t a 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 v a r . d e n t a t a becomes S. g o r m a n i i . and *>• 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 becomes S. O c c i d e n t a l i s . i i Table Of Contents ABSTRACT . .. . . .* . '/ i LIST OF TABLES i v LIST OF FIGURES . .:. ....... .. .... .. .......,>. y* w.-, .-.V.-. . V AC K NOWL E EG E H ENT S .................... ,<••• • • .... . ..... i x INTRODUCTION ......... .... . . . . . . . . . . . . . , , . . . . . . . / 1 MATERIALS AND METHODS .v.w. .V. .V. »V >v>./. .V. V. «^ . Y 6 Chromosomal Studies 13 Hybridization Studies ................................... 14 Numerical Studies ... .............. ...................... . 15 Breeding Systems ....... .... .... ...... •. .-......... 17 Habitat And P o l l i n a t i o n Studies 18 RESULTS AND DISCUSSION 19 Morphology .....».........• 19 Chromosomal Studies ......... ............................ 42 Hybridization .............................. ...... ....... 63 Numerical Studies ....................................... 87 Breeding System Observations • • • • 114 i i i Bagging T e s t s •».......*.........* *.•.«.*.... •.. 114 F l o r a l O b s e r v a t i o n s 118 Geography, H a b i t a t And P o l l i n a t i o n E c o l o g y O b s e r v a t i o n s . 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 . ..... . . . . . . . . . . . • .. ... ..... • • 161 I n s e c t Specimens ........................................ 161 i v LIST OF TABLES T a b l e I , L i s t of C o l l e c t i o n S i t e s 9 T a b l e I I . L i s t o f M o r p h o l o g i c a l C h a r a c t e r s Used f o r Computer A n a l y s i s . . . v . . v . . . . v . . .............. 16 T a b l e I I I . , Summary o f Chromosome Number f o r N a t u r a l P o p u l a t i o n s ...................... .... ...... ............ 44 T a b l e IV. P o l l e n F e r t i l i t y i n A r t i f i c i a l H y b r i d s ........76 T a b l e V. Baw P r i n c i p l e Components f o r PCA A n a l y s i s ........99 T a b l e V I . , Bagging T e s t s f o r Automatic S e l f i n g and S e l f C o m p a t i b i l i t y ..................,. 115 Tab 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 T a b l e V I I I . Average S o i l Depth Beneath P l a n t s ............. 125 T a b l e IX. 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 of I n s e c t V i s i t o r s .....•....•.•........••128 V LIST OF FIGURES F i g u r e 1 : Map of C o l l e c t i o n S i t e s . .......................7 F i g u r e 2 : Cut-away F l o w e r and F r u i t Drawings o f J?» 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 . ••/.... .............. 20 F i g u r e 3 : Cut-away Flower Drawings o f S , . o c c i d e n t a l i s v a r . o c c i den t a l i s, S. r u f i d u l a . s.--, o c c i d e n t a l i s : Var. d e n t a t a , S. i n t e q r i f o l i a v a r . c l a y t o n i i f o l i a . S. i n t e a r i f o l i a v a r i i n t e q r i f o l i a , and S. i n t e q r i f o l i a v a r . l e p t o p e t a l a . ................................... 23 F i g u r e 4 : Sketch o f A r t i f i c i a l l y Produced S . , t a f i d u l a x S. o c c i d e n t a l i s H y b r i d and .Representative P a r e n t a l P l a n t s . ... ................. ,•>• 27 F i g u r e 5 : Sk e t c h o f A r t i f i c i a l l y Produced S. r u f i d u l a X S. Occident a l l s - v a r . dgntata H y b r i d and 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 . ......... • 29 F i g u r e 6 : Sk e t c h o f A r t i f i c i a l l y Produced S. r u f i d u l a x J . i n t e q r i f o l i a H y b r i d and 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 . ,....-»....... * .................................... 31 F i g u r e 7 : S k e t c h o f A r t i f i c i a l l y Produced S. r u f i d u l a x S. o c c i d e n t a l i s v ar. l a t i p e t i o l a t a . H y b r i d and 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 . .................... y...33 F i g u r e 8 : S k e t c h o f A r t i f i c i a l l y Produced S. 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 v a r . d e n t a t a H y b r i d aud 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 . .,......,....,.,........ 35 F i g u r e 9 : Leaf O u t l i n e s o f S. r u f i d u l a , S. o c c i d e n t a l v a r . 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 v a r . l a t i p e t i o j a t a . and S. i n t e g r i f p j j - i a v a r . i n t e q r i f o l i a v i v. . .... 38 F i g u r e 10 : L e a f O u t l i n e s of D i p l o i d and T e t r a p l o i d S. o c c | d e n t a l i s v a r . d e n t a t a . ........................40 F i g u r e 11 : Camera L u c i d a Drawings Showing V a r i a t i o n i n Chromosome Number f o r One P o p u l a t i o n o f S. r u f i d u l a i n t he Columbia R i v e r Gorge. ..........................47 F i g u r e 12 : Camera L u c i d a Drawings Showing V a r i a t i o n i n Chromosome Number f o r One P o p u l a t i o n o f S. r u f i d u l a i n the Columbia R i v e r Gorge 49 F i g u r e 13 : Camera L u c i d a Drawings Showing M e i o s i s i n C r y p t i c H y b r i d s and Normal C e l l s from P o p u l a t i o n s o f S. r u f i d u l a i n t h e Columbia R i v e r Gorge. .............. 51 F i g u r e 14 : Camera L u c i d a Drawings o f M e i o t i c C e l l s c f 5. r u f i d u l a and 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 . .53 F i g u r e 15 : Camera L u c i d a Drawings Showing V a r i a t i o n i n Chromosome Number f o r S. o c c i d e n t a l i s v a r . d e n t a t a . .55 F i g u r e 16 : Camera L u c i d a Drawings Showing H e i o t i c Chromosome Numbers f o r 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 and S. i n t e q r i f o l i a Re l a t i v e s . «,...... y .59 F i g u r e 17 : Camera L u c i d a Drawings o f M e i o t i c C e l l s c f S. oregana and S. i n t e q r i f o l i a v a r . c l a v t c n i i f c l i a P o p u l a t i o n s . ...........................• •.... .61 F i g u r e 18 : Camera L u c i d a Drawings Showing V a r i a t i o n i n Chromosome Number o f M e i o t i c F i g u r e s i n A r t i f i c i a l H y b r i d s . .. .... v . . . . . . . . . v . . . . ..........................66 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 A x i s 1 vs. 2 D e p i c t i n g A r t i f i c i a l 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 . .68 F i g u r e 20 : P r i n c i p a l Components A n a l y s i s A x i s 1 vs. 3 v i i D e p i c t i n g A r t i f i c i a l 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 . .70 F i g u r e 21 : P r i n c i p a l Components A n a l y s i s A x i s 2 v s . 3 D e p i c t i n g A r t i f i c i a l 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 . , . 7 2 F i g u r e 22 ; C r o s s i n g Success C h a r t . ....................... 74 F i g u r e 23 : S c a t t e r P l o t D e p i c t i n g C h a r a c t e r I n t r o g r e s s i o n Between S. i n t e g r i f o l j a v a r . c l a y t o n i i f o 1 i a and 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 . d e n t a t a . ........ ....80 F i g u r e 24 : S c a t t e r P l o t D e p i c t i n g C h a r a c t e r I n t r o g r e s s i o n Between S. i n t e q r i f c l i a v a r . c l a y t c n i i f o l i a and Sym p a t r i c S. r u f i d u l a P l a n t s . .... ...... ................82 F i g u r e 25 : S c a t t e r P l o t D e p i c t i n g C h a r a c t e r I n t r o g r e s s i o n Between S. r u f i d u l a from t h e Columbia R i v e r Gorge and S. O c c i d e n t a l l s v a r . o c o i d e n t a l i s . ..................84 F i g u r e 26 : Dendrogram of T o t a l Sample Osed i n C l u s t e r i n g Program, UBC-CGBOUP. ..86 F i g u r e 27 : P r i n c i p a l Components A n a l y s i s A x i s 1 vs. 2 f r c o Samples of N a t u r a l P o p u l a t i o n s . ........................93 F i g u r e 28 : P r i n c i p a l Components A n a l y s i s A x i s 1 vs. 3 from Samples o f N a t u r a l P o p u l a t i o n s . ..95 F i g u r e 29 : P r i n c i p a l Components A n a l y s i s A x i s 2 vs. 3 from Samples of N a t u r a l P o p u l a t i o n s . ........................97 F i g u r e 30 : Hyperspace P l o t of Grouping R e l a t i o n s h i p s c f C a n o n i c a l V a r i a b l e s i n SDF A n a l y s i s Program and P o i n t Cloud B e p r e s e n t a t i o n Of S. i n t e q r i f c l i a v a r . leptop„etala . ......................................... 101 F i g u r e 31 : I n d i v i d u a l P o i n t s i n Hyperspace Cloud R e l a t i o n s h i p s about the Group Means f o r Sk i n t e g r i f o l j a v a r . c l a y t o n i i f c l i a and S. i n t e q r i f o l i a v a r . v i i i l e p t e p e t a l a . • •.. .•.......•................. .......... 104 F i g u r e 32 : I n d i v i d u a l P o i n t s i n Hyperspace C l o u d R e l a t i o n s h i p s about the Group Means f o r H e x a p l o i d H y b r i d P o p u l a t i o n and 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 . ..106 F i g u r e 33 : I n d i v i d u a l P o i n t s i n Hyperspace C l o u d R e l a t i o n s h i p s about the Group Means f o r S. c c c i d e n t a l i s v a r . d e n t a t a and 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 . .................. ...... . . . . . . . . . ... .......108 F i g u r e 34 : I n d i v i d u a l P o i n t s i n Hyperspace C l o u d R e l a t i o n s h i p s about the Group Means f o r Columbia R i v e r Gorge S. i n t e q r i f c l i a and Vancouver I s l a n d S. i n t e q r i f o l i a . ......................... ... .......... 110 F i g u r e 35 : I n d i v i d u a l P o i n t s i n Hyperspace C l o u d R e l a t i o n s h i p s about the Group Means f o r S. r u f i d u l a from Vancouver I s l a n d and t h e Olympic Mountains and S. r u f i d u l a from t h e Columbia R i v e r Gorge w i t h A r t i f i c i a l H y b r i d s . ..........................,.«>......112 F i g u r e 36 : S p e c u l a t i v e P o l y p l o i d R e l a t i o n s h i p s i n •S. r u f i d u l a and R e l a t e d Taxa. ......................... 151 F i g u r e 37 : D i s t r i b u t i o n Map of S. g o r m a n i i . S» o c c i d e n t a l i s m and 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 H y b r i d P o p u l a t i o n . ... •........... .... ..... ............. 162 F i q u r e 38 : D i s t r i b u t i o n Map o f S. r u f i d u l a and S. l a t i p e t i o l a t a . ..... .y«. ..... . • •>•;#•• .. «... ........... .164 ix ACKNOWLEDGEMENTS "The j o u r n e y need n ot be a l o n e a t a l l moments. He can and do spa r k one a n o t h e r , and c a r r y each o t h e r on." P a u l u s Berensohn To many who have h e l p e d a l o n g t h e way s i m p l e thanks a r e i n s u f f i c i e n t , b u t , a l a s , they a r e a l l I have t o demonstrate my a p p r e c i a t i o n . My w i f e , Mary Dee, knows a l l too 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 t o t h i s p a p e r . 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 , d ata r e c o r d e r , computer o p e r a t o r , i l l u s t r a t o r , f r i e n d and mother 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 o f t h i s l a b o r of l o v e . My d a u g h t e r , R a c h e l , p r o v i d e d i n t a n g i b l e j o y s and much needed d i v e r s i o n s when she a r r i v e d d u r i n g the f i n a l phases of t h i s work. Kay Beamish has been a most h e l p f u l 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 t h i s s t u d y . Funds f o r t r a v e l , equipment and expenses were p r o v i d e d by an N.R.C, Grant (No. ,67-6901) t o Dr. Beamish, a O.B.C. , Graduate F e l l o w s h i p , C a r e e r s »75 S t i p e n d , and a U.B.C. Student T r a v e l S t i p e n d t o the auth o r . The members of my a d v i s o r y committee d e s e r v e thanks f o r t h e i r t h o u g h t f u l s u g g e s t i o n s a l o n g the way and c a r e f u l r e a d i n g of t h e f i n a l m a n u s c r i p t s , The f l o w e r c l o s e - u p d r a w i n g s were done by my good f r i e n d , Joan M i l l e r . Gary B r a d f i e l d g e n e r o u s l y p r o v i d e d h i s s t a t i s t i c a l e x p e r t i s e , computer program, and t i m e i n r e a d i n g c e r t a i n s e c t i o n s o f t h i s paper. Johann van Seenen i d e n t i f i e d t he i n s e c t specimens and Jo h n P i n d e r - M o s s a s s i s t e d i n numerous herbarium 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 s t u d y . So many 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 and encouragement a l c n g t h e way i t i s d i f f i c u l t t o thank each one he r e . Of t h e non-human c o h o r t s , t h e UBC computer 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 mostly t h e r e a r e the p l a n t s . 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. x i 1 S o r t o f a Song L e t the snake wai t under h i s weed and the w r i t i n g be o f words, slow and g u i c k , s h a r p t o s t r i k e , g u i e t t o w a i t , s l e e p l e s s . Through metaphor t o r e c o n c i l e t h e people and the s t o n e s . Compose. (No i d e a s b u t i n t h i n g s ) I n v e n t ! S a x i f r a g e i s my f l o w e r t h a t s p l i t s the r o c k s . - W i l l i a m C a r l o s W i l l i a m s (1883-1963) 1 INTBODOCTION The genus S a x i f r a g a L. i s composed of herbaceous, 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 , a l p i n e and r o c k y p l a c e s i n temperate r e g i o n s . The genus i s c i r c u m b o r e a l and r e s t r i c t e d m o stly t o the N o r t h e r n Hemisphere. The name, S a x i f r a g a , comes from t h e L a t i n words, s a j s i meaning r o c k and f r a g e r e meaning t o break or fragment, and i s p r o b a b l y a r e f e r e n c e t o t h e h a b i t a t s which many members occupy. A l e s s l i k e l y 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 t o the f o l k use o f some members o f t h e genus f o r t h e t r e a t m e n t o f k i d n e y s t o n e s (Spongberg 1972). E t h n o b o t a n i c a l uses of Saxi,f raga s p e c i e s a r e as a l e a f exudate 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 1974), as a r o o t e x t r a c t used f o r the t r e a t m e n t of a wide range of 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 p a i n s t o r e l i e f o f d y s e n t e r y (Watt 1972), as a l p i n e r o c k garden p l a n t s , and as t h e hanging b a s k e t h o u s e p l a n t , S. sarmentosa S c h r c e d e r , known a s s t r a w b e r r y b e g o n i a , s t r a w b e r r y geranium, or mot h e r - o f - t h o u s a n d s . A l t h o u g h s e v e r a l e a r l y 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 t h e knowledge o f t h e genus ( L i n n e a u s 1753, Don 1822, Hooker 1833), the comprehensive monograph o f E n g l e r and I r m s c h e r (1916) brought t o g e t h e r t h e e a r l i e r t axonomic 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 of p l a n t s . In North America the s e c t i o n B o r a p h i l a of E n g l e r and Irmscher (1916) has two te m p e r a t e c e n t e r s of s p e c i e s d i v e r s i t y , one i n t h e E a s t e r n A p p a l a c h i a n Region and t h e o t h e r i n t h e Rocky Mountain Region of t h e P a c i f i c Northwest (Spongberg 1972). Both 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 g l a c i a t i c n s 2 and the e v o l u t i o n a r y h i s t o r y of the p l a n t s i n t h o s e a r e a s i s , i n many r e s p e c t s , c l o s e l y l i n k e d w i t h t h e h i s t o r y of g l a c i a l advance and r e t r e a t . T h i s i s e s p e c i a l l y e v i d e n t i n the r a p i d l y e v o l v i n g s p e c i e s complexes o f t h o s e a r e a s . I n such complexes s i n g l e m o r p h o l o g i c a l c h a r a c t e r i s t i c s which s e p a r a t e one e n t i t y 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 f i n d . , The name, r u f i d u l a , was f i r s t used t o d e s c r i b e a s p e c i e s o f t h e genus M i c r a n t h e s by S m a l l and Hydberg (1905). I t was s u b s e q u e n t l y t r e a t e d as a p a r t o f t h e genus S a x i f r a g a by James Macoun (1906) whose f a t h e r , J o h n , c o l l e c t e d the t y p e m a t e r i a l from Mt. F i n l a y s o n on Vancouver I s l a n d , B r i t i s h Columbia i n 1887. P e r s i s t e n t c o n f u s i o n has surrounded t h e r e l a t i o n s h i p s among S a x i f r a g a r u f i d u l a (Small) James Macoun and i t s c l o s e P a c i f i c Northwest r e l a t i v e s i n t h e s e c t i o n B o r a p h i l a s u b s e c t i o n K j v a l i - v i r g i n i e n s i s ( E n g l e r and I r m s c h e r , 1916) up t o the p r e s e n t t i m e . E a r l y monographic t r e a t m e n t s d e a l t w i t h t h e v a r i a b i l i t y e x t a n t w i t h i n t h i s c l o s e l y r e l a t e d group by r e c o g n i z i n g a l a r g e number of s p e c i f i c e n t i t i e s ( S m a l l and Bydberg 1905, Johnson 1S23). More r e c e n t l y , t h e s e have been reduced t o two o r t h r e e b r o a d l y d e f i n e d s p e c i e s complexes w i t h 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 1944, H i t c h c o c k et a l . 1961, H i t c h c o c k and C r o n g u i s t 1973, Krause and Beamish 1972,1973). The c h a n n e l i n g of the Columbia R i v e r through i t s gorge has p e r m i t t e d a zone o f c o n t a c t between d r i e r i n t e r i o r f l c r i s t i c e lements and more mesic c o a s t a l s p e c i e s . The i n t i m a t e c o n t a c t o f o t h e r w i s e g e n e r a l l y e c o l o g i c a l l y and g e o g r a p h i c a l l y i s o l a t e d p l a n t s i n s i t u a t i o n s where e n v i r o n m e n t a l l y i n t e r m e d i a t e o r 3 perhaps unique m i c r o - h a b i t a t s 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 c o m p l i c a t e d p a t t e r n s of v a r i a b i l i t y and e v o l u t i o n w i t h i n s e v e r a l groups of p l a n t s , i n c l u d i n g t h e Clay_tonia p e r f p l i a t a p o l y p l o i d complex ( M i l l e r 1976), and t h e S i s v r i n c h i u m § 3 1 gg|?*i?-53ffl- j-dahoense- l i t t o r a l e d u o d e c a p l o i d s p e c i e s group (Henderson 1976) . S e v e r a l a u t h o r s have commented on t h e e x t e n t of m o r p h o l o g i c a l i n t e r m e d i a r y between S. r u f i d u l a and S. o c c i d e n t a l i s S.,Wats. B a c i g a l u p i , i n Shram* s f l o r a (1944), 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 two s u b s p e c i f i c t a x a ; s u b s p e c i e s o c c i d e n t a l i s and r u f i d u l a . H i t c h c o c k e t a l . (1961), and H i t c h c o c k and C r o n g u i s t (1S73) t r e a t e d S. r u f i d u l a as a v a r i e t y o f S. o c c i d e n t a l i s and c o n s i d e r e d v a r i e t y r u f i d u l a t r a n s i t i o n a l t o v a r i e t y d e n t a t a ( E n g l , and Irmsch.) C.L. H i t c h c o c k and v a r i e t y a l l e n i i ( Small)C.L. H i t c h c o c k i n t h e Columbia B i v e r Gorge and a d j a c e n t Oregon. , H i t c h c o c k a l s o i n c l u d e d v a r i e t y i d a h o e n s i s ( P i p e r ) C . L . H i t c h c o c k i n S. o c c i d e n t a l i s and d e s c r i b e d a new v a r i e t y , l a t i p e t i o l a t a C . L . H i t c h c o c k . S a x i f r a g a r u f i d n l a was c o n f i r m e d by Krause and Beamish (1973) as a s p e c i e s and S, o c c i d e n t a l i s was c o n s i d e r e d as an e x t r e m e l y v a r i a b l e e n t i t y w i t h o u t s u b s p e c i f i c t a x a . They noted 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. r u f i d u l a and S. o c c i d e n t a l i s i n t h e Columbia B i v e r Gorge was l a r g e l y u n e x p l o r e d . The e n t i t y i d a h o e n s i s had been t r e a t e d by Krause and Beamish (1972) as a s u b s p e c i e s o f S. m a r s h a l l i i but t h e y r e c o g n i z e d t h a t S. m a r s h a l l i i subsp. i d a h o e n s i s has some i n t r o g r e s s a n t c h a r a c t e r i s t i c s of ,S. o c c i d e n t a l i s . E l v a n d e r (1975) c o n f i r m e d the c l a s s i f i c a t i o n by H i t c h c o c k e t j | l . (1961) , 4 and H i t c h c o c k and C r o n g u i s t (1973) of S. o c c i d e n t a l i s v a r i e t i e s i d a h o e n s i s , r u f i d u l a . and l a t i p e t i o l a t a . He c o n s i d e r e d t h e v a r i e t i e s d e n t a t a and a l i e n i i t o l a c k m o r p h o l o g i c a l d i s t i n c t i o n s and v a r y c o n t i n u o u s l y w i t h v a r i e t y o c c i d e n t a l i s and c o n s e g u e n t l y , he t r e a t e d them as synonyms of v a r i e t y o c c i d e n t a l i s . None o f t h e s e p r e v i o u s s y s t e m a t i c i n v e s t i g a t i o n s have addressed the problem o f t h e c o n f u s i o n t h a t e x i s t s among the r e l a t i v e s o f S. r u f i d u l a i n the Columbia B i v e r Gorge. Eeamish (1961,1967) and Krause and Beamish (1972,1973) f o c u s e d t h e i r 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 i n B r i t i s h Columbia a l t h o u g h they r e p o r t e d s e v e r a l chromosome c o u n t s o f the m a t e r i a l from t h e Columbia B i v e r Gorge and o t h e r a r e a s of Oregon and Washington. E l v a n d e r (1975) based h i s s t u d y cn n u m e r i c a l , c y t o l o g i c a l , and c h r o m a t o g r a p h i c e v i d e n c e i n p l a n t s l a r g e l y from Idaho and Rontana. He b r i e f l y d i s c u s s e d S. r u f i d u l a . S. o c c i d e n t a l i s v a r . d£JLt§il# and 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 b u t d i d not i n c l u d e d a t a from p o p u l a t i o n s o f t h o s e e n t i t i e s i n h i s b a s i c s t u d y . The r e l a t i o n s h i p s among S. r u f i d u l a . 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 . S , , o c c i d e n t a l i s v a r . d e n t a t a and 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 i n t h e Columbia B i v e r Gorge and o t h e r a r e a s e x t e n d i n g northward i n t o Southwestern B.C. a r e d e a l t w i t h i n t h e p r e s e n t paper. C o r r e l a t i o n s o f h y b r i d i z a t i o n s t u d i e s and e c o l o g i c a l 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 m o r p h o l o g i c a l a n a l y s e s based on t h r e e d i f f e r e n t n u m e r i c a l approaches a r e p r e s e n t e d 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 c l a s s i f i c a t i o n . S i n c e S. r u f i d u l a a n d i t s c l o s e r e l a t i v e s are 5 o f t e n found growing s y m p a t r i c a l l y w i t h members of S. I n t e g r i f o l i a ( s e c t . I n t e g r i f c l i a e o f E n g l e r and I r m scher 1916), th e v a r i e t i e s i n t e q r i f o l i a . c l a y t o n i i f o l i a , and l e p t o p e t a l a a r e a l s o i n c l u d e d as p o t e n t i a l s o u r c e s o f h y b r i d i z a t i o n and c h a r a c t e r i n t r o g r e s s i o n i n the s t u d i e s p r e s e n t e d here. S e v e r a l t a x a p e r i p h e r a l t o the main t h r u s t of t h i s s t u d y may have some c l o s e r e l a t i o n s h i p s w i t h the t a x a d i s c u s s e d i n t h i s paper. E v i d e n c e t h a t S. oreqana has c l o s e t i e s t o S.•..occidentalis v a r . l a t i p e t i o l a t a i s brought f o r t h aisd as a r e s u l t i t i s i n c l u d e d i n t h e key t o s p e c i e s . A s y s t e m a t i c s t u d y of S. i n t e g r i f o l i a and S. oreqana sensu l a t o i s c u r r e n t l y i n p r o g r e s s a t t h e U n i v e r s i t y o f U a s h i n g t o n ( E l v a n d e r , PhD d i s s e r t a t i o n , i n p r e p a r a t i o n ) . , S a x i f raga • o c c i d e n t a l i s v a r . d e n t a t a may i n t r o g r e s s t o S. m a r s h a l l i i subsp. m a r s h a l l i i . Other t a x a mentioned here i n c l u d e S. m a r s h a l l i i subsp. i d a h o e n s i s , S. r e f l e x a , S. n i v a l i s , S« c a l i f c r n i c a * J» i n t e g r i f o l i a v a r . c o l u m b i a n a . S. rhomboidea. S, f e r r u g i n e a and a group o f E a s t e r n North American s p e c i e s . S a x i f r a j a i s i i f l f i f s i i a v a r . c o l u m b i a n a i s t r e a t e d here as e q u i v a l e n t t o S. j , n t e g r j f c l j a v a r . , l e p t p p e t a l a . The p r e s e n t paper uses the t e r m i n o l o g y of 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 but 5 . r u f i d u l a and S. m a r s h a l l i i subsp. , i d a h p e n s i s a r e t r e a t e d as r e c o g n i z e d by Krause and Beamish (1972, 1973), and ,S. o c c i d e n t a l i s v a r . a l l e n i i a s synonymous w i t h v a r . c c c i . d e n t a j . i s . Proposed changes i n t h e c l a s s i f i c a t i o n and r e v i s e d c o m e c c l a t u r e o f the t a x a s t u d i e d appear i n the Taxonomy and C o n c l u s i o n s s e c t i o n s o f t h i s paper. 6 MATERIALS AMD METHODS Live plants, pressed specimens, flower buds and several s o i l samples were co l l e c t e d from populations from the mouth of the Columbia River east to beyond the Columbia River Gorge. Some c o l l e c t i o n s were also made in areas extending north and south i n the lower Columbia River area and adjacent Coast Range. Populations from the Olympic Mountains, Southeastern Vancouver Island, and Cascades in Washington and the Southern Mainland of B r i t i s h Columbia (Fig. 1, Table I) were also sampled. Many co l l e c t i o n s came from l o c a l i t i e s with two sympatric taxa. Fifteeen to twenty rosettes from a large number of the populations sampled were transplanted to ccldframes f i l l e d with a peat-vermiculite bedding mixture which were located at the University of B r i t i s h Columbia. Mass c o l l e c t i o n s of buds i n meiosis were sampled. Buds from i n d i v i d u a l plants were coll e c t e d and a large number of plants were pressed as voucher material. Specimens are on deposit at the university of B r i t i s h Columbia, Specimens studied for morphological comparison, c l a s s i f i c a t i o n , annotation, and geographic d i s t r i b u t i o n included a t o t a l of 1,507 herbarium sheets representing about 5,000 plants from the following herbaria: Brigham Young University (BEY), University of Minnesota (MIN), U.S. National Herbarium (NA) ,University of Oregon (ORE) , Oregon State University (OSC) , University of B r i t i s h Columbia (UBC), University of C a l i f o r n i a at Berkely (UC), B r i t i s h Columbia P r o v i n c i a l Museum (V) w Washington State University (MS) , and University of Washington (WTU). Several hundred additional herbarium sheets were also 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 p r e s e n t study. One c o l l e c t i o n from B e r t h o u d P a s s , C o l o r a d o , i s not shown. 9 • Table L. 7 C o l l e c t i o n s f o r Study. C o l l e c t i o n Tjaxpn Code Number l o c a t i o n S. r o f i d u l a 616 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. of V i e n t o S t . Pk., Hood B i v e r Co., Ore. 608 T r o u t d a l e , 1.3 mi s. o f Sandy B. B r i d g e , Multnomah Co., Ore. 610 Yeon Pk., c a . 1.5 mi a l o n g t r a i l t o McCord F a l l s , Multnomah Co. Ore. , 612A Mayer Pk., 0.8 mi n. o f Mayer S t . Pk., Wasco Co., Ore. 613A The D a l l e s , 2. 0 mi w. o f Chynoweth C r . on Old U.S. , Hwy. 30, Wasco Co., Ore. 637 M o s i e r , 1.0mi e. c f H o s i e r on O l d Columbia 8. Hwy., Wasco Co., Ore, 673 Lake 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. . 675A Mt. P l e a s a n t , 1.9 mi e. o f t h e Clar k - S k a m a n i a Co. bdry. on S t , Hwy. 14, C l a r k Co. , Wash. 647A Washougal,1.0 mi w. o f C l a r k -Skamania Co. bdry. on S t . Hwy. 14, C l a r k Co., Hash. 76-1 Marmot P a s s , c a . 5 mi s. of Camp Myst e r y , s.e. o f Marmot P a s s , J e f f e r s o n Co,, Wash. 617A Bingen, 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 Co., Wash. 669A Bingen Lk., 7.4 mi e. of w. bdry., of K l i c k a t a t Co. on S t . Hwy. 14, Wash. 672 Cock, 9.8 mi w, o f e. bdry, o f Skamania Co. on S t . Hwy. 14, Bash., 6 18A Skamania Co., 3 .5 mi e. o f C l a r k -Skamania Co. bdry. on S t . Hwy. 1 Wash, 645A C o l l i n s , 0.9 n i e. of C o l l i n s Depot Bd. on S t . Hwy. 14, Skamania Co., Wash. NMO-8 Nanaimo, c a , 5 m i ' s , o f Nanaimo cn White Rapids Rd., V . I . , B.C. 10 T a b l e It JconJMt) Taxon S. o c c i d e n t a l i s var. denata C o l l e c t i o n Code Number NOOS-B UCL-R 623A 624B 627A 629A 606 630 607 S. o c c i d e n t a l i s 629C var. 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 605 var. o c c i d e n t a l i s B M R 626 666 COEN 687 S. i n t e q r i f o l i a 641 var. c l a y t o n i i f o l i a Nanoose H i l l , c a. 15 mi n. o f Nanaimo on Nanoose H i l l , V . I . , B.C. Upper Campbell Lk., 7.4 mi w. o f Campbell Lake B r i d g e , V . I . , B.C Sooke, s. s i d e of Sooke B. a t Sooke P o t h o l e s Pk., V.I.,B.C.. H i l l H i l l , 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 Prov. Pk., V . I . , B.C. Saddle Mt., cn 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. of D e l e n a , Columbia Co. Ore, T i l l a m o o k Co., 2.0 mi w. o f e. bdry. o f T i l l a m o o k Co. cn S t , Hwy. 6, Ore. Kalama B., 3 mi e. o f I n t e r s t a t e 5 on Kalama B. Bd., C o w l i t z Co., Wash. Saddle Mt., on t r a i l t o summit near t o p , C l a t s o p Co. , Ore. C h e h a l i s 8. , (hybrid) , 3.3 mi w. o f L i t t e l , L e wis Co. Wash. , Mt. Baker 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. and n. of Y a l e below Can. Hwy. 1, B.C. Liumchen B i d g e , s. o f S a r d i s , B » C» C o r n w a l l Lookout, s e . o f Hat Creek, B.C. B o o t a n i e V a l l e y , Skwaha Mt. area n. o f L y t t o n , B.C. G i l i a m Co., 1.7 mi e. of w, bdry. G i l i a m Co. on I n t e r s t a t e 80 n., Ore. 11 T a b l e I . (con't) C o l l e c t i o n HiLSSLE Code Number L o c a t i o n 609 T r c u t d a l e , 1.3 mi s. o f Sandy 8. B r i d g e , Multnomah Co., Ore. 611 Bowena, 2,4 mi w. of Bowena ov e r p a s s on I n t e r s t a t e 80-N,, Hasco c o., Ore, 612B Mayer Pk., 0.8mi n. of Mayer S t . Pk., Wasco Co., Ore. 613B The D a l l e s , 2.0 mi w, o f Chynoweth Cr. on Old U.S. Hwy. 30, Wasco Co. Ore. 667 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. 30, Sherman Co., Ore. 692 Stephen's P a s s , 11 mi e. o f Stephen's Pass S k i Lodge on U.S. Hwy. 2, Chelan Co., Hash. 675B Mti 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. bd r y . on St.Hwy. 14, C l a r k Co., l a s h . 668 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 Co., Wash. 669B Bingen Lk. , 7.4 mi e. o f w. bdry. o f K l i c k a t a t Co. :on S t . Hwy. 14, Wash. ., 642 Benchmark, U.S.G.S. V425,1S68, 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 Co., Wash. 617B B i n g e n , 5.6 mi e. of B i n g e n , on S t . Hwy 14, K l i c k i t a t Co., Bash. 619 C l a r k Co. l i n e , b d r y . w i t h Skamania Co. on St.,Hwy, 14, Sash. 5 A i n t e g r i f o l i a 671 G r i z z l y Lk., about 0.5 mi v a r . i n t e g r i f c ^ i a • downstream from G r i z z l y Lk. S i s k i y o u Co., C a l i f o r n i a MIMA Mima Mounds, w. o f L i t t l e Bock on Bd. t o Mima, T h u r s t o n Co., Wash. 6 23B Sooke, s . s i d e of Sooke B. a t P o t h o l e s Pk., V . I . , B.C. 624B M i l l H i l l , M i l l H i l l 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. 627B 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. (con* t) C c l l e c t i c n Tax on Code Number L o c a t i o n NJ30-B Nanaimo, c a , 5 mi s, o f Nanaimo on White B a p i d s Bd. , V. I . , E. C, NOOS-B Nanoose H i l l , ca. 15 mi n. o f Nanaimo on Nanoose H i l l , V . I . , B.C. 625 Y a l e , c a . 1.5 mi e. and n. o f Y a l e below Can. Hwy 1, B.C., 617B B i n g e n , 5.6 mi e. of Bingen on S t . Hwy. 14, K l i c k a t a t Co., Wash. 78-1 H a r r i s o n Lk., 1.5 mi w. of H a r r i s o n , Hot S p r i n g s , B.C. 682 E l k B a l l s , below Campbell Lk. Dam, V . I . , B.C. S. i n t e g r i f o l i a 648 P r i n c e t o n , o f f Hwy. e. o f va r . l e p t o p e t a l a P r i n c e t o n , road t c g o l f c o u r s e , B. C. S*. orjejgajja BEBH Berthoud Pass, Bdry. of Grand Co. C o l o r a d o 13 i n s p e c t e d from m a t e r i a l on l o a n to t h e U n i v e r s i t y o f Washington., Chrcmosomal S t u d i e s Squashes o f a n t h e r t i s s u e nere made by f i r s t f i x i n g bud m a t e r i a l i n C a r n o y f s 3 : 1 , a b s o l u t e e t h a n o l : g l a c i a l a c e t i c a c i d f l u i d . Buds were t h e n s t a i n e d u s i n g Snow*s (1963) b u l k method and warmed i n a 40 C oven f o r about 24 hours. S l i d e p r e p a r a t i o n s were p r e s e r v e d i n Hoyer*s permanent medium ( f i l e x o p c u l c s and Beneke 1952) . P o l l e n s t a i n a b i l i t y o f l i v e , p r e s e r v e d and d r i e d f l o w e r s was measured by c o u n t i n g up t o 200 p o l l e n g r a i n s which had been s t a i n e d and mounted i n l a c t o p h e n o l - a n i l i n e b l u e s t a i n (Sass 1958). L a r g e , d a r k - s t a i n i n g g r a i n s were c o n s i d e r e d f e r t i l e . P r e l i m i n a r y i n v e s t i g a t i o n s of p o l l e n g r a i n s i z e c o n f i r m e d S o k o l o v s k a y a • s (1958) r e p o r t s t h a t p o l l e n s i z e does not appear t o be c o r r e l a t e d w i t h chromosome number and f u r t h e r i n v e s t i g a t i o n was d i s c o n t i n u e d . 14 H Y b r i d l z a t i G j p S t u d i e s P l a n t s used as female p a r e n t s were e m a s c u l a t e d , bagged w i t h w a t e r - r e s i s t a n t parchment bags p r i o r t o f l o w e r i n g and a r t i f i c i a l l y c r o s s - p o l l i n a t e d under a d i s s e c t i n g m i c r o s c o p e . Only c r o s s e s i n v o l v i n g s i m u l t a n e o u s l y blooming p l a n t s were s u c c e s s f u l . S t i g m a t i c r e c e p t i v i t y was e s t i m a t e d from t h e o n s e t of a s l i g h t watery appearance and p a p i l l a t e c o n d i t i o n of the s t i g m a t i c s u r f a c e . I n c r e a s e d s t a i n a b i l i t y u s i n g l a c t o p h e n o l - a n i l i n e blue s t a i n (Sass 1958) a l s o i n d i c a t e d s t i g m a t i c r e c e p t i v i t y . Whole a n t h e r s were used t c t r a n s f e r p o l l e n masses t o t h e s t i g m a s . Samples o f h y b r i d and o u t c r o s s e d seed were p l a n t e d i n a w e l l - d r a i n e d sandy s o i l m i x t u r e i n e a r l y December i n rows beneath about 1-2mm o f sand and a 1cm l a y e r o f p e a - s i z e d g r a v e l i n large-mouthed p o t s s e t i n a s h a l l o w pan of water. P l a n t i n g s were p l a c e d o u t s i d e under ambient temperature and l i g h t c o n d i t i o n s and m i s t e d p e r i o d i c a l l y t o m a i n t a i n a moist environment. By A p r i l s e e d l i n g s o f n e a r l y e v e r y c r o s s had germinated and l a t e r were t r a n s p l a n t e d t o sandy p r o p a g a t i o n beds where t h e y were a l l o w e d t o grow f o r one season. I n t h e s p r i n g o f the second season s e e d l i n g s were t r a n s p l a n t e d i n t o p o t s . As they f l o w e r e d , samples were examined f o r t h e i r morphology and 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. 15 N u m e r i c a l S t u d i e s F o r t y - t h r e e q u a n t i t a t i v e c h a r a c t e r s o r c h a r a c t e r c o m b i n a t i o n s (Table I I ) were measured on 263 l i v e , f l o w e r i n g p l a n t s (from n a t u r a l p o p u l a t i o n s and a r t i f i c a l l y produced h y b r i d s ) f o r n u m e r i c a l taxonomic t r e a t m e n t s . P e r i o d i c i n f e s t a t i o n s of r u s t , a p h i d s , mildews, s l u g s and r o o t w e e v i l s not o n l y i n c r e a s e d p l a n t 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 the de v e l o p m e n t a l morphology of some growing p l a n t s . Such i n d i v i d u a l s were a v o i d e d wherever p o s s i b l e i n the s t u d y but a c e r t a i n amount of s p u r i o u s v a r i a t i o n may be t h e r e s u l t o f re s p o n s e s t o minor i n f e s t a t i o n or subsequent p e s t i c i d e t r e a t m e n t . P o p u l a t i o n samples c o n s i s t e d of 8-10 p l a n t s chosen a t random from w e l l - e s t a b l i s h e d t r a n s p l a n t a t i o n s grown i n c o l d frames. L e a f measurements were t a k e n o f a mature l e a f o f each p l a n t a f t e r i t was c a r e f u l l y removed from t h e o u t e r r o s e t t e w h o r l and p r e s s e d . F l o w e r measurements were s t a n d a r d i z e d by c h o o s i n g f l o w e r s as 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 m i d p o i n t o f a n t h e r d e h i s c e n c e and by t r y i n g t o o b t a i n f l o w e r s 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 c o n v e r t e d t o m e t r i c v a l u e s , counted v a r i a b l e s were t r a n s f o r m e d u s i n g the square r o o t t r a n s f o r m a t i o n and the r e s u l t a n t d a t a m a t r i x was s u b j e c t e d t o t h r e e d i f f e r e n t a v a i l a b l e computer programs. 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 program which was used i s c a l l e d UBC-CGBOUP, a v a i l a b l e a t the U n i v e r s i t y o f B r i t i s h Columbia Computing C e n t e r . The a l o g a r i t h m used i s t h a t of Hard (1963) and the c o d i n g i s m o d i f i e d from Veldman (1967). ft P r i n c i p a l Components a n a l y s i s (PCA) program c a l l e d PBINCOMPS 16 T a b l e I I X C h a r a c t e r s used i n N u m e r i c a l S t u d i e s No. C h a r a c t e r 1. p e t a l I n . 3. s e p a l I n , 5. g l a n d wd, 7, s t y l e h t . 9. p e r i a n t h wd. 11. f i l a m e n t wd. a t bottom 13, h a i r no. a t l e a f t i p 15, h a i r no, on p e d i c e l 17, a n g l e o f l o w e s t b r a c t t o stem 19, h a i r no. a t l o w e s t branch 21. 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 ln./wd. r a t i o 25. scape diam. a t l o w e s t branch 27. p l a n t h t . 29. i n f l o r e s c e n c e ln./wd. r a t i o 31. i n f l o r e s c e n c e I n . from w i d e s t p a r t t o top 33. l e a f ln./wd. r a t i o 35. h a i r I n . p e t i o l e 37. a n g l e l e a f s h o u l d e r 39. p e t i o l e ln./wd. r a t i o 41. s t y l e I n . • g l a n d h t . / gynoecium I n . 43, l e a f I n . / p e t i o l e I n . r a t i o No. C h a r a c t e r 2. p e t a l ln./wd. r a t i o 4. s e p a l ln./wd. r a t i o 6. g l a n d ln./wd. 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. gynoecium h t . 12. p e d i c e l I n . 1 4 . , h a i r I n . l e a f t i p 16.,angle of l o w e s t 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 scape 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 angle 28. f i l a m e n t I n . 30. p l a n t I n . / I n . t o l o w e s t 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 of l e a f t i p 38. p e t i o l e wd. 40. In. of t o o t h a t l e a f t i p 42. g l a n d wd. sguared * f i l a m e n t I n . s g u a r e d / ( s t y l e I n . + g l a n d In.) sguared. a b b r e v i a t i o n s : ht.=h e i g h t , l n . = l e n g t h , no.-number, wd.=width 17 ( a v a i l a b l e from Brad f i e l d . U n i v e r s i t y o f B r i t i s h C o l u m b i a , Department of Botany) was used t o o b t a i n a s t a n d a r d i z e d c o r r e l a t i o n m a t r i x which was t h e n o p e r a t e d upon t o form an e i g e n v e c t o r m a t r i x . F i r s t v s . second, f i r s t v s . t h i r d , and second v s . . t h i r d e i g e n v e c t o r p l o t s were a l s o produced. Ten groups i n c l u d i n g h y b r i d and i n t e r m e d i a t e groups were a s s i g n e d and a n a l y s e d u s i n g a s t e p w i s e d i s c r i m i n a t e f u n c t i o n a n a l y s i s program (BMD07M, Dixon 1970) t o determine c a t e g o r i z i n g f u n c t i o n s , i m p o r t a n t d i s c r i m i n a t i n g v a r i a b l e s and group hyperspace r e l a t i o n s h i p s . B r e e d i n g Systems T e s t s f o r autogamous p o l l i n a t i o n were c a r r i e d out by; 1) bagging f l o w e r buds f o r t h e f l o w e r i n g season and 2) hand p o l l i n a t i n g e m a s c u l a t e d , bagged f l o w e r s w i t h p o l l e n from o t h e r f l o w e r s on the same p l a n t . T e s t s f o r a s e x u a l seed p r o d u c t i o n which a l s o s e r v e d as p r o c e d u r a l c o n t r o l s f o r hand p o l l i n a t i o n e x p e r i m e n t s were conducted by a l l o w i n g e m a s c u l a t e d , bagged buds t o mature w i t h i n the p o l l i n a t i o n bags. U n d e h i s c e d seed and u n f e r t i l i z e d o v u l e s were r e c o r d e d i n a subsample o f each i n f l c r e s e n c e . Seed number was e s t i m a t e d where hig h numbers of seeds were produced. The number o f seeds i n immature c a r p e l s was c o u n t e d from s e v e r a l p l a n t s o f each t a x o n . Repeated c o u n t s on a s i n g l e i n f l o r e s c e n c e were grouped a c c o r d i n g t o the d e t e r m i n a t e o r d e r and p o s i t i o n of the f l o w e r s on the i n f l o r e s c e n c e b r a n c h e s . 18 Habitat find P o l l i n a t i o n Studies Observations of insect behavior on Sax.jfraqa flowers were taken over a period of 21 hours on 5 days at 3 locations. The locations included two on Vancouver Island 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. occidentalis population at Yale, B r i t i s h Columbia. Observations of behaviour included the time spent foraging (for nectar and/or pollen) or resting, the numbers of and distances between flowers and inflorescences v i s i t e d , and i f possible, the v i s i t o r s ' preferences for flowers in various stages of anthesis. Insects observed v i s i t i n g Sasifraga flowers were co l l e c t e d and i d e n t i f i e d tc 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 . Pollen was washed or scraped from the c o l l e c t e d insects to determine the amount of gaxifraga pollen present. S o i l depth to rock substrate (or i n rare cases to a oaximum depth of 30 cm) was measured from beneath 15 to 30 plants chosen at random and representing populations on Vancouver Island, the Lower Mainland of B r i t i s h Columbia, and a transect of stations along the Columbia Biver Gorge. About 20 cc samples of s o i l were taken from beneath 15 plants chosen at random from 7 populations |6 cn Vancouver Island, 1 at Yale, B r i t i s h Columbia), Three locations received repeated sampling over the course of one flowering season. The samples from each population were homogenized, weighed and dried in a 60 C vacuum oven for 24 hours before dry weight measurements were taken. 19 RBSCLTS ftMP DISCUSSIGN H o r p h o l o g i c a l f e a t u r e s which d i s t i n g u i s h between the t a x a i n c l u d e both f l o r a l and v e g e t a t i v e c h a r a c t e r i s t i c s i n most taxonomic t r e a t m e n t s , /The p o s i t i o n o f t h e o v a r y has been used as an i m p o r t a n t key 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 f e a t u r e which has c r e a t e d problems f o r t a x c n o m i s t s i s t h e d i s p a r i t y between t h e degree o f ovary i n f e r i o r i t y a t e a r l y a n t h e s i s (judged by m i d p o i n t of a n t h e r d e h i s c e n c e ) and l a t e r a t m a t u r i t y o f t h e f r u i t . F o r i n s t a n c e , i n 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 development proceeds from a g r e a t e r than h a l f i n f e r i o r o v a r y as i n S. i n t e q r i f o l i a and S, oregana t o an a l m o s t c o m p l e t e l y s u p e r i o r o v a r y i n f r u i t ( F i g . 2 f t , B ) . E v i d e n t l y t h e d i f f e r e n c e between f l o w e r i n g and f r u i t i n g c o n d i t i o n s i s d i f f i c u l t t o d e t e c t cn p r e s s e d specimens and has 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 taxonomic t r e a t m e n t s i f an e a r l y f r u i t i n g c o n d i t i o n where p e t a l s a r e p e r s i s t e n t i s i n t e r p r e t e d as a n t h e s i s . P r e s s i n g and d r y i n g p r o c e s s e s may d i s t o r t p r e c i s e o v a r y r e l a t i o n s h i p s . Even i n l i v e 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 u s i n g m i d p o i n t o f a n t h e r d e h i s c e n c e as a p r e c i s e event i n a n t h e s i s , v a r i a b i l i t y i n s t y l a r e l o n g a t i o n , r e c e p t a c l e w i d t h and g l a n d p o s i t i o n i n g c r e a t e d i f f i c u l t i e s i n g u a n t i t i v e measurement. N o n e t h e l e s s , as a c h a r a c t e r i s t i c which s e p a r a t e s the major s p e c i e s complexes, o v a r y p o s i t i o n i s u s e f u l p r o v i d e d t h a t a n t h e s i s i s c l e a r l y 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 a n t h e r 20 F i g u r e 2: Drawing o f S. o c c i d e n t a l i s v a r , l a t i p e t i c l a t a f l o w e r (A) and f r u i t (£). Flower i s a t m i d p o i n t o f a n t h e r d e h i s c e n c e . 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 -shaped segment of ovary a r e removed t o show th e p o s i t i o n of t h e ov a r y . Note the 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 gland 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 herbarium m a t e r i a l i t i s o f t e n p o s s i b l e t o i n t e r p r e t f l o w e r s which a re pa s t a n t h e r d e h i s c e n c e as b e i n g t h o s e from which p o l l e n has been removed by p o l l i n a t o r s . The b r e a d t h and shape of t h e n e c t a r g l a n d a re u s e f u l c h a r a c t e r i s t i c s i n s e p a r a t i n g some s p e c i e s and s p e c i e s complexes. ., • I n S. r u f i d u l a and many p o p u l a t i o n s 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 i t i s a s m a l l r i n g l i k e band e n c i r c l i n g the l o w e r p o r t i o n o f t h e ovary ( F i g . 3,A,B). The gl a n d and i t s s e c r e t i o n s a r e hidden a t t h e base o f t h e appressed p e t a l s and f i l a m e n t s i n many of these p l a n t s . I n S. o c c i d e n t a l i s v a r . d e n t a t a ( F i g . 3,C), S. o c c i d e n t a l i s v a r . l a t i pe t i e 1 a t a ( F i g . 2A), and members o f t h e S. i n t e g r i f o l i a - oregana- complex ( F i g . 3,D-F) the g l a n d at a n t h e s i s i s an o b c o n i c d i s c which c o v e r s a c o n s i d e r a b l e p o r t i o n o f t h e t o p of t h e o v a r y and exudes n e c t a r i n d i f f u s e , g l i s t e n i n g d r o p l e t s . F i l a m e n t shape v a r i e s w i d e l y among r e l a t i v e s o f S. o c c i d e n t a l i s v a r . , 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 and 5. o c c i d e n t a l i s v a r . , d e n t a t a f i l a m e n t s a r e l i n e a r or s u b u l a t e . A few p l a n t s from h i g h mountain p o p u l a t i c r s of S» r u f i d u l a on Vancouver I s l a n d and the Olympic Mountains have s l i g h t l y c l a v a t e f i l a m e n t s . , I n 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 t h e f i l a m e n t s a r e u s u a l l y a t l e a s t scuewhat c l a v a t e b u t on some p r e s s e d specimens s h r i n k a g e i n d r y i n g produces a p p a r e n t l y l i n e a r f i l a m e n t s . A l l members c f t h e S. i n t e q r i f o l i a -oregana complex have l i n e a r o r s u b u l a t e f i l a m e n t s ( F i g . 3D-F) . I t i s i n t e r e s t i n g t h a t i n o t h e r r e l a t i v e s of S. O c c i d e n t a l j s such as S. m a r s h a l l i i , , which have y e l l o w o r green p e t a l s p o t s the f i l a m e n t s a r e b r o a d l y c l a v a t e and may have ure 3: Drawing 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 . , d e n t a t a ( C ) , §* i n t e q r i f o l i a v a r . c l a y t o n i i f o l i a (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 v a r . l g g t o e e t a l a ( F ) , f l o w e r s a t m i d p o i n t o f a n t h e r d e h i s c e n c e . N e a r e s t 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 -shaped seqment o f the o v a r y w a l l are removed t o show the p o s i t i o n o f t h e ov 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 , o v a r y p o s i t i o n , f i l a m e n t s t r u c t u r e 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 the bases o f t h e f i l a m e n t s . 25 a s p e c i a l f u n c t i o n i n p o l l i n a t o r a t t r a c t i o n . C o r r e l a t i o n s o f t h e s e f e a t u r e s as w e l l as g l a n d c h a r a c t e r s w i t h d i v e r s e p o l l i n a t i o n s t r a t e g i e s c c u l d perhaps p r o v i d e an i n t e r e s t i n g e v o l u t i o n a r y s t o r y , , P e t a l s i z e , c o l o r , and shape vary c o n s i d e r a b l y among the t a x a ( F i g . 2,3). S a x i f r a g a 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 d i f f e r s from i t s r e l a t i v e s i n t h i s study by h a v i n g u s u a l l y e l l i p t i c p e t a l s which o f t e n a r e somewhat narrowed i n t o a c l a w l i k e base. C e r t a i n dwarfed a l p i n e forms of 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 as w e l l as S. r u f i d u l a may be n e a r l y a p e t a l o u s w i t h d a r k l y a n t h o c y a n i c i n f l o r e s c e n c e s and f l o w e r p a r t s . S a x i f r a g_a r u f i d u l a . 5. o c c i d e n t a l i s v a r . d e n t a t a and S. o c c i d e n t a l i s v a r . l a t i p e t j.olat a u s u a l l y have o v a t e p e t a l s w i t h broad bases. V a r i e t i e s o f S. intggx^fojLia range from a p e t a l o u s or w i t h s m a l l g r e e n i s h - p e t a l e d forms t o l a r g e , w h i t e - p e t a l e d f o r m s , S a x i f r a g a r u f i d u l a has a f l a t - t o p p e d or b r o a d l y convex, d i f f u s e - f l o w e r e d i n f l o r e s c e n c e ( F i g . 4, ft, 5 #B) . Some Columbia B i v e r Gorge p l a n t s have b r o a d l y o b c o n i c i n f l o r e s c e n c e s . I n c o n t r a s t , 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 ( F i g . 4,B), 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 ( F i g . 7 . f t ) . S. o c c i d e n t a l i s v a r . d e n t a t a ( F i g . 5,ft) and most v a r i e t i e s of S. i n t e g r i f o l i a ( F i g . 6,B) u s u a l l y have c o n i c or i n t e r r u p t e d - c o n i c i n f l o r e s c e n c e s which range i n f l o w e r d e n s i t y from open t o c o n g e s t e d , Montane forms of 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 as w e l l as S. i n t e g r i f o l i a v a r . a p e t a l a are u s u a l l y f e w - f l o w e r e d , dense, c a p i t a t e p a n i c l e s . Flower number ranges per i n f l o r e s c e n c e from few (4-42) i n S. ...-rufidula, e s p e c i a l l y those 26 on Vancouver I s l a n d and the Olympic Mountains, t o s e v e r a l (13-81) i n 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 , S. o c c i d e n t a l i s v a r . d e n t a t a and •• ~s. i n t e q r i f o l i a v a r . i n t e q r i f c l i a . S a x i f r a g a 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 and S. i n t e q r i f o l i a v a r . c l a v t o n i i f o l i a have a l a r g e number o f f l o w e r s , u s u a l l y e x c e e d i n g 75. S a x i f r a g a o c c i d e n t a l i s v a r . d e n t a t a and * j§. i n t e q r i f o l i a v a r . c l a v t o n i i f o l i a have f i n e , b r i t t l e , v e r t i c a l l y p e n e t r a t i n g networks o f r h i z o i r e s . Such networks may a l s o be p r e s e n t i n S. j a r s h a l I i i subs p. mar s h a l l i i and S. i n t e q r i f o l i a v a r . l e p t o p e t a l a . The o t h e r t a x a have s h o r t , s t o u t , h o r i z o n t a l r h i z omes which do not form deep n e t w o r k s , a l t h o u q h r o s e t t e r e placement from s h o r t branches and b r a n c h l i k e b a s a l b u l b l e t s appears t o be a common means of v e q e t a t i v e qrowth i n t h i s qroup c f p l a n t s . Many f e a t u r e s o f the l e a v e s i n most t a x a v a r y w i t h i n a r o s e t t e . The t e r m i n a l l e a v e s are u s u a l l y more pubescent w i t h fewer t e e t h and a r e much s m a l l e r i n s i z e t h a n t h e l e a v e s from the o u t e r whorl o f t h e r o s e t t e . The l e a v e s may a l s o demonstrate a p l a s t i c r esponse t o shaded c o n d i t i o n s i n which they become e l c n q a t e and have a q r e a t e r s u r f a c e a r e a . Seme l e a f c h a r a c t e r i s t i c s are o f taxonomic s i q n i f i e a n c e . S. o c c i d e n t a l i s v a r . 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 , has broad, s h o r t , r a t h e r i n d i s t i n c t p e t i o l e s ( F i q . 9D). I n t h a t f e a t u r e i t resembles S. oreqana and d i f f e r s from t h e o t h e r t a x a which have d i s t i n c t l y narrowed, e v i d e n t p e t i o l e s ( F i q . 9,A-C,E). The t r a n s i t i o n from b l a d e t o p e t i o l e i s most abrupt i n S, i n t e q r i f o l i a v a r . . c l a v t o n i i f o l i a and S. o c c i d e n t a l i s v a r . 2 7 F i g u r e 4: H a b i t s k e t c h c f a r t i f i c i a l h y b r i d 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 l a n t s . 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 . o c c i d e n t a l i s IB), and an a r t i f i c i a l l y produced S. r u f i d u l a x 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 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 F i g u r e 5: H a b i t s k e t c h o f a r t i f i c i a l h y b r i d 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 l a n t s . 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 . , 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 a r t i f i c i a l l y produced S. r u f i d u l a x S. o c c i d e n t a l i s v a r . d e n t a t 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. 31 F i g u r e 6: H a b i t s k e t c h o f a r t i f i c i a l h y b r i d 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 l a n t s . D i p l o i d S. r u f i d u l a - ( f l ) . t e t r a p l o i d S. i n t e q r i f o l i a ( B ) , and an a r t i f i c i a l l y 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. . 3 2 gure 7 : H a b i t s k e t c h of a r t i f i c i a l h y b r i d 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 l a n t s . N=38 S . o c c i d e n t a l i s v a r . 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 produced S. r u f i d u l a x 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 FI 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 F i g u r e 8: H a b i t s k e t c h of a r t i f i c i a l h y b r i d 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 l a n t s . , 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 . d e n t a t a (A), t e t r a p l c i d S. o c c i d e n t a l i s (B) , and an a r t i f i c i a l l y 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 v a r . d e n t a t 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. 36 37 d e n t a t a ( F i g . 10) and u s u a l l y more g r a d u a l f o r members c f the o t h e r groups ( F i g . 9 ) . The r a t h e r d i s t i n c t rounded or somewhat squared t e e t h of S . r u f i d u l a and S. o c c i d e n t a l i s make t h e b l a d e - p e t i o l e t r a n s i t i o n e v i d e n t but t h e a c t u a l a n g l e c f the b l a d e base i s u s u a l l y r a t h e r o b t u s e i n t h e s e t a x a ( F i g . 10). M a r g i n a l t e e t h of S i r u f i d u l a t e n d t o be d e e p l y rounded and a c u t e whereas 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 t e e t h a r e u s u a l l y r i g h t - a n g l e d o r o b t u s e . However. S. r u f i d u l a p l a n t s from the Columbia B i v e r Gorge a r e a p p a r e n t l y i n t r o g r e s s a n t f o r t h i s c h a r a c t e r ( F i g . 9 ,A-C). Leaves of S. r u f i d u l a a r e u s u a l l y r e d d i s h t i n g e d on the l o w e r s u r f a c e s w h i l e t h o s e 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 and S. o c c i d e n t a l i s v a r . d e n t a t a v a r y from green t o r e d d i s h . S a x i f r a g a o c c i d e n t a 1 i s v a r . l a t i p e t i o l a t a l e a v e s a r e u s u a l l y l i g h t green on both s u r f a c e s , a f e a t u r e r e s e m b l i n g members of the S. i n t e g r i f o l i a -oreqana group. Seme m o r p h o l o g i c a l f e a t u r e s which r e q u i r e a b r o a d e r s u r v e y and f u r t h e r a t t e n t i o n i n c l u d e : l e a f t h i c k n e s s i n c r o s s s e c t i o n , stcmate c h a r a c t e r i s t i c s , f i n e s t r u c t u r a l d i f f e r e n c e s i n seed c o a t s c u l p t u r i n g , and p a p i l l c s i t y of p e t a l s , f i l a m e n t s , and n e c t a r g l a n d s . Although v a r i a t i o n was observed i n t h e s e f e a t u r e s , d e f i n i t e c o r r e l a t i o n s w i t h most groups were not p o s s i b l e on t h e b a s i s o f l i m i t e d m a t e r i a l and data c o n c e r n i n g t h e e x t e n t o f v a r i a b i l i t y i n e t h e r members of S a x i f r a g a . These c h a r a c t e r i s t i c s may be of c o n s i d e r a b l e u s e f u l n e s s i n d e f i n i t i o n of s u b s e c t i o n s w i t h i n t h e genus. I t was noted t h a t S. i n t e q r i f c l i a v a r . l e p t o p e t a l a a p p a r e n t l y i s unique amonq the observed t a x a i n t h a t i t s p e t a l s a r e w i t h o u t a d a x i a l 38 F i g u r e 9: L e a f o u t l i n e d r a wing shewing v a r i a b i l i t y i n shape and margin c h a r a c t e r i s t i c s i n samples o f d i 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 j . q e n t a l i s v a r . o c c i d e n t a l i s (B) , t e t r a p l o i d and h e x a p l o i d 5. r u f i d u l a i n t h e Columbia B i v e r Gorge ( C ) , n=38 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 (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 v a r . i n t e g r i f o l i a (E) l e a v e s . Each sample i s from one p o p u l a t i o n . Each l e a f i s taken from t h e l o w e r w h o r l o f l e a v e s from 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 d i f f e r e n c e s i n p e t i o l e l e n g t h and w i d t h , p e t i o l e - b l a d e t r a n s i t i o n and d e n t i t i o n . i n c r e a s e d v a r i a b i l i t y o f Columbia B i v e r Gorge p o l y p l o i d S. r u f i d u l a p l a n t s and resemblance 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 drawing showing v a r i a b i l i t y i n shape and d e n t i t i o n i n samples of 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 v a r * 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 t a k e n from th e lower w h o r l o f l e a v e s from 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 the 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 seed c o a t s u r f a c e s i s w i t h o u t r i d g e s o r p r o j e c t i o n s . I t would be of i n t e r e s t t o s t u d y d i f f e r e n c e s i n f l o r a l u l t r a - v i o l e t r e f l e c t a n c e p a t t e r n s f o r t h i s t a x c n and i t s r e l a t i v e s . Use o f t h e s e c h a r a c t e r i s t i c s may prove e s p e c i a l l y h e l p f u l i n s t u d i e s o f E a s t e r n and Western North American s p e c i e s - p a i r s . Chromosomal S t u d i e s P o l y p l o i d - a n e u p l c i d s e r i e s a r e common f o r many groups o f p l a n t s (Tobgy 1943, Lewis and Raven 1958, J a c k s o n 1962, Kyhos 1965, C a r r 1975, S u b h a s i 1975) i n c l u d i n g S a x i f r a g a { D a m b c l t and P c d l e c h 1965, Dambolt 1968, E l v a n d e r 1975) and a r e c o n f i r m e d i n the p r e s e n t s t u d y as w e l l . D i p l o i d s w i t h 10 p a i r s , t e t r a p l o i d s w i t h 19 p a i r s and h e x a p l o i d s w i t h 29 c r 28 p a i r s have been r e p o r t e d f o r 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 and 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 i n 5. f e r r u g i n e a a 10 - p a i r e d d i p l o i d and 1 9 - p a i r e d reduced t e t r a p l o i d s i t u a t i o n fRandhawa and Eeamish 1972) has e v o l v e d i n an a p p a r e n t l y p a r a l l e l } p o l y p l o i d - a n e u p l c i d p a t t e r n t o the one seen i n S. r u f i d u l a and i t s r e l a t i v e s . The same s i t u a t i o n may a l s o be the case i n S. i n t e q r i f o l i a v a r . i j j t e j r j , f o l i a where 1 9 - p a i r e d p o p u l a t i o n s a r e common. Chromosome counts f o r S a x i f r a g a p o p u l a t i o n s a r e summarized i n T a b l e 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 from Vancouver I s l a n d , t h e Olympic P e n i n s u l a and t h e Opper Willamette R i v e r dxainaqe a r e , 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 p a i r s of chromosomes (see F i g . 14C). Krause and Beamish (1913) 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 (n=29) a t E l k F a l l s on Vancouver I s l a n d . A ttempts t c r e l o c a t e t h a t p o p u l a t i o n were u n s u c c e s s f u l , p o s s i b l y because o f l o c a l e x t i n c t i o n r e s u l t i n g from h y d r o e l e c t r i c water d i v e r s i o n . . S a x i f r a g a r u f i d u l a p o p u l a t i o n s from t h e Columbia R i v e r Gorge a r e more complex and v a r i a b l e i n t h e i r chromosome numbers. D i p l o i d i n d i v i d u a l s (n=10) o c c u r i n some mixed p o p u l a t i o n s w i t h t h e more common t e t r a p l o i d (n=19) ( F i g . 11). Other p o p u l a t i o n s had o n l y t e t r a p l o i d or 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 . 1 2 ) , but t h e p o s s i b i l i t y o f r a r e d i p l o i d i n d i v i d u a l s cannot be r u l e d c u t c o n s i d e r i n g t h e s m a l l number of c o u n t s made. , Some of t h e S. r u f i d u l a p l a n t s from most p o p u l a t i o n s a l o n g t h e Columbia R i v e r Gorge undergo an abnormal m e i o s i s t y p i c a l of h y b r i d s between p l a n t s of d i f f e r e n t l e v e l s of p o l y p l o i d y ( F i q . 13A,B). D i p l o i d , t e t r a p l o i d and h e x a p l o i d p l a n t s and t h e i r p u t a t i v e n a t u r a l h y b r i d s a r e m o r p h o l o g i c a l l y a l m o s t i d e n t i c a l . A 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 Southwestern Washington p r e v i o u s l y r e f e r r e d t o S. o c c i d e n t a l i s (Krause and Maze No. 690001 i n Krause and Beamish 1972) i s 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 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 . 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 (n=29, F i q . 13C,D). T h i s p o p u l a t i o n p r o b a b l y r e p r e s e n t s a r a r e , s t a b i l i z e d a l l o p o l y p l o i d h y b r i d e n t i t y . Attempts t o l o c a t e s i m i l a r p o p u l a t i o n s e l s e w h e r e i n t h e area were u n s u c c e s s f u l , a l t h o u q h t h e r e are resemblances t c c e r t a i n S. i n t e g r i f o l i a specimens o f the F o r t L e w i s and Olympia, Washinqton r e q i o n s . One specimen frcm near Mima, Washinqton had an u n c e r t a i n chromosome c o u n t of 44 T a b l e 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.) o c c i d e n t a l i s No. No. P l a n t s C e l l s M e i o t i c Chromosome No. I l l o_r II+ I ) * Y a l e , B.C. 2 6 19 (626) C o r n w a l l Lookout, B.C. 3 10 19 (COBN) Ht. B a k e r , Hash. 2 5 19 (BAK) a t a Saddle Mt., Oregon 3 8 10 (629A) T i l l a m o o k , Oregon 11 44 10 (630) Kalama B i v e r , Hash. 2 7 20 (607) D e lena, Oregon (606) 8 23 20(6) ,19+2(2) . d u l a M t . F i n l a y s o n , V . I . , 3 20 10 B.C. (627A) Nanoose H i l l , V . I . , 4 7 10 B.C. (NOOS-B) Nanaimo,V.I.,B.C. , 4 31 10 (NMO-R) Upper Campbell L a k e , 3 5 10 V.I.,B.C. (UCL-R) l a k e C r e s c e n t , Wash. 4 16 10 (673) Bingen Lake, Hash. , 2 10 29 (669A) H t . P l e a s a n t , Wash. 1 3 10 (675A) Yecn Park, Ore. 6 21 10(2) ,19(3) ,ca.20 (610) V i e n t c , Ore., 5 13 1 9 ( 4 ) , ca.19 (616) T r o u t d a l e , O r e . 5 7 c a . 2 0 * 1 , ca.27+2. (608) 29, 19+12, 19+13 Bayer P a r k , Ore. 4 11 ca.19,ca.16+8, <612A) ca.29, ca.20+5 The D a l l e s , Ore. 4 16 ca.29 (3) ,ca. 26+6 Skamania Co., Wash. 4 26 19"(3) ,ca. 14+8 (618A) •Numbers i n p a r e n t h e s e s i n d i c a t e the number of 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 T a b l e 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 n o A ) Ho. No. M e i o t i c Chromosome No P l a n t s C e l l s • j l l o r I I + I ) * 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 Saddle Mt., Ore. 5 (629C) i n t e g r i f o l i a M t . F i n l a y s o n , V . I . , 3 B.C. (627) Nanaimo, V.I.,B.C, 1 (NMO-I) El k F a l l s , V.I.,B.C. 1 (682) H a r r i s o n Lake, B.c. 1 (78-1) Y a l e , B.C. 1 (625) Mayer Park, Ore. 4 (612B) G r i z z l y Lake, Ca. 1 (671) B i n g e n , Hash. 1 (617) Mima Bounds, Hash. 1 (MIMft) S l S l t c n i i f o l i a T r o u t d a l e , Ore. 5 (609) Bowena, Ore. 5 (611) Mayer Pk., Ore. 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 Horn, Hash. 1 (646) L y l e , Hash. 1 B i n g e n , Hash. 6 (617) oreaa^a Berthoud P a s s , C o l o . (BEE) 11 23 3 2 3 3 2 20 4 1 4 16 12 27 6 55 20 2 10 1 30 28,29 (3) ca.36,38j(3) ,ca.40 19 19 19 19 19 19,ca. 17+5(3) 19 19 ca.47 10(4) ,19 10,8 + 11 (4) 10 (3) ,9+5 10 10 10 10 10 10 10 (2) ,14+10, ca.14+13, ca. 18+9, ca.23+10 c a . 36 46 n=ca. 47 (see F i g . 16D) and may be r e l a t e d t o t h e C h e h a l i s h y b r i d p o p u l a t i o n o r t o S. oregana ( 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 ) , The chromosome c o u n t s f o r 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 agree w i t h t h e m a j o r i t y o f e a r l i e r c o u n t s (Beamish 1961, Krause and Beamish 1972) o f n=19 f o r a l l p o p u l a t i o n s sampled i n t h e p r e s e n t s t u d y ( F i g . 14A,B,0)• P a c k e r (1968) has r e p o r t e d a d i p l o i d n=10 p o p u l a t i o n from B l a k e s t o n Mt., A l b e r t a and E l v a n d e r (1975) r e c o r d s two mixed d i p l o i d and t e t r a p l o i d p o p u l a t i o n s from T r a p p e r Peak i n t h e B i t t e r r o o t Mts., Montana and from the Storm Lake Pass i n the Anaconda Range, Montana, where i n d i v i d u a l s a p p a r e n t l y a r e i n t e r m e d i a t e between S. m a r s h a l l i i sufesp. i d a h o e n s i s and S. o c c i d e n t a l i s v a r , o c c i de ja t a l i s . Krause (Krause and Beamish 1972) has r e c o r d e d h i g h e r numbers of n=28 or 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 Columbia and t h e mountains of Idaho. P l a n t s i d e n t i f i e d as S. o c c i d e n t a l i s v a r . d e n t a t a ( H i t c h c o c k e t a l , 1973) were c o l l e c t e d from f o u r l o c a t i o n s west of the Columbia R i v e r Gorge and t o w a r d t h e mouth of the C c l u m b i a R i v e r . P l a n t s o f t h e C o a s t Range of Oregon, i n c l u d i n g t h o s e from Saddle Mountain, C l a t s o p County, are d i p l o i d (n=10) ( F i g . 15C-E). P l a n t s from Columbia County (EP606) # Oregon, and C o w l i t z Co.,Washington (EP607), have 20 o r sometimes 19 p a i r s o f chromosomes ( F i g . 15A,B). These t e t r a p l o i d s a s w e l l as o t h e r p l a n t s from the W i l l a m e t t e R i v e r V a l l e y p r o b a b l y r e p r e s e n t a l l o t e t r a p l o i d h y b r i d i z a t i o n s between S. i n t e q r i f o l i a v a r . c l a v t o n i i f o l i a and d i p l o i d S. o c c i d e n t a l i s v a r . d e n t a t a as judqed by t h e i r m o r p h o l o g i c a l resemblance and the p r o x i m i t y o f 47 F i g u r e 11: Camera l u c i d a drawing o f p o l l e n 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 grey c i r c l e s . Drawings here a r e from f o u r p l a n t s i n one p o p u l a t i o n a l o n g the Columbia R i v e r Gorge. P a r t s D and E a r e from t h e same p l a n t . ; &=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 metaphase 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 12: Camera l u c i d a drawings of PMC* s. S a x i f raga r u f i d u l a p l a n t s from the T r o u t d a l e p o p u l a t i o n . A=PMC at 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 b i v a l e n t s + 1 u n i v a l e n t 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 13: Camera l u c i d a drawings o f PMC's., C r y p t i c h y b r i d p l a n t s from T r o u t d a l e (A,B) and C h e h a l i s R i v e r (C,D)• a=PMC a t l a t e metaphase 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 m u l t i v a l e n t s ? ) B=PMC a t l a t e metaphase I ; S, r j i f i d u l a c r y p t i c h y b r i d (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 = p c l l e n g r a i n m i t o s i s ; metaphase; 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 (EP605-26) n=29 53 Figure 14: Camera lucida drawings cf PMC»s. Plants from Yale, Mt. Baker and Cornwall populations of S. .occidentalis var, o c c i d e n t a l i s and Mt. Finlayson population of S. ru f i d u l a . Upper scale i s for c e l l s A-C, lower scale i s f o r c e l l s D and E. A=PMC at diplotene-diakinesis; S. oc c i d e n t a l i s var. occide 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. occidentalis (BAKEB) n= 19 C=PMC at diplotene-diakinesis; S. rjjfidaia (EP627) n=10 D=PMC at metaphase I; S. occidentalis var. occide n t a l i s lEPCOBN) n=19 ~E=PMC at~~metaphase I; S. rufid u l a (EP618-17) n=19 55 F i g u r e 15: Camera l u c i d a drawings o f PMC*s. S. o c c i d e n t a l i s v a r , d e n t a t a p l a n t s from Kalama B i v e r , D e l e n a , T i l a m o o k , 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 ; 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 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 . de 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 d i a k i n e s i s and 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 (EP629A-25) n=10 57 S. i n t e q r i f o l i a v a r . c l a y t q n i i f o l i a p l a n t s . However, seme of th e S. o c c i d e n t a l i s v a r . d e n t a t a p l a n t s from the W i l l a m e t t e B i v e r i n t e r g r a d e i n f e a t u r e s such as c l a v a t e f i l a m e n t s t o S . , m a r s h a l l i i subsp. a a r s h a l l i i b u t the f i l a m e n t s are l e s s expanded and p e t a l - l i k e t han t h o s e o f S. m a r s h a l l i i subsp. m a r s h a l l i i . P l a n t s known as S. o c c i d e n t a l i s v a r . , 1 a t i p e t i o l a t a ( H i t c h c o c k e t a l , 1973) from i s o l a t e d peaks of the N o r t h e r n Oregon Coast Bange (Chambers 1974), were found t o have a h i q h nuaber o f chromosomes. Counts v a r i e d somewhat, w i t h most b e i n g around 38 o r 39 p a i r s ( F i g . 16A,C). T h i s c o r r e s p o n d s w i t h c o u n t s of S. oreqana from a t . Hood, Oregon (n=ca. 38, E l v a n d e r , p e r s o n a l communication) and from Berthoud P a s s , C o l o r a d o (n=ca. 36) ( 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 19 chromosome p a i r s on the Lower Mai n l a n d o f B r i t i s h C o l u m b i a , o r Vancouver I s l a n d ( F i q . 16,B,E) and southward i n t o Washinqton. P l a n t s from t h e Columbia B i v e r Gorqe and 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 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 a r e n e c e s s a r y t o d e t e c t c o r r e l a t e d chromosomal d i f f e r e n c e s . At l e a s t some o f t h e p l a n t s from the Columbia B i v e r Gorge have t h e 1 9 - p a i r e d complement. P o p u l a t i o n s from the p r a i r i e s s o u t h o f Tacoaa and Olympia Washington may be r e l a t e d t o S. oregana and one i n d i v i d u a l had a h i g h number o f chromosomes (2n=ca. 47,51) (my c o u n t s . Fig.16,D and 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 ) . S a x i f r a g a i n t e g r i f o l i a v a r . clay, t o n j i f o l i a p o p u l a t i o n s have been r e p e a t e d l y counted as d i p l o i d s w i t h 10 chrcuoscme p a i r s ( F i g . 17,B-F). One 1 9 - p a i r e d i n d i v i d u a l was r e c o v e r e d from 58 t h e o t h e r w i s e 1 0 - p a i r e d p o p u l a t i o n a t T r o u t d a l e , Oregon. P l a n t s from near B i n g e n , Washington were mostly s t e r i l e h y b r i d forms w i t h abnormal m e i o s i s a l t h o u g h a few 1 0 - p a i r e d i n d i v i d u a l s were found. F u r t h e r c y t e l o g i c a l and m o r p h o l o g i c a l a n a l y s e s o f t h e E a s t e r n Columbia R i v e r Gorge p o p u l a t i o n s a r e n e c e s s a r y t o u n r a v e l the 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 p l a n t s and t h e c l o s e l y r e l a t e d S. i n t e g r i f o l i a v a r . i n t e q r i f o l i a and S. i n t e g r i f o l i a v a r . l e p t p p e t a l a which a l s o o c c u r t h e r e . Northward i n i t s range S. i n t e q r i f o l i a v a r . l € P t o p e t a , l a has 19 p a i r s o f chromosomes but p o p u l a t i o n s from the E a s t e r n Columbia R i v e r Gorge may be more complex chromosomally where t h e y i n t e r g r a d e w i t h 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 and l . i n t e g r i f o l i a v a r . i n t e q r i f o l i a , I n S a x i f r a g a c l o s e d i p l o i d r e l a t i v e s w i t h 9 c h r c i o s c m e p a i r s a r e n o t a b l y a b s e n t from l i t e r a t u r e r e p o r t s and frcm t h e p r e s e n t s t u d y . The l a c k o f such e n t i t i e s i n n a t u r e argues a g a i n s t t h e p r o d u c t i o n o f 1 9 - p a i r e d p l a n t s d i r e c t l y from chromosome d o u b l i n g o f a h y b r i d between 10 and 9 - p a i r e d p a r e n t a l p l a n t s . A more p l a u s i b l e h y p o t h e s i s f o r the 10,19,20,28,29,38 p o l y p l o i d - a n e u p l o i d seguence based on t h e a v a i l a b l e e vidence i s t h a t 1 0 - p a i r e d p r o g e n i t o r s gave r i s e t o p o l y p l o i d o f f s p r i n g w i t h 20 p a i r s of chromosomes f o l l o w e d by subsequent l o s s of a chromosome p a i r . Some p a i r i n g i n s t a b i l i t y i s apparent i n one 2 0 - p a i r e d 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 v a r . d e n t a t a which may be an i n d i c a t i o n t h a t i t i s of r e c e n t o r i g i n and may e v e n t u a l l y conform to the 1 9 - p a i r e d p o l y p l o i d - a n e u p l o i d p a t t e r n found i n many p o p u l a t i o n s o f the o t h e r t a x a . C o n t a c t and c r o s s i n g between 19 and 1 0 - p a i r e d p l a n t s f o l l o w e d by d o u b l i n g o f chromosome 59 F i g u r e 16: Camera l u c i d a drawings of PMC ,s of 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 and S. i n t e g r i f oljj, a p i a nt 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 v a r . l ^ i i e e t i o l a t a <EP629C) n=ca. 38 B=P0C a t metaphase I; S. i n ^ a E i J p J L i a v a r . 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 v a r . l a t i p e t i o l a t a (E0629C) n= 3 8 D=PMC a t d i a k i n e s i s ; S.- . i n t e g r j f o ^ j a - S . oreqana? (EPMIHft) n=ca.47 E=PHC a t d i a k i n e s i s ; 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 A. B. M X 60 c. D. y \ a 3 ^ 10JJ 61 F i g u r e 17; Camera l u c i d a drawings of PHC*s. P l a n t s from Berthoud 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 , and Bowena p o p u l a t i o n s . The upper s c a l e i s f o r c e l l s A-C, the lowe r s c a l e i s f o r c e l l s D-F. A=PMC a t anaphase I ; S. oregana (BPBEfi) n=ca. 36 Upper p o l e shows 34 b o d i e s o f which a t l e a s t 2 a r e o v e r l a p p i n g . l o w e r p o l e shows 35 bo d i e s o f which a t l e a s t 2 a r e o v e r l a p p i n g , 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 a t d i a k i n e s i s ; 5. / 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 (EP668-14) n=10 D=PMC a t metaphase I ; S. i n t e g r i f o l i a v a r . c j j j j t o n i i f o l i a (EP617-21) n=10 E=PMC a t d i a k i n e s i s ; S. i n t e g r j - f o l i a v a r . c l a j t o n i i f o l i a (EP6C9-310) n=10 F=PHC a t l a t 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 y t o n i i f o l i a (EP611) n=10 62 63 number produced 2 9 - p a i r e d p o p u l a t i o n s , and t h e n=28 c o u n t s may be f u r t h e r a n e u p l o i d r e d u c t i o n s . , P r o b a b l y c o u n t s of 38 p a i r s r e p r e s e n t h y b r i d c o m b i n a t i o n s of two 1 9 - p a i r e d e n t i t i e s o r c o m b i n a t i o n s i n v o l v i n g d i p l o i d and h e x a p l o i d components. A l t h o u g h the p o l y p l o i d p r o c e s s e s i n t h i s group appear t o i n v o l v e mostly a l l o p o l y p l o i d y , t h e p o s s i b i l i t y of a u t o p c l y p l o i d y cannot be e l i m i n a t e d . R e p o r t s i c S. r e f l e x a (Krause and Beamish 1973) where d i p l o i d s , t e t r a p l o i d s , and h e x a p l c i d s were found i n one p o p u l a t i o n seem to i n d i c a t e t h a t a u t o p o l y p l o i d y o c c u r s i n r e l a t e d t a x a . Moore (1959) and T a y l o r and M u l l i g a n (1968) r e p o r t an a p p a r e n t l y a u t o p o l y p l o i d d e r i v a t i o n f o r p o p u l a t i o n s of S. t a y l o r i . an endemic to t h e Queen C h a r l o t t e I s l a n d s and t h e Brooks P e n i n s u l a on Vancouver I s l a n d , B r i t i s h C o lumbia. M l b r i d i z a t i o n Most a r t i f i c i a l F1 h y b r i d s among S. r u f i d u l a . 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 . S. o c c i d e n t a l i s v a r . d e n t a t a , and S. i n t e g r i f o l i a , a r e g e n e r a l l y 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 the two p a r e n t a l e n t i t i e s { F i g . 4-8). However, the m u l t i v a r i a t e computer a n a l y s e s p l a c e d c e r t a i n h y b r i d s c l o s e r t o t h e i r p o l y p l o i d p a r e n t s than t o t h e i r d i p l o i d p a r e n t s , e s p e c i a l l y • S. i n t e g r i f o l i a • and S. o c c i d e n t a l i s ( F i g . 19-21). P o l l e n f e r t i l i t y of h y b r i d s i s summarized i n T a b l e IV. A r t i f i c i a l h y b r i d i z a t i o n s were s u c c e s s f u l among a l l c o m b i n a t i o n s o f t a x a which were attempted ( F i g . 2 2 ) . D i f f e r e n c e s i n seed s e t per 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 about by i n e f f i c i e n t c r o s s i n g t e c h n i q u e s as w e l l as s m a l l sample 64 s i z e s . S i m i l a r v a r i a t i o n i n seed g e r m i n a t i o n r a t e s (.18-.90) may be the r e s u l t o f d i f f i c u l t i e s i n assessment o f seed m a t u r i t y , i m proper g e r m i n a t i o n c o n d i t i o n s , o r s m a l l sample s i z e s , a p p a r e n t l y i t i s p o s s i b l e t o o b t a i n F1 h y b r i d s from c r o s s e s among most p l a n t s i n t h i s r e l a t e d group w i t h o u t d i f f i c u l t y . The a r t i f i c i a l F1 h y b r i d s which s u r v i v e d t o m a t u r i t y demonstrated v a r i o u s m e i o t i c b e h a v i o u r s . B i v a l e n t f o r m a t i o n was r e s t r i c t e d t o a f r a c t i o n of the genome and m u l t i v a l e n t s were absent i n n e a r l y a l l c r o s s e s ( F i g . 18). a p o s s i b l e e x c e p t i o n i s th e c r o s s between S. r u f i d u l a (n=19) and 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 (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 a s s o c i a t i o n s , c o u l d Jse seen i n seme c e l l s . B i v a l e n t f o r m a t i o n was h i g h (15-19 p a i r s ) , 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 v a r . l a t i p e t i o l a t a had a h y b r i d o r i g i n i n v o l v i n g t e t r a p l o i d S. r u f i d u l a . The m o r p h o l o g i c a l a f f i n i t i e s and p l o i d y l e v e l (n=ca. 38) o f 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 s u g g e s t t h a t S. oregana (n=19?.ca. 36.ca. 38) was p r o b a b l y t h e o t h e r p a r e n t . a n a l y s i s o f m e i o s i s i n a r t i f i c i a l i n t e r v a r i e t a l 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 has been c o n s i d e r a b l e genomic d i v e r g e n c e among the t a x a . When d i p l o i d S. r u f i d u l a i s c r o s s e d 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 , the r e s u l t a n t h y b r i d has 10 or fewer b i v a l e n t s . H y b r i d s between S. r u f i d u l a (n=19) and -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=19) produced about 9 b i v a l e n t s i n d i c a t i n g t h a t t h e y have o n l y a p a r t i a l chromosome complement i n common ( F i g . 18,C). I n c o n t r a s t , t h e c r o s s between pcGi den t a l i s v a r . , O c c i d e n t a l i s and S. o c c i d e n t a l i s v a r . d e n t a t a (n=20) produced F1 i n d i v i d u a l s which were almost c o m p l e t e l y s t e r i l e and f a i l e d t o undergo m e i o t i c d i v i s i o n . When 65 S» r u f i d u l a (n=10) i s c r o s s e d w i t h S. i n t e q r i f o l i a (n=19) u s u a l l y 10 o r fewer b i v a l e n t s a r e f o r c e d (Fig* 1 8 ) . The r e s u l t a n t h y b r i d s c l o s e l y resemble the s t e r i l e i n t e r m e d i a t e s found i n many f i e l d s i t u a t i o n s (see F i g . 6 ) . The c r o s s between 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 and p o l y p l o i d S. r u f i d u l a produced s e v e r a l v i g o r o u s F1 h y b r i d s which had a h i g h e r p o l l e n f e r t i l i t y t h a n o t h e r a r t i f i c i a l h y b r i d s (Table I V ) . O c c a s i o n a l l y an a r t i f i c i a l F1 h y b r i d d e v i a t e d e n t i r e l y from e i t h e r p a r e n t a l or i n t e r m e d i a t e m o r p h o l o g i e s ( F i g . 19-21, p l a n t no. 3*). C r o s s e s between S. i n t e q r i f o l i a and S. i n t e q r i f o l i a v a r . l e p t c p e t a l a resembled S. i n t e g r i f o l i a i n p e t a l c h a r a c t e r i s t i c s and had from 6 t o n e a r l y 19 b i v a l e n t s but even i n t h e h i g h e r p a i r i n g i n d i v i d u a l s anaphase a b n o r m a l i t i e s produce h i g h l y s t e r i l e p o l l e n . S. r u f i d u l a x S. o c c i d e n t a l i s F1 h y b r i d s resembled §» o c c i d e n t a l i s i n i n f l o r e s c e n c e shape and s i z e (see F i g . 4 ) . A l t h o u g h b i o l o g i c a l r e l a t i o n s h i p s and a f f i n i t i e s may be e l u c i d a t e d by h y b r i d i z a t i o n e x p e r i m e n t s , taxonomic d e c i s i o n s i n t h i s group p r o b a b l y s h o u l d be based on t h e more pragmatic s i m i l a r i t i e s and d i s c o n t i n u i t i e s of m o r p h o l o g i c a l and merphcraetrie d a t a . , S i n c e a l l p o s s i b l e c r o s s e s were not completed ( F i g . 2 2 ) , many a s p e c t s o f t h e h y b r i d r e l a t i o n s h i p s a r e as yet u n c l e a r . Even i f a l l h y b r i d s were a v a i l a b l e f o r a n a l y s i s c o n s i d e r a b l e c a u t i o n i n i n t e r p r e t a t i o n would be n e c e s s a r y i n l i g h t o f l i t e r a t u r e r e p o r t s t h a t i n wheat, o a t s , r y e , and e t h e r s , homeclogous p a i r i n g i s under g e n e t i c c o n t r o l ( a l l e y 1960 # R i l e y and Law 1965, R ajhathy and Thomas 1972, H o s s a i n 1977) and t h a t b i v a l e n t o r m u l t i v a l e n t f o r m a t i o n may be a f u n c t i o n ef the 66 F i g u r e 18: Camera l u c i d a drawings o f PMC*s from a r t i f i c i a l h y b r i d s . The upper s c a l e i s f o r c e l l s A-B, t h e lower s c a l e i s f o r c e l l s C-D. A=PMC a t metaphase I ; S. o c c i d e n t a l ' s v a r . 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 metaphase 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 u n i v a l e n t s C=PMC a t metaphase I ; S. r u f i d u l a t e t r a p l o i d 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 . d e n t a t a t e t r a p l o i d 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 u n i v a l e n t s 68 Figure 19: P r i n c i p a l components analysis graph of axis one vs. two showing a r t i f i c i a l l y produced hybrid i n d i v i d u a l s and representative parental populations., Symbols are as follows: D= S. occi d e n t a l i s var. / dentata : 1= S. I n t e q r i f o l i a var. 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. occidental's B= S. r u f i d u l a (from Vancouver Island and the Olympic Mountains) r= S. r u f i d u l a (from the Columbia Biver 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. occidentalis (o) 3= S, occidentalis (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, occidentalis var. dentata (D) 5= S. r u f i d u l a (r) x S. occidentalis var. dentata (D) 6= S. r u f i d u l a (B) x S. occidentalis var. 1§M BftA-ol§ia (L) 3* occupies a point to the right of i t s placement within the plot. I t i s probably an aberrant i n d i v i d u a l . , The percentage of the t o t a l variance accounted for by the f i r s t three axes i s 21.36, 11.73, and 7.87 percent respectively. Eigenvector 2 U) O U> H 5 1 + "2° S3 ro-ro ro ro ro m CD < CD o-n <-+ O ~5 Or D M M O M O O M ~ 01 * o ro 0) ro 05 ( J l o o o M Of LO ro-D O 7 0 F i g u r e 20: 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. t h r e e 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 and t o t a l v a r i a n c e f i g u r e s a r e as i n F i g u r e 19. .71 .24-c_ O +-> u £ 0 § . Li R R R 1 r 4 r R R 1 o r 6 -.2-h D I L D x l 1 L D o D °I D o o D D D D D D D D D D D D • 1 0 Eigenvector 1 3 72 F i g u r e 21: P r i n c i p a l components a n a l y s i s graph c f a x i s two vs. t h r e e 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 . 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 as i n as 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 the lower r i g h t c o r n e r than drawn here. ! L L 2 L c I D Ri 1 1 R o R R R K I D r r 2 1 ° r~ 3 o° r 6 £ J 5 R L 1 L D 23 LR 0 0 • 0 4 L D D D D D D D DD 0 _ ^Eigenvector 2 74 F i g u r e 22: C r o s s i n g s u c c e s s c h a r t f o r t h e t a x a c o n s i d e r e d i n the p r e s e n t 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 r e l a t i v e s u c c e s s of seed s e t . Numbers o u t s i d e t h e 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 the l i n e s a r e the average number o f seeds s e t per attempt.,Numbers 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 s e t f o l l o w e d 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 as f o l l o w s : C=S, o c c i d e n t a l i s • • v a r , d e n t a t a . 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 v a r , o c c i d e n t a l i s , and B=S, r u f i d u l , a . The absence o f c o n n e c t i n g l i n e s i n d i c a t e s t h a t those c r o s s e s were not attempted. 75 Cros s ing 19 76 T a b l e IVj. H y b r i d P o l l e n F e r t i l i t y . C r o s s Between: r u f . (10) r u f . 119) r u f . (10) r u f . (19) occ. (19) l a t i . (ca. i n t . (19) i n t . (19) occ. (19) o c c . (19) dent.|19) dent.(19) dent. (19) 38) x r u f . x r u f . (10) * x l e p t . ( 1 9 ) * No. I n d i v i d u a l s 4 4 3 1 3 3 4 2 (19) average J l e r t i l i t i 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. ?) * These c r o s s e s were made e a r l i e r by K.I. Beamish and a r e i n c l u d e d here f o r comparison. V a r i e t a l names are a b b r e v i a t e d t o the f i r s t t h r e e or 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 or gene c o m b i n a t i o n s . F u r t h e r m o r e , numerous a u t h o r s have shown t h a t m e i o t i c e v e n t s i n a n t h e r t i s s u e can be e n v i r o n m e n t a l l y m o d i f i e d {Skovsted 1934, Sax 1937, Singh 1975, Beamish 1961) . O r n d u f f (1969) has p r e s e n t e d s e v e r a l c a s e s which argue a g a i n s t r i g i d s p e c i e s d e f i n i t i o n s based on c r o s s a b i l i t y and/or i n t e r f e r t i l i t y i n p l a n t h y b r i d i z a t i o n s t u d i e s . N a t u r a l l y o c c u r r i n g h y b r i d s between l o c a l l y s y m p a t r i c S. r u f i d u l a and S. -, i n t e < j r i f o l i a v a r . i n t e j r . i f . p l i a on Vancouver I s l a n d and i n p l a c e s a l o n g t h e Columbia B i v e r Gorge can be t e n t a t i v e l y r e c o g n i z e d by t h e i r i n t e r m e d i a t e morphology which r e s e m b l e s the a r t i f i c i a l l y produced h y b r i d ( F i g . 6 ) , abnormal m e i o s i s , and t h e i r h i g h degree of p o l l e n s t e r i l i t y . H y b r i d i n t e r m e d i a t e s c o n s i s t e n t l y f a i l t o s e t good seed but a r e v e g e t a t i v e l y v i g o r o u s and may e x i s t i n n a t u r e f o r many y e a r s . A r t i f i c a l F1 h y b r i d s between S . r j i f i d u l a and S. i n t e g r i f o l i a have f l o u r i s h e d under c u l t i v a t i o n i n t h e c o l d frames a t t h e U n i v e r s i t y o f B r i t i s h Columbia f o r 12 y e a r s . J u d g i n g from a l a r g e p o p u l a t i o n s o u t h o f Nanaimo, B.C., s t e r i l e i n t e r m e d i a t e i n d i v i d u a l s a r e , i n n a t u r e , u s u a l l y r a r e i n c omparison t o t h e i r p a r e n t a l e n t i t i e s . Only about 30 p u t a t i v e h y b r i d s were seen compared t o about 3,000 S. r u f i d u l a p l a n t s and c o n s i d e r a b l y more S. i n t e q r i f o l i a v a r . i n t e g r i f o l i a 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 i n d i v i d u a l s r e s e m b l i n q S. r u f i d u l a i n h a b i t a t and morpholoqy a r e perhaps more common. The e x a c t n a t u r e o f h y b r i d r e l a t i o n s h i p s i s much more d i f f i c u l t t o d etermine 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 and up t o t h r e e v a r i e t i e s o f S. i n t e g r i f o l i a may occ u r i n c l o s e p r o x i m i t y . V a r i a b i l i t y i n l e v e l s o f p l o i d y w i t h i n p o p u l a t i o n s o f S, r u f i d u l a as w e l l as S. i n t e g r i f o l i a v a r . c l a y t o n i i f o l i a a l s o confound p r e c i s e d e t e r m i n a t i o n s o f h y b r i d o r i g i n s . I n d i v i d u a l s which a r e m o r p h o l o g i c a l l y c l o s e t o one o r the o t h e r o f t h e p a r e n t a l e n t i t i e s may prove t o be m e i o t i c a l l y i r r e g u l a r and p o l l e n s t e r i l e . Samples f o r c e r t a i n Columbia R i v e r Gorge l o c a t i o n s , n o t a b l y t h e S. i n t e g r i f o l i a p o p u l a t i o n n e a r B i n g e n , Washington, c o n t a i n a l a r g e p r o p o r t i o n of t h e s e c r y p t i c h y b r i d s . J u d g i n g from herbarium m a t e r i a l , h y b r i d swarms a r e common i n s e v e r a l a r e a s . , Wilhelm S u k s d o r f i n 1916 made a mass c o l l e c t i o n of one such area o f E a s t e r n Washington near Spokane and Spangle where p u t a t i v e S, o c c i d e n t a l i s {or S. m a r s h a l l i i subsp., i d a h o e n s i s ) x S. i n t e g r i f o l i a v a r . l e p t o p e t a l a {or v a r . columbiana?) h y b r i d s were f o u n d . P a r e n t a l p l a n t s were e i t h e r r a r e o r i n a d e g u a t e l y c o l l e c t e d . Many o f t h e p u t a t i v e h y b r i d p l a n t s had reduced p o l l e n f e r t i l i t y as w e l l as b e i n g somewhat i n t e r m e d i a t e i n appearance. Another a r e a w i t h h i g h numbers of p u t a t i v e h y b r i d i n d i v i d u a l s between 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 and S. i n t e g r i f o 1 i a v a r . l e p t o p e t a l a i s t h e Petes P o i n t a r e a of the Wallowa Mountains o f n o r t h e a s t e r n Oregon. H y b r i d 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 v a r . c l a y t o n i i f o l i a and S. m a r s h a l l i i subsp. m a r s h a l l i i have been r e c o r d e d f o r t h e Mary's Peak area o f t h e Coast Range and Opper W i l l a m e t t e r e g i o n o f western and c e n t r a l Oregon. I n the s e a r e a s s h o r t - t e r m h y b r i d f i t n e s s , l o n g e v i t y , v e g e t a t i v e r e p r o d u c t i o n , and l a r g e a r e a s o f p o s s i b l y i n t e r m e d i a t e h a b i t a t may e x p l a i n the 79 apparent abundance of h y b r i d forms and s c a r c i t y o f p a r e n t a l forms., F e r t i l e p o l y p l o i d p o p u l a t i o n s of probable h y b r i d o r i g i n a r e common i n s e v e r a l l o c a t i o n s i n t h e Lower Columbia R i v e r a r e a . Two t e t r a p l o i d p o p u l a t i o n s o f S. o c c i d e n t a l i s v a r . d e n t a t a which conform t o E n g l e r and I r m s c h e r ' s (1916) d e s c r i p t i o n and which are near the t y p e l o c a l i t y f o r S. o c c i d e n t a l i s v a r . 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 v a r . d e n t a t a and S. i n t e g r i f o l i a v a r . c l a y t o n i i f o l i a ( F i g . 2 3 ) . These resemble the type specimens o f §. o c c i d e n t a l i s v a r . d e n t a t a from E l k Rock, near Oswego, Clackamas County, Oregon (the f i r s t specimen c i t e d i n E n g l e r and Ir m s c h e r 1916, i s H e l l e r 10059) ., Krause and Beamish (1972) have p r e v i o u s l y r e f e r r e d t h e C l a t s o p Co. P l a n t s t o S. o c c i d e n t a l i s but synonymized S. o c c i d e n t a l i s - v a r . d e n t a t a w i t h S. m a r s h a l l i i subsp. , m a r s h a l , l i j . I n t r o g r e s s i o n o f S. i n t e j r i f o l i a v a r . c l a v t o n i i f o l i a 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 the Columbia R i v e r Gorge can be seen by comparing S. r of id,a l a p l a n t s from s y m p a t r i c and non-sympatrie s i t e s ( F i g . 24) . S i n c e the i n t r o g r e s s a n t §• E2£i&£±& p l a n t s a re a l s o c o m p l e t e l y i n t r o g r e s s a n t t o S. o c c i d e n t a l i s ( F i g . 25) i t i s u n c l e a r whether i n t e r m e d i a c y r e s u l t e d from p a s t h y b r i d i z a t i o n between S. r u f i d u l a and S. o c c i d e n t a l i s f o l l o w e d by t h e l o c a l e x t i n c t i o n o f S. o c c i d e n t a l i s from t h e Columbia R i v e r Gorge area o r i s t h e r e s u l t of 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 genes, e s p e c i a l l y t h o s e o f 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 . i n t o S. r u f i d u l a . The h y b r i d o r i g i n of S. r u f i d u l a i n t h e Columbia B i v e r Gorge may 80 F i g u r e 23: S c a t t e r diagram showing i n t r o g r e s s i c n o f c e r t a i n c h a r a c t e r i s t i c s between S. i n t e g r i f o l i a v a r . c l a y t o n i i f o l i a (sguares) and t e t r a p l o i d 5. o c c i d e n t a l i s v a r . d e n t a t a p l a n t s ( c i r c l e s ) . D i p l o i d S. o c c i d e n t a l i s v a r . d e n t a t a p l a n t s a r e t r i a n g l e s . 81 50" £ 25 + C 05 CL 5 + H y y + Gland Wcl.(mm) O > .6 4~.5 9 < .39 A A A A A A A A + 2 3 Petal Ln.(mm) 82 F i g u r e 24: S c a t t e r diagram showing i n t r o g r e s s i o n o f c h a r a c t e r i s t i c s between S. j n t e g r . j f o l i a v a r . c l a i + ^ i i i f o l i a (squares) and sym pa t r i e " S. r u f i d u l a ( c i r c l e s ) i n d i v i d u a l s . P l a n t s i n lower r i g h t a r e J ' S J f i d u i a p l a n t s from a nearby p o p u l a t i o n which i s not l o c a l l y s y m p a t r i c w i t h S. i n t e q r i f o l i a v a r . 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). ~ 8 3 5 1 T E U i ' 2 7 c _CTJ CL Gland Wcl.(mm) • ^.5 mm ; © .3- .49 mm O K29mm. © o 3 + 12 2.5 A A A A ^ A A A A . •3.7 Petal Ln.(mm) 84 F i g u r e 25: S c a t t e r diagram showing i n t e r m e d i a c y o f c e r t a i n c h a r a c t e r i s t i c s between 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 <sguares, EP626,Yale, B.C.) and Columbia B i v e r Gorge S. r u f i d u l a p l a n t s ( c i r c l e s ) . T r i a n g l e s r e p r e s e n t presumably n o n - i n t r o g r e s s a n t d i p l o i d S. r u f i d u l a p l a n t s from Vancouver I s l a n d . Leaf Ln.(cm) 0-2.9 0 3-4.5 O 4.6-7.6 O A O A A A + 75 125 Inf lorescence Angle (degrees) 8 6 have l e d t o a seco n d a r y resemblance between S. r u f i d u l a and S. o c c i d e n t a l i s . The more v a r i a b l e i n t r o g r e s s a n t morphs o f S» O f i d u l a a r e m o s t l y t h o s e o f h i g h e r p l o i d y l e v e l s and t h u s 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 o f the chromosome complements of s t e r i l e h y b r i d i n d i v i d u a l s . V a r i a b i l i t y w i t h i n S. o c c i d e n t a l i s v a r . d e n t a t a c a n be demonstrated t o c o r r e l a t e w i t h a h i g h e r p l o i d y l e v e l which p r o b a b l y r e f l e c t s a h y b r i d o r i g i n . T e t r a p l o i d s appear t o be c o m p l e t e l y i n t r o g r e s s a n t w i t h S. i n t e g r i f o l i a v a r . c l a y t o n i i f o l i a ( F i g . 23). S c a t t e r diagrams i n t h i s group a re perhaps l e s s u s e f u l t h a n m u l t i v a r i a t e a n a l y s e s as t o o l s t o document i n t r o g r e s s i o n s i n c e t h e y u t i l i z e a r a t h e r s m a l l number of c h a r a c t e r i s t i c s . F u r t h e r e v i d e n c e o f i n t r o g r e s s i o n i s p r o v i d e d by both P r i n c i p l e Components A n a l y s i s and Stepwise D i s c r i m i n a n t F u n c t i o n programs. The programs show t h a t t h e Columbia B i v e r Gorge S. 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 than d i p l o i d S. r u f i d u l a p o p u l a t i o n s from Vancouver I s l a n d and t h e Olympic Mountains, The Columbia R i v e r p l a n t s a l s o form i n t e r m e d i a t e s p a t i a l r e l a t i o n s h i p s between t h e S, o c c i d e n t a l i s o r S. i n t e q r i f o l i a groups and t h e Vancouver 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 i n t r o g r e s s a n t w i t h e i t h e r s y m p a t r i c S. i n t e q r i f o l i a - v a r . i n t e g r i f o l i a o r a l l o p a t r i c 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 . 87 N u m e r i c a l Studi.es I d e n t i f i c a t i o n o f major groups and p o p u l a t i o n s was p o s s i b l e u s i n g t h e h i e r a r c h i c a l c l u s t e r i n g program c a l l e d OBC-CGBOUP. R e l a t i o n s h i p s among l a r g e r groups i n t h e phenogram { F i g . 26) were s i m i l a r r e g a r d l e s s o f whether a r t i f i c i a l h y b r i d s a r e i n c l u d e d o r e x c l u d e d from the a n a l y s i s with the e x c e p t i o n t h a t a l l S. i n t e q r i f o l i a p l a n t s group t o g e t h e r a t g r e a t e r l e v e l s o f s i m i l a r i t y i n the absence of h y b r i d s . A r t i f i c i a l h y b r i d s were u s u a l l y c l u s t e r e d w i t h i n one p a r e n t a l group a l t h o u g h a few c l u s t e r e d i n d e p e n d e n t l y o r with a n o n p a r e n t a l group. S. r u f i d u l a s e p a r a t e d i n t o n o r t h e r n and Columbia R i v e r Gorge p o p u l a t i o n s w h i l e S. o c c i d e n t a l i s grouped w i t h the l a r g e r body o f S. r u f i d u l a p l a n t s f o r t h e Columbia R i v e r Gorge. D i p l o i d S. r u f i d u l a p o p u l a t i o n s c l u s t e r e d t o g e t h e r as a group w i t h p h e n e t i c s i m i l a r i t y t o t h e i r Columbia R i v e r Gorge c o u n t e r p a r t s . 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 i s v a r . d e n t a t a and §• O c c i d e n t a l i s v a r . , 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 t o each o t h e r and t o v a r i e t i e s o f S. i . n t e q r j | g l i a t h a n t c the S . r j i f i d u l a and 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 group. Perhaps t h i s i s because of the s e v e r a l c h a r a c t e r i s t i c s which J . 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 and S.„. o c c i d e n t a l i s v a r . d e n t a t a have i n common w i t h S. i n t e g r i f o l i a - such as g l a n d f e a t u r e s and i n f l o r e s c e n c e s h a p e . ; A l t e r n a t i v e l y i t may be a 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 p h e n e t i c a l l y d i s s i m i l a r groups o f OTU»s ( C r o v e l l o 1S70). More p r e c i s e measurement o f o v a r y p o s i t i o n c o u l d perhaps have i n c r e a s e d the p h e n e t i c s i m i l a r i t y between S. o c c i d e n t a l i s v a r . d e n t a t a and t h e ure 26: 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 i n d i v i d u a l s were used. P l o t i s t h e l o g o f t h e e r r o r v a l u e a t a g i v e n c l u s t e r i n g s t e p . I n d i v i d u a l s a r e c l u s t e r e d under one l i n e below an e r r o r v a l u e of 15. Be f e r e n c e t o t h e numbers i s g i v e n i n the accompanying key. V a r i e t a l names are a b b r e v i a t e d t o the f i r s t 3 o r 4 l e t t e r s . B . C = B r i t i s h C o l u m b i a , 0.N.—Olympic M o u n t a i n s , C.E.G.=Columbia R i v e r Gorge, V .I.=Vancouver I s l a n d , Hybs.=hybrids. Key t o l i n e numbers: 1. r u f i d u l a x o c c i d e n t a l i s , 2. 2 a t i £ e t i c l a t a x r j j i i d u l a 3. r u f i d u l a x d e n t a t a 4. i n t e g r i f o l i a x O l i i u i a 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. i n t e g r i f c l i a x r u f i d u l a 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 : d e n t a t a x r u f i d u l a 10. o c c i d e n t a l i s x d e n t a t a 11• r u f i d u l a x o c c i d e n t a l i s 12. o c c i d e n t a l i s x d e n t a t a 13. r u f i d u l a x o c c i d e n t a l i s 14. 2 r u f i d u l a x o c c i d e n t a l i s 15. r u f i d u l a x o c c i d e n t a l i s 16. d e n t a t a 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. o c c i d e n t a l i s : r u f i d u l a 18• o c c i d e n t a l i s 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. i n t e g r i f o l i a 23. o c c i d e n t a l i s x d e n t a t a 24. r u f i d u l a 25. r u f i d u l a 26. r u f i d u l a 27. r u f i d u l a ; . i n t e g r i f o l i a 28. r u f i d u l a 29. o c c i d e n t a l i s x d e n t a t a 30. 2 d e n t a t a " 31. ifltga£ifclla x I s J i o ^ g t a l a 32. i n t e g r i f o l i a x l e p t o p e t a l a ; l e p t c p e t a l a x d e n t a t a 33. r u f i d u l a x "(EP605) C h e h a l i s fi. 34. r u f i d u l a 35. d e n t a t a 36. r u f i d u l a x d e n t a t a 37. (EP605) C h e h a l i s 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 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 groups. R e s u l t s o f t h e h i e r a r c h i c a l g r o u p i n g a n a l y s i s o f S. o c c i d e n t a l i s - v a r . d e n t a t a and 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 e a r l y i n d i c a t e t h a t t h e s e two groups d e s e r v e s p e c i f i c s t a t u s . S a x i f r a g a in it ge,gr.j.fo ijLia -var. l e p t o p e t a l a f a i l e d t o s e p a r a t e frcm 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 . p r o b a b l y because o f t h e e x t e n s i v e numbers of i n t e r m e d i a t e Columbia R i v e r Gorge p l a n t s i n t h e sample.. I n c o n t r a s t , S . . i n t e q r i f o l i a v a r . c l a y t o n i i f o l i a p o p u l a t i o n s form a f a i r l y d i s t i n c t group from S. i n t e g r i f o l i a i n s p i t e of c l o s e c o n t a c t s i n t h e Columbia R i v e r Gorge ( F i g . 2 6 ) . The p o p u l a t i o n from C h e h a l i s , Washington (EP605) c l u s t e r e d i n d e p e n d e n t l y w i t h some a f f i n i t i e s f o r t h e group o f mixed 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 var., l e p t o p e t a l a . S. i n t e g r i f o l i a v a r . c l a y t o n i i f o l i a , and h y b r i d s . I t s morphology and chromosome number (n=29) suggest an a l l o h e x a p l o i d o r i g i n between S. o c c j , d e n t a j . j s a n d S. i n t e g r i f o l i a . The r e l a t i o n s h i p s of groups i n t h e 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 compare o n l y r o u g h l y w i t h p r e s e n t taxonomic t r e a t m e n t s (Krause and Beamish 1972,1973, H i t c h c o c k e t a l , 1961, H i t c h c o c k and C r o n g u i s t 1973, E l v a n d e r 1975) . S a x i f raga 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 and S. Occident a l l s v a r . , d e n t a t a do not appear t o c l u s t e r as good s u b s p e c i f i c e n t i t i e s 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 . The a l l o h e x a p l o i d p o p u l a t i o n from near C h e h a l i s , Washington does not group w i t h the S. o c c i d e n t a l i s o r •S. O f i d u l a i n d i v i d u a l s a l t h o u g h i t i s c o n s i d e r e d w i t h i n S. o c c i d e n t a l i s by Krause and Beamish (1972) and i s p r o b a b l y a h y b r i d i n v o l v i n g S. o c c i d e n t a l i s . The c l u s t e r i n g of t h i s 91 p o p u l a t i o n as w e l l as t h a t of a number of a r t i f i c i a l h y b r i d i n d i v i d u a l s i s s i m i l a r to t h e c l u s t e r i n g p a t t e r n d e s c r i b e d by Smith (1969) i n V a c c i n i u m where n a t u r a l h y b r i d s formed c l u s t e r s t h a t were d i s t i n c t from c l u s t e r s of t h e i r presumed p a r e n t s . The u s e f u l n e s s o f a c l u s t e r i n g t e c h n i q u e i n the p r e s e n t s t u d y appears l i m i t e d t c t h e d e f i n i t i o n of l a r g e r n a t u r a l groups on the b a s i s o f m u l t i p l e c h a r a c t e r i s t i c s . D e t e r m i n a t i o n of b i o l o g i c a l r e l a t i o n s h i p s , p h e n e t i c d i s t a n c e among t h e g r o u p s , h y b r i d a f f i n i t i e s , and c o r r e l a t i o n s t o chromosome number are a l l g u e s t i o n a b l e u s i n g t h i s t e c h n i g u e . However, t h e s u r p r i s i n g l y h i g h g r o u p i n g e r r o r l e v e l between S. o c c i d e n t a l i s v a r . d e n t a t a and 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 s t r o n g l y s u q q e s t s t h a t t h e y a r e m o r p h o l o g i c a l l y d i s t i n c t from S. o c c i d e n t a l i s v a r . g c c i d e n t a l i s and S. r u f i d u l a . The p r e s e n t s t u d y c o n f i r m e d r e p o r t s by o t h e r s ( E y l e s and B l a c k i t h 1965, R i s i n g 1968, Hhitehouse 1969, S c h i l l i n g and H e i s e r 1976) t h a t P r i n c i p a l Components A n a l y s i s (PCA) t r e a t s h y b r i d r e l a t i o n s h i p s and o t h e r f a c t o r s i n c l u d i n g c o r r e l a t i o n s w i t h c y t o l o g i c a l d a t a i n a more p r e c i s e manner t h a n s t a n d a r d h i e r a r c h i c a l c l u s t e r i n g methods. The r e c o g n i z e d taxonomic groups p r e s e n t s p a t i a l o v e r l a p s w i t h i n t e r m e d i a t e i n d i v i d u a l s among s e v e r a l groups ( F i g . 2 7 - 2 9 ) . ; I n c o n t r a s t t o a r e p e a t e d m u l t i v a r i a t e s t u d y of i n t r o g r e s s i o n which i n v o l v e s c r o s s o v e r s i n h y b r i d s and b a c k c r o s s i n g t o one parent at t h e d i p l o i d l e v e l (Bloom 1976) the i n t r o g r e s s i o n w i t h i n the p r e s e n t group appears t o i n v o l v e p r o c e s s e s o f p o l y p l o i d i z a t i o n and c o n s e q u e n t l y i n c r e a s e d v a r i a b i l i t y w i t h i n t h e p a r e n t a l e n t i t i e s . C o n s i d e r i n g t h e p r o b a b l e h y b r i d o r i g i n s of such p o l y p l o i d t a x a as 92 S. o c c i d e n t a l i s , S. j j i t e ^ K i f a l i a , 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 . and c e r t a i n p o p u l a t i o n s of S. r u f i d u l a and S« . o c c i d e n t a l i s v a r . d e n t a t a . i n t e r a e d i a c i e s r e p r e s e n t r e a s o n a b l e assessments of t h e ccmplex r e l a t i o n s h i p s i n v o l v e d . S a x i f r a j g a r u f i d u l a p o p u l a t i o n s s e p a r a t e d i n t o two c l o u d s 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 group o f d i p l o i d , t e t r a p l o i d , and h e x a p l o i d i n d i v i d u a l s from the Columbia B i v e r Gorge ( F i g . 27,28) . 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 . d e n t a t a p o p u l a t i o n s showed g r e a t e r v a r i a b l i l i t y and tended t o group i n i n t e r m e d i a t e h y p e r s p a c e s c l o s e t o S. o c c i d e n t a l i s , and S. o c c i d e n t a l i s v a r . , J=ati£etiolata, S. i n t e g r i f o l i a and t h e a l l o h e x a p l o i d 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 h y b r i d s a r e i n c l u d e d i n t h e PCA, t h e i r i n t e r m e d i a c y and v a r i a b i l i t y i s c l e a r l y r e p r e s e n t e d (see F i g . 19-21). O c c a s i o n a l s e g r e g a t e s ( F i g . 19-21, p l a n t 3*) appear d i f f e r e n t and a r e p r o b a b l y e x t r e m e l y unbalanced m o n s t r o s i t i e s which would not 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 show c l o s e r a f f i n i t i e s t o the Columbia R i v e r Gorge S. r u f i d u l a . S. o c c i d e n t a l i s or S. i n t e g r i f o l i a p a r e n t t h a n t o the o t h e r p a r e n t a l group. The raw p r i n c i p a l component ( e i g e n v e c t o r ) s o l u t i o n s ( T a b l e V, h y b r i d s excluded) a r e s i m i l a r whether or not a r t i f i c i a l h y b r i d s a r e i n c l u d e d i n t h e a n a l y s i s . Ten p a r e n t a l and h y b r i d groups d e f i n e d by h i e r a r c h i c a l g r o u p i n g and p r i n c i p a l components a n a l y s e s and s t a n d a r d taxonomic t r e a t m e n t s were r e c o g n i z e d f o r use i n t h e 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 (SDF) a n a l y s i s ( F i g . , 30-35). The s i x d i s c r i m i n a t i n g v a r i a b l e s and F v a l u e s f o r i n c l u s i o n ( c r i t i c a l F p r o b a b i l i t y 0.05), i n t h e o r d e r used, were: p l a n t h e i g h t ( 5 1 . 9 ) , 93 F i g u r e 27: P r i n c i p a l components qraph o f a x i s one vs. two showing i n d i v i d u a l s frcm s e v e r a l n a t u r a l t a x a 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 = d i p l o i d 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 Columbia B i v e r Gorge S. uiiiLaJLa D= S. o c s i d s n t a l l s v a r . deflt&ta 0= S. occj,< Jgntaiis v a r . sssiijgfltalis L= S. o c g i i g n t a l i s v a r . l a J i j a e t l o J a i a X = h e x aploid h y b r i d p o p u l a t i o n from C h e h a l i s B i v e r EP6C5 1= S. i n t e g r i f o l i a v a r . i n t e g r i f o l i a c= s. i a t s a t i i c i i a v a r . c j ^ i t s n i U e i i a . M= S. i f i t e g £ i f c l i a v a r . l&eio.pe.tala. The percentage o f 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 t h r e e axes i s 19.95, 12.17, and 6,87 p e r c e n t r e s p e c t i v e l y . Eigenvector 2 to o LO 33 3 3 33 S L 3 3 ^ -i 33 ^V3 50 33 S3 S3 S3 S3 S3 33 S3 S3 * S 3 ^ - , siV _S3 S3 ~> -J S3. -J -J m CD < , CD o-f-o o o o o o o D O O M M 7 H D M M 0 D H ° o o a o x o o o o M M M M M O O O O O o o o n o n O H o ^ o o n ° o o n o o o o o o o ^ o o_c9 o X o ^ r - O X X o o o o o c o c o o n o 95 F i g u r e 28; P r i n c i p a l components graph o f a x i s one v s . t h r e e showing i n d i v i d u a l s from s e v e r a l n a t u r a l t a x a 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 i n F i g u r e 26. 96 .3-00 £_ o +-» u (D 0 c CD L u -.3-O I I M °I I I C M M M M C C IC M C M R „ r r 1 r r r r r R r r r r P R p B R R RR R r R r r R R K R r R R R _ r R r R r R r r R r r r x I 1 1 I x ° .o ix. c 0 c X X c L D C o ib D- o i O OO 0 L C • c c D ^ C D C I n C r D D u L C D D L D M C DC C c C C D D D D D D _ D D D D D C CC D H_ 0 Eigenvector 1. .2 97 F i g u r e 29: P r i n c i p a l components graph showing i n d i v i d u a l s frcm s e v e r a l 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 ar e as i n F i g u r e 26, P l a n t *C i s t h e r i g h t i n t h i s r e p r e s e n t a t i o n . o f a x i s two vs. t h r e e n a t u r a l t a x a and not i n c l u d e d . Symbols d i s p l a c e d s l i g h t l y t o 98 00 L. o •+-> u 0) o > c <D D) Lu c*c c c c C C c c M 1 i 1 M M M M M M M I L .M r r O r X 0_ X I C I D i c R OR R R r R & R" R I B C R °R C C ° c c c ' c c D Si r 0 rXx r M n L r r r r L r r 1 r r r X r D r r -j-%r X r Oo r OO r 0 ° r O. L D O O O L r D D. D 1 D D r D L L D D , D D D D 1 Eigenvector 2 99 T a b l e V.. The raw p r i n c i p a l component s o l u t i o n . * V a r i a b l e No. J s t 2nd 3rd 4 t h 1. -0.1174 0.2560 0.0 362 0.0733 2. 0.0941 0.1533 0.0639 0.0 938 3. -0.0110 0.1943 0.1528 0.0401 4. 0.1087 0.123 8 -0.0 270 0.2 052 5. 0.2609 -0.0537 0.1897 -0.0 963 6. -0.0485 -0.0380 -0.0228 -0.1*23 7. -0.1232 0.2034 0.0256 0.2571 8. 0.1223 0.0767 -0.0233 0.0258 9. 0. 1798 -0.0644 0.3628 -0.0733 10. -0.0812 0.2692 -0.0656 0.1438 11. -0.0397 0.0204 0.1641 -0.2045 12. 0.0768 0.1146 -0.3414 0.1300 13. 0.0114 0.3009 0.0120 -0.0150 14. -0.0368 0.1845 0.0738 0.0048 15. 0.1485 0.2158 0.1051 -0.0034 16. 0.1247 0.1240 C, 10 93 0.0663 17. -0.0268 0.0197 -0.0664 -0.0991 18. 0.0922 0.1000 0.0863 -0.1326 19. 0.1828 0.1409 0.1536 0.1225 20. 0.1451 0,1982 0.1312 0.0790 21. -0.1307 0.2130 0.0324 -0. 1724 22. 0.1143 0.0221 -0.2373 -0.1960 23. 0.0214 -0.0568 -0.1147 -0.0713 24. 0.1383 0.1932 -0.2079 -0.OCC1 25. 0.2367 0.0714 -0.1214 -0.2404 26. -0.2871 0.0182 -0.0S38 -0.1099 27. 0.2956 -0.0747 -0.0520 -0.0546 28. -0.0455 0.2652 0.1701 0.1720 29. 0.0639 0.1619 -0.1869 -0.0798 30. 0.2615 -0.0067 -0.2043 -0.0916 31. 0.2743 -0.0533 -0,1184 0.1539 32. 0.1939 -0.0791 0.0 717 0.3015 33. 0.1864 0.2205 -0.1632 -0.0470 34. -0. 1018 0.2619 0.0607 -0.0 926 35. 0.G595 0.0153 -0.2388 0.1640 36. 0.2190 -0.002 5 0.2913 0.0396 37. 0.0748 0.1716 0.1181 -0.3577 38. 0. 1621 -0.1472 -0.1351 0.3517 39. -0.1733 0.2272 -0.2316 0.0896 40. 0.2593 0.0451 -0.1651 -0.1621 41. 0.1418 0.0209 0.1137 0.0500 42. -0.0074 0.0292 C.C793 0.2037 43. 0.0821 0.1830 -0.0541 -0.1629 44. 0.0821 0.1830 -0.0541 -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 of the 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 w i t h h y b r i d s (above). **• V a r i a b l e s are as coded i n Table I I . 100 leaf blade angle at petiole (23.16), gland height/width (21.09), petiole length/width (17.81), gynoecium height (15.52), and leaf length (12.76). Group c l a s s i f i c a t i o n functions show that plant height (Variable 27) e f f e c t i v e l y discriminates S. occidentalis and d i p l o i d , polyploid, and hybrid S. rufi(|ula i n d i v i d u a l s from the remaining taxa and S. i n t e g r i f o'\ja. var. c l a y t o n i i f o l i a from a l l the others.. Leaf shoulder angle (Variable 37) separates S. occidentalis var. l a t i p e t i o l a t a and S. i n t e q r i f o l i a var. IgPjopetala from the other groups while gland height/width r a t i o (Variable 6) discriminates d i p l o i d S. r u f i d u l a , S. occidentalis var. l a t i p e t i o l a t a t and S. occidentalis var. dentata groups. Petiole length/width r a t i o (Variable 39) distinguishes S. i n t e q r i f o l i a var. c l a y t o n i i f o l i a and S« i n t e g r i f o l i a var. leptopetala and gynoecium height (Variable 10) separates S. o c c i d e n t a l i s and the S. occidentalis hexaploid hybrid population. Leaf length (Variable 32) separates both groups of S. r u f i d u l a from the remaining group. Of the t o t a l cases, 65.79% were c o r r e c t l y c l a s s i f i e d by SDF analysis including over 65% of 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 , J . 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 . S. o c c i d e n t a l i s var. 2aii£§M2l3iS 31) , 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 , and Vancouver Island and Olympic Mountains S. r u f i d u l a i n d i v i d u a l s . Some 63.7% of the S. j l n t e g r i f o l i a var. leptopetala plants were grouped with S. i n t e g r i f o l i a (Fig. 31,B), while i n d i v i d u a l s of the allohexaploid population (FP605) were scattered among S. oc c i d e n t a l i s (20%), t e t r a p l o i d or hybrid S. r u f i d u l a (30%), and d i p l o i d S. ru f i d j i l a (20%) groups (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 of t h e f i r s t t h r e e c a n o n i c a l v a r i a b l e s . The f i r s t two c a n o n i c a l 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 produce a s t r a t i f i e d t h r e e - d i m e n s i o n a l p l o t which c l a r i f i e s 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 . T h i s r e p r e s e n t a t i o n d e p i c t s r e l a t i o n s h i p s of group means as s p h e r e s on s u p p o r t i n g l i n e s . Group means symbols are as f o l l o w s : A= S. r u f i d u l a p l a n t s from t h e Columbia R i v e r Gorge and h y b r i d s B - S. i n t e g r i f o l i a Columbia R i v e r Gorge O S. i n t e q r i f c l i a D= S. o c c i d e n t a l i s L= S. o c c i d e n t a l i s M = S. i n t e g r i f o l i a 0= S. o c c i d e n t a l i s B = J« r u f i d u l a p l a n t s from Vancouver I s l a n d and the Olympic Mountains I = S. i n t e g r i f o l i a v a r . i n t e g r i f o l i a p l a n t s frcm Vancouver I s l a n d X=Hexaploid 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 p l a n t s from C h e h a l i s , Hash. v a r . i n t e g r i f o l i a p l a n t s from the va r . c l a j f t o n i i f o l i a v a r . d e n t a t a v a r . l3ti£etiolata v a r . l e n t o p e t a l a v a r . o c c i d e n t a l i s 103 32,A). Of S. o c c i d e n t a l i s v a r . d e n t a t a i n d i v i d u a l s , 28% were p l a c e d i n t o t he S. i n t e q r i f o l i a group ( F i g . 33,A). Columbia B i v e r Gorge S. i n t e g r i f o l i a p l a n t s ( F i q . 34,A) d e m c c s t r a t e i n t e r m e d i a c i e s w i t h S. i n t e q r i f o l i a v a r . c l a v t o n i i f o l i a (19%) ( F i q . 34,B) but p r o b a b l y t h e y have i n s i q n i f i c a n t d i f f e r e n c e s f r c m t h e Vancouver I s l a n 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 qroup w i t h which 12% a r e c l a s s i f i e d . S e v e r a l Columbia B i v e r Gorge 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 a r t i f i c i a l h y b r i d s ( F i g . 35,B) a r e i n t e r m e d i a t e between S. i n t e q r i f o l i a v a r . i n t e g r i f o l i a ( 5 % ) , S. o c c i d e n t a l i s ( 1 2 . 5 % ) . S. r u f i d u l a (10%) and S. o c c i d e n t a l i s v a r . d e n t a t a ( 5 % ) . Such i n t e r m e d i a c i e s and i n c r e a s e d v a r i a b i l i t y i n SDF a n a l y s i s r e l a t i o n s h i p s have been used as 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 i n n a t u r a l p o p u l a t i o n s ( S c h u e l e r and R i s i n g 1976). I t has been s u g g e s t e d t h a t SDF a n a l y s i s i s a p r e f e r r e d method f o r s e p a r a t i o n o f p a r e n t a l s p e c i e s g r o u p s , e s p e c i a l l y where they 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 means (Danick and Burns 1975, Namkoong 1966). The p r o b a b l e h y b r i d n a t u r e 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 v a r . d e n t a t a p l a n t s , Columbia B i v e r Gorge S. r u f i d u l a groups ( F i g . 35,B), and a h e x a p l o i d p o p u l a t i o n from C h e h a l i s , Washington i s documented by SDF a n a l y s i s as i s t h e uniqueness o f 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 morphology. Taken i n c o n j u n c t i o n w i t h r e s u l t s o f t h e 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 program, the SDF program r e s u l t s agree f a v o r a b l y w i t h a trea t m e n t which c l a s s i f i e s S. r u f i d u l a . S. o c c i dent a11s v a r . lsii£§ij=2JsSta, and S. o c c i d e n t a l i s 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 from S. o c c i d e n t a l i s . A c c o r d i n g t o t h e r u l e s of b o t a n i c a l nomenclature 104 F i g u r e 31: 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 t h r e e c a n o n i c a l v a r i a b l e s . The f i r s t two c a n o n i c a l 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 produce a s t r a t i f i e d t h r e e - d i m e n s i o n a l p l o t which c l a r i f i e s 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 are shown as i n the 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 p o i n t s f o r the S. i n t e g r i f o l i a v a r . c l a y t o n j i f g l j a c l o u d as t r i a n g u l a r symbols and P a r t B shows s i m i l a r h y perspace 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 v a r . l e p t o p e t a l a c l o u d . Symbols are as i n F i g u r e 29. 105 106 F i g u r e 32: St 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 t h r e e c a n o n i c a l v a r i a b l e s . The f i r s t two c a n o n i c a l 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 produce a s t r a t i f i e d t h r e e - d i m e n s i o n a l p l o t which c l a r i f i e s 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 are shown as i n the 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 p o i n t s f o r the S. 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 ( C h e h a l i s R i v e r , EP605) c l o u d 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 as i n F i g u r e 29. 5 G tr Canonical Variable 2 108 r e 33: 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 the f i r s t t h r e e c a n o n i c a l v a r i a b l e s . The f i r s t two c a n o n i c a l 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 produce a s t r a t i f i e d t h r e e - d i m e n s i o n a l p l o t w hich c l a r i f i e s 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 . Group means are shown as i n the 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 p o i n t s f o r the S. o c c i d e n t a l i s v a r . d e n t a t a c l o u d as t r i a n g u l a r symbols and P a r t B shows s i m i l a r hyperspace p o i n t s f o r the 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 c l c u d , Symbcls are as i n F i g u r e 29. Canonical Variable 2 6br 110 F i g u r e 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 t h r e e c a n o n i c a l v a r i a b l e s . The f i r s t two c a n o n i c a l 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 produce a s t r a t i f i e d t h r e e - d i t i e n s i c n a l p l o t which c l a r i f i e s 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 ar e shown as 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 p o i n t s f o r t h e §. i n t e g r t f g l i a v a r . i n t e g r i f o l i a c l o u d frcm the Columbia R i v e r Gorge as t r i a n g u l a r symbols and P a r t B shows s i m i l a r hyperspace p o i n t s f o r the S. i n t e g r i f o l j . a v ar. i n t e g r i f o l i a c l o u d from Vancouver I s l a n d and the B.C. M a i n l a n d . Symbols are as i n F i g u r e 29. i n 112 F i g u r e 35: 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 of t h e f i r s t t h r e e c a n o n i c a l v a r i a b l e s . The f i r s t two c a n o n i c a l 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 produce a s t r a t i f i e d t h r e e - d i m e n s i o n a l p l o t which c l a r i f i e s 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 as 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 p o i n t s f o r the S. r u f i d u l a c l o u d from Vancouver I s l a n d and the Olympic Mountains as t r i a n g u l a r symbols and P a r t B shows s i m i l a r hyperspace p o i n t s f o r the S. r u f i d u l a c l o u d from the Columbia B i v e r Gorge and t h e S. r u f i d u l a h y b r i d c l o u d . Symbols are as i n F i g u r e 29. C a n o n i c a l V a r i a b l e 2 114 S. r u f i d u l a becomes 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 v a r . l a t i p e t i o l a t a 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 v a r . d e n t a t a becomes S. g o r m a n i i . and S. .. o c c i d e n t a l i s becomes 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 i n Taxonomy s e c t i o n ) . B r e e d i n g System O b s e r v a t i o n s Baacfing, T e s t s The r e s u l t s o f t e s t s f o r a u t o m a t i c s e l f i n g and s e l f i n c o m p a t i b i l i t y a r e p r e s e n t e d i n T a b l e V I , Seed s e t from 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, u n t r e a t e d c o n d i t i o n i s f r e q u e n t l y h i g h e r than seed s e t i n the h a n d - s e l f e d c o n d i t i o n . T h i s p r o b a b l y r e s u l t e d from damage i n h a n d l i n g e s p e c i a l l y d u r i n g 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 p o l l i n a t i o n s t o c o i n c i d e w i t h s t i g m a t i c r e c e p t i v i t y . A l l t a x a are c l e a r l y s e l f - c o m p a t i b l e and the r e l a t i v e s o f S. r u f i d u l a , e.g. S. o c c i d e n t a l i s v a r . d e n t a t a and S. o c c i d e n t a l i s as w e l l as S. i n t e q r i f o l i a v a r . l e p t o p e t a l a may have a h i g h e r degree o f p o t e n t i a l s e l f - f e r t i l i z a t i o n t han S, i n t e j r i J a l i a v a r , 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 • v a r . c l a y t o n i i f o l i a . o r S. o c c i d e n t a l ' s v a r , l a t i p e t i o l a t a . E l v a n d e r ( p e r s o n a l communication) r e p o r t s t h a t he o b t a i n e d no seed s e t f o r members of t h e S. i n t e q r i f o l j a -S. oregana complex i n bagged, u n t r e a t e d c o n d i t i o n s but s i m i l a r seed s e t i n bagged h a n d - s e l f e d t e s t s t o those r e p o r t e d here f o r S. i n t e g r i f o l i a and i t s r e l a t i v e s . D i f f e r e n c e s i n t h e r e s u l t s c i t e d here are p o s s i b l y due t o the bagging method used i n t h i s 115 Ta b l e VIj, •, Bagging t e s t s f o r s e l f i n g . S e l f e d by. Hand Ro. T o t a l B r u i t s Seed ( P l a n t s ) A b b r e v i a t e d Taxa r u f . ( d i p l . ) r u f . ( p o l y . ) d e n t , ( d i p l . ) d e nt, { t e t r . ) o c c . ( t e t r . ) 57 (6) 33 (5) 82(4) 35(3) 40(3) l a t i . 112(3) (n=38) i n t . ( t e t r . ) c l a y , ( d i p l . ) l e p t . ( t e t r . ) 13(2) 67(3) 8(1) % Good* Seed 1402 646 1313 1312 1719 1087 646 603 37 55.2 25. 1 55.9 59.1 27.9 6.3 13.9 16.9 21.6 Bagged U n t r e a t e d No. T o t a l F r u i t s Seed ( P l a n t s ) 19(3) 4(1) 7(1) 11(1) 27(3) 59(2) 5(2) 20(1) 37(3) % Good Seed 990 230 228 254 1516 530 407 179 470 48. 7.8 0.0 76.4 48.9 5.8 0.4 15. 1 35.5 * "Good" seed d e f i n e d as l a r g e , s w o l l e n seed v e r s u s s m a l l , s h r i v e l e d seed which 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 seed. 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 and a p o m i x i s . O u t c r o s s e d Ml l a n d O u t c r o s s i n g C c n t r c l . O n t r e a t e d Emasculated Bagged No. T o t a l % Good* No., T o t a l % Good* No. T o t a l % Good F r u i t s Seed Seed F r u i t s Seed Seed F r u i t s Seed Seed ( P l a n t s ) ( P l a n t s ) ( P l a n t s ) a b b r e v i a t e d Taxa r u f . 10(2) 505 59.6 315(8) 7698 66.3 ( d i p l . ) r u f . , 20(3) 932 56. 1 103(5) 4054 38.9 ( p c l y . ) dent. 17(1) 206 33.5 355(5) 3304 69.4 ( d i p l . ) dent. 17(1) 675 71.0 61 (4) 2253 63.3 ( t e t r . ) o c c , 21(2) 1210 .6-2.5 41 (3) 2767 45.5 ( t e t r . ) l a t i . 52(2) 707 39.6 (n=38) i n t . ./ ( t e t r . ) 6(1) 348 0.0 35(5) 1852 75.1 401 0.0 c l a y . 22 (2) 1271 74.3 41 (1) ( d i p l . ) l e p t . 8(4) 681 82. 5 12(1) 544 1.3 ( t e t r . ) 10 (2) 608 0.0 21(4) 621 0.5 6 (1) 149 0.0 47(7) 2554 0.2 15(2) 676 0.0 30(6) 1691 0.0 32(2) 410 O.O 44(6) 1 175 0.3 * Gocd seed d e f i n e d i n Ta b l e V I . 1 1 7 s t u d y which may have a l l o w e d a u t o d e p o s i t i o n as a r e s u l t o f mec h a n i c a l j a r r i n g of 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 (Table VII) was u s u a l l y h i g h e r i n J3. i n t e g r i f o l i a r e l a t i v e s , or n e a r l y e g u a l i n S. o c c i d e n t a l i s r e l a t i v e s , t o seed s e t i n the s e l f i n g t e s t s . In a few c a s e s t h e o u t c r o s s e d seed s e t was lo w e r t h a n i n s e l f i n g t e s t s . T e r - A v a n e s i a n (1978) has r e p o r t e d t h a t i n s e v e r a l d i f f e r e n t f l o w e r i n g p l a n t g r o u p s , low numbers o f p o l l e n g r a i n s d e p o s i t e d 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 g e n e r a l f a i l u r e o f f e r t i l i z a t i o n . T h i s 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 to s e t seed where e f f o r t s t o d e p o s i t l a r g e amounts of p o l l e n were u n s u c c e s s f u l . 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 . i a t i£ e t i o 1 a t a , S. i n t e g r i f o l i a v a r . , c l a y t o n i i f o l i a . and S. i n t e q r i f o l i a v a r . l e p t o p e t a l a f a i l e d t o s e t l a r g e amounts of seed when exposed t c p o l l i n a t o r s , perhaps as a r e s u l t of premature h a r v e s t i n g or i n f e r t i l i t y i n those p a r t i c u l a r i n d i v i d u a l s . I n a l l c a s e s seed s e t was a b s e n t or a t low background l e v e l s ( l e s s than 1.0%) i n em 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 ) . A l t h o u g h a p o m i x i s which r e g u i r e s p o l l i n a t i o n i s p o s s i b l e , the wide range o f v a r i a b i l i t y o f o f f s p r i n g from i n d i v i d u a l p l a n t s and t h e f a c t t h a t h y b r i d i z a t i o n s produce o f f s p r i n g which d i f f e r markedly from the femal e p a r e n t argues a g a i n s t i t s o c c u r r e n c e at l e a s t as a major means of seed p r o d u c t i o n . 118 f l o j E i l i O b s e r v a t i o n s Nectar p r o d u c t i o n b e g i n s w i t h a n t h e r d e h i s c e n c e and c o n t i n u e s f o r s e v e r a l days u n t i l the a p p a r e n t l y r e c e p t i v e , p a p i l l a t e , s t i g m a t i c s u r f a c e l o s e s i t s c h a r a c t e r i s t i c wetness. Some UV-absorbing and r e - e m i t t i n g s u b stances a r e p r e s e n t i n S a x i f r a g a n e c t a r which may be v i s u a l s t i m u l i t o f o r a g i n g bees o r f l i e s . S a x i f r a g a f l o w e r s are c h a r a c t e r i s t i c a l l y p r o t a n d r c u s but t h e r e i s an o v e r l a p i n f i n a l a n t h e r d e h i s c e n c e and o n s e t of s t i g m a t i c 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 s e p a r a t i o n e x i s t s between d e h i s c e d a n t h e r s and s t i g m a s ( F i g . , 2 , 3 ) , f i l a m e n t and s t y l e e l o n g a t i o n and c u r v a t u r e may b r i n g the a n t h e r s and s t i g m a s i n t o c l o s e p r o x i m i t y so t h a t a u t o - d e p c s i t i o n i s p o s s i b l e p r o v i d e d t h a t p o l l e n has not been p r e v i o u s l y removed by f o r a g i n g i n s e c t s and t h a t s l i g h t e x t e r n a l f o r c e s b r i n g t h e a n t h e r and s t i g m a i n t o c o n t a c t . There i s a l s o t h e p o s s i b i l i t y t h a t p o l l e n may f l o a t a c r o s s a f i l m of water from a n t h e r t o s t i g m a , as o b s e r v e d f o r o t h e r p l a n t s by Hagerup (1950, 1951) or be blown about by wind (Hyde 1950,1969, Hyde and W i l l i a m s 1961, P r o c t o r and Yeo 1973) , e s p e c i a l l y s i n c e p o l l e n was o b s e r v e d t o l o s e i t s s e l f - a d h e r e n t p r o p e r t i e s a f t e r p r o l o n g e d c o n t a c t w i t h a i r . No t e s t s f o r w i n d - p o l l i n a t i o n o r r e p e a t e d t e s t s f o r water-mediated s e l f i n g were c a r r i e d out i n t h e p r e s e n t study. However, c a s u a l o b s e r v a t i o n s demonstrate t h a t f r e s h S a x i f r a g a p o l l e n i s h ydrophobic and d r o p l e t s from a l i g h t m i s t i n g can accumulate i n S a x i f r a g a f l o w e r s p r o d u c i n g a c o n t i n u o u s water s u r f a c e between a n t h e r s and s t i g m a on which r a f t s of p o l l e n were observed t o f l o a t and a p p a r e n t l y c o n t a c t the s t i g m a . A l s o a r e d u c t i o n was 119 noted i n t h e adherence o f p o l l e n g r a i n s t o each o t h e r and t o t h e t a p e t a l w a l l a f t e r p r o l o n g e d exposure t o the atmosphere. Such a r e d u c t i o n i n o i l y adherence may enhance t h e chances of l i m i t e d anemophily f o r t h e s e open, d i s h - t y p e f l o w e r s i n t h e absence o f a n i m a l p o l l e n v e c t o r s . Temporal s e p a r a t i o n o f f l o w e r i n g t i m e s i s e v i d e n t between s y m p a t r i c t a x a . However, i n most c a s e s t h e r e i s a s h o r t p e r i o d of 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 p o l l i n a t i o n s a re p o s s i b l e . No such p o l l i n a t i o n s were r e c o r d e d d u r i n g p e r i o d s o f i n s e c t o b s e r v a t i o n but t h e presence of h y b r i d i n d i v i d u a l s t e s t i f i e s t o t h e o c c a s i o n a l o c c u r r e n c e of h y b r i d i z i n g e v e n t s . Whether t h e r e a r e p h e n o l o g i c a l d i f f e r e n c e s c o r r e l a t e d w i t h d i f f e r e n t 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 as Lewis and Suda (1976) have d e s c r i b e d f o r C l a y t o n i a v i r q i n i c a i s u n c l e a r from t h e p r e s e n t samples. D e t e r m i n a t i o n o f such d i f f e r e n c e s would r e q u i r e more e x t e n s i v e and t e m p o r a l l y s e p a r a t e d s a n p l i n q of i n d i v i d u a l p o p u l a t i o n s which e x h i b i t two or more l e v e l s o f p o l y p l o i d y . I t i s i n t e r e s t i n q t o n o t e t h a t where d i f f e r e n t d i p l o i d and p o l y p l o i d t a x a occur s y m p a t r i c a l l y , the p o l y p l o i d s tend t o f l o w e r l a t e r i n t h e season than t h e d i p l o i d s . S e a s o n a l i s o l a t i o n a l s o appears t o o c c u r t o some e x t e n t w i t h i n t a x a over broad q e o g r a p h i c o r t o p o g r a p h i c r a n g e s . t Southern l o w l a n d 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 t h o s e a t h i g h e r l a t i t u d e s or a l t i t u d e s , even under u n i f o r m t r a n s p l a n t c o n d i t i o n s . The number of o v u l e s per c a r p e l v a r i e s c o n s i d e r a b l y w i t h i n an i n f l o r e s c e n c e . The e a r l i e r - m a t u r i n g f l o w e r s which occupy p o s i t i o n s c l o s e r t o t h e c e n t r a l branches produce more o v u l e s than the l a t e r m a t u r i n g ones which a r e a t p r o g r e s s i v e l y more 120 t e r m i n a l p o s i t i o n s on t h e i n f l o r e s c e n c e branches. Some d i f f e r e n c e s i n o v u l e / c a r p e l number among t a x a e x i s t . When the e a r l i e s t , most c e n t r a l f l o w e r s a r e compared, S. o c c i d e n t a l i s v a r . d e n t a t a . 5. r u f i d u l a . S. i n t e g r i f o l i a v a r . i n t e q r i f o l i a and 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 p l a n t s form a s e r i e s f r c m l o w e s t t o h i g h e r numbers of o v u l e s per c a r p e l w i t h the h i g h e s t numbers a p p e a r i n g i n S. i n t e g r i f o l i a v a r . l e p t o p e t a l a . S. i n t e g r i f o l i a v a r . 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 i n t h a t o r d e r . ,• These t r e n d s do not seem t o be c o r r e l a t e d w i t h i n f l o r e s e n c e s i z e , e x t e n t o f b r a n c h i n g , o r f l o w e r number. Al t h o u g h d e t a i l e d o b s e r v a t i o n s a r e l a c k i n g , o t h e r f l o r a l f e a t u r e s do n o t seem t o show s u c h a marked r e d u c t i o n i n numbers or s i z e w i t h p o s i t i o n i n the i n f l o r e s c e n c e . F u r t h e r s t u d i e s a r e needed to d e t e r m i n e t h e s y s t e m a t i c and b i o l o g i c a l s i g n i f i c a n c e of t h e s e 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 number. E l v a n d e r C1978) has o b s e r v e d t h a t some p o p u l a t i o n s o f S. i n t e g r i f o l i a v a r . i n t e q r i f o l i a a r e g y n o d i o e c i o u s , an o b s e r v a t i o n which i s s u p p o r t e d by t h e f i n d i n g of m a l e - s t e r i l e p l a n t s , a p p a r e n t l y s e e d - f e r t i l e on Vancouver I s l a n d . He has noted a p o s i t i v e c o r r e l a t i o n between the degree of c o m p a c t i o n of t h e i n f l o r e s c e n c e and t h e degree of f l o w e r i n g synchrony among v a r i e t i e s o f 5. i n t e g r i f o l i a . He concluded t h a t i n f l o r e s c e n c e compaction and a s s o c i a t e d f l o w e r i n g s y n c h r o n y promotes o u t c r o s s i n g i n p o p u l a t i o n s w i t h compacted or c o n g e s t e d i n f l o r e s c e n c e s . E v i d e n t l y S. i n t e g r i f s l i a v a r . , i n t e g r i f o l i a p o p u l a t i o n s w i t h l e s s compacted i n f l o r e s c e n c e s a r e t h e ones which a r e g y n o d i o e c i o u s . E v i d e n c e of m a l e - s t e r i l e i n d i v i d u a l s and e v i d e n c e f o r s i m i l a r c o r r e l a t i o n s i n v o l v i n g degree o f 121 compaction and gynodioecy i n the t a x a o t h e r than S. i n t e g r i f o l i a i s l a c k i n g from the p r e s e n t s t u d y . The r e a s o n s f o r complete seed and p o l l e n s t e r i l i t y i n some i n d i v i d u a l s a r e u n c l e a r , but one p o s s i b i l i t y i s t h a t t h e y r e p r e s e n t v e g e t a t i v e l y v i g o r o u s s e g r e g a n t h y b r i d o r mutant forms which c l o s e l y resemble one o r t h e o t h e r p a r e n t i n morphology and h a b i t a t p r e f e r e n c e , Another e x p l a n a t i o n may i n v o l v e the degree of i n s t a b i l i t y 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 g e o l o g i c a l l y r e c e n t o r i g i n s . I n c e r t a i n s t e r i l e i n d i v i d u a l s what appear t o be m y c e l i a l fragments may be seen among the a b o r t i v e p o l l e n o r p o l l e n mother c e l l s . Fungus-induced s t e r i l i t y may ind e e d 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 v e g e t a t i v e l y v i g o r o u s , s t e r i l e i n d i v i d u a l s may compose a l a r g e p o r t i o n o f c e r t a i n p o p u l a t i o n s (Beamish 1961). Although t h e bagging t e s t s and f l o r a l o b s e r v a t i o n s r e v e a l i n t e r e s t i n g b i o l o g i c a l 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 u s e f u l i n u n c o v e r i n g c h a r a c t e r i s t i c s t o s e p a r a t e the c l o s e l y r e l a t e d taxonomic e n t i t i e s i n v o l v e d i n t h i s s t u d y , . Geoggaghy• H a b i t a t And P o l l i n a t i o n Ecology O b s e r v a t i o n s 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 v a r . o c c i d e n t a l ' s i s a widespread e n t i t y which i s found on mountaintops and c l i f f s i d e s i n t h e Bocky Mountains and Cascades from s o u t h e r n A l a s k a and t h e Yukon t o Montana and Idaho. I t grows on exposed mossy or b a r e grcund i n ro c k c r e v i c e s and cn r o c k y peaks i n v e r n a l l y mesic h a b i t a t s which become r a p i d l y x e r i c as t h e season p r o g r e s s e s . I t i s not found on Vancouver I s l a n d , t h e Olympic Mountains o r i n 122 the Columbia B i v e r Gorge. S a x i f r a g a r u f j u l u l a grows commonly on l o s l a n d c l i f f s and o t h e r s i t e s where e a r l y s p r i n g r u n o f f i s abundant. I t i s found from Lane County i n Oregon n o r t h i n t o t h e Columbia B i v e r Gorge and then i n t h e Olympic Mountains and northward on the e a s t s i d e o f Vancouver I s l a n d . I t o c c u r s on mountaintop a r e a s i n t h e Olympic Mountains and on Vancouver I s l a n d as w e l l as at l o w e r e l e v a t i o n s . L o c a l p o p u l a t i o n e x t i n c t i o n caused by i n c r e a s e d g r a z i n g p r e s s u r e w i t h the r e c e n t i n t r o d u c t i o n o f mountain g o a t s may e x p l a i n f a i l u r e to f i n d S. r u f i d u l a p l a n t s which have been r e p o r t e d on H u r r i c a n e Ridge and i n the O b s t r u c t i o n Pass Area o f the Olympic Mountains. V e r b a l 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 on t h e B r i t i s h Columbia mainland were a l s o i n v e s t i g a t e d but o n l y one o c c u r r e n c e has been documented. L i t e r a t u r e r e p o r t s (Welsh 1974) of 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 w i t h 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 o r S. n i v a l i s s i n c e t h e r e a r e no specimens o f §• 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 . r u f i d u l a i n the c o l l e c t i o n s a t the U n i v e r s i t y o f A l a s k a , t h e S t a t e Museum a t Juneau, i n t h e Anderson c o l l e c t i o n a t Ames, Iowa, o r among Welsh's specimens from BYU. The specimen r e p o r t e d f o r s o u t h e r n A l a s k a by E n g l e r and I r m s c h e r (1916) i s a l s o p r o b a b l y a s m a l l 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 ' s p l a n t s i n c e i t was f i r s t l a b e l e d a s S. y j . r a l l i e n s i s , a l t h o u g h i t was not seen d u r i n g t h e p r e s e n t s t u d y . The h a b i t a t s o f S. o c c i d e n t a l i s v ar. d e n t a t a a re s i m i l a r t c t h o s e o f S. r u f i d u l a but S. o c c i d e n t a l i s v a r . d e n t a t a i s found west o f t h e Columbia R i v e r Gorge i n the Columbia R i v e r and W i l l a m e t t e R i v e r l o w l a n d s and h i g h e r i n t o the C o a s t Range o f 123 Oregon. I t i s commonly found on s h a l l o w s o i l o f v o l c a n i c r o c k f a c e s i n open o r p a r t l y shaded seepage a r e a s where e a r l y s p r i n g m o i s t u r e i s abundant. I t s e x t e n s i v e rhizome system a p p a r e n t l y a l l o w s i t to o b t a i n 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 and propagate by v e g e t a t i v e means. I t i s i n t e r e s t i n g t h a t Chambers ( p e r s o n a l communication) has c o l l e c t e d m a t e r i a l from a sunny l o c a t i o n on Saddle Mountain which c l o s e l y resembles §* r u f i d u l a w h i l e o t h e r p l a n t s i n shady a r e a s from t h e same l o c a l i t y a r e n e a r e r t o S. o c c i d e n t a l i s v a r . d e n t a t a . , E x a m i n a t i o n o f p l a n t s from open, sunny a r e a s a t t h e same l o c a l i t y i n t h e p r e s e n t s t u d y i n d i c a t e s t h a t t h e s n a l l e r , h e l i o p h i l i c forms a r e d i p l o i d S. o c c i d e n t a l i s v a r . d e n t a t a 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 r e v e a l t e t r a p l o i d o r h e x a p l o i d S. r u f i d u l a p l a n t s i n t h e v i c i n i t y . S a x i f r a g a 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 i s endemic t c a few i s o l a t e d v o l c a n i c p l u g s and mcuntaintop b a l d s o f C l a t s o p and T i l l a m o o k C o u n t i e s , Oregon. I t grows i n open, r a t h e r g r a s s y a r e a s i n somewhat deeper and perhaps d r i e r s o i l . I t blooms l a t e r i n t h e season t h a n S. o c c i d e n t a l i s var, d e n t a t a which i s s y m p a t r i c w i t h i t on Saddle Mountain and p r o b a b l y a t o t h e r s i t e s as w e l l . , The v a r i e t i e s o f S. i n t e q r i f o l i a a r e g e o g r a p h i c a l l y s e g r e g a t e d over much o f t h e i r r a n g e , but i n the Columbia S i v e r Gorge they o f t e n are found growing t o g e t h e r i n mixed p o p u l a t i o n s . S e a s o n a l d i f f e r e n c e s i n f l o w e r i n g t i m e s and p o s s i b l e m i c r o h a b i t a t d i f f e r e n c e s a p p a r e n t l y i s o l a t e the v a r i e t i e s which a r e s y m p a t r i c a l t h o u g h t h e r e i s e v i d e n c e t h a t h y b r i d i z a t i o n and i n t r o g r e s s i o n are common i n t h e Columbia E i v e r 124 Gorge. S a x i f r a g a i n t e g r i f o l i a v a r . i n t e g r i f o l i a o c c u r s on open g r a s s y s o i l p o c k e t s and l e d g e s of r o c k y headlands and o u t c r o p s from t h e Upper f r a s e r V a l l e y and C e n t r a l Vancouver I s l a n d s o u t h t h r o u g h the Puget Sound and then e a s t i n t o the Columbia B i v e r Gorge and southward as f a r as s o u t h c e n t r a l Oregon. S. i n t e g r i f o l i a v a r . l e p t c p e t a l a i s commonly found i n open r o c k y p l a c e s or g r a s s y savannah beneath P i n u s ccnderosa i n the B r i t i s h Columbia i n t e r i o r southward t o Montana, I d a h o , and Oregon and west as f a r as t h e e a s t e r n p o r t i o n of the Columbia B i v e r Gorge. 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 o c c u r s i n s h a l l o w and o f t e n g r a v e l l y or d i s t u r b e d s o i l a l o n g r o a d c u t s and strearabanks as w e l l as on l e d g e s of r o c k y o u t c r o p s and c l i f f s i d e s from t h e Washington Cascades s o u t h t o t h e Columbia B i v e r Gorge and from n o r t h e a s t Oregon and western Idaho west and s o u t h t o N o r t h e r n C a l i f o r n i a . O b s e r v a t i o n s on s o i l depth i n d i c a t e a s i g n i f i c a n t d i f f e r e n c e between the S. r u f i d u l a o r S, o c c i d e n t a l i s p l a n t s and t h e l o c a l l y s y m p a t r i c v a r i e t i e s of S. i n t e g r j l f o l j a ( T a b l e V I I I ) . D i f f e r e n c e s i n s o i l depth p r e f e r e n c e s f o r one l a r g e r sample of s y m p a t r i c S. i n t e g r i f o l i a v a r . c l a y t o n i i f o l i a and S. / i n t e g r i f o l i a • • v a r . i n t e g r i f o l i a p l a n t s from t h e Columbia B i v e r Gorge were l e s s o b v i o u s . S i m i l a r l y no c l e a r d i f f e r e n c e s i n depth of s o i l t o r o c k y s u b s t r a t e emerge between n o r t h e r n and s o u t h e r n S. r u f i d u l a p o p u l a t i o n 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 • may have a deeper s o i l depth p r e f e r e n c e than does S. 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 a r e n e c e s s a r y . N a t u r a l l y o c c u r r i n g h y b r i d s between S. r u f i d u l a and S. i n t e g r i f o l i a were sampled a t one l o c a l i t y and average s o i l depth i s i n t e r m e d i a t e 125 T a b l e V i l l i Average s o i l depth (cm) beneath 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 from l o c a l i t i e s i n t h e Columbia B i v e r Gorge and B r i t i s h C olumbia. r u f i d u l a i£tejrilglia No. . Ave. No. , Ave. IfOca 1 i t i e s H a n t s Depth P l a n t s Depth Columbia B i v e r Gorge T r o u t d a l e 10 4.3 10 4.9 M t . P l e a s a n t 9 3.6 9 6.7 C l a r k Co. L i n e 9 3.3 10 9.6 H o s i e r 10 4.4 10 8.6 Mayer Park 10 4.9 10 14.0 The D a l l e s 10 4.5 20 11.6* B r i t i s h Columbia H t . F i n l a y s o n 15 6.2 15 10.7 Sooke 15 3.3 15 6.5 Nanaimo** 15 3.6 15 6.5 Nanoose 15 3.1 15 5.7 * A subsample o f i n t e g r i f o l i a v a r . i n t e g r i f o l i a (n=20) from t h i s l o c a l i t y had an average depth of 14.9cm. Other Columbia B i v e r Gorge 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 a r e v a r . i n t e g r i f o l i a . ** An i n t e r m e d i a t e h y b r i d subsample (n=15) from t h i s s i t e had an average s o i l depth o f 5.5 cm. 126 T a b l e I J U S o i l m o i s t u r e i n g. H20(g. d r y w e i g h t ) - 1 under p l a n t s a t f i v e s y m p a t r i c l o c a l i t i e s . S i t e Date r u f i d u l a i n t e g r i f o l i a Nanoose March 1.039* .565 A p r i l .801 .675* May .514 .216 Nanaimo March 1.117 .487 May 1.014* .517* Ht. F i n l a y s o n March .995* .638 Sooke March .752* .725 o c c i d e n t a l i s i n t e g r i f o l i a l a l e A p r i l .614* ~* .426 * These d a t e s c l o s e s t t c h e i g h t of bloom f o r t h a t p l a n t taxon a t each l o c a l i t y . , 127 f o r the h y b r i d p l a n t s . A few s y m p a t r i c l o c a t i o n s were sampled f o r s o i l m o i s t u r e t o document d i f f e r e n c e s i n water r e l a t i o n s between S. r u f i d u l a and S. i n t e q r i f o l i a and between S. o c c i d e n t a l i s and S. i n t e g r i f o l i a (Table I X ) . E s p e c i a l l y d u r i n g peak f l o w e r i n g t i m e s t h e s o i l beneath S. r u f i d u l a p l a n t s c o n t a i n s more m o i s t u r e than s o i l beneath S. i n t e g r i f o l i a p l a n t s . I f f i e l d g e r m i n a t i o n o c c u r s i n about A p r i l o r May, as i t does f o r s e e d l i n g s i n c u l t i v a t i o n under n a t u r a l t e m p e r a t u r e and l i g h t c o n d i t i o n s , s i m i l a r d i f f e r e n c e s i n s o i l m o i s t u r e l e v e l s may be i m p o r t a n t f a c t o r s i n h a b i t a t s e l e c t i o n a t the s e e d l i n g e s t a b l i s h m e n t phase. Such d i f f e r e n c e s i n water r e l a t i o n s h e l p t o e x p l a i n h i g h e r observed m o r t a l i t y r a t e s i n t r a n s p l a n t e d S. r u f i d u l a specimens, assuming t h a t t h e a r t i f i c i a l w a t e r i n g regime d u p l i c a t e d S. i n t e q r i f o l i a s o i l m o i s t u r e parameters but f a i l e d to m a i n t a i n adequate or s u s t a i n e d s o i l water l e v e l s f o r S. r u j i d a l a . I n s e c t s p e c t r a a r e summarized i n Table X* , F l o w e r s of a l l t a x a s t u d i e d were v i s i t e d 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 f l i e s a l t h o u g h bees and o c c a s i o n a l l y wasps are a l s o a t t r a c t e d t o t h e f l o w e r s and c o l l e c t both p o l l e n and n e c t a r . The f l i e s a r e d i v i s i b l e by t h e i r b e h a v i o r i n t o m o t i l e and more s e d e n t a r y c a t e g o r i e s w i t h t h e l a r g e r a n t h o p h i l o u s S y r p h i d s , B o m b y l i d s , and d e e r f l i e s much more l i k e l y t o e f f e c t c r o s s - p o l l i n a t i o n t h a n t h e i r s m a l l e r ^ g r o u n d - l o v i n g c o h o r t s . One 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 the t a x a o b s e r v e d c o u l d be t h e g r e a t e r dependence o f S. i n t e q r i f o l i a v a r . i n t e g r i f o l i a upon f l i e s as p o l l i n a t i n g a g e n t s (Table X). " Numerous a u t h o r s have suggested t h a t bees i n p a r t i c u l a r show 128 T a b l e X A Summary o f major groups o f i n s e c t v i s i t o r s observed i n 4 p o p u l a t i o n s . Numbers i n d i c a t e the 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 . No. Bees, % Bees, No. T o t a l gasps Wa s£s p j p t e r a V i s i t o r s §jt O f i d u l a 14 32.53 29 43 (NOOS-B) 10.5 hrs.;4,5 A p r i l S j , r u f i d u l a 10 19.0% 43 53 (NMO-B) 5.25 hrs.;21,22 A p r i l Sj. o c c i d e n t a l i s 13 54.2% 11 24 (Yale-626) 2 h r s . ; 19 A p r i l S A i n t e g r i f o l i a 2 5.8% 3 2 34 (NHO-I) 4 h r s . ; 10,11 May 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 d e c l i n e s ( B r i t t a i n and Newton 1933, Stephens 1956, Simpson and Duncan 1956, Lewis 1961, Free 1963,1968, L e v i n and Anderson 1970) . C o m p e t i t i o n f o r Hymenopteran p o l l i n a t o r s i n th e 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 v a r . i n t e g r i f o l i a c c u l d be more i n t e n s e j u d g i n g from t h e g r e a t e r v a r i e t y and abundance o f c o n c u r r e n t l y f l o w e r i n g s p e c i e s which are perhaps more a t t r a c t i v e to bees than S a x i f r a g a f l o w e r s . S l i g h t s e l e c t i o n p r e s s u r e due t o 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 may f a v o r more pronounced s e a s o n a l d i f f e r e n c e s i n f l o w e r i n g t ime ( W c N e i l l y and A n t o v o n i c s 1967, 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 s y m p a t r i c S. r u f i d u l a p o p u l a t i o n s ( A n t o n o v i c s 1968). A s h i f t i n v i s i t o r s p e c t r a between s y m p a t r i c S. i n t e g r i f o l i a and 5. r u f i d u l a t y p e p l a n t s , i f f u r t h e r , more d e t a i l e d o b s e r v a t i o n s c o n f i r m t h i s s p e c u l a t i o n , may have been an i m p o r t a n t e v o l u t i o n a r y i n f l u e n c e i n t h e development o f such p o s s i b l y f l y - a d a p t e d s t r u c t u r e s a s th e e n l a r g e d n e c t a r g l a n d which p r e s e n t s d i f f u s e , g l i s t e n i n g , u l t r a - v i o l e t a b s o r b i n g n e c t a r { P e r c i v a l 1965) as w e l l as reduced p e t a l s i z e , p a p i l l a e , and showiness i n S. i n t e g r i f o l i a v a r . l e p t o p e t a l a . The e v o l u t i o n of gynodioecy i n S. i n t e g r i f o l i a may a l s o be c o r r e l a t e d t o such a p o l l i n a t o r s h i f t i f s e l e c t i o n 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 compensate 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 ( P e r c i v a l 1965, F a e g r i and van der P i j l 1966). 130 TAXQNGJ3Y SJBfcies D e f i n i t i o n In T h i s Complex The d e f i n i t i o n o f a s p e c i e s i n t h i s complex group c f t a x a must a l l o w f o r o v e r l a p s i n v a r i a b i l i t y and the m o r p h o l o g i c a l i n t e r m e d i a c y o f many specimens. Hence, i t must f o c u s on t h e c l u s t e r s o f v a r i a b i l i t y which e x i s t around c e r t a i n m o r p h o l o g i c a l c h a r a c t e r i s t i c s . I s o l a t i n g mechanisms which r e s t r i c t gene f l o w are c o n s i d e r e d of secondary importance t o m o r p h o l o g i c a l t r e n d s i n a r r i v i n g a t a w o r k a b l e c l a s s i f i c a t i o n i n t h i s p a r t i c u l a r group., I f the presence of i n t e r m e d i a t e s and e v i d e n c e of gene f l o w are used as main c r i t e r i a f o r t r e a t i n g the t a x a a t a s u b s p e c i f i c l e v e l , the r e s u l t i n g s p e c i e s a r e e x t r e m e l y l a r g e and v a r i a b l e w i t h unmanageable ranges i n morphology, e c o l o g y , c y t o l o g y , e t c , When t h e s e e n t i t i e s a re i n c l u d e d i n r e g i o n a l f l o r a s , perhaps a f u n c t i o n a l approach would be t o t r e a t them as s p e c i e s a g g r e g a t e s i n the main key w i t h a s e p a r a t e , more p r e c i s e key t o t h e a g g r e g a t e t a x a . 131 Key To The S p e c i e s The f o l l o w i n g key does not d i s c r i m i n a t e v a r i e t a l o r s u f c s p e c i f i c t a x a w i t h i n S . ; i n t e q r i f o l i a o r S. oregana sensu H i t c h c o c k and C r o n q u i s t (1973) or the t a x a p e r i p h e r a l l y r e l a t e d t o 5. o c c i d e n t a l i s such as s u b s p e c i e s o f S. m a r s h a l l i i , S. r e f l e x a . o r the S. n i v a l i s - t e n u i s complex • sensu Krause and Beamish (1972,1973). S a x i f r a g a i n t e g r i f o l i a and S. oregana a r e not i n c l u d e d i n the f o l l o w i n g d e s c r i p t i o n s . A s h o r t e r , s i m p l i f i e d key i s p r e s e n t e d i n t h e Appendix. I t i s designed f o r i d e n t i f i c a t i o n of t h e m a j o r i t y o f specimens, but many i n t e r m e d i a t e o r h y b r i d specimens w i l l not i d e n t i f y e a s i l y u s i n g t h a t key. , A. Ovary l e s s than 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 scapes ,5m o r l e s s t a l l ( n e a r l y t o about h a l f i n f e r i o r i n seme h y b r i d e n t i t i e s of S. o c c i d e n t a l i s and S. g o r m a n i i : p e t i o l e d i s t i n c t , u s u a l l y more than 2x as l o n g as wide; l e a f n a r g i n s from s h a l l o w l y s i n u a t e - d e n t a t e t o , more commonly, rounded o r s q u a r e - d e n t a t e ( i f t e e t h markedly reduced o r near absent see l e a d AA); p e t a l s u s u a l l y 2.5mm o r g r e a t e r (sometimes s m a l l , g r e e n i s h , r e d d i s h or l a c k i n g i n a l p i n e forms o f S. , o c c i d e n t a l i s . C and S. a e g u i d e n t a t a . C C ) . B. N e c t a r q l a n d a t a n t h e s i s a s w o l l e n , c y l i n d r i c o r douqhnut-shaped r i n q t h a t almost c o v e r s t h e o v a r y ; r h i z o m e s 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 or as a few s m a l l r e ddish-brown f r a q m e n t s i n p r e s s e d m a t e r i a l ; l e a v e s t a p e r i n g a b r u p t l y i n t o an e l e n q a t e p e t i o l e ; o v a r y u s u a l l y 132 from 1/3 t o about 1/2 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 , •,...............>,«.•••• •v..,.. S a x i f r a g a g o r m a n i i 1. BB. Nectar g l a n d 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 ; l e a v e s t a p e r i n g more o r l e s s g r a d u a l l y from t h e b l a d e i n t o a p e t i o l e ; ovary 1/3 or l e s s 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 . C. I n f l o r e s e n c e u s u a l l y from c o n i c a l t o a t i g h t l y c l u s t e r e d h e a d l i k e p a n i c l e ; f i l a m e n t s u s u a l l y c l a v a t e t o sometimes n a r r o w l y o b l a n c e o l a t e ; t e e t h on l e a f margins s g u a r e - d e n t a t e ; p e t a l s u s u a l l y t a p e r e d i n t o a somewhat narrowed base; g l a n d from i n c o n s p i c u o u s t o a somewhat broadened band. ................... ................... S. O c c i d e n t a l i s 2. CC., I n f l o r e s e n c e u s u a l l y f l a t topped t o obtuse c o n i c a l , not t i g h t l y c l u s t e r e d i n t o a dense h e a d l i k e p a n i c l e ; f i l a m e n t s l i n e a r or s u b u l a t e , r a r e l y s l i g h t l y c l a v a t e o r o b l a n c e o l a t e i n montane f o r m s ; t e e t h on l e a f margins s i n u a t e - d e n t a t e t o s g u a r i s h - d e n t a t e ; p e t a l s w i t h a r a t h e r broad base; g l a n d an i n c o n s p i c o u s , narrow band. .... .. S. a e g n i d e n t a t a 3, ,• Ah. Ovary h a l f or more i n f e r i o r a t ant h e r d e h i s c e n c e (becoming s u p e r i o r i n f r u i t i n some cases) o r , i f l e s s than h a l f i n f e r i o r , t hen scapes e x c e e d i n g .5 meters; p e t i o l e from almost l a c k i n g , t o e l o n g a t e and narrow; l e a f margins e n t i r e or m i n u t e l y d e n t i c u l a t e t o s h a l l o w l y o r unevenly s i n u a t e - d e n t a t e ; p e t a l s about 2.5mm o r l e s s ( g r e a t e r i n 133 S. i n t e g r i f o l i a * see DD and S. l a t i p e t i o l a t a . £ ) . D, P e t i o l e s a l m o s t l a c k i n g , s h o r t and broad u s u a l l y l e s s t h a n 2x as broad as l o n g ; l e a f b l a d e s t a p e r i n g g r a d u a l l y i n t o p e t i o l a r r e g i o n ; margins uneven u n d u l a t e d e n t a t e t o s h a l l c w l y and unevenly s i n u a t e d e n t a t 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 o f t e n w i t h l o n g , c l e a r or merely p i n k i s h g l a n d u l a r h a i r s . , E, Scapes u s u a l l y l e s s t h a n 0.5m; p l a n t s i n mo i s t g r a s s y s o i l of Northwest Oregon C o a s t a l Range mountaintop «balds". ..................................... S. l a t i f i e t i s l a t a 4 . EE. Scapes o f t e n 0.5m o r g r e a t e r ; p l a n t s i n boggy o r swampy p l a c e s from t h e S i e r r a Nevada n o r t h t o t h e Cascades o f c e n t r a l H a s h i n g t o n and west t h r o u g h o u t the R o c k i e s Mountains., ..........•................... ............. S. oregana. CD. P e t i o l e s e l o n g a t e even i f reduced i n l e n g t h , d i s t i n c t l y narrowed and e v i d e n t ; l e a f b l a d e s t a p e r i n g g r a d u a l l y o r a b r u p t l y i n t o p e t i o l a r r e g i o n ; margins v a r i o u s , commonly n e a r l y 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 w i t h dark r e d d i s h - t i p p e d g l a n d s . 134 1» 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 T o r r e y a 23:106. 1923. (8.W. Gorman 4081, E l k Bock, Multnomah Co., Oregon, June 2,1917 HSI) S. m a r s h a l l i i f. d e n t a t a E n g l . & Irmsch. P f l a n z e n r . IV,117,1:36. 1916. { H e l l e r 10059, E l k Bock, near Oswego, Clackamas Co., Oregon) S. o c c i d e n t a l i s Wats. v a r . d e n t a t a ( E n g l . & Irmsch.) C L . H i t c h c . , Vase. P I . Pac. Borthw., U n i v . Wash. P r e s s , H i t c h c o c k e t a l . I l l : 149. 1961. , P e r e n n i a l , r o s e t t e - f o r m i n g herb w i t h d a r k , r e d d i s h , d e l i c a t e , d e e p l y - g r o w i n g and b r a n c h i n g r h i z o m e s . Leaves s i m p l e , e x s t i p u l a t e ; b l a d e s e l l i p t i c t o o v a t e , t a p e r i n g a b r u p t l y o r somewhat g r a d u a l l y i n t o a d i s t i n c t p e t i o l e u s u a l l y mere than 2x as 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 s p a r s e l y r n s t y v i l l o u s below; margins from s u b e n t i r e ( i n some t e t r a p l o i d i n t r o g r e s s a n t s w i t h S. ' n t e g r i f o l i a v a r . c l a y t o n i i f o l i a ) t o s i n u a t e d e n t a t e ; t e e t h n e a r e s t apex about 0.5 mm l o n g . I n f l o r e s c e n c e s e v e r a l - f l o w e r e d (20-81), open, c o n i c a l p a n i c u l a t e ; r a c h i s i n d e f i n i t e o r e v i d e n t . 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 or a s c e n d i n g ; p e t a l s about 2.5 mm l o n g or l o n g e r , w h i t e , t a p e r i n g g r a d u a l l y t o a broad base, mostly d e c i d u o u s i n f r u i t ; a n t h e r s orange t o y e l l o w ; f i l a m e n t s l i n e a r to s u b u l a t e ( r a r e l y c l a v a t e i n i n t e r m e d i a t e s w i t h S. m a r s h a l l i i subsp. m a r s h a l l i i : c a r p e l s b r o a d l y p y r i f o r m a t a n t h e s i s ; g l a n d s w o l l e n , doughnut-shaped, s u r r o u n d i n g t h e upper 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 i n f e r i o r a t anther d e h i s c e n c e ; g l a n d remnant o f t e n a l i n e a r r i d g e e n c i r c l i n g t h e f r u i t . , (n=10,19,20) March t o A p r i l . , Bocky c l i f f s i d e s i n v e r n a l l y wet seeps or m o i s t p l a c e s from C l a t s o p 135 County, Oregon e a s t t o C o w l i t z Co, Washington and s o u t h t o L i n c o l n and Marion C o u n t i e s i n Oregon. £S£E£§§J3iative Specimens OREGON: C l a t s o p Co.: exposed bare 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 Saddle Mt., c a . 3225*, 3 June 1973, K.L. Chambers 3752 • (WT0,0BC); r o c k c r e v i c e s and b l u f f s , n ear A s t o r i a , 6 May 1933, G.P. Baker s . n . ( Q B E ) ; L i n c o l n Co.: O t t e r C r e s t , 26 March 1930. anonymous s.n. I QBE): T i l l a m o o k Co.: exposed knobs, r i d g e s above burned f o r e s t , g r a v e l l y s l o p e s , T i l l a m o o k Burn a r e a , N. W i l s o n B i v e r Hwy. , e x a c t l y 3 mi s e . of Blue Lake, 3000*, 6 June 1S75, K.L. Chambers 4065 (OSC). The f o l l o w i n g specimens t e n d t o r esemble S. i n t e q r i f p l i , a v a r . c l a y t o n i i f o l i a i n t h e i r q r e a t e r p l a n t s i z e {ca. 20 cm), more n e a r l y h a l f i n f e r i o r o v a r i e s , and reduced t e e t h : OREGON: Columbia Co.: damp r o c k y b l u f f s , about w a t e r f a l l s , s. f o r k o f C l a t s k a n i e C reek, 10 mi above C l a t s k a n i e , 15 May 1927, J..W. IMlJson 24j47 (STD) ; Lane Co.: H i l l ' s C r eek, 300', 29 May 1938, Dgtlina 2783 (ORE). T h i s specimen i s p r o b a b l y a h y b r i d between q o r m a n i i and •S. i n t e g r i f o l i a v a r . c l a y t o n i j . f o l i a ( p o l l e n i s 100% s t e r i l e ) : OREGON: Clackamas Co.; near Milwaukee, E l k Rock, 28 March 1885, 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. Acad. 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):14 4. ; 1905. 136 ( J . Macoun. N.c. Y a l e Mt., B.C. May 17,1875. CAN) M i c r a n t h e s saximcntana S m a l l , N. Am. F l . 22 (2) :145. 19C5. (A. & E. N e l s o n , 5917, Yancey«s, Y e l l o w s t o n e N a t l . Park J u l y 17, 1889 MINN!) M e r anthes la£a S m a l l , N. Am. F l . 22 ( 2 ) : 145. 1905. (John Macoun, l y t t o n , B.C. A p r i l 16,1889) M i c r a n t h e s a l l e n i i S m a l l , N. Am. F l . 22(2):144.,1905. (O.D. A l l e n 242, Goat Mountains, Sash. June 27, 1896) S. l a t a Fedde, J u s t Bot. J a h r e s b . 31(1):613. 1906. S. a l l e n i i Fedde, J u s t Bot. J a h r e s b . 3 1 (1):613. 19C6. S. m i c r o c a r p a Johnson, Minn. S t . P I . S c i . 4:25. 1923. (M.F• E l r o d 98a, M i s s o u l a , Mont. MINN!) S. o c c i d e n t a l i s v a r . w a l l o w e n s i s Peck, L e a f 1 . West. Bot. , 5:60. 1947. (Peck 18542, above I c e Lake, Wallowa Mts., Wallowa Co., Oregon J u l y 4, 1894) S. o c c i d e n t a l i s v a r . a l l e n i i C.L. H i t c h c . , Vase. P i . Pac. Northw., Univ. Wash. P r e s s , H i t c h c . £l a l . , 111:49. 1961. P e r e n n i a l , r o s e t t e - f o r m i n g herb w i t h s h o r t , s t o u t , s e l d o m - b r a n c h i n g rhizome. Leaves s i m p l e , e x s t i p u l a t e ; b l a d e s 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 i n t o a d i s t i n c t p e t i o l e , u s u a l l y g l a b r o u s above, o r r a r e l y p u b e r u l e n t , s p a r s e l y t o d i s t i n c t l y r u s t y v i l l o u s below; margins u s u a l l y s g u a r e d e n t a t e ; t e e t h n e a r e s t apex u s u a l l y c a . 5mm l o n g o r l o n g e r . I n f l o r e s e n c e s e v e r a l - f l o w e r e d (13-50), p a n i c u l a t e t o d e n s e l y c l u s t e r e d h e a d l i k e p a n i c u l a t e , c o n i c a l t o s p h e r i c a l i n more h e a d l i k e i n d i v i d u a l s ; r a c h i s i n d e f i n i t e o r d i s t i n c t . 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 t o souewhat r e f l e x e d ; p e t a l s about 2 . 5 mm l o n g , w h i t e , u s u a l l y t a p e r i n g 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; a n t h e r s orange t o y e l l o w ; f i l a m e n t s u s u a l l y a t l e a s t s l i g h t l y c l a v a t e t o d e f i n i t e l y c l a v a t e ( a l t h o u g h not p e t a l o i d ) ; c a r p e l s e l o n g a t e , 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 e n c i r c l i n g t h e o v a r y w a l l ; o v a r y 1/3 o r l e s s (sometimes al m o s t 1/2) 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 , S t y l a r beaks r e c u r v e d i n f r u i t ; g l a n d remnants 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 t o August, Bocky o u t c r o p s , c l i f f s i d e s and mountaintops a d j a c e n t t o v e r n a l s t r e a m l e t s , seeps, wet r o c k f a c e s or moist areas i n s h a l l o w s o i l (about 7 cm deep), widespread i n the Bocky Mountain r e g i o n and n o r t h Cascades from Washington, Idaho, and Montana n o r t h t o S c u t w e s t e r n A l a s k a and a d j a c e n t Yukon, ISlU§e£iaili§ Specimens ALEEBTA: r o c k y s l o p e above B e r t h a Lake, Waterton Lakes N a t l . P a r k , 21 June 1930, jB.-C. M c C a l l a 3618 (MIN); ERITISH COLUMBIA: among r o c k s , A t l i n Hot S p r i n g s , 2250', 5 J u l y 1914, I.J. Anderson 2420 (V) ; L i l l o o e t , 13 Hay 1916, E . J H . Andersfin 2419 (V); r o c k o u t c r o p s , Manning Park, B l a c k w a l l , 14 J u l y 1960, K . I . Beamish. F. Vrugtman 60805 (V,UBC); c a l c a r e o u s f o o t o f g l a c i e r , Cougar V a l l e y , S e l k i r k Mts., 1600», 18 J u l y 1908, F.K, B u t t e r s and E.W.D. Holway ,340 (MIN) : meadow, Mt. Bobson P a r k , Snowbird P a s s , 9 Aug. 1975, C.C, Chuanjg 75J[399 (V) ; g l a c i a l moraines. Sphinx G l a c i e r f o r e l a n d . G a r i b a l d i P a r k , 500 0*, 26 Aug, 1965, B, E r a s e r s.n. (UBC); r o c k s l i d e above r i v e r on e. 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 1934, E.T. HcCabe 758 (UC,WTU); C r o n i n Mt., 10 mi s. o f S m i t h e r s , 1 J u l y 1967, G. 138 Mendel 127 (V); wet c r e v i c e s . C a t h e d r a l L a k e s , Mt. Bcmford, A s h n c l a D i s t r . , 7000', 12 J u l y 1951, T.M.C. T a y l o r 1346 (8S.PC): Marble Mts., Lake B o o t a h n i e , 5000', -20,25 June 1938, J . j j l . anji E.M. Thompson 70 (UC.WS.MIN); McComal Creek Quadrangle, c a . 2 mi w. of sw, c o r n e r o f Thutade, S t i k i n e Mt., 4500*, 14 June 1969, S.L. HeIsfa, K. Riqby 9109 (BB1). IDAHO; K o o t e n a i Co.: June 1892, J.B. H i e b e r q s.n. (OBE); C u s t e r Co.: among r o c k s i n seepage from snowbank, common, Mt. ssw. o f A l t u r u s L a k e , S t a n l e y , n=10, 11 Aug. 1969* D.L. Krause 68 (UEC)MONTANA: Beaver Head Co.: t o p o f O d e l l Peak, P i o n e e r Bange, 24 J u l y 1946, C.L. H i t c h c o c k and C.V. Muhlicjc 14925 (HS) ; Deerlodge Co.: c a . 18 mi s.w. o f Anaconda on e. s l o p e above t r a i l between Storm Lake P a s s and Goat F l a t j u s t on Goat F l a t s i d e o f a l a r g e l i m e s t o n e o u t c r o p p i n g , 9100», 7 J u l y 1974. P. Elvander 443 (WTO). G l a c i e r Co.: g r a s s l a n d on w e l l d r a i n e d s o i l , 2.6 mi n. of S t . Mary, 4 J u l y 1950, D. Lynch 6284 (HS) : NEVADA: E l k o Co.: i n f r e g u e n t i n c r e v i c e s of r o c k s a l o n g stream bank, 1/2 mi above Thomas Canyon Campgrounds, La M o i l l e Canyon, 15 June 1941, A.H. Holmgren 1122 (UC). OBEGON: Grant Co.: r o c k y s l o p e , ne. o f s u m i i t of S t r a w b e r r y Mt., 8900»,1 Aug. 1953, A. Crongui.st 7703 (WS^HTD) : Wallowa Co.; s t e e p exposed s l o p e s o f P e t e ' s P o i n t , 19 J u l y 1962, G» l a s o n 5428 (OSC). WASHINGTON: Snohomish Co. : under d r i p p i n g c l i f f s , Mt. D i c k e r s o n , Cascade Mts.„ 5000», 17 J u l y 1932, J.H. 2h£J£Son 8853 (WTO) ; Okanagan Co.: r o c k y o u t c r o p s , below S l a t e Peak a t t h e head o f t h e S l a t e F o r k o f t h e Paysatan B i v e r , 28 J u l y 1940, J . Ownby., E . G . Meyer 2222 (HS,0BI,OSC,MIN,0C); P i e r c e Co.: r o c k y s l o p e , Mt. B a i n i e r N a t l . P a r k , C r y s t a l Mt. I n d i a n Henry's, 2 J u l y 1928, F.A. Warren 785 (WS); Skamania 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 , 4000*, 7 June 1945, D.C. Inqrahm .1877 (HS,OSC) ; Spokane Co.: damp r o c k y h i l l s i d e , near L a t a h C r e e k , s e . of Spangle, 12 flay, 27 June 1916, J.S. Suksdorf 8616 (WS); Whatcom Co.: s h a l l o w s o i l on r o c k y s l o p e s a t t i m b e r l i n e , at. Baker, 6200', 11 J u l y 1922, H.J..- ;>-Mason 3871 {DC). WYOMING: Grand Teton N a t l . P ark: moist mossy h i l l s i d e . Cascade Canyon, 7 5 0 0 f , 19 June 1933. I . W i l l i a m s J135 {CSC). 3. / S a x i f r a g a a e g u i d e n t a t a (Small) Bosend. J.n E n g l . Bot. J a h r b . 37, B e i b l . 83:70, 1905. S a x i f r a g a r u f i d u l a (Small) James Macoun Ottawa Nat., 20:162. 1906. J i S f a n t b e s r j i f i d u l a S m a l l , N. Am. F l . 22(2) : 140. 1905. (John Macoun, Mt. F i n l a y s c n , Vancouver I s l a n d May 17, 1887 HY) M i c r a n t h e s a e g u i d e n t a t a S m a l l . N. Am. F l . , 22(2):145. 1905. (Suksdorf 967, Lower C a s c a d e s , Skamania Co., Wash. WS) S. r u f i d u l a f . 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. 1916. S. r u f i d u l a f . minor E n g l . S I r m s c h . , P f l a n z e n r . IV,117,1:39. 1916. '§.* J t l i c k i t a t e n s i s Johnson, Minn. Stud, P I . S c i , 4: 25. 1923. {Suk s d o r f , K l i c k i t a t Co., Wash. A p r i l 9 and May 1883 HS!) S. o c c i d e n t a l i s Wats, subsp. r u f i d u l a B a c i g a l u p i i n Abrams, 111. F l . Pac. S t . 2:366.,1944. S. o c c i d e n t a l i s v a r . r u f i d u l a H i t c h c , V a s e P I . Pac. Northw., Oniv. Wash. P r e s s , B i t c h c . E% a l , , 111:49. 1961. P e r e n n i a l r o s e t t e - f o r m i n g herb w i t h s h o r t , s t o u t , few 140 b r a n c h i n g , h o r i z o n t a l r h i z o m e s . Leaves s i m p l e , e x s t i p u l a t e ; b l a d e s e l l i p t i c t o o v a t e , t a p e r i n g somewhat g r a d u a l l y t o a b r u p t l y i n t o a d i s t i n c t p e t i o l e , g l a b r o u s above, r u s t y tcmentose t o r u s t y v i l l o u s below; margins d e e p l y s i n u a t e d e n t a t e to somewhat s h a l l o w l y s i n u a t e d e n t a t e ; t e e t h n e a r e s t apex u s u a l l y g r e a t e r t h a n .5mm l o n g . I n f l o r e s c e n c e few- (4-42) t o s e v e r a l - f l o w e r e d (as h i g h as 7 4 ) , open, s p r e a d i n g , f l a t - t o p p e d , convex o r o b t u s e l y c o n i c a l ; r a c h i s u s u a l l y i n d e f i n i t e . F lowers 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 o r a s c e n d i n g ; p e t a l s 2.5 mm l o n g o r l o n g e r , w h i t e , t a p e r i n g g r a d u a l l y t o a u s u a l l y broad base, mostly d e c i d u o u s i n f r u i t ; a n t h e r s from dark r e d t o y e l l o w ; f i l a m e n t s l i n e a r o r o b l a n c e o l a t e t o s l i g h t l y c l a v a t e i n some montane fo r m s ; c a r p e l s narrow, b o t t l e - s h a p e d a t a n t h e s i s ; g l a n d 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 e n c i r c l i n g t h e ovary w a l l ; o v a r y 1/3 or l e s s 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 . F r u i t i n g s t y l a r beaks r e c u r v e d ; g l a n d remnant i n c o n s p i c u o u s i n f r u i t . (n=10,19,28 c a , 28,29) Mid February t o J u l y . On s h a l l o w s o i l <ca. 3-4 cm) o f r o c k y o u t c r o p s and c l i f f s i d e s i n v e r n a l l y m o i s t , o f t e n d r i p p i n g s e e p s , washes or r i v u l e t s from the Upper W i l l a m e t t e B i v e r area n o r t h and e a s t i n t o t h e Columbia B i v e r Gorge, t h e n found from t h e Olympic Mountains to e a s t c e n t r a l Vancouver I s l a n d . , Be p r e s e n t a t i v e Specimens BBITISH COLOMBIA: A l b e r n i , Vancouver I s l a n d , A p r i l 1914, C a r t e r CI.59- ( V ) ; r o c k y b l u f f s , Cowichan Lake, B a l d Mt. , Vancouver I s l a n d , 24 a a r c h 1940, I.M. Cowan s.n. (V); i n 141 c r e v i c e s o f r o c k s , s o i l damp, n. end o f Shawnigan L a k e , Vancouver I s l a n d , (n=10), 20 Feb. 1971, D.L. Krause. 1-71 (UBC); mossy wet r o c k s . H i l l H i l l , Vancouver I s l a n d , 17 Karch 1895, J . B. Anderson 84 (V). CBEGON: Clackamas Co.; E l k Bock C l i f f s 2 mi n. of Oswego Lk., (n=10) , 17 Harch 1971, D.L. Krause 9-7.1 (UBC,UBC) ; Hood B i v e r Co.: open b a s a l t i c k n o l l s w i t h S e l a g j n e l l a w a l l a c i j , 12.5 mi w . o f Hood B i v e r a l o n g p o r t i o n O l d Columbia B i v e r Hwy., 11 A p r i l 1958. C L . H i t c h c o c k and C. V. Mul'ck 21501 (SS) ; Lane Co.: s t e e p , g r a v e l l y , wet s o i l , G»Leary H t n . , 2800», 28 June 1938, L.E. D e t l i n g 30.82. (OBE) ; L i n n Co.: moss mats on e. f a c i n g c l i f f , S antiam fi., 20 mi e. of Sweethome, 7 A p r i l 1951, JU C r o n ^ u i s t 6828 (HS); Marion Co.: c l i f f . S i l v e r Creek F a l l s , 9 A p r i l 1940, M. H r i j h t s.n., (OSC); Multnomah Co.: on r o c k s and r o c k y c l i f f s near E l k Bock, 12 A p r i l 1903. M.». Gorman s.n. (herbarium no. 22637) (WTB); a l o n g Sandy B i v e r a t j u n c t i o n of e., S t a r k e Bd. and e. Columbia Hwy., 2 2 March 1926, H.E. J?£ck 1452J[ (OSC); moist s l o p e s , near Multnomah F a l l s , 18 A p r i l 1935, J.jf. Thompson 17370 (BS) ; Uasco Co.; wet c l i f f s , 6 mi w« o f The D a l l e s cn Columbia B i v e r H w y . , ( p o l l e n f e r t i l i t y , s t e r i l e - 1 4 , f e r t i l e - 1 9 0 , E. P e r k i n s ) , 27 March 1946, H.H. Baker 274 (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. o f C l a r k - S k a m a n i a Co. l i n e , S t . Hwy. 14, n=10, 14 March 1971, D.L. Krause 2-7J (DEC); Greys Harbor Co.: wet c l i f f s o f Mt. C o l o n e l Bob, 3500», 12 J u l y 1930, J.JJ. Thompson 9404 (PC) : J e f f e r s o n Co.: r o c k y c r e s t of Constance B i d g e , 5500*, 30 May 1931, J.H., Thompson 6583 (OSC); K l i c k i t a t Co.: 3 mi e. o f Bi n g e n , 300«, 26 A p r i l 1950, L.S. Rose 50073 (OC); Mason Co.; r o c k o u t c r o p s where p r o t e c t e d i n narrow canyons, near summit of 142 fit. E l i n o r , 11 June 1940, E.G. J e j e r J783 (§S) ; Skamania Co.: wet c l i f f s . Cape Horn, 10 A p r i l , 27 May 1920, 1. Suksdorf 10365 (WS,UC) . The f o l l o w i n g specimens, u s u a l l y o f h i g h e r mountain a r e a s , show s l i g h t resemblances i n c l a v a t e f i l a m e n t s and c l u s t e r e d p a n i c u l a t e i n f l o r e s c e n c e s t o S. o c c i d e n t a l i s : BRITISH COLUMBIA: S t r a t h c o n a P a r k , Mt. B o o s t e r Comb, Vancouver I s l a n d , J u l y 1937, N.C. Stewart 10481 (V) . WASHINGTON: J e f f e r s o n Co.: r i d g e , head of D o s e w a l l i p s B., 6000*, 29 J u l y 1921, l . P . T a y l o r s.n. (UCJ; C l a l l a m Co.: m o i s t r o c k y banks, Ht. An g e l e s , 5000*, 16 J u l y 1931, J.W. Thompson 739S (UC, OSC). These specimens approach S. m a r s h a l l i i - i n t h e i r c l a v a t e f i l a m e n t s , l o n g e r p e d i c e l s and o f t e n r e f l e x e d s e p a l s : OREGON: L i n n Co.: e. f a c i n g c l i f f a l o n g Santiam R. 20 mi e. o f Sweethome, 7 A p r i l 1951. A. C r o n q u i s t 6828 (UC) : Lane Co.; s t e e p n. s l o p e , m o i s t , Mt. 0»Leary, 4 800*, 28 June 1938. L.E. P e t l i n g 3082 (UC); Marion Co.: c l i f f , t o p o f House Mt., 31 May 1926, M . J . Peck 14638 (OSC). These specimens a r e t e n a t i v e l y c l a s s i f i e d here as S. h o w e l l i i Greene but t h i s t a x o n i s d o u b t f u l l y d i s t i n c t from S« r u f i d u l a . F u r t h e r work i s n e c e s s a r y on t h i s problem. OREGON: Jo s e p h i n e Co.: d r i e d up but l a t e l y moist b l u f f s . E i g h t D o l l a r Mt. near Selma, 26 March 1926, L.F. Henderson .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 , 10 mi B e s t o n Rd., 24 Feb. 1973, M. W i l l i a m s s.n. (OBE) . 4. S a x i f r a g a l a t i p e t i o l a t a (C.L. H i t c h c o c k ) P e r k i n s and E l v a n d e r . 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 . . Vase. P I . P a c , Northw., Univ. Hash. P r e s s , H i t c h c . e t a l . , 111:49. 1961. (fl.H.Gorman 3561, S a d d l e Mt., C l a t s o p Co., Oregon, June 20, 1915. WTU!) P e r e n n i a l , r o s e t t e - f o r m i n g herb w i t h s h o r t , s t o u t , seldom b r a n c h i n g rhizome., Leaves s i m p l e , e x s t i p u l a t e b l a d e s c i l i a t e above, s p a r s e l y l o n g v i l l o u s below, w i d e l y o v a t e , t a p e r i n g g r a d u a l l y t o a s h o r t , b r o a d , c i l i a t e , p e t i o l a r r e g i o n , p u b e r u l e n t above and below t o f a i n t l y r u s t y s e r i c e o u s below; margins d e n t a t e t o u n d u l a t e e r o s e . I n f l o r e s e n c e u s u a l l y many-flowered (46-231), c o n g e s t e d , c o n i c p a n i c l e , t o a somewhat c o n g e s t e d , p a n i c u l a t e ( o c c a s i o n a l l y corymbiform) head; c e n t r a l r a c h i s e v i d e n t t o i n d i s t i n c t . 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 r e f l e x e d ; p e t a l s about 2.5 mm l o n g o r l o n g e r , w h i t e , t a p e r i n g g r a d u a l l y t o a broad base, p e r s i s t e n t i n t o f r u i t ; a n t h e r s y e l l o w ; f i l a m e n t s l i n e a r ; c a r p e l s o b c o n i c o r umbonate o b c o n i c ; g l a n d a f l a t t e n e d d i s c c o v e r i n g the t o p c f the ovary a t a n t h e r d e h i s c e n c e , g r a d i n g i n t o t h e s t y l a r t i s s u e ; o v a r y 1/2 o r more i n f e r i o r at a n t h e r d e h i s c e n c e , becoming s u p e r i o r i n f r u i t . F r u i t i n g s t y l a r beak r e f l e x e d ; g l a n d remnant a l i n e a r r i d g e e n c i r c l i n g t h e f r u i t . (n=ca. 38) L a t e May t o e a r l y J u l y . S h a l l o w s e l l o f h i g h e r v o l c a n i c p l u g s and mountaintop " b a l d s " i n m o i s t , g r a s s y a r e a s of C l a t s o p and n o r t h e r n T i l l a m o o k C o u n t i e s , Oregon. fie prese nt a t j v e Specimens OBEGGN: C l a t s o p Co.: r o c k y s l o p e s . Saddle Mt.,2800-3300*, 144 20 June 1915. a . l . Gorman 3561ISOTYPE (WS); mo i s t open s l o p e s . Saddle Mt., 28 June 1952, J.T. H o w e l l s.n. (0C); Douglas f i r - s p r u c e f o r e s t , r o c k c r e v i c e s nw. e x p o s u r e , moderate shade, Saddle Mt., 2200', 19 June 1932, L.E. B e t l i n g 7906 ( ORE). The f o l l o w i n g a r e c l a s s i f i e d as S . i nt eg r i f o l i a but resemble S. l a t i p e t i o l a t a i n l e a f shape, broad s h o r t p e t i o l e s , and w h i t e pubescence cn the upper l e a f s u r f a c e : OREGON: P c l k Co.: v e r y wet p l a c e s , Monmouth, 20 May 1893 . W.J. Spil1man 78 <WS); Marion Co.: g r a v e l l y s o i l , common, Salem, B r o o k s P a s t u r e , 5 A p r i l 1919, fl.W. Gorman 4J4JJ (WS) ; WASHINGTON: T h u r s t c n Co.: Rock P r a i r i e , 12 May 1934, I.£. Q$i§ JL§93 (IS) . cojsayjsioNS The p r e s e n t s t u d i e s i n d i c a t e t h a t a reassessment c f the S» o c c i d e n t a l i s s p e c i e s complex i s necessary. The v a r i e t a l t a x a which H i t c h c o c k e t a l . (1961) and H i t c h c o c k and C r o n q u i s t (1973) l i s t f o r S. o c c i d e n t a l i s . namely r u f i d u l a , d e n t a t a , and l a t i p e t i p l a t a , a r e n o t merely e n t i t i e s which r e p r e s e n t d i s t i n c t e v o l u t i o n a r y t r e n d s w i t h i n S. o c c i d e n t a l i s - - 1 . Jhej s r are I S * PJipJ^SJ-SaAA X# 9S030fifei£§llX# and-- cy^plpg^icaj.ly s e p a r a t e d o v e r much o f t h e i r r a n g e s . Althpush c h a r a c t e r inter<jr.adation does o c c u r among some t a x a i n c e r t a i n a r e a s , e s j p e c i a l l y i n . p o l y p l o i d i n d i v i d u a l s and p o p u l a t i o n s , t h e ^|.^t4]|.qtjQBS;-;b.ased-,on-c e r t a i n c o m b i n a t i o n s o f c h a r a c t e r s a r e s u f f i c i e n t tp m a i n t a i n each o f t h e s e , J.anx- --.tlsej-taxenoiiic -^dif f 4euitiej§^®sn.a----these -t a xa can be a t t r i b u t e d t o h y b r i d i z a t i o n and a 11 p p p i y g 1 o i d y i n v o l v i n g v a r i o u s members o f t h e S. i n t e q r i f o l i a s p e c i e s 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 grcup appears to be o f l e s s i m p o r t a n c e but i t i s p o s s i b l e t h a t n e a r l y i d e n t i c a l d i p l o i d and t e t r a p l o i d o r t e t r a p l o i d and h e x a p l o i d S.. a e g u i den t a t a p l a n t s from t h e same l o c a t i o n have a r i s e n d i r e c t l y and w i t h o u t h y b r i d i z a t i o n w i t h o t h e r t a x a . S a x i f r a g a a e g u i d e n t a t a i s c o n f i r m e d as a c y t o l o g i c a l l y and m o r p h o l o g i c a l l y v a r i a b l e s p e c i e s i n agreement w i t h s e v e r a l p r e v i o u s taxonomic t r e a t m e n t s ( S m a l l and Bydberg 1905, Hacoun 1906, E n g l e r and I r m s c h e r 1916, Krause and Beamish 1S73). Some i n d i v i d u a l s from t h e Columbia B i v e r Gorge a r e a resemble S. o c c i d e n t a l i s m o r p h o l o g i c a l l y but t h e resemblance may be the r e s u l t c f complex h y b r i d i z a t i o n and p o l y p l o i d i z a t i o n p r o b a b l y 146 i n v o l v i n g S. i n t e g r i f o l i a v a r . c l a y t o n i i f c l i a o r i t s r e l a t i v e s i n t h a t a r e a . There i s e v i d e n c e t h a t some i n t r o g r e s s i o n between S. i n t e g r i f o l i a v a r . c l a y t o n i i f o l i a and S. a e g u i d e n t a t a i s o c c u r r i n g i n s y m p a t r i c p o p u l a t i o n s a l o n g t h e Columbia B i v e r Gorge. The t e t r a p l o i d 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 the r e s u l t o f a n c i e n t h y b r i d i z a t i o n s and p o l y p l o i d y perhaps between S. o c c i d e n t a l i s and s y m p a t r i c S. j.ntegrifpjLj,a 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 v a r . l e p t o p e t a • l a . Even i f resemblance i s the 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 w i t h S. p c c i d e n t a l i s i t appears t h a t S. a e g u i d e n t a t a i s p r e s e n t l y 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 w i t h s e v e r a l c h a r a c t e r i s t i c m o r p h o l o g i c a l f e a t u r e s . A r t i f i c i a l h y b r i d s between 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 p o l l e n and seed s t e r i l e and show no g r e a t e r c y t o l o g i c a l s i m i l a r i t i e s t h a n c r o s s e s i n v o l v i n g presumably more d i s t a n t l y r e l a t e d e n t i t i e s . N u m e r i c a l t r e a t m e n t s a l s o tend t o d i s t i n g u i s h a S. a e g u i d e n t a t a group. Columbia B i v e r Gorge S. a e g u i d e n t a t a p o p u l a t i o n s show c l o s e r s i m i l a r i t i e s t o S. o c c i d e n t a l ' s and o t h e r p o l y p l o i d o r h y b r i d i n d i v i d u a l s than do S. a e g u i d e n t a t a p l a n t s from Vancouver I s l a n d and t h e Olympic Mountains. The r e l a t i o n s h i p between S. a e g u i d e n t a t a and t h e m o r p h o l o g i c a l l y s i i i l a r and c y t o l c g i c a l l y unknown e n t i t y , S. h o w e l l i i . o f s o u t h w e s t e r n Oregon and northwe s t C a l i f o r n i a d e s e r v e s f u r t h e r s t u d y . 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 a v a r i a b l e , montane taxon which shows e v i d e n c e o f h y b r i d i z a t i o n i n s e v e r a l a r e a s o f i t s range. D e t a i l e d e x a m i n a t i o n o f i t s r e l a t i o n s h i p s w i t h S. n i v a l i s and S. t e n u i s as w e l l as w i t h S. m a r s h a l l i i subsp. 147 M i s h a l l i i and subsp. i d a h o e n s i s i s n e c e s s a r y . F u r t h e r work u s i n g a r t i f i c i a l h y b r i d i z a t i o n s among d i p l o i d S. o c c i d e n t a l i s , S. r e f l e x a . S. nj.vaJJLs, s u b s p e c i e s of S. .ffiargfoaAlii- d i p l o i d S. a e g u i d e n t a t a . and S. g o r m a n j i would be a u s e f u l a d d i t i o n t o t h e p r e s e n t s t u d y . , A major problem w i t h such s t u d i e s would be t h e s t r o n g s e a s o n a l 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 t h e s e p l a n t s which are adapted t o f l o w e r i n g regimes i n d i v e r s e a l t i t u d e s and l a t i t u d e s . 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. o c c i d e n t a l i s v a r . i s t r e a t e d here as a s e p a r a t e s p e c i e s f r c m 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 . f o r m e r l y S. r u f i d u l a • , w i t h a d i s t i n g u i s h a b l e morphology and a g e o g r a p h i c a l d i s t r i b u t i o n i n t h e Coast Range and Lower W i l l a m e t t e R i v e r o f Oregon e x t e n d i n g n o r t h t o C l a r k County, Washington. The t y p e l o c a t i o n i s E l k Rock, Oregon and specimens frcm t h a t s i t e c l o s e l y resemble nearby t e t r a p l o i d p l a n t s (n=20,19). T e t r a p l o i d s group w i t h d i p l o i d p o p u l a t i o n s i n t h e n u m e r i c a l s t u d i e s . A r t i f i c i a l h y b r i d s between t e t r a p l o i d S. o c c i d e n t a l i s and t e t r a p l o i d S. g e r m a n i i a r e s t e r i l e and c o n s i s t e n t l y f a i l t o undergo m e i o t i c d i v i s i o n s i n a n t h e r t i s s u e , i n c o n t r a s t t o c r o s s e s i n v o l v i n g S. a j S I i a j a i i and S. a e g u i d e n t a t a where m e i o s i s i n the F1 h y b r i d does o c c u r but i s i r r e g u l a r and p o l l e n f e r t i l i t y i s low., As a group, S. g o r m a n i i shows c l o s e r p h e n e t i c a f f i n i t i e s t o v a r i e t i e s o f S. i n t e g r i f o l i a and S. l a t i p e t i o l a t a . f o r m e r l y S. g c c i d e n t a l j s v a r . l a t i p e t i o l a t a . than t o S. ocjej, d e n t a l i s o r S. a e g u i d e n t a t a . Some specimens t e n d toward S. m a r s h a l l j i subsp. m a r s h a l l i i and 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 t h e W i l l a m e t t e R i v e r a r e a a r e needed to c l a r i f y t h i s problem. 148 Evidence i s p r e s e n t e d t h a t S. l a t i p e t j . o l a t a ( C L , H i t c h c o c k ) P e r k i n s and E l v a n d e r i s a l s o a s e p a r a t e s p e c i e s from S. o c c i d e n t a l i s . , S a x i f r a g a l a t i p e t i o l a t a has s e v e r a l s i m i l a r i t i e s t o S. oregana and t h e S. i n t e q r i f o l i a - c o m p l e x . M o r p h o l o g i c a l , c y t o g i c a l <n=ca.38), and n u m e r i c a l a n a l y s i s a l l i n d i c a t e a h y b r i d o r i g i n and a f f i n i t i e s w i t h S.oregana (n=3 8) . Other s y s t e m a t i c e v i d e n c e from s t u d i e s o f the S. i n t e g r i f o l i a group* S. rhomboidea, and C a l i f o r n i a n r e l a t i v e s o f S. i n t e q r i f o l i a ( E l v a n d e r , p e r s o n a l c ommunication, 1978) c o n f i r m t h a t i t i s p r o b a b l y more c l o s e l y r e l a t e d t o S. oregana. However i t i s e c o l o g i c a l l y and g e o g r a p h i c a l l y i s o l a t e d from S. oregana and has a number o f d i s t i n c t i v e m o r p h o l o g i c a l f e a t u r e s . T h e r e f o r e i t i s t r e a t e d here as a s e p a r a t e s p e c i e s . S t e r i l e i n t e r m e d i a t e s o c c u r i n i n t e r m e d i a t e h a b i t a t s i n most a r e a s where S. a e g u i d e n t a t a comes i n c l o s e s y a p a t r i c c o n t a c t w i t h S. i n t e g r i f o l i a o r S. i n t e g r i f o l i a v a r . c l a y t o n i i f o l i a . These can be r e c o g n i z e d m o r p h o l o g i c a l l y and c y t c l o g i c a l l y o r 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 . H y b r i d swarms occ u r i n s e v e r a l a r e a s , seme o f which may i n v o l v e S. a e g u i d e n t a t a and S, j n t e g r a j g l i a v a r , i n t e g r i f o l i a a s p a r e n t a l e n t i t i e s but o t h e r s a r e more l i k e l y the r e s u l t o f p r e v i o u s c o n t a c t between S. o c c i d e n t a l i s o r S. m a r s h a l l i i subsp, i d a h o e n s i s and S. i n t e g r i f o l i a ? a r , , l e p t o p e t a l a . C r y p t i c n a t u r a l h y b r i d s which resemble one p a r e n t a l e n t i t y and undergo abnormal m e i o s i s or e x h i b i t reduced f e r t i l i t y a r e not uncommon, e s p e c i a l l y i n the Columbia H i v e r Gorge a r e a among p o p u l a t i o n s o f mixed d i p l o i d and t e t r a p l o i d o r t e t r a p l o i d and h e x a p l o i d S. a e g u i d e n t a t a p l a n t s , but a l s o i n e t h e r a r e a s and 149 f o r o t h e r t a x a such as S, i n t e q r i f o l i a v a r . c l a y t o n i i f o l i a . I t i s d i f f i c u l t t o t e l l whether these i n d i v i d u a l s , e s p e c i a l l y i n t h e Columbia a i v e r Gorge, a r e the r e s u l t of c r o s s e s between two s i b l i n q e n t i t i e s w i t h d i f f e r e n t p l o i d y l e v e l s o r s e g r e g a t e s o f i n t e r s p e c i f i c c r o s s e s which c l o s e l y resemble the p a r e n t i n morphology and h a b i t a t p r e f e r e n c e s . , One p o p u l a t i o n west of C h e h a l i s , Washington (EP605), (n=29) i s a p p a r e n t l y an i n t e r m e d i a t e between 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 . The most r e a s o n a b l e t r e a t m e n t o f t h i s p r o b l e m a t i c a l p o p u l a t i o n appears t o be t o c l a s s i f y i t w i t h §• o c c i d e n t a l i s and f u r t h e r document i t s unigue background. The p o s s i b i l i t y cannot be e x c l u d e d t h a t i t i s a r e l i c t u a l p o p u l a t i o n of a once more widespread e n t i t y . I t s apparent c l o s e r e l a t i o n s h i p t o S. i n t e q r i f o l i a p l a n t s from t h e Mima and F t . L e w i s , Washington, r e g i o n needs f u r t h e r i n v e s t i g a t i o n . I t i s t r e a t e d here as b e l o n g i n g t o S, o c c i d e n t a l i s . The g e n e r a l p a t t e r n o f p o l y p l o i d e v o l u t i o n i n t h e group has most l i k e l y been one o f s e v e r a l p r o b a b l y independent d i p l o i d h y b r i d i z a t i o n s f o l l o w e d by chromosome d o u b l i n g and c o u p l e d w i t h a n e u p l o i d r e d u c t i o n ( i n the absence o f an a v a i l a b l e n i n e - p a i r e d p a r e n t a l s p e c i e s ) . T h i s c y c l e has a p p a r e n t l y r e p e a t e d i t s e l f i n s e v e r a l a r e a s t o g i v e r i s e t o h i g h e r p l o i d y l e v e l s and i n t r o g r e s s a n t forms. I n S. a e g u i d e n t a t a p o p u l a t i o n s which e x h i b i t mixed p l o i d y l e v e l s , h i g h e r l e v e l s o f p o l y p l o i d y may have o c c u r r e d through the c o m b i n a t i o n o f r a r e unreduced gametes from the same p o p u l a t i o n but h y b r i d i z a t i o n w i t h o t h e r t a x a , f o l l o w e d by chromosome 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 c h a r t of t h e r e l a t i o n s h i p s i n v o l v i n g S. aegui.deBta,|a and i t s 150 a l l i e s i s p r e s e n t e d i n F i g u r e 36. The combined d i f f i c u l t i e s o f l i m i t e d a r t i f i c i a l h y b r i d c o m b i n a t i o n s , p o s s i b l e d i f f e r e n c e s i n g e n e t i c c o n t r o l o f s y n a p s i s , and the p o s s i b i l i t y o f e x t i n c t p a r e n t a l e n t i t i e s l i m i t s 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 t h a t t h e p l a n t s i n v e s t i g a t e d a re most l i k e l y f a c u l t a t i v e s e x u a l o u t c r o s s e r s . There i s l i t t l e e v i d e n c e i n f a v o r o f a p o m i c t i c seed p r o d u c t i o n . The somewhat e a r l i e r f l o w e r i n g members of the o c c i d e n t a l i s group { S. S£gJdentajLj.s, S, a e g u i d e n t a t a . and S, gorma n j i ) a r e c a p a b l e of s e t t i n g more autogamous seed and may be f r e q u e n t e d by r e l a t i v e l y h i g h e r numbers of Hymenopteran v e r s u s D i p t e r a n p o l l i n a t o r s than t h e i n t e g r i f o l i a r e l a t i v e s (S. i n t e g r i f o l i a v a r . i s t e g r i f o l i a , S. i n t e g r i f o l i a v a r . c l a y t o n i i f o l i a . §* JB£ eggJf93- i a v a r . l e p t o p e t a l a . and S. l a t i p e t i o l a t a ) . These o b s e r v a t i o n s may 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 f l o r a l morphology between t h e two groups as w e l l as the oc c u r r e n c e o f gynodioecy i n c e r t a i n p o p u l a t i o n s o f S. i n t e a r i f o l i a v a r . , i n t e g r i f o l i a . A l t h o u g h the r e s u l t s o f bagging t e s t s and p o l l i n a t i o n s t u d i e s show i n t e r e s t i n g d i f f e r e n c e s between the major s p e c i e s complexes s t u d i e d , w i t h i n 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 s p e c i e s group, t h e y a r e l e s s u s e f u l as a taxonomic t o o l . The p o s t - g l a c i a l h i s t o r y o f t h e group i s complex and a p p a r e n t l y v a r i e d . Bandhawa and Beamish (1972) r e v i e w e d t h e ev i d e n c e f o r g l a c i a l r e f u g i a i n n o r t h w e s t e r n North America. The r e f u g i a l a r e a s a p p a r e n t l y c o n t a i n r e l i c t u a l d i p l o i d p o p u l a t i o n s i n c o n t r a s t t c widespread p o l y p l o i d c o l o n i z e r s o f s u r r o u n d i n g g l a c i a t e d a r e a s , Bandhawa and Beamish (1972) used the 151 F i g u r e 36: S p e c u l a t i v e p o l y p l o i d f o r m a t i o n s w i t h i n t h e 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 (the synonym- r u f i d u l a i s used i n t h e f i g u r e ) l i n e a g e w i t h emphasis on the Columbia R i v e r Gorge r e l a t i o n s h i p s . S e v e r a l o t h e r c o n s t r u c t i o n s a r e a l s o l o g i c a l l y p o s s i b l e . Numbers i n p a r e n t h e s i s a r e n o t s u p p o r t e d i n the l i t e r a t u r e . J o i n i n g l i n e s i n d i c a t e p o s s i b l e a l l o p o l y p l o i d r e l a t i o n s h i p s but i n c e r t a i n i n s t a n c e s a u t o p o l y p l o i d y 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 . 1 5 2 ntegrifol ia ca 5 5 SPECULATIVE RELATDNSHIPS oregana latipetiolata 38 38 occidentalis 29->28 rufidula 2 9 ^ 2 8 in tegr i fo l i a (20)-> 19 leptopetala. (20) ->19 / occidentalis gormanii 19 "20-> 19 rufidula . (20 )^19 (integrifolia) 10? rufidula 10 Proto integrifolia (10) Proto occidentalis (10) 153 d i s t r i b u t i o n o f 10 and 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 as an example of such a p a t t e r n . The narrow s o u t h e r n and western d i s t r i b u t i o n o f d i p l o i d s and w i d e s p r e a d ranges o f p o l y p l o i d s w i t h i n S. o c c i d e n t a l i s and some members o f S. i n t e g r i f o l i a conform w e l l t o the p a t t e r n e x h i b i t e d by S. f e r r u g i n e a . A l t h o u g h d i p l o i d S. a e g u i d e n t a t a p o p u l a t i o n s have r e c c l o n i z e d f o r m e r l y g l a c i a t e d a r e a s o f t h e Olympic Mountains and Vancouver I s l a n d , 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 are abundant i n a r e a s s o u t h c f the g l a c i a l boundary presumably r e s u l t i n g from c o n t a c t s between c o a s t a l and i n t e r i o r f l o r i s t i c elements i n t h e Columbia R i v e r Gorge. S i m i l a r complex h y b r i d i z a t i o n and p o l y p l o i d i z a t i o n p r o c e s s e s are a l s o e v i d e n t i n t h e h i s t o r y o f S. g o r m a r i i and S. l a t i p e t i o l a t a . Zones o f complex h y b r i d a c t i v i t y such as t h e Columbia R i v e r Gorge may p r o v i d e t h e raw 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 which s e l e c t i o n can o p e r a t e i n 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 ( S t e b b i n s 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 a r e a s 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 o r such r e s t r i c t i o n may be the r e s u l t of narrow e c o g e o g r a p h i c sympatry between two t e t r a p l o i d o r t e t r a p l c i d - d i p l o i d p r o g e n i t o r s which have s i n c e become l o c a l l y e x t i n c t . The absence of S. oregana from the Coast Range o f Oregon app e a r s t c be i n the l a t t e r c a t e g o r y w i t h r e s p e c t t o i t s p r o b a b l e p a r e n t a l r e l a t i o n s h i p t o S. j L a t j p e t i o l a t a . A 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 and i t s r e l a t i v e s i n c l u d i n g S. ., oregana. S. rhomboidea, and S. c a l i f o r n i c a , the l a t t e r two perhaps h a v i n g r a t h e r c l o s e r t i e s w i t h S. o c c i d e n t a l i s and S. 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 of the r e l a t i o n s h i p s between d i s j u n c t Western and E a s t e r n N o r t h American s p e c i e s p a i r s (Spongberg 1972) such as S. o c c i d e n t a l i s - 5 . v i r g i n i e n s i s , §• oregana-S. p e n s y l v a n i c a . S. f e r r u g j n e a - S . m i c b a u x i i . S. r e f l e x a - S. m i c r a n t h i d i f d j i a and S. mags[bap. 1 j i - S . c a r e l i n i a n a . Chromosome c o u n t s are i n c o m p l e t e f o r many t a x a , e s p e c i a l l y t h o s e where t h e f i e l d season c o n f l i c t s w i t h i n s t i t u t i o n a l s c h e d u l e s and t h o s e 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 s h o u l d be g i v e n t c 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 and areas 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 B i v e r a r e a , t h e Wallowa Mountains df n o r t h e a s t e r n Oregon, the Spokane a r e a of e a s t e r n Washington, and the h i g h e r mountain a r e a s o f n o r t h e r n C a l i f o r n i a and s o u t h e r n Oregon.. 155 LITERATURE C££ED A l e x o p o u l o s , C , J . , and E.S.Beneke. 1952* L a b o r a t o r y manual f o r i n t r o d u c t o r y mycology. 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Ah a p p l i c a t i o n o f c a n o n i c a l a n a l y s i s t o 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, 161 APPENDIX I n s e c t Specimens L i s t of i n s e c t s c a p t u r e d w h i l e v i s i t i n g S a x i f r a g a f l o w e r s : P=with g a x i f r a g a p o l l e n on body or 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: Vespidae P ( 2 ) : P o l i s t e s f u s c a t u s (1) ; F o r m i c i d a e ( 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 ) ; Andrenidae P f 1 1 ) . COLEOPTEBA: E l a t e r i d a e ( 1 ) . DIPTEBA: Anthomyidae ( 3 ) : Scatophaga spp., P ( 6 ) ; T a c h i n i d a e P ( 8 ) ; B o m b y l i i d a e : Bombvlius major P ( 2 ) ; Empidae P ( 7 ) ; C a l l i p h o r i d a e ( 1 ) ; Muscidae: Musea- dpmestica ( 1 ) ; Ceratopogonidae ( 2 ) ; M y c e t o p h i l i d a e P<2); Chironomidae {H); S y r p h i d a e ( 3 ) : Metasyrphus spp. (5) ; Dasvsvrohus s p . ( 1 ) ; Sp h a e r o p h o r i a sp. (1) ; Agrcmyzidae (1) . 162 F i g u r e 37: 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 Oregcn, Washington, and s o u t h w e s t e r n B r i t i s h Columbia o n l y ) and S* g c r m a n i i . The s i n g l e sguare i s t h e h e x a p l o i d i n t e r m e d i a t e p o p u l a t i o n <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 d i s t r i b u t i o n o f S, a e g u i d e n t a t a (the synonym, S. r u f i d u l a i s used i n the f i g u r e ) and S. l a t i p e t i o l a t a . D o t t ed l i n e i n d i c a t e s the w estern l i m i t of S. oreqana 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 ABBREVIETED KEY A. Ovary 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 s i s ; l e a f m a r g i n s s h a l l o w l y s i n u a t e - d e n t a t e t o e v e n l y d e n t a t e B. N e c t a r g l a n d a doughnut-shaped r i n g a t a n t h e s i s ; 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 brown n e t w o r k 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 band; r h i z o m e s s h o r t , s t o u t , h o r i z o n t a l C. I n f l o r e s e n c e f l a t - t o p p e d o r o b t u s e - c o n i c a l , f i l a m e n t s s u b u l a t e S_. a e g u i d e n t a t a CC. I n f l o r e s e n c e n o t f l a t - t o p p e d , u s u a l l y a c l u s t e r e d , h e a d l i k e t o n a r r o w l y c o n i c a l p a n i c l e , f i l a m e n t s c l a v a t e S_. o c c i d e n t a l i s AA. Ovary h a l f o r more i n f e r i o r a t a n t h e s i s ; l e a f , m a r g i n s e n t i r e t o u n e v e n l y s i n u a t e - d e n t a t e ( 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 i n t o b l a d e s ; v e s t i t u r e i n i n f l o r e s e n c e w i t h l o n g , c l e a r o r p i n k i s h - g l a n d u l a r h a i r s E. :Scapes u s u a l l y 0.5m or. .less-. .^ S. l a t i p e t i o l a t a EE. Scapes u s u a l l y more t h a n 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 more or l e s s a b r u p t l y i n t o b l a d e s ; v e s t i t u r e i n i n f l o r e s e n c e w i t h r e d d i s h g l a n d s S_. i n t e g r i f o l i a
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Systematics of Saxifraga rufidula and related species from the Columbia River gorge to southwestern British Columbia
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