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Form and structure of variation within lineages of Wyethia and Balsamorhiza Robson, Kathleen Anne 1985

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e FORM AND STRUCTURE OF VARIATION WITHIN LINEAGES OF WYETHIA AND BALSAMORHIZA by KATHLEEN ANNE ROBSON B.S., U n i v e r s i t y of C a l i f o r n i a at Davis, 1983 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in THE FACULTY OF GRADUATE STUDIES Botany Department We accept t h i s t h e s i s as conforming to the r e q u i r e d standard THE UNIVERSITY OF BRITISH COLUMBIA December 1985 © Kathleen Anne Robson, 1985 In p resen t ing this thesis in partial fu l f i lment of the requ i rements for an a d v a n c e d d e g r e e at t h e Univers i ty o f Br it ish C o l u m b i a , I agree that t h e Library shall m a k e it f reely avai lable for re fe rence a n d s tudy . I further agree that p e r m i s s i o n for ex tens ive c o p y i n g o f this thesis fo r scho lar ly p u r p o s e s may be g ranted by the h e a d o f m y d e p a r t m e n t o r by his o r her representat ives , it is u n d e r s t o o d that c o p y i n g o r p u b l i c a t i o n o f this thesis fo r f inanc ia l gain shall no t b e a l l o w e d w i t h o u t m y w r i t t e n p e r m i s s i o n . D e p a r t m e n t of T h e Un ivers i ty o f Brit ish C o l u m b i a 1956 M a i n M a l l V a n c o u v e r , C a n a d a V 6 T 1Y3 DE-6(3 /81) i i A b s t r a c t T h i s study i l l u s t r a t e s a method of comparing m o r p h o l o g i c a l aspects of w i t h i n - s p e c i e s v a r i a t i o n to s p e c i a t i o n events w i t h i n l a r g e r l i n e a g e s i n an e f f o r t t o compare the v a r i a t i o n which has a r i s e n as a r e s u l t of ontogeny to that which has been expressed d u r i n g phylogeny. The source and s t r u c t u r e of v a r i a t i o n at these d i f f e r e n t l e v e l s of b i o l o g i c a l o r g a n i z a t i o n i n the genera Wyethia and Balsamorhiza (Asteraceae, Heliantheae) were s t u d i e d using d i f f e r e n t a n a l y t i c a l t echniques. T h i s approach attempts to i l l u s t r a t e how a deeper understanding of the nature of b i o l o g i c a l change over time might be gained when these h i e r a r c h i c a l l y r e l a t e d l e v e l s of v a r i a t i o n are analyzed with a p p r o p r i a t e techniques and compared. T h i s study demonstrates a new and p o t e n t i a l l y v a l u a b l e a p p l i c a t i o n of d i f f e r e n t methodologies to the examination of e v o l u t i o n a r y p r o c e s s e s . The r e l a t i o n s h i p s w i t h i n and between Wyethia and Balsamorhiza have presented problems i n the past and an attempt was made to c l a r i f y the e v o l u t i o n a r y p a t t e r n s among s p e c i e s . The methods of p h y l o g e n e t i c s y s t e m a t i c s were employed in order to gain a b e t t e r understanding of these h i s t o r i c a l r e l a t i o n s h i p s . I t was found that taxonomic outgroups were v a l u a b l e mainly i n t h e i r a b i l i t y to d e l i m i t small c l a d e s , which were then used as f u n c t i o n a l outgroups. The use of f u n c t i o n a l outgroups p e r m i t t e d a l a r g e r number of v a r i a b l e s to be used, i n c l u d i n g a more d e t a i l e d set of m o r p h o l o g i c a l c h a r a c t e r s and data d e r i v e d from f l a v o n o i d c hemistry. More data and the r e s t of the taxa are r e q u i r e d before r e l a t i o n s h i p s among a l l s p e c i e s can be f u l l y c l a r i f i e d . Small monophyletic groups of s p e c i e s were i d e n t i f i e d and confirmed from the r e s u l t s of the c l a d i s t i c a n a l y s e s . These c l a d e s were used to pro v i d e a framework w i t h i n which a the r e s u l t s of the a n a l y s i s of w i t h i n - s p e c i e s v a r i a t i o n c o u l d be viewed. The second p a r t of t h i s study examined morphological v a r i a t i o n based on measurements taken from i n d i v i d u a l p l a n t s . Techniques based on p r i n c i p a l components a n a l y s i s were employed to explore t h i s v a r i a t i o n f o r s t r u c t u r e and source. V a r i a t i o n i n separate data s e t s d e s c r i b i n g heads, shoots and leaves was apportioned to taxonomic l e v e l s by a n a l y s i s of v a r i a n c e of PCA sc o r e s . Most of the v a r i a t i o n was a t t r i b u t a b l e to groups of r e l a t e d s p e c i e s (genera and s e c t i o n s ) or to the s p e c i e s themselves. P o p u l a t i o n s accounted f o r l i t t l e of the o v e r a l l v a r i a t i o n . 95% c o n f i d e n c e e l l i p s e s t a t i s t i c s were c a l c u l a t e d f o r po i n t swarms d e s c r i b i n g each s p e c i e s . The e l l i p s e r e s u l t s were t r a n s l a t e d i n t o graphic r e p r e s e n t a t i o n s of v a r i a t i o n and v a r i a b l e c o r r e l a t i o n s t r u c t u r e from the head, shoot and l e a f data f o r each s p e c i e s . These g r a p h i c s were then p r o j e c t e d onto the branch t i p s of cladograms so that w i t h i n - s p e c i e s v a r i a t i o n c o u l d be compared among s p e c i e s and l i n e a g e s composed of s p e c i e s . When w i t h i n - s p e c i e s v a r i a t i o n was examined f o r changes i n source and s t r u c t u r e i t was found that much of i t appears to be a t t r i b u t a b l e to the h i s t o r y of a c l a d e . In a d d i t i o n , some unique and unexpected aspect of v a r i a t i o n w i t h i n each s p e c i e s i s always apparent and may be r e l a t e d to the s p e c i a t i o n event i t s e l f . That i s , the nature and source of ontogenetic v a r i a t i o n i v appears to be r e l a t e d to p h y l o g e n e t i c v a r i a t i o n . The v a r i a t i o n q u a n t i f i e d from d i f f e r e n t h i e r a r c h i c a l l e v e l s of b i o l o g i c a l o r g a n i z a t i o n appears to demonstrate s i m i l a r p r o p e r t i e s of h i s t o r i c a l c onservatism and emergence. These r e s u l t s are examined i n the l i g h t of c u r r e n t t h e o r i e s which address the nature of b i o l o g i c a l change over time and are found to be best e x p l a i n e d by a new theory of e v o l u t i o n based on n o n e q u i l i b r i u m thermodynamics. V Table of Contents A b s t r a c t i i L i s t of Tables v i L i s t of F i g u r e s v i i Acknowledgement v i i i Chapter I INTRODUCTION 1 Chapter II MATERIALS AND METHODS 6 1 . P l a n t s 6 2. Data . . .'• 9 A. Measured V a r i a b l e s 9 i . C a u l i n e Leaf V a r i a b l e s 10 i i . Shoot V a r i a b l e s 11 i i i . Head V a r i a b l e s 11 B. C a t e g o r i c a l V a r i a b l e s 12 i . M o r p h o l o g i c a l V a r i a b l e s 12 i i . F l a v o n o i d Chemistry 13 3. Analyses 17 A. Phy l o g e n e t i c Systematics 17 B. E x p l o r a t o r y Data A n a l y s i s 19 Chapter I I I RESULTS AND DISCUSSION 24 v i 1. P h y l o g e n e t i c Systematics 24 A. R e s u l t s 24 B. D i s c u s s i o n 27 2. E x p l o r a t o r y Data A n a l y s i s 33 A. Sources of V a r i a t i o n 33 B. Nature of Wi t h i n - S p e c i e s V a r i a t i o n 36 Chapter IV SUMMARY AND EXPLANATIONS 43 1 . Summary 43 A. Phy l o g e n e t i c Systematics 43 B. E x p l o r a t o r y Data A n a l y s i s 44 2. E x p l a n a t i o n s 46 Chapter V EPILOG 56 Chapter VI TAXONOMIC DESCRIPTIONS 61 I . Wyethia 61 I I . Balsamorhiza * 74 I I I . Taxonomic Outgroups 81 LITERATURE CITED 89 APPENDIX A - PLANT COLLECTIONS 95 v i i L i s t of Tables Table T . - Species and I n d i v i d u a l s 105 Table I I . - C h a r a c t e r s f o r Taxonomic Outgroups 106 Table I I I . - C h a r a c t e r s f o r A r t o r h i z a 107 Table IV. - C h a r a c t e r s f o r Wyethia r e t i c u l a t a / e l a t a ...108 Table V. - L i s t of F l a v o n o i d s 109 Table VI. - R e s u l t s of PCA 110 Table V I I . - Sources of V a r i a t i o n 111 v i i i L i s t of F i g u r e s F i g u r e 1. - EDA Measurements 112 F i g u r e 2. - F l a v o n o i d C l a s s e s and S u b s t i t u t i o n s 113 F i g u r e 3. - Cladogram with H e l i a n t h e l l a 114 F i g u r e 4. - Cladogram with E n c e l i a 115 F i g u r e 5. - Cladogram with H e l i o p s i s 116 F i g u r e 6. - Cladogram with F l o u r e n s i a 117 F i g u r e 7. - Cladogram with Rudbeckia 118 F i g u r e 8. - Cladogram with A r t o r h i z a 119 F i g u r e 9. - Cladogram with Wyethia r e t i c u l a t a / e l a t a ...120 F i g u r e 10. - Dry F l i e s of Species 121 F i g u r e 11. - Dry F l i e s of Species 122 F i g u r e 12. - Dry F l i e s of Species 123 F i g u r e 13. - Dry F l i e s and Cladograms . ... ...... 1 24 F i g u r e 14. - Dry F l i e s and Cladograms 125 ix Acknowledgement I am extremely g r a t e f u l to my s u p e r v i s o r , Jack Maze, and committee members Dan Brooks and Gary B r a d f i e l d f o r t h e i r h e l p , t h e i r senses of humor and, e s p e c i a l l y , f o r t h e i r r e c o g n i t i o n and acceptance of the f a c t that I am one of those people who, l i k e the song says, "need t h e i r r e i n s l a i d on 'em l o o s e " . I t i s obvious to me that my f a t e i n t h i s department would have been grim indeed had Jack not been here. He has been a genuine f r i e n d and mentor. Dan's remarkable d i s c o v e r y that one can put two and two together i n b i o l o g y and come up with emergence has been of c e n t r a l importance to my work. As well,- t h i s study would not have been p o s s i b l e without Rob Scagel's a n a l y t i c a l wizardry. Jack and Rob p r o v i d e d the p e r f e c t atmosphere by combining great r e s p e c t f o r s c i e n c e and c r e a t i v i t y with healthy iconoclasm. I t ' s a p i t y such a t t i t u d e s are so r a r e . I would l i k e to thank W i l f N i c h o l l s f o r h i s h e l p i n the f i e l d , Susan McCormick f o r her s k i l l as a chemist, and both of them f o r t h e i r support d u r i n g t r y i n g times. C V . Finnegan's H i s t o r y and Philosophy of B i o l o g y p r o v i d e d a haven of t h o u g h t f u l n e s s and wisdom in the shallow ocean of "modern" b i o l o g y . His was the s i n g l e most worthwhile course I have ever attended. I would a l s o l i k e to thank a l l the graduate students i n Botany and Zoology who have now become my good f r i e n d s . T h e i r w i l l i n g n e s s to d i s c u s s a n y t h i n g , t h e i r admirable c a p a c i t y f o r beer and t h e i r genuine i n t e r e s t i n X s c i e n c e have made my time here not only bearable, but great fun. The people acknowledged here have shown p a t i e n c e with my bad temper and t o l e r a n c e toward my q u e s t i o n a b l e s a n i t y . I cannot express my d e l i g h t i n having c r o s s e d paths with them a l l . 1 I. INTRODUCTION T h i s study has attempted to q u a n t i f y , r e l a t e and e x p l a i n the v a r i a t i o n found w i t h i n and among s p e c i e s and c l a d e s of the genera Wyethia and Balsamorhiza (Asteraceae, H e l i a n t h e a e ) . For these purposes, d i f f e r e n t m u l t i v a r i a t e techniques were used i n order to assess the v a r i a t i o n present at d i f f e r e n t l e v e l s of b i o l o g i c a l o r g a n i z a t i o n so that a l l r e s u l t s c o u l d be compared. The conceptual b a s i s of such an approach r e s t s on the b e l i e f that e v o l u t i o n a r y phenomena are manifested through these h i e r a r c h i c a l l y r e l a t e d l e v e l s of p h y l o g e n e t i c and ontogenetic d i f f e r e n t i a t i o n . The p a t t e r n s expressed at and above the l e v e l of s p e c i e s were summarized and c l a r i f i e d f o l l o w i n g the methods of p h y l o g e n e t i c s y stematics (Wiley, 1981). A separate assessment of w i t h i n - s p e c i e s v a r i a t i o n was d e r i v e d using methods based on p r i n c i p a l components a n a l y s i s . The comparison of r e s u l t s based on these d i f f e r e n t methods i l l u s t r a t e s a new and p o t e n t i a l l y worthwhile approach to the study of v a r i a t i o n at d i f f e r e n t l e v e l s of b i o l o g i c a l o r g a n i z a t i o n . The advantage of t h i s combined use of techniques i s that i t permits e v o l u t i o n a r y processes to be s t u d i e d i n g r e a t e r depth than any one method used alone c o u l d a c h i e v e . T h i s approach may permit more d i r e c t q u e s t i o n s to be asked of the r e l a t i o n s h i p s between ontogeny and phylogeny. One i s s u e addressed i n t h i s study regards the monophyly of Wyethia with respect to Balsamorhiza. J u s t i f i c a t i o n f o r the s e p a r a t i o n of these two taxa has been questioned i n the p a s t , 2 most notably by Weber (1946). In h i s monograph of Wyethia, Weber i n c l u d e d a d i s c u s s i o n of Balsamorhiza and commented on i t s apparent c l o s e n e s s to Wyethia. The s p e c i e s i n c l u d e d i n these genera are a l l herbaceous, taprooted p e r e n n i a l s r e s t r i c t e d to areas west of the Rocky Mts. from southern Canada to northern Mexico. They are a l s o s i m i l a r i n that they possess p i s t i l l a t e , f e r t i l e ray f l o r e t s , a c o n d i t i o n that i s not common i n most other c l o s e r e l a t i v e s . Weber's study i n c l u d e d chromosome counts from r e p r e s e n t a t i v e s of both genera. He found n=l9 or 2n=38 f o r a l l s p e c i e s surveyed. P r i o r to Weber's study, Sharp (1935) had i n c l u d e d Balsamorhiza i n h i s d e s c r i p t i o n of s e v e r a l epappose composite genera. Weber m o d i f i e d t h i s taxonomic grouping somewhat, arguing that the lac k of a pappus was not, by i t s e l f , s u f f i c i e n t to d e f i n e a genus. He concluded that two of Sharp's s p e c i e s d i d not belong i n Balsamorhiza. Balsamorhiza invenusta and B. b o l a n d e r i were subsequently removed from that genus and p l a c e d i n Weber's s e c t i o n Agnorhiza of Wyethia. Although a l l other members of Wyethia possess some s o r t of a pappus, other f e a t u r e s of morphology and d i s t r i b u t i o n suggested to him that the i n c l u s i o n of these s p e c i e s i n h i s s e c t i o n Agnorhiza was a more n a t u r a l grouping. Weber l e f t Wyethia i n t a c t as a genus separate from Balsamorhiza, but he q u e s t i o n e d the n a t u r a l n e s s of t h i s grouping: " 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 Wyethia i s more c l o s e l y a k i n to Balsamorhiza than to any other e x i s t i n g genus, d i f f e r i n g only i n the leafy-stemmed h a b i t . In f a c t , i f the data now at hand had been a v a i l a b l e at the time of the o r i g i n a l 3 d e f i n i t i o n of the genera, i t i s q u i t e p o s s i b l e that the two would have been regarded as a s i n g l e genus with two very w e l l -marked subgenera. The presence of connecting l i n k s between the pappose and epappose c o n d i t i o n i n B. s a g i t t a t a (Sharp,1935) and B. macrophylla would support such an i n t e r p r e t a t i o n , and the s i m i l a r karyotype would be a d d i t i o n a l c o n f i r m i n g evidence." For the p a r t of t h i s study which examined r e l a t i o n s h i p s among s p e c i e s , the methods of p h y l o g e n e t i c s y s t e m a t i c s were employed to c o n s t r u c t and t e s t e v o l u t i o n a r y hypotheses. The f o u r t e e n s p e c i e s of Wyethia were i n c l u d e d i n these a n a l y s e s , along with s p e c i e s of Balsamorhiza. S e v e r a l taxonomic outgroups were used in i n i t i a l attempts to p o l a r i z e the c h a r a c t e r s and to i d e n t i f y n a t u r a l groups w i t h i n the genera. Some of the c l a d e s which were confirmed as monophyletic in the i n i t i a l a n a l y s es and which appeared to be r e l a t i v e l y p l e s i o m o r p h i c , were used as f u n c t i o n a l outgroups in l a t e r runs (Watrous and Wheeler,1984). T h i s p e r m i t t e d more c h a r a c t e r s to be i n c l u d e d i n attempts to achieve b e t t e r r e s o l u t i o n of p a t t e r n s of s p e c i a t i o n . The use of a d i f f e r e n t set of data, based on measurements taken fron i n d i v i d u a l s , and methodology f a l l i n g under the g e n e r a l heading of e x p l o r a t o r y data a n a l y s i s (Hoaglin, et_ a l . , 1983,1985). per m i t t e d the assessment of v a r i a t i o n at a d i f f e r e n t l e v e l . The data were separated i n t o three groups of v a r i a b l e s d e s c r i b i n g f l o w e r i n g heads, the shoots upon which they develop and the leaves which a r i s e from those same shoots. These s e t s of data were analyzed s e p a r a t e l y so that t h e i r r e s u l t s c o u l d be compared. T h i s p a r t of the study employed p r i n c i p a l components 4 a n a l y s i s , a n a l y s i s of v a r i a n c e and the c a l c u l a t i o n of 95% confi d e n c e e l l i p s e s t a t i s t i c s ( J o l i c o e u r , 1968; J o l i c o e u r and Mosimann, 1960; Scagel, et a l . , 1985), based on the r e s u l t s of the PCAs. A n a l y s i s of v a r i a n c e was used to ap p o r t i o n the v a r i a t i o n present i n the s e t s of morphological data to l e v e l s i n a taxonomic h i e r a r c h y . That i s , I wished to know where, given the l e v e l s of genus, s e c t i o n , s p e c i e s or p o p u l a t i o n , the m a j o r i t y of the v a r i a t i o n o c c u r s . L e v e l s which account f o r l a r g e amounts of v a r i a t i o n i n a data set may a l s o be viewed as sources of d i f f e r e n t i a t i o n , or as p l a c e s where o r g a n i z a t i o n changes and organisms d i v e r g e . These are the l e v e l s where e v o l u t i o n a r y processes are assumed to be most s t r o n g l y manifested, given the p l a n t s t r u c t u r e s represented and the v a r i a b l e s d e f i n i n g them. Using these same se t s of data, f u r t h e r analyses based on 95% c o n f i d e n c e e l l i p s e s t a t i s t i c s were performed to examine other aspects of w i t h i n - s p e c i e s v a r i a t i o n . The r e s u l t s of these a n a l y s e s were made graphic and p r o j e c t e d onto the branch t i p s of cladograms d e f i n i n g l i n e a g e s of s p e c i e s . V a r i a t i o n w i t h i n s p e c i e s was viewed w i t h i n the context of v a r i a t i o n among s p e c i e s w i t h i n and among s m a l l , monophyletic groups. I wished to know what p a t t e r n s of v a r i a t i o n or o r g a n i z a t i o n were r e v e a l e d w i t h i n and among s p e c i e s when the r e s u l t s of these d i f f e r e n t techniques were compared. F i n a l l y , I wished to compare the explanatory c a p a b i l i t i e s of d i f f e r e n t t h e o r i e s c l a i m i n g to address the causes of b i o l o g i c a l o r g a n i z a t i o n and change over time when presented with 5 these r e s u l t s . 6 I I . MATERIALS AND METHODS 1. PLANTS The s p e c i e s concept r e c o g n i z e d here i s d e f i n e d a f t e r Wiley (1981) as a group of organisms which share membership i n a unique l i n e a g e and maintain a group i d e n t i t y . Groups of p l a n t s were excluded from t h i s study i f the c h a r a c t e r i s t i c s d e f i n i n g them seemed unc l e a r or i n d i s p u t e . Although h y b r i d s were commonly encountered i n the f i e l d , they were e a s i l y r ecognized and were not i n c l u d e d i n any of the a n a l y s e s . The numbers of i n d i v i d u a l s r e p r e s e n t i n g each s p e c i e s from which a c t u a l measurements were taken are shown i n Table I. D e s c r i p t i o n s of c o l l e c t i n g s i t e s may be found i n Appendix A. Vouchers f o r a l l s p e c i e s have been d e p o s i t e d i n the U n i v e r s i t y of B r i t i s h Columbia Herbarium (UBC). A minimum of f i v e p l a n t s were c o l l e c t e d from each s i t e (a " p o p u l a t i o n " , as d e f i n e d here) and were assumed to be i n t e r b r e e d i n g . These samples may have been too small to allow a f u l l assessment to be made of v a r i a t i o n w i t h i n and among i n d i v i d u a l s and p o p u l a t i o n s . Nonetheless, the c o l l e c t i o n s were assumed to provide g e n e r a l l y s u f f i c i e n t m a t e r i a l f o r the analyses performed here. Each i n d i v i d u a l was represented by one f l o w e r i n g shoot at a n t h e s i s . Basal leaves were i n c l u d e d , when present, but data r e g a r d i n g them were not i n c l u d e d i n the analyses based on PCA. A n t h e s i s was d e f i n e d as f u l l y opened t e r m i n a l heads with anthers and s t y l e branches ex e r t e d i n at l e a s t the outermost whorl of d i s k f l o r e t s . T h i s r e s t r i c t i o n was p l a c e d on the 7 m a t e r i a l sampled so that a l l shoots would be at approximately the same stage of development. In most cases, c o l l e c t i o n s were made such that the heads t e r m i n a t i n g the c e n t r a l a x i s of the shoot met these c r i t e r i a . In a few i n s t a n c e s , however, the c e n t r a l heads were j u s t past f l o w e r i n g . T h i s had an e f f e c t only on the v a r i a b l e d e s c r i b i n g numbers of ray f l o r e t s and i s d i s c u s s e d f u r t h e r i n the s e c t i o n on measured v a r i a b l e s . A l l s p e c i e s s t u d i e d are taprooted p e r e n n i a l s with s i m i l a r root systems, so v a r i a b l e s d e s c r i b i n g r o o t s were excluded here. A c o l l e c t i o n i n c l u d i n g whole p l a n t s with roots would have been u n e c e s s a r i l y cumbersome to s t o r e and t r a n s p o r t . As w e l l , I had no d e s i r e to k i l l i n d i v i d u a l s , e s p e c i a l l y when they were members of a r a r e or endemic s p e c i e s . During the sampling, an attempt was made to represent d i f f e r e n t areas of each s p e c i e s range. For some of the widespread taxa t h i s was not d i f f i c u l t and i s r e f l e c t e d i n the number of p o p u l a t i o n s r e p r e s e n t i n g them. Most of the s p e c i e s of Wyethia and Balsamorhiza were represented by two to seven p o p u l a t i o n s (see Table I and Appendix A). A l l s p e c i e s of Wyethia were c o l l e c t e d , but u n f o r t u n a t e l y , W. ovata, W. invenusta and W. scabra were only encountered once. These taxa are t h e r e f o r e represented by only f i v e i n d i v i d u a l s each. Balsamorhiza was represented by seven s p e c i e s , which c e r t a i n l y d i d not account f o r the range of v a r i a t i o n i n the genus (see Taxonomic D e s c r i p t i o n s ) . I t was assumed, however, that the c o l l e c t i o n s were s u f f i c i e n t to assess w i t h i n - s p e c i e s v a r i a t i o n and to gain some f u r t h e r understanding of the 8 ph y l o g e n e t i c r e l a t i o n s h i p s among s p e c i e s of Wyethia and Balsamorhiza. The three s p e c i e s p l a c e d i n s e c t i o n A r t o r h i z a were i n c l u d e d here, but B. d e l t o i d e a and B. careyana were represented by only f i v e i n d i v i d u a l s each. Four s p e c i e s of s e c t i o n Balsamorhiza were examined, a l l with a sample s i z e g r e a t e r than f i v e except B. s e r r a t a . One of the c o l l e c t i o n s i n c l u d e d as Balsamorhiza incana was obtained from a s i t e suppossedly to the south of the range of t h i s taxon,(see Taxonomic D e s c r i p t i o n s and Appendix A) but these p l a n t s were m o r p h o l o g i c a l l y i n d i s t i n g u i s h a b l e from the s p e c i e s . The four other taxa l i s t e d i n Table I represent some of the c l o s e r e l a t i v e s of Wyethia and Balsamorhiza. These s p e c i e s , with the exception of H e l i a n t h u s c u s i c k i i represent genera which were i n c l u d e d as taxonomic outgroups i n the p h y l o g e n e t i c a n a l y s e s . Data from the outgroup s p e c i e s l i s t e d i n Table I were i n c l u d e d in the assessment of w i t h i n - s p e c i e s v a r i a t i o n . T h i s was done i n order to compare v a r i a t i o n expressed w i t h i n s p e c i e s of the study group to v a r i a t i o n w i t h i n r e p r e s e n t a t i v e outgroup s p e c i e s . The ch o i c e of these p a r t i c u l a r outgroup taxa was made based s o l e l y on t h e i r convenience of c o l l e c t i o n . Species which are sympatric with Wyethia and Balsamorhiza were c o l l e c t e d when encountered. These outgroups s p e c i e s , as w e l l as the a d d i t i o n a l outgroup genera used i n the p h y l o g e n e t i c a n a l y s e s , are a l l o u t l i n e d i n Taxonomic D e s c r i p t i o n s . 9 2 . DATA A l l v a r i a b l e s used i n t h i s study belong to one of two major groups, d e f i n e d as f o l l o w s . Measured v a r i a b l e s c o n s i s t e d of those t h i n g s that c o u l d be counted or measured on organisms. They i n c l u d e both continuous and d i s c r e t e v a r i a b l e s and were used f o r the e x p l o r a t o r y data a n a l y s i s s e c t i o n of t h i s study. C a t e g o r i c a l v a r i a b l e s d i f f e r e d i n that c h a r a c t e r s t a t e s were chosen and the presence or g e n e r a l c o n d i t i o n of an organ or s t r u c t u r e was a s s i g n e d to one of these. V a r i a b l e s of t h i s type were used to c o n s t r u c t a p a r t i a l p h y l o g e n e t i c h y p o t h e s i s f o r the taxa of i n t e r e s t . The measured v a r i a b l e s were converted to c a t e g o r i c a l v a r i a b l e s by a s s i g n i n g a s t a t e of 0 or 1 to e i t h e r s i d e of t h e i r means in one of the c l a d i s t i c s runs. T h i s a n a l y s i s was not i n c l u d e d i n the r e s u l t s of t h i s study, however, because outgroups were represented by only a few s p e c i e s . T h i s was c o n s i d e r e d an i n s u f f i c i e n t sample upon which to base c h a r a c t e r assessment of outgroup genera. As w e l l , I d i d not have t h i s kind of d e t a i l e d m o r p h o l o g i c a l data f o r the other taxonomic outgroups employed in the p h y l o g e n e t i c a n a l y s i s . A. MEASURED VARIABLES These data c o n s i s t e d of continuous ( l e n g t h s and widths, in c e n t i m e t e r s ) and d i s c r e t e (numbers of t h i n g s ) v a r i a b l e s . 10 V a r i a b l e s of t h i s kind were used f o r the assessment of v a r i a t i o n based on p r i n c i p a l components a n a l y s i s . Information r e g a r d i n g b a s a l leaves or organs borne on a x i l l a r y shoots was excluded, s i n c e many s p e c i e s lack these s t r u c t u r e s . A l l i n d i v i d u a l s used in t h i s p a r t of the study had at l e a s t two leaves a t t a c h e d to the main stem of the f l o w e r i n g shoot, and only these were compared d i r e c t l y (as i l l u s t r a t e d i n F i g u r e 1). Measurements of p h y l l a r i e s were taken only from the flower head a r i s i n g d i r e c t l y from the apex of the main shoot, when more than one i n f l o r e s c e n c e was present. T h i s r e s t r i c t i o n was a l s o p l a c e d on the v a r i a b l e d e s c r i b i n g numbers of ray f l o r e t s , u n l e s s those of the c e n t r a l head had f a l l e n and been l o s t d u r i n g the c o l l e c t i n g . For heads, as f o r l e a v e s , the attempt was made to a v o i d any assumptions of homology between i n f l o r e s c e n c e s t e r m i n a t i n g a c e n t r a l shoot and those a r i s i n g from a x i l l a r y buds. The data gathered from the f l o w e r i n g shoots was d i v i d e d i n t o three p a r t s f o r analyses i n v o l v i n g apportionment of v a r i a t i o n and the d e s c r i p t i o n s of w i t h i n - s p e c i e s v a r i a t i o n . D e s c r i p t i o n s of v a r i a b l e s and t h e i r acronyms f o l l o w . i . C a u l i n e Leaf V a r i a b l e s MDLN1 and MDLN2: Lengths of blades of leaves l o c a t e d approximately midway up the a x i s of the shoot, i f more than two leaves were pr e s e n t . These were measured from the apex to the most proximal p o i n t of attachment of the blade to the p e t i o l e . MDWD1 and MDWD2: The width of the same two c a u l i n e leaves at 11 t h e i r widest p o i n t s . MDPT1 and MDPT2: The lengths of the p e t i o l e s of the same two c a u l i n e l e a v e s . i i . Shoot V a r i a b l e s These v a r i a b l e s were i n c l u d e d as a d i s c r e t e set of data i n the analyses s i n c e I wished to examine the v a r i a t i o n i n numbers of organs which arose from a p a r t i c u l a r shoot s e p a r a t e l y from the measurements of the organs themselves. I t d i d not seem reasonable to assume that the number of l e a f or a x i l l a r y shoot meristems i n i t i a t e d on a p a r t i c u l a r a x i s was n e c e s s a r i l y s t r o n g l y c o r r e l a t e d with the v a r i a b l e s d e s c r i b i n g the mature organs which had a r i s e n from those meristems. #OPP: The t o t a l number of o p p o s i t e l y arranged leaves on the shoot. #ALT: The t o t a l number of a l t e r n a t e l y arranged leaves on the shoot. STHT: The height of the main stem a x i s from i t s p o i n t of attachment at the caudex to the base of the c e n t r a l flower head. #HDS: The t o t a l number of flower heads present on a shoot, i n c l u d i n g both t e r m i n a l and a x i l l a r y i n f l o r e s c e n c e s . i i i . Head V a r i a b l e s With the exce p t i o n noted below, a l l measurements were taken from the c e n t r a l head t e r m i n a t i n g the main a x i s of the f l o w e r i n g shoot. HDWD: The width of the t e r m i n a l head, measured acr o s s the base of the r e c e p t a c l e . R A Y # : The number of ray f l o r e t s present 12 on the t e r m i n a l head. In a couple of i n s t a n c e s , these were l o s t d u r i n g c o l l e c t i n g because the c e n t r a l heads were almost past a n t h e s i s . P l a n t s i n such c o n d i t i o n are noted i n the c o l l e c t i n g notes i n Appendix A. In these cases, ray counts were taken e i t h e r from a x i l l a r y heads, i f present, or means were taken from the r e s t of the i n d i v i d u a l s i n the c o l l e c t i o n . PHLN1, PHLN2, PHNL3: The l e n g t h from base to apex of three of the outermost p h y l l a r i e s of the t e r m i n a l head. PHWD1, PHWD2, PHWD3: The width at t h e i r widest p o i n t f o r the same three p h y l l a r i e s . PH#,: The t o t a l number of p h y l l a r i e s f o r the t e r m i n a l head. B. CATEGORICAL VARIABLES i . M o r p h o l o g i c a l V a r i a b l e s An i n i t i a l set of mor p h o l o g i c a l v a r i a b l e s was used f o r the study group and f i v e outgroup genera. These are l i s t e d and d e s c r i b e d i n Table I I . Because of u n c e r t a i n a f f i n i t i e s and l i m i t e d data each outgroup was employed independently so that the r e s u l t i n g t r e e s c o u l d be compared (Donoghue and Cantino, 1984). U n f o r t u n a t e l y , the number of c h a r a c t e r s t h a t c o u l d be used f o r a l l members of the study group, as w e l l as f o r outgroups, was q u i t e s m a l l . I t was o f t e n the case that c h a r a c t e r s which v a r i e d among ingroup s p e c i e s seemed to' vary s i m i l a r l y w i t h i n outgroup genera. Because of t h i s , assessments of p o l a r i t y were o f t e n i m p o s s i b l e to make, given the l i m i t e d 13 data at hand. These problems are d i s c u s s e d i n g r e a t e r depth i n a l a t e r s e c t i o n . I n i t i a l a n a l y s es employing taxonomic outgroups were s u f f i c i e n t t o i d e n t i f y small monophyletic groups of s p e c i e s w i t h i n Wyethia and Balsamorhiza. In an attempt to improve the r e s o l u t i o n of the r e l a t i o n s h i p s among s p e c i e s , these c l a d e s were employed as f u n c t i o n a l outgroups (Watrous and Wheeler,1981) i n f u r t h e r a n a l y s e s . T h i s p e r m i t t e d the use of a l a r g e r data set c o n s i s t i n g of both morphological and chemical v a r i a b l e s . F u n c t i o n a l outgroups c o n s i s t e d of small c o l l e c t i o n s of taxa which were r e p e a t e d l y confirmed as monophyletic, r e g a r d l e s s of the o v e r a l l outcome of a given run. The taxa and c h a r a c t e r s used are shown i n Table III and Table IV. i i . F l a v o n o i d Chemistry For the purposes of t h i s study, only non-polar f l a v o n o i d aglycones (those l a c k i n g a t t a c h e d sugars) were examined. These were ob t a i n e d by washing l e a f m a t e r i a l i n non-polar s o l v e n t s so that only the contents of the g l a n d u l a r trichomes or the c u t i c l e would be d i s s o l v e d . The p o l a r f l a v o n o i d g l y c o s i d e s which r e s i d e in vacuoles are not e x t r a c t e d by t h i s method. These p o l a r compounds were b r i e f l y examined f o r one taxon only (Balsamorhiza  s a g i t t a t a ) and were found to c o n s i s t of g l y c o s i d e s of q u e r c e t i n and kaempferol. These f l a v o n o i d s are c o n s i d e r e d to be r a t h e r u b i q u i t o u s among angiosperms ( F a i r b r o t h e r s , et a l . , 1975). S t r u c t u r a l d i f f e r e n c e s of s u r f a c e f l a v o n o i d s among ta x a appeared 1 4 to provide more i n t e r e s t i n g and u s e f u l c h a r a c t e r s . Because of t h i s , the examination of i n t e r n a l l y sequestered f l a v o n o i d g l y c o s i d e s was not c o n s i d e r e d to be worthwhile and was not pursued f u r t h e r . F l a v o n o i d aglycones were e x t r a c t e d from a i r - d r i e d l eaves i n e i t h e r dichloromethane or c h l o r o f o r m . The e x t r a c t s were taken to dryness and chromatographed over P o l y c h l a r AT columns using dichloromethane:MeOH (3:1) and g r a d u a l l y i n c r e a s i n g amounts of MeOH. F r a c t i o n s from these columns were f u r t h e r separated on Sephadex LH-20 (MeOH) columns and on polyamide t h i n - l a y e r chromatograms. Compounds were clea n e d over Sephadex LH-20 p r i o r to s p e c t r a l a n a l y s i s . S t r u c t u r a l d e t e r m i n a t i o n s were made based on UV, 1HNMR and MS data, as w e l l as by co-chromatography a g a i n s t standard compounds. S e v e r a l compounds i s o l a t e d from some of the s p e c i e s of Wyethia were found to be new n a t u r a l products. The d e t a i l s of the s p e c t r a l i n f o r m a t i o n l e a d i n g to the c h a r a c t e r i z a t i o n of these new compounds i s given elsewhere (McCormick, et a l . , 1985a; McCormick, et a_l. , 1985b; McCormick, et a l . , i n p r e s s ) . These w i l l not be d i s c u s s e d f u r t h e r here, except as regards t h e i r use as c h a r a c t e r s . B a s i c s t r u c t u r e s r e p r e s e n t i n g c l a s s e s of f l a v o n o i d s and major s u b s t i t u t i o n p a t t e r n s are i l l u s t r a t e d i n F i g u r e 2. A l i s t i n g of the s p e c i e s of Wyethia and Balsamorhiza f o r which f l a v o n o i d s have been c h a r a c t e r i z e d i s shown in Table V. Some chemical c h a r a c t e r s may p r o v i d e good i n f o r m a t i o n with r e s p e c t to r e l a t i o n s h i p s , e s p e c i a l l y among s p e c i e s of Wyethia. 15 These i n c l u d e f l a v o n o i d s with b a s i c s t r u c t u r a l d i f f e r e n c e s , such as the B - r i n g attachment of i s o f l a v o n e s . The d i f f e r e n c e s i n bonding, p r o t o n a t i o n and h y d r o x y l a t i o n of A - r i n g carbons 1 and 2 , such as those seen i n f l a v o n o l s as opposed to flavanones, may a l s o be important. The a b i l i t y to p r o v i d e a f l a v o n o i d s k e l e t o n with a c e r t a i n s u b s t i t u e n t and the s p e c i f i c p a t t e r n s of met h y l a t i o n or p r e n y l a t i o n of compounds may provide u s e f u l i n f o r m a t i o n as w e l l . U n f o r t u n a t e l y , time and circumstances d i d not permit the f u l l e l u c i d a t i o n of f l a v o n o i d aglycones made by a l l s p e c i e s of Wyethia and Balsamorhiza. However, s i n c e r e p r e s e n t a t i v e s from a l l s e c t i o n s of both genera have been examined, g e n e r a l comparisons were made by co-chromatography f o r the remaining s p e c i e s . The only c h a r a c t e r s that c o u l d be obtained by t h i s method were the presence of flavanones and p r e n y l a t e d compounds. If any flavanones are present, one w i l l appear as a pink spot on a TLC a f t e r a p p r o p r i a t e treatment. I f p r e n y l a t e d compounds are present, some of these w i l l be flavanones and a s e r i e s of pink spots w i l l appear. Because of t h e i r c o l o r s , these compounds are not e a s i l y confused with o t h e r s , but some important chemical c h a r a c t e r s cannot be ev a l u a t e d i n t h i s way. I s o f l a v o n e s and methylated f l a v o n o l s are very d i f f e r e n t compounds, yet they may both appear as black spots at the non-polar end of a TLC p l a t e . I f e l t that chemical data of t h i s kind c o u l d not be i n c l u d e d in the present study, but t h i s i n f o r m a t i o n should prove v a l u a b l e when f l a v o n o i d s from the remaining taxa are f u l l y i d e n t i f i e d . An attempt was a l s o made to gather i n f o r m a t i o n on the 1 6 f l a v o n o i d s of outgroups from the l i t e r a t u r e , but there was not a great amount a v a i l a b l e . With the s i n g l e exception of F l o u r e n s i a , outgroup genera appear to produce only methylated f l a v o n o l s . Some s p e c i e s of F l o u r e n s i a produce p r e n y l a t e d flavanones (Bohlmann and Jakupovik, 1979) but only a few members of the genus have so f a r been examined. A v a i l a b l e i n f o r m a t i o n on outgroup chemistry seemed too sketchy, so these c h a r a c t e r s were excluded from use i n c l a d i s t i c a n a l y s i s employing taxonomic outgroups. 1 7 3. ANALYSES A. PHYLOGENETIC SYSTEMATICS The c h o i c e of outgroups f o r use i n c l a d i s t i c analyses may be a source of d i f f i c u l t y i n b o t a n i c a l s t u d i e s (Funk, 1981; Donoghue and Cantino, 1984). I t i s o f t e n i m p o s s i b l e to choose a s i n g l e , most c l o s e l y r e l a t e d outgroup. At best, the numerous p o s s i b i l i t i e s can be narrowed down to a few, and t h i s study proved to be no e x c e p t i o n . T e s t i n g the monophyly of Wyethia with re s p e c t to Balsamorhiza and f i n d i n g p o l a r i t i e s f o r c h a r a c t e r s r e q u i r e d the use of s e v e r a l outgroups and t h i s approach was s t i l l not f u l l y s u c c e s s f u l . I g e n e r a l l y chose outgroup taxa from among those which have been c o n s i d e r e d c l o s e to the study group by most workers. R e q u i r i n g that outgroups be members of the same s u b t r i b e as the taxa of i n t e r e s t would have been the simplest approach. U n f o r t u n a t e l y , the l i m i t s of the s u b t r i b e s of the H e l i a n t h e a e i s a s u b j e c t p r e s e n t l y i n d i s p u t e . In the two most recent treatments of the t r i b e H e l i a n t h e a e , Wyethia and Balsamorhiza remain t o g e t h e r , but i n d i f f e r e n t s u b t r i b e s . Stuessy (1977) p l a c e s them in h i s H e l i a n t h i n a e , while Robinson ( 1 9 8 1 ) t h i n k s they belong i n h i s E c l i p t i n a e . There are a few genera which have been r e p e a t e d l y p l a c e d c l o s e to Wyethia and Balsamorhiza by past and present workers. H e l i a n t h e l l a , H e l i o p s i s , E n c e l i a , F l o u r e n s i a and Rudbeckia are c o n s i d e r e d to 18 be c l o s e l y r e l a t e d to the taxa of i n t e r e s t . A l l were used here fo r outgroup comparison employing generic outgroups. I found that I was o f t e n very l i m i t e d i n the c h o i c e of c h a r a c t e r s which c o u l d be used with taxonomic outgroups. Presence, absence or c o n d i t i o n of the pappus, f o r i n s t a n c e , v a r i e s s i m i l a r l y among s p e c i e s of both ingroup and outgroup genera. I suspect that most s p e c i e s possess the i n f o r m a t i o n necessary to make some s o r t of a pappus, but the developmental c o n t r o l of t h i s c h a r a c t e r seems to vary g r e a t l y w i t h i n outgroups and study groups a l i k e . T h i s problem a l s o e x i s t s with other p o t e n t i a l l y important c h a r a c t e r s , such as l e a f arrangement, number of heads per shoot, presence of b a s a l l e a v e s , l e a f venation p a t t e r n s and o t h e r s . For Wyethia and Balsamorhiza, these c h a r a c t e r s tend to be s t a b l e enough to be i n c l u d e d as p a r t of the g e n e r i c or s e c t i o n a l d e f i n i t i o n . However, g e n e r i c d e s c r i p t i o n s f o r some of the outgroups i n c l u d e statements such as, "leaves o p p o s i t e , at l e a s t below", meaning that t h e i r arrangement may change to a l t e r n a t e toward the shoot apex. In such a s i t u a t i o n i t becomes very d i f f i c u l t to decide on a p l e s i o m o r p h i c s t a t e without being completely a r b i t r a r y . C h a r a c t e r s such as these were coded so that change d u r i n g e v o l u t i o n c o u l d be shown to have gone in e i t h e r d i r e c t i o n . T h i s i s obvious from Table I I , which d e s c r i b e s c h a r a c t e r s used in the i n i t i a l group of analyses employing taxonomic outgroups. Two f u n c t i o n a l outgroups were a l s o used in separate analyses so that a d d i t i o n a l c h a r a c t e r s c o u l d be i n c l u d e d . The f i r s t c o n s i s t e d of s e c t i o n A r t o r h i z a of Balsamorhiza and the 19 second was made up of the s p e c i e s p a i r Wyethia r e t i c u l a t a and W. e l a t a . Each of these f u n c t i o n a l outgroups had been r e p e a t e d l y confirmed as monophyletic by e a r l i e r a n a l y s e s . As w e l l , one or the other c l a d e u s u a l l y appeared to share the fewest number of synapomorphies with the other ingroup taxa, a c c o r d i n g to the r e s u l t s from the taxonomic outgroup comparisons. The employment of f u n c t i o n a l outgroups p e r m i t t e d the use of c h a r a c t e r s which c o u l d not be used with generic outgroups (Table III and Table IV ) . A l l p h y l o g e n e t i c analyses were performed using the systematics package PHYSYS ( F a r r i s and M i c k e v i c h , 1983). I wished to examine r e l a t i o n s h i p s among s p e c i e s i n c l u d e d i n these analyses and to see i f the ingroup genera and s e c t i o n s , as they are p r e s e n t l y d e f i n e d , appeared to be monophyletic. In a d d i t i o n , the second p a r t of t h i s study r e q u i r e d that small l i n e a g e s of s p e c i e s be i d e n t i f i e d and confirmed as monophyletic based on the r e s u l t s of the c l a d i s t i c a n a l y s e s . B. EXPLORATORY DATA ANALYSIS The a n a l y s e s i n c l u d e d under t h i s heading had two main purposes. The f i r s t was to i d e n t i f y taxonomic l e v e l s to which the v a r i a t i o n i n the d i f f e r e n t data s e t s c o u l d be a t t r i b u t e d . A second set of a n a l y s e s were performed i n an attempt to b e t t e r understand the r e l a t i o n s h i p s between s p e c i a t i o n and development 20 by d e s c r i b i n g the morp h o l o g i c a l v a r i a t i o n w i t h i n s p e c i e s and comparing i t to the v a r i a t i o n among them. In these a n a l y s e s , the data s e t s used were based on measurements taken from i n d i v i d u a l p l a n t s , u n l i k e the p h y l o g e n e t i c a n a l y s e s , which were based on c h a r a c t e r s possessed by a l l members of each s p e c i e s . The combined use of these methodologies has allowed an assessment to be made of the v a r i a t i o n present at d i f f e r e n t l e v e l s of b i o l o g i c a l o r g a n i z a t i o n . Time c o u l d not be d i r e c t l y r e l a t e d to developmental change i n the present study. N e v e r t h e l e s s , i t was assumed that ontogenetic processes are not completely d i v o r c e d from the f i n a l product of mature organism. I t seemed worthwhile, t h e r e f o r e , to expl o r e w i t h i n - s p e c i e s v a r i a t i o n i n as much d e t a i l as the a v a i l a b l e data would a l l o w . The data were d i v i d e d i n t o t h r e s s e t s of v a r i a b l e s d e s c r i b i n g shoots, leaves and heads. T h i s was done because I d i d not wish to assume that these s t r u c t u r e s , which a r i s e from d i f f e r e n t meristems (induced, l e a f and f l o r a l , r e s p e c t i v e l y ) would n e c e s s a r i l y demonstrate s i m i l a r p a t t e r n s of v a r i a t i o n . As w e l l , there are meristematic areas w i t h i n d e v e l o p i n g organs, such as l e a v e s , which may c o n t r i b u t e to the f i n a l form of the organ i n a r e l a t i v e l y independent f a s h i o n . Recent work suggests that v a r i a t i o n among d i f f e r e n t p l a n t s t r u c t u r e s may be a t t r i b u t e d to d i f f e r e n t sources, such as the i n d i v i d u a l p l a n t or the s t r u c t u r e i t s e l f ( S c a g e l , et a l . , 1985; Scagel,l984; Maze, et a_l., i n p r e s s ; Robson, e_t a l . , i n p r e p . ) . T h i s p a r t of the study centered around the use of p r i n c i p a l components a n a l y s i s (PCA) of a c o r r e l a t i o n m a t r i x . The 21 s t a t i s t i c a l package MIDAS (Fox and Gu i r e , 1976) and the program ANOVAR (Greig and O s t e r l i n , 1978), p r o v i d e d through the U n i v e r s i t y of B r i t i s h Columbia Computing Centre, were used f o r these a n a l y s e s . The three d i f f e r e n t data s e t s d e s c r i b i n g heads, shoots and lea v e s were analyzed independently so that t h e i r r e s u l t s c o u l d be compared. Normality of d i s t r i b u t i o n was checked using a graphic technique, cumulative frequency d i s t r i b u t i o n of st a n d a r d i z e d v a r i a b l e s (Zar, 1974). Four of the v a r i a b l e s , PHLN3, PHWD1, PHWD2 and PHWD3 were not normally d i s t r i b u t e d . Improvement of these d i s t r i b u t i o n s was achieved when a square root t r a n s f o r m a t i o n was performed on the f i r s t v a r i a b l e and a log t r a n s f o r m on the l a s t t h r e e . The PCA a x i s s c o r e s , f o r axes with eigenvalues g r e a t e r than one, were s u b j e c t e d to a n a l y s i s of v a r i a n c e i n order to ap p o r t i o n the v a r i a t i o n present to d i f f e r e n t taxonomic l e v e l s . I wished to assess the congruence between a c t u a l data p o l a r i t y , as expressed i n PCA a x i s s c o r e s , and v e c t o r s r e l a t e d to d i f f e r e n t h i e r a r c h i c a l l e v e l s . The model used f o r the a n a l y s i s was Y=A+B(A)+C(AB)+D(ABC)+e, where Y=PCA a x i s s c o r e s , A=the e f f e c t of genera, B=the e f f e c t of s e c t i o n s , C=the e f f e c t of s p e c i e s , D=the e f f e c t of p o p u l a t i o n s and e=the r e s i d u a l term. ANOVA was not used here f o r the purpose of t e s t i n g any p a r t i c u l a r hypotheses, but i n s t e a d to a p p o r t i o n v a r i a t i o n to these d i f f e r e n t h i e r a r c h i c a l l e v e l s u sing r a t i o s of sums of squares f o r PCA axes with eigenvalues g r e a t e r than 1. I a l s o wished to examine i n some d e t a i l the s t a t i s t i c s d e s c r i b i n g v a r i a t i o n and c o v a r i a t i o n w i t h i n the p o i n t swarm 22 r e p r e s e n t i n g each s p e c i e s . E c c e n t r i c i t y , theta and area f o r 95% confi d e n c e e l l i p s e s were c a l c u l a t e d (Scagel, et a l . , 1985; J o l i c o e u r , 1968; J o l i c o e u r and Mosimann, 1960) f o r each p o i n t swarm that d e s c r i b e d a s p e c i e s . T h i s was done s e p a r a t e l y f o r each data set r e p r e s e n t i n g f l o w e r i n g heads, stem leaves and shoots. Theta d e s c r i b e s the angle between the major v e c t o r through the p o i n t swarm f o r a s p e c i e s and a r e f e r e n c e a x i s r e l a t i v e to the e n t i r e data set (the f i r s t PCA a x i s ) . Theta o f f e r s an estimate of v a r i a b l e i n t e r - c o r r e l a t i o n s by c a l c u l a t i n g the p o s i t i o n of each p o i n t swarm i n v a r i a b l e space r e l a t i v e to a l l o t h e r s . The c a l c u l a t i o n of e c c e n t r i c i t y p r o v i d e s an estimate of the shape of the p o i n t swarm about the v e c t o r which summarizes i t . T h i s assesses the s t r e n g t h of v a r i a b l e c o r r e l a t i o n s , with 0 i n d i c a t i n g no c o r r e l a t i o n and 1 i n d i c a t i n g strong c o r r e l a t i o n s . The area of p o i n t swarms may be d e s c r i b e d as the r e l a t i v e s i z e of each, estimated w i t h i n the o v e r a l l v a r i a b l e space. R e l a t i v e l y l a r g e areas imply g r e a t e r v a r i a t i o n among those same v a r i a b l e s f o r a given p o i n t swarm, while small areas i n d i c a t e l i t t l e o v e r a l l v a r i a t i o n . These s t a t i s t i c s allow f u r t h e r d e s c r i p t i o n s of the data, as they give some i n d i c a t i o n of c o r r e l a t i o n p a t t e r n s of the v a r i a b l e s d e s c r i b i n g s i m i l a r s t r u c t u r e s among d i f f e r e n t s p e c i e s . They a l s o a l l o w some estimate to be made regarding the p a t t e r n of v a r i a b l e r e l a t i o n s h i p s f o r d i f f e r e n t s t r u c t u r e s w i t h i n a s i n g l e s p e c i e s . Using these techniques, r e l a t i v e d i f f e r e n c e s i n the i n t e r a c t i o n of v a r i a b l e s d e s c r i b i n g mature p l a n t s t r u c t u r e s 23 were assessed and compared. These estimates of the amount and nature of v a r i a t i o n w i t h i n s p e c i e s were represented g r a p h i c a l l y as "dry f l i e s " , which present a summary of these s t a t i s t i c s f o r each s p e c i e s as a s i n g l e i l l u s t r a t i o n . T h i s was done so that p a t t e r n s c o u l d be compared v i s u a l l y . The d e t a i l s of t h e i r c o n s t r u c t i o n are d e s c r i b e d i n the next chapter. I wished to see what comparisons c o u l d be made among d e s c r i p t i o n s of c o r r e l a t i o n s t r u c t u r e w i t h i n s p e c i e s when shown g r a p h i c a l l y at the branch t i p s of a proposed phylogeny. Only s m a l l , monophyletic groups of s p e c i e s were used i n these comparisons i n an attempt to determine how some of the w i t h i n -s p e c i e s v a r i a t i o n might show p a t t e r n or s t r u c t u r e w i t h i n the context of the h i s t o r y of a s m a l l l i n e a g e . Comparisons i n v o l v i n g l a r g e areas of cladograms, even when w e l l confirmed, would have been i n a p p r o p r i a t e because w i t h i n - s p e c i e s v a r i a t i o n was q u a n t i f i e d based only on mature s t r u c t u r e s . As w e l l , t h e t a and e c c e n t r i c i t y a s s i g n v a l u e s on a bounded s c a l e and t h e i r r e s u l t s , when shown a g a i n s t a l a r g e or complex phylogeny, c o u l d imply f a l s e r e v e r s a l s or p a r a l l e l i s m s . For these reasons, only small c l a d e s were used to p r o v i d e the framework of s p e c i a t i o n events w i t h i n which the v a r i a t i o n among mature p l a n t s t r u c t u r e s c o u l d be viewed. The d e s c r i p t i o n s of w i t h i n - s p e c i e s v a r i a t i o n and o r g a n i z a t i o n were compared among s p e c i e s and among small l i n e a g e s composed of s p e c i e s . 24 I I I . RESULTS AND DISCUSSION 1 . PHYLOGENETIC SYSTEMATICS A. RESULTS The r e s u l t s of these analyses are not c o n c l u s i v e with respect to the o v e r a l l r e l a t i o n s h i p s of Wyethia and Balsamorhiza. F i g u r e s 3-7 i l l u s t r a t e cladograms r e s u l t i n g from the use of taxonomic outgroups, while F i g u r e s 8 and 9 show the d i f f e r e n t r e s u l t s that were obtained when f u n c t i o n a l outgroups were employed. Tree lengths and c o n s i s t e n c y i n d i c e s are shown with each cladogram. The valu e s f o r c o n s i s t e n c y , or r a t i o s of the minimum number of t r a n s f o r m a t i o n s r e q u i r e d f o r e x p r e s s i o n of the raw data s e t s to the a c t u a l number of changes shown on the t r e e s , are not high f o r any of these cladograms. The lowest c o n s i s t e n c y i n d i c e s were generated with the use of f u n c t i o n a l outgroups, implying that changes i n c h a r a c t e r s t a t e s f o r d i f f e r e n t v a r i a b l e s are not s t r o n g l y c o r r e l a t e d . In g e n e r a l , the use of of each outgroup generated only a s i n g l e t r e e . In cases where more than one t r e e was generated, F - r a t i o s were c a l c u l a t e d and only the cladogram g i v i n g the lowest value i s shown (Brooks, et a l . , i n re v i e w ) . Based on the use of taxonomic outgroups and morp h o l o g i c a l data, the r e s u l t s suggest that Wyethia and Balsamorhiza are p a r a p h y l e t i c , implying that some members of Wyethia may have c l o s e r r e l a t i v e s among s p e c i e s of Balsamorhiza. Analyses based on the use of H e l i a n t h e l l a , E n c e l i a , H e l i o p s i s , F l o u r e n s i a and 25 Rudbeckia as taxonomic outgroups are shown in Figures 3-7. Figures 8 and 9 i l l u s t r a t e cladograms generated with functional outgroups. Section Artorhiza of Balsamorhiza i s shown as the outgroup, in Figure 8. In Figure 9 the functional outgroup was derived from the species pair Wyethia  reticulata/W. e l a t a . These two small clades were repeatedly confirmed as monophyletic based on results from analyses using taxonomic outgroups. In addition, one or the other functional outgroup was usually represented as having arisen early in the hypothesized lineage and i s shown close to the taxonomic outgroup. Analyses using functional outgroups permitted the use of more detailed morphological data. The presence of flavanones and prenyl substituents were also included as characters (Tables III and IV). The ov e r a l l results of analyses based on functional outgroups are quite d i f f e r e n t from those using taxonomic outgroups and the members of the present genera, with few exceptions, now appear to share a more recent common ancestry. In the cladogram which used the Wyethia reticulata/W. elata combination as a functional outgroup, (Figure 9) the members of Balsamorhiza do appear to form a monophyletic group, but part of Wyethia i s s t i l l paraphyletic with respect to them. In the cladogram using section Artorhiza as the outgroup,' (Figure 8) Wyethia and Balsamorhiza do not appear to be paraphyletic, although many of the relationships are s t i l l vague. The addition of new data, especially chemical characters, seems to provide stronger evidence of synapomorphies among many species presently 26 i n c l u d e d i n Wyethia. The r e s t of t h i s i n f o r m a t i o n i s appa r e n t l y needed before p h y l o g e n e t i c r e l a t i o n s h i p s among a l l the taxa can be c l a r i f i e d . Although a f u l l phylogeny c o u l d not be completed, many s p e c i e s are re p e a t e d l y shown to form monophyletic groups w i t h i n the present genera. For Wyethia, at l e a s t , these groups do not e n t i r e l y agree with the present s e c t i o n a l c l a s s i f i c a t i o n s . None of the s e c t i o n s appear to d e f i n e monophyletic groups w i t h i n Wyethia. I suspect that two of these s e c t i o n s , Wyethia and Agnorhiza, have been d e f i n e d almost e x c l u s i v e l y by the r e l a t i v e s i z e of the ranges of the s p e c i e s i n v o l v e d . The p o s s e s s i o n of a l a r g e range may provide a good c h a r a c t e r , but i t has been weighted too h e a v i l y . Other shared c h a r a c t e r s must be allowed to prov i d e evidence of r e l a t e d n e s s , as w e l l . Endemism c e r t a i n l y makes a u s e f u l key c h a r a c t e r , but i t does n e c e s s a r i l y r e v e a l much about e v o l u t i o n a r y r e l a t i o n s h i p s . I d i d not have r e p r e s e n t a t i v e s from a l l s p e c i e s p r e s e n t l y i n c l u d e d i n Balsamorhiza s e c t i o n Balsamorhiza, and so cannot present a f u l l e r p h y l o g e n e t i c hypothesis f o r t h i s genus. I d i d not wish to make guesses r e g a r d i n g s p e c i e s d e f i n i t i o n s and c o u l d not have defended the use of one treatment of s e c t i o n Balsamorhiza over another. I t i s not e n t i r e l y c l e a r why the d e f i n i t i o n s of s p e c i e s i n t h i s s e c t i o n have remained so vague, but a great d e a l of c o n f u s i o n c e r t a i n l y e x i s t s . The frequent h y b r i d i z a t i o n of any taxon i n the genus with any other one i s r e s p o n s i b l e f o r some of t h i s d i f f i c u l t y and some s p e c i e s may have a r i s e n i n this.way, adding to the c o n f u s i o n . In a d d i t i o n , 27 i t seems that many small p o p u l a t i o n s of p l a n t s now i n s e c t i o n Balsamorhiza e x h i b i t a great d e a l of v a r i a t i o n with r e s p e c t to other p o p u l a t i o n s of the same supposed s p e c i e s . Species d e f i n i t i o n s have been attempted s e v e r a l times (Sharp, 1935; Weber, 1946; Ownbey and Weber, 1943; C r o n q u i s t , 1955), but the c r i t e r i a by which s p e c i e s are d e f i n e d seems to vary g r e a t l y among aut h o r s . Some phenomenon, perhaps s p e c i a t i o n - i n - p r o g r e s s , may be underway here, but i t r e q u i r e s examination i n f a r more depth before p o s s i b l e e x p l a n a t i o n s can be o f f e r e d . I f e l t that I c o u l d not r e l y on the l i t e r a t u r e and d i d not wish to be r e s t r i c t e d to the use of herbarium m a t e r i a l a l o n e . C a r e f u l and ex t e n s i v e sampling of known numbers of i n d i v i d u a l s from v a r i o u s p o p u l a t i o n s or d i s j u n c t s are r e q u i r e d before t h i s problem can be d e a l t with p r o p e r l y . B. DISCUSSION In s p i t e of the f a i l u r e of t h i s study to d e f i n e the o v e r a l l r e l a t i o n s h i p s among a l l s p e c i e s i n the genera, s e v e r a l s m a l l e r monophyletic groups were i d e n t i f i e d based on reasonably strong evidence. These are f u r t h e r confirmed by the use of s e c t i o n A r t o r h i z a and the Wyethia r e t i c u l a t a / W . e l a t a s p e c i e s p a i r as f u n c t i o n a l outgroups. Because they are so uncommon, i s o f l a v o n e s are noted i n the f o l l o w i n g d i s c u s s i o n when they have been i d e n t i f i e d i n a s p e c i e s , even though t h e i r presence c o u l d not be used' as a c h a r a c t e r i n the present study. 28 Wyethia m o l l i s i s re p e a t e d l y shown to share a more recent common a n c e s t r y with the endemic s p e c i e s W. invenusta and W. ovata than with other members of s e c t i o n Wyethia. Wyethia  m o l l i s i s a widespread s p e c i e s , found i n Northern C a l i f o r n i a and throughout the S i e r r a Nevada at mid to high e l e v a t i o n s . The two r e l a t e d s p e c i e s , while having only very l i m i t e d ranges, are near the area covered by W. m o l l i s . Wyethia m o l l i s and W. invenusta both make methylated f l a v o n o l s , p r e n y l a t e d flavanones and i s o f l a v o n e s . Wyethia ovata makes i s o f l a v o n e s and methylated f l a v o n o l s . W. b o l a n d e r i , another S i e r r a n endemic, makes only methylated f l a v o n o l s (McCormick et a_l. , 1985b). Two other endemics, Wyethia r e t i c u l a t a and W. e l a t a , are shown i n many of the c l a d i s t i c a n a l y s es to share a r e l a t i v e l y c l o s e a n c e s t r y with the other S i e r r a n s p e c i e s j u s t d i s c u s s e d . These two taxa appear to form a c l o s e s p e c i e s p a i r and share s e v e r a l synapomorphies. These i n c l u d e a genuine lac k of any b a s a l leaves a r i s i n g (independently of f l o w e r i n g shoots) from separate underground c a u d i c e s . The p l a n t s possess only c a u l i n e l e a v e s , which are w e l l developed, t r i a n g u l a r i n shape and a l t e r n a t e l y arranged. Wyethia r e t i c u l a t a produces i s o f l a v o n e s and methylated f l a v o n o l s , while Wyethia e l a t a produces only methylated f l a v o n o l s . Both s p e c i e s are endemic to very small areas along the west slope of the S i e r r a Nevada, but n e i t h e r seems to be o b v i o u s l y r e s t r i c t e d by geography or h a b i t a t d i f f e r e n c e s and both are extremely a g g r e s s i v e w i t h i n t h e i r ranges. The only other s p e c i e s which completely l a c k s b a s a l leaves 29 i s Wyethia scabra. In many r e s p e c t s , however, t h i s taxon i s an anomaly and appears to possess many autapomorphies. Although t h i s s p e c i e s makes the i s o f l a v o n e s and p r e n y l a t e d flavanones c h a r a c t e r i s t i c of some other members of Wyethia, i t s leaves are l i n e a r and narrow, with a g e n e r a l l y (but not always) a l t e r n a t e arrangement. In a l l of the analyses employing g e n e r i c outgroups and m o r p h o l o g i c a l c h a r a c t e r s , W. scabra i s always shown as having a monophyletic r e l a t i o n to Balsamorhiza h o o k e r i . However, in the cladograms using f u n c t i o n a l outgroups and a l a r g e r number of v a r i a b l e s , W. scabra i s seen to share a c l o s e r a n c e s t r a l h i s t o r y with some of the s p e c i e s i n s e c t i o n Wyethia. R e s u l t s based on the use of g e n e r i c outgroups u n i t e Wyethia scabra and Balsamorhiza hookeri on what may be the homoplasous c h a r a c t e r s of l i n e a r l e a v e s and p h y l l o t a x y , which may vary from o p p o s i t e to a l t e r n a t e on the same stem. The analyses using f u n c t i o n a l outgroups imply that these c h a r a c t e r s c o n s t i t u t e autapomorphies f o r both s p e c i e s . Wyethia scabra i s n a t i v e mostly to the d e s e r t s of Utah and i t s placement with respect to the other members of Wyethia i s s t i l l q u e s t i o n a b l e . Balsamorhiza hookeri and the u n i d e n t i f i e d s p e c i e s which may be c l o s e to i t r e q u i r e f u r t h e r a t t e n t i o n as w e l l . S e c t i o n A l a r c o n i a i n c l u d e s only the two taxa Wyethia  h e l e n i o i d e s and W. g l a b r a , both of which produce i s o f l a v o n e s , flavanones and p r e n y l a t e d compounds. These two are seen to form a s p e c i e s p a i r or two members of a trichotomy i n a l l of the c l a d i s t i c a n a l y s e s . There does not, however, appear to be any s t r o n g j u s t i f i c a t i o n f o r s e p a r a t i n g them out as a s e c t i o n . The 30 r e s u l t s imply that these two s p e c i e s share a r e l a t i v e l y c l o s e a n c e s t r y with at l e a s t some members of s e c t i o n Wyethia, and perhaps to p a r t of Agnorhiza as w e l l . They have been given s e c t i o n a l s t a t u s because of t h e i r l a r g e , l e a f y p h y l l a r i e s , w e l l developed pappus and huge achenes. These c h a r a c t e r s c e r t a i n l y u n i t e the s p e c i e s as a p a i r , but may not j u s t i f y s e c t i o n a l r e c o g n i t i o n , u n l e s s we are to do the same f o r a l l good s p e c i e s p a i r s . The placement of a l l the taxa now i n c l u d e d i n s e c t i o n Wyethia remains u n c e r t a i n . One s p e c i e s i n p a r t i c u l a r , Wyethia  h e l i a n t h o i d e s , i s shown to share a more recent h i s t o r y with some members of Balsamorhiza than with other s p e c i e s of Wyethia, based on s e v e r a l of the cladograms. T h i s r e l a t i o n i s suggested repe a t e d l y when taxonomic outgroups were used. Wyethia  h e l i a n t h o i d e s produces methylated f l a v o n o l s , as do a l l members of Balsamorhiza so f a r surveyed, but there i s no r e l i a b l e i n f o r m a t i o n yet a v a i l a b l e on i t s a b i l i t y to produce i s o f l a v o n e s . In the c l a d i s t i c analyses employing f u n c t i o n a l outgroups, W. h e l i a n t h o i d e s lack of flavanones and p r e n y l a t e d compounds i s seen as e i t h e r symplesiomorphic or as autapomorphic r e v e r s a l s . E i t h e r way, these cladograms imply W. h e l i a n t h o i d e s i s pa r t of a monophyletic group i n c l u d i n g most or a l l of the present s p e c i e s of Wyethia. Some other members of s e c t i o n Wyethia seem to be c l o s e l y r e l a t e d to each other, but the d e t a i l s of the r e l a t i o n s h i p s w i t h i n t h i s group, as w e l l as i t s a n c e s t r y with r e s p e c t to other taxa remains ambiguous. Wyethia a m p l e x i c a u l i s , W. a n g u s t i f o l i a , 31 W. l o n g i c a u l i s and W. a r i z o n i c a , appear in v a r i o u s combinations on most cladograms. Chemically, these four taxa share synapomorphies with some other members of Wyethia i n t h e i r p r o d u c t i o n of p r e n y l a t e d flavanones (and i s o f l a v o n e s , at l e a s t i n the case of W. a n g u s t i f o l i a ) . The two present s e c t i o n s of Balsamorhiza are r e p e a t e d l y confirmed as o u t l i n i n g monophyletic groups of s p e c i e s , r e g a r d l e s s of the outgroup employed. The only e x c e p t i o n to t h i s was the anomalous taxon Balsamorhiza h o o k e r i , whose exact r e l a t i o n s h i p to the other s p e c i e s i s s t i l l u n c e r t a i n . S e c t i o n A r t o r h i z a and the other a v a i l a b l e members of s e c t i o n Balsamorhiza each possess a number of synapomorphies, although t h e i r r e l a t i o n s to each other are s t i l l u n c l e a r . The use of f u n c t i o n a l outgroups i n the systematic s e c t i o n of t h i s study has p e r m i t t e d the use of more d e t a i l e d m o r phological and chemical data, and t h i s has proven v a l u a b l e . Although t h i s a d d i t i o n a l data lends f u r t h e r support to the a p p a r e n t l y monophyletic groups w i t h i n the two genera, t h e i r o v e r a l l r e l a t i o n s h i p s with r e s p e c t to each other were seen to change d r a m a t i c a l l y . To me t h i s suggests that most of the taxonomic outgroups share too d i s t a n t of an a n c e s t r y with the study groups. The use of g e n e r i c outgroups was necessary f o r the i d e n t i f i c a t i o n of monophyletic c o l l e c t i o n s of s p e c i e s w i t h i n the study groups, but were of l i t t l e use i n f u r t h e r c l a r i f i c a t i o n of r e l a t i o n s h i p s , s i n c e so few c h a r a c t e r s c o u l d be employed. The main value of taxonomic outgroups, i n t h i s study at l e a s t , was fo r the i d e n t i f i c a t i o n of monophyletic f u n c t i o n a l outgroups, 32 which c o u l d then be used i n the f u r t h e r c l a r i f i c a t i o n of r e l a t i o n s h i p s . Such f u n c t i o n a l outgroups have the a d d i t i o n a l advantage of being c o n s t r u c t e d from data t h a t , at the outset of the study, was e q u i v a l e n t to the data used f o r the ingroups. Chemical c h a r a c t e r s appear to pro v i d e strong evidence of synapomorphies f o r many members of Wyethia. The presence of such r e l a t i v e l y r a r e f l a v o n o i d s as i s o f l a v o n e s and p r e n y l a t e d flavanones i n the remaining taxa should prove to be v a l u a b l e c h a r a c t e r s . The f u l l e l u c i d a t i o n of f l a v o n o i d s f o r a l l ingroup taxa, as w e l l as b e t t e r i n f o r m a t i o n r e g a r d i n g outgroups i s r e q u i r e d before good use can be made of t h i s d ata. The problems w i t h i n and between Wyethia and Balsamorhiza have not been f u l l y r e s o l v e d here. A great deal more i n f o r m a t i o n i s needed before these r e l a t i o n s h i p s can be c l a r i f i e d . The taxa i n c l u d e d i n s e c t i o n Balsamorhiza must be p r o p e r l y d e f i n e d , among other t h i n g s . A l l of the subspecies p r e s e n t l y i n c l u d e d i n Wyethia scabra are i n need of examination as w e l l . Areas of cladograms which c o n s i s t e n t l y demonstrated monophyletic groups of s p e c i e s , r e g a r d l e s s of t h e i r r e l a t i o n s h i p s to other taxa, were used to form a b a s i s f o r the second p a r t of t h i s study. I wished to examine the nature of v a r i a t i o n among i n d i v i d u a l s w i t h i n each s p e c i e s i n order to see what p a t t e r n s might emerge when these d e s c r i p t i o n s were viewed w i t h i n the context of small l i n e a g e s of s p e c i e s . The r e s u l t s of the p h y l o g e n e t i c a n a l y s e s , while incomplete, p r o v i d e d a s u f f i c i e n t l y s t r o n g and necessary s t r u c t u r a l context w i t h i n which a f i n e r l e v e l of v a r i a t i o n c o u l d be nested. 33 2. EXPLORATORY DATA ANALYSIS A. SOURCES OF VARIATION These a n a l y s e s were based on the three data s e t s d e s c r i b i n g the heads, shoots and leaves of the t w e n t y - f i v e s p e c i e s l i s t e d i n Table I. Table VI shows the PCA r e s u l t s f o r these three s e t s of data. E i g e n v a l u e s and the percent v a r i a n c e accounted fo r by the f i r s t two axes are l i s t e d along with l o a d i n g s f o r v a r i a b l e s d e s c r i b i n g each s t r u c t u r e . I wished to i d e n t i f y the major sources of v a r i a t i o n given l e v e l s of s t r a t i f i c a t i o n d e s c r i b i n g genera, s e c t i o n s , s p e c i e s and p o p u l a t i o n s . Those l e v e l s which account f o r the major amount of v a r i a t i o n on a given PCA a x i s are c o n s i d e r e d here to be the most s t r o n g l y d i f f e r e n t i a t e d l e v e l s . In other words, the p o l a r i t y expressed by a swarm of data p o i n t s can be compared with v e c t o r s r e l a t e d to d i f f e r e n t l e v e l s i n a h i e r a r c h y . A s t r o n g source of v a r i a t i o n can a l s o be viewed as a source of d i f f e r e n t i a t i o n , or as a p l a c e where a change i n o r g a n i z a t i o n i s seen (Maze, et a l . , 1983; Maze and Parker, 1983; Wheeler and G u r i e s , 1982). L e v e l s which account f o r l a r g e r amounts of v a r i a t i o n should a l s o r e f l e c t r e a l sources of b i o l o g i c a l d i f f e r e n t i a t i o n . R e s u l t s of these analyses are shown i n Table V I I . For v a r i a b l e s d e s c r i b i n g heads, the l e v e l of s p e c i e s accounts f o r the m a j o r i t y of the v a r i a t i o n i n both p r i n c i p a l component axes. T h i s i n d i c a t e s that the v e c t o r s r e l a t e d to s p e c i e s agree best with the major axes through the p o i n t swarm c o n s i s t i n g of head 34 d a t a . I t f u r t h e r i m p l i e s t h a t , because s p e c i e s account f o r the m a j o r i t y of t h i s v a r i a t i o n , they are a l s o i t s primary source. In other words, one m a n i f e s t a t i o n of s p e c i a t i o n among these organisms appears to be d i f f e r e n t i a t i o n among v a r i a b l e s d e s c r i b i n g f l o w e r i n g heads. S e c t i o n s a l s o account f o r some of the v a r i a t i o n i n the head data, implying that monophyletic groups account f o r changes i n o r g a n i z a t i o n as w e l l . For l e a v e s and shoots, the m a j o r i t y of the v a r i a t i o n appears at the l e v e l of genus and i n the r e s i d u a l term which c o n s i s t s of the v a r i a t i o n w i t h i n p o p u l a t i o n s (between and w i t h i n i n d i v i d u a l s ) and measurement e r r o r . The amount of v a r i a t i o n accounted f o r by genera i n the shoot and l e a f data may be l a r g e due to the r e l a t i v e l y great extremes i n s i z e , arrangement and numbers of c a u l i n e leaves accounted for by the present g e n e r i c groupings. The PCA would i n d i c a t e t h i s by the r e l a t i v e value of lo a d i n g s f o r these v a r i a b l e s . In other words, the genera agree to some extent with data p o l a r i t y . In the p h y l o g e n e t i c analyses employing f u n c t i o n a l outgroups, a great d e a l of the p o l a r i t y i n the data a l s o agreed with the present g e n e r i c groupings. As w e l l , the v a r i a t i o n a t t r i b u t e d to the l e v e l of s e c t i o n s i n the head data i s l i k e l y due to some of the present groupings of s p e c i e s . T h i s may be e s p e c i a l l y apparent when s p e c i e s p a i r s are given s e c t i o n a l s t a t u s , such as i n A l a r c o n i a . The r e s u l t s of the c l a d i s t i c a n a l y s e s have shown that the present s e c t i o n s of Balsamorhiza appear to be mainly monophyletic. Monophyletic groups should reasonably account f o r much of the v a r i a t i o n i n a data s e t , s i n c e each l i n e a g e would d e f i n e a p o i n t of 35 d i f f e r e n t i a t i o n . I do not p r e s e n t l y have a completed a l t e r n a t e p h y l o g e n e t i c hypothesis with which to make comparisons. I would expect, though, that monophyletic groupings of organisms should show g r e a t e r agreement with data p o l a r i t y than p o l y p h y l e t i c or p a r a p h y l e t i c ones. P o p u l a t i o n s account f o r more of the o v e r a l l v a r i a t i o n among shoots and leaves than do s p e c i e s i n PCA 1, but t h i s i s r e v e r s e d f o r PCA 2. I t i n t e r e s t i n g to note, however, that at no time do p o p u l a t i o n s account fo r as much v a r i a t i o n as does the r e s i d u a l term f o r any a x i s through any data s e t . T h i s i m p l i e s that p o p u l a t i o n s , at l e a s t as they are d e f i n e d here, do not c o n s t i t u t e a p a r t i c u l a r l y important l e v e l of b i o l o g i c a l o r g a n i z a t i o n or d i f f e r e n t i a t i o n (source of v a r i a t i o n ) . I t i s p o s s i b l e t hat the sample s i z e s upon which the p o p u l a t i o n s were based were i n s u f f i c i e n t to account f o r d i f f e r e n c e s among p o p u l a t i o n s . Nonetheless, the r e s u l t s presented here imply that p o p u l a t i o n s do not represent a s t r o n g source of d i f f e r e n t i a t i o n f o r these organisms. Although they may be " s i g n i f i c a n t l y " d i f f e r e n t ( t h i s hypothesis was not t e s t e d ) , the d i f f e r e n c e s among p o p u l a t i o n s appear to be t r i v i a l when compared to the v a r i a t i o n present i n the e n t i r e data s e t . T h i s r e s u l t agrees with s e v e r a l other recent s t u d i e s (Maze, 1984; Scagel, 1984; Yeh and El-Kassaby, 1980; Robson, e_t a_l. , i n p r e p . ) . I would a l s o add that p o p u l a t i o n s may have had the b e n e f i t of the doubt in t h i s p a r t of the study, s i n c e they are e q u i v a l e n t to s p e c i e s f o r s e v e r a l taxa due to small sample s i z e s . Perhaps p o p u l a t i o n d i f f e r e n t i a t i o n i s an o r g a n i s m - s p e c i f i c phenomenon that i s not 36 p r e s e n t l y manifested w i t h i n most of the s p e c i e s d e s c r i b e d here. T h i s p o s s i b i l i t y w i l l be addressed when s e c t i o n Balsamorhiza i s examined -in g r e a t e r depth, for i t may be that d i f f e r e n c e s among d i s j u n c t p o p u l a t i o n s are a source of the c o n f u s i o n surrounding some of these taxa. B. NATURE OF WITHIN-SPECIES VARIATION Specie s , and the v a r i a t i o n w i t h i n and among the i n d i v i d u a l s i n c l u d e d w i t h i n them seemed to be l e v e l s worth e x p l o r i n g i n g r e a t e r d e t a i l . In my o p i n i o n , both i n d i v i d u a l s and s p e c i e s may be viewed as h i e r a r c h i c a l l y r e l a t e d e n t i t i e s . That i s , they c o n s t i t u t e r e a l l e v e l s of b i o l o g i c a l o r g a n i z a t i o n and are interdependent, but not e q u i v a l e n t . I f t h i s i s so, then the nature of the r e l a t i o n s h i p s between these h i e r a r c h i e s i s worth f u r t h e r e x p l o r a t i o n . I t may be p o s s i b l e to b e t t e r understand e v o l u t i o n a r y processes through a search f o r p a t t e r n s common to these d i f f e r e n t l e v e l s . I wished to ask what e f f e c t s p e c i a t i o n ( p h y l o g e n e t i c v a r i a t i o n ) has had upon ontogenetic v a r i a t i o n w i t h i n each s p e c i e s and i f any of t h i s can be assessed by o b s e r v i n g s t r u c t u r e s of mature i n d i v i d u a l s . N i n e t y - f i v e percent c o n f i d e n c e e l l i p s e s t a t i s t i c s ( J o l i c o e u r , 1968; J o l i c o e u r and Mosimann, 1960) were c a l c u l a t e d f o r each data set d e s c r i b i n g heads, shoots and leaves f o r each of the t w e n t y - f i v e s p e c i e s . Included i n these analyses were 3 outgroup s p e c i e s , H e l i a n t h e l l a c a l i f o r n i c a , H. u n i f l o r a and 37 Rudbeckia o c c i d e n t a l i s . H e l i a n t h u s c u s i c k i i , a s p e c i e s from another c l o s e l y r e l a t e d genus, was a l s o i n c l u d e d . Because t h i s p a r t of the study was based upon measurements taken from i n d i v i d u a l s , i t was hoped that v a r i a t i o n w i t h i n s p e c i e s c o u l d be estimated. I d i d not wish to d e s c r i b e i n f i n e r d e t a i l the s p e c i f i c c h a r a c t e r s t a t e changes or a sequence of such events by which one s p e c i e s d i f f e r s from another. That was the purpose f o r which c l a d i s t i c a n a l y s i s was used. The purpose f o r which my v e r s i o n of e l l i p s e d e s c r i p t i o n a n a l y s i s was used (or abused) was to probe the w i t h i n - s p e c i e s v a r i a t i o n f o r the d e t a i l s of i t s s t r u c t u r e . T h i s was done i n order to see i f w i t h i n - s p e c i e s v a r i a t i o n changed with s p e c i a t i o n and, i f so, to determine i f any of t h i s c o u l d be r e l a t e d to d i f f e r e n t h i e r a r c h i c a l l e v e l s and h i s t o r i c a l c o n s t r a i n t s . The p r o d u c t i o n of these e l l i p s e s t a t i s t i c s and the g r a p h i c s by which they are represented are f a r removed by a b s t r a c t i o n from the o r i g i n a l raw data. T h i s i s necessary i n order to made c e r t a i n comparisons where p a t t e r n and s t r u c t u r e cannot be p e r c e i v e d from a raw data matrix. Techniques s i m i l a r to those used here have been employed i n recent s t u d i e s d e s c r i b i n g v a r i a t i o n among and w i t h i n i n d i v i d u a l s and p o p u l a t i o n s (Maze, et a l . , 1986b; Scagel, e_t a_l. , 1985; Robson, e_t a_l. , i n p r e p . ) . Values f o r theta estimate the placement of a swarm of data p o i n t s d e s c r i b i n g a s p e c i e s i n v a r i a b l e space, r e l a t i v e to a l l other s p e c i e s represented. In t h i s sense, t h e t a p r o v i d e s an assessment of the nature of the d i f f e r e n c e s i n c o r r e l a t i o n s among p o i n t swarms f o r each s p e c i e s . I t says nothing, however, 38 about the r e l a t i v e q u a n t i t y of v a r i a t i o n or the degree of i n t e r -r e l a t e d n e s s of the v a r i a b l e s i n c l u d e d i n the a n a l y s i s . Theta g i v e s an a b s t r a c t e d estimate of the nature of o r g a n i z a t i o n of s t r u c t u r e s making up i n d i v i d u a l s w i t h i n each s p e c i e s r e l a t i v e to a l l o t h e r s . Estimates of e l l i p s e area give an i n d i c a t i o n of o v e r a l l amount of v a r i a t i o n f o r s t r u c t u r e s w i t h i n a s p e c i e s r e l a t i v e to those same s t r u c t u r e s among a l l s p e c i e s . T h i s permits a q u a n t i t a t i v e assessment to be made of that v a r i a t i o n . For i n s t a n c e , I have asked of the data how much v a r i a t i o n e x i s t s among c a u l i n e leaves i n one s p e c i e s compared to v a r i a t i o n among a l l c a u l i n e l e a v e s . Within the c o n s t r a i n t of the s p e c i e s l i m i t s , do leaves vary widely, or are they a l l r a t h e r s i m i l a r to each other? I wished to ask about the v a r i a t i o n d i f f e r e n t taxa show fo r homologous s t r u c t u r e s i n order to see i f the sources c o u l d be i d e n t i f i e d and the v a r i a t i o n e x p l a i n e d . E c c e n t r i c i t y c a l c u l a t i o n s allowed the r e l a t i v e measurement of v a r i a b l e c o r r e l a t i o n s (inter-dependency) to be made f o r the same s t r u c t u r e s among the s p e c i e s . Returning to the l e a f example, I have asked i f a change i n l e n g t h i s always accompanied by some e q u i v a l e n t change i n width f o r each l e a f , r e g a r d l e s s of o v e r a l l d i f f e r e n c e s between l e a v e s . Both t h e t a and e c c e n t r i c i t y , as estimates of c o r r e l a t i o n s , e v a l u a t e d i f f e r e n c e s i n w i t h i n - s p e c i e s c o r r e l a t i o n s . E l l i p s e s s t a t i s t i c s f o r each s p e c i e s have been made graphic ( F i g u r e s 10-12) so that p a t t e r n s c o u l d be more e a s i l y noted. The three v a l u e s which d e s c r i b e some aspect of an e l l i p s e have been 39 given a common o r i g i n and appear as "dry f l i e s " . The l e f t h a l f shows values f o r t h e t a , with a s c a l e running from 80 to -80. The r i g h t s i d e of the dry f l y c o n s i s t s of two p a r t s . The upper r i g h t quadrant g i v e s an estimate of area and i t s s c a l e runs from 40 to 0, with the h o r i z o n t a l l i n e marking 0. Values c l o s e r to zero i n d i c a t e s m a l l e r r e l a t i v e amounts of v a r i a t i o n . The lower r i g h t quadrant d e p i c t s e c c e n t r i c i t y , where the h o r i z o n t a l l i n e i s given a value of 1, i n d i c a t i n g complete c o r r e l a t i o n among v a r i a b l e s . I t i s given a low value of .6, s i n c e v a r i a b l e i n t e r -dependency w i t h i n these p l a n t s t r u c t u r e s i s r a r e l y any c l o s e r to 0 than t h i s . The extremes i n s c a l e s shown f o r these dry f l i e s are based on t h i s data s e t . Even though some sample s i z e s were small (only f i v e p l a n t s ) these s t a t i s t i c s are assumed to be reasonably a c c u r a t e s i n c e they were based, u l t i m a t e l y , on a PCA of the e n t i r e data s e t . That PCA i s l i k e l y s t a b l e , and so the s t a t i s t i c s d e r i v e d from i t are a l s o s t a b l e ( S c a g e l , et a_l. , 1985). When shown simply as a group of dry f l i e s , i t i s d i f f i c u l t to p i c k out any p a t t e r n s or trends of v a r i a t i o n w i t h i n s p e c i e s which may be due to the a n c e s t r y of the s p e c i e s . I wished to compare these dry f l i e s w i t h i n the context of monophyletic groups d e r i v e d from p h y l o g e n e t i c a n a l y s i s . The attempt was made to see where v a r i a t i o n appeared to be history-dependent and where i t seemed unique w i t h i n and among small l i n e a g e s of s p e c i e s . Among most of the ingroup taxa, t h e t a f o r shoots never drops below zero on the s c a l e . Almost a l l of the s p e c i e s i n 40 Wyethia and Balsamorhiza have val u e s f o r t h e t a somewhere above zero, where exact values seem to be at l e a s t p a r t i a l l y dependent on r e l a t e d n e s s . Only Wyethia scabra and Balsamorhiza h o o k e r i , among ingroup taxa, show th e t a values f o r shoots below zero. T h i s r e s u l t i s seen f o r the two outgroup s p e c i e s of H e l i a n t h e l l a , as w e l l . I t seems l i k e l y that the d i f f e r e n c e i n t h e t a seen f o r these taxa has to do with the nature of c o n s t r a i n t s w i t h i n which nodes develop on the stems. In these four s p e c i e s , p h y l l o t a x y may be a l t e r n a t e or o p p o s i t e , or both c o n d i t i o n s may e x i s t on a s i n g l e shoot. T h i s s i t u a t i o n i s never the case f o r any of the other ingroup taxa. C a u l i n e l e a v e s of a l l other s p e c i e s of Wyethia and s e c t i o n A r t o r h i z a of Balsamorhiza are always a l t e r n a t e . In a l l of the other members of s e c t i o n Balsamorhiza f o r which I have data, c a u l i n e leaves c o n s i s t of a s i n g l e o p posite p a i r . The two ingroup taxa which show values f o r theta more s i m i l a r to those of H e l i a n t h e l l a have been d e s c r i b e d as anomalous in many other r e s p e c t s . These r e s u l t s f u r t h e r i l l u s t r a t e the c o n f u s i n g v a r i a t i o n found w i t h i n these s p e c i e s . U n t i l the p h y l o g e n e t i c r e l a t i o n s h i p s are c l a r i f i e d , the p o s s i b l e h i s t o r i c a l s i g n i f i c a n c e of these d i f f e r e n c e s i n v a r i a t i o n s t r u c t u r e f o r shoots of Wyethia scabra and Balsamorhiza hookeri remain obscure. Three s p e c i e s of s e c t i o n A r t o r h i z a , a monophyletic group, are shown in F i g u r e 13 and are compared with the two s p e c i e s of H e l i a n t h e l l a . The shoots of the Balsamorhizas a l l show low va l u e s f o r area and values c l o s e to one f o r e c c e n t r i c i t y . T h i s i m p l i e s that shoot v a r i a b l e s are h i g h l y i n t e r - c o r r e l a t e d and 41 that d i f f e r e n c e s among shoots do not pro v i d e a great source of v a r i a t i o n . Other s t r u c t u r e s , however, appear to be l e s s c o n s t r a i n e d i n t h e i r v a r i a t i o n . Among flower heads, Balsamorhiza  d e l t o i d e a demonstrates lower e c c e n t r i c i t y than do the other members of t h i s group. T h i s i s probably most s t r o n g l y i n f l u e n c e d by p h y l l a r i e s , which may vary g r e a t l y i n s i z e among flower heads, p r o v i d i n g a unique source of v a r i a t i o n f o r t h i s s p e c i e s compared to the r e s t of the l i n e a g e . The dry f l i e s r e p r e s e n t i n g the two a v a i l a b l e s p e c i e s of the outgroup H e l i a n t h e l l a are q u i t e d i f f e r e n t from A r t o r h i z a . T h i s i s most s t r o n g l y r e f l e c t e d i n t h e t a , area and e c c e n t r i c i t y f o r shoots. The o r g a n i z a n i z a t i o n of shoots i n these s p e c i e s d i f f e r from A r t o r h i z a i n both the nature and q u a n t i t y of t h e i r v a r i a t i o n . Area and e c c e n t r i c i t y f o r shoots i s g r e a t e r i n H e l i a n t h e l l a . As w e l l , shoots of the two outgroup s p e c i e s are q u i t e d i f f e r e n t from each other, implying that o r g a n i z a t i o n of shoots i s not h i g h l y c o n s t r a i n e d e i t h e r w i t h i n or among s p e c i e s . Within A r t o r h i z a , however, heads and leaves appear to p r o v i d e g r e a t e r sources of v a r i a t i o n than they do among the outgroup spec i e s . Other l i n e a g e s of s p e c i e s may a l s o be compared. F i g u r e 14 shows three members of s e c t i o n Balsamorhiza, B. incana, B. macrophylla and B. s e r r a t a next to a p a i r of endemics, Wyethia e l a t a and W. r e t i c u l a t a . The two endemics appear very s i m i l a r to each other, r e l a t i v e to the Balsamorhiza s p e c i e s , except i n e c c e n t r i c i t y f o r l e a v e s . T h i s d i f f e r e n c e i n w i t h i n -s p e c i e s v a r i a b l e c o r r e l a t i o n s f o r lea v e s suggests that 42 s p e c i a t i o n events may be p a r t l y manifested as changing sources and i n t e n s i t i e s of o r g a n i z a t i o n a l c o n s t r a i n t d u r i n g ontogeny. Some unique p a t t e r n of w i t h i n - s p e c i e s v a r i a t i o n appears to e x i s t f o r each s p e c i e s assessed. As w e l l , the nature, degree and source of t h i s new v a r i a t i o n , seen here as the r e s u l t of many ontogenies, seems to be c l o s e l y r e l a t e d to l i n e a g e s composed of s p e c i e s . H i s t o r i c a l c o n s t r a i n t s w i t h i n monophyletic groups appear to show str o n g p a t t e r n s of o r g a n i z a t i o n and v a r i a t i o n , e s p e c i a l l y when compared with other c l a d e s of somewhat d i f f e r e n t a n c e s t r y . 43 IV. SUMMARY AND EXPLANATIONS 1 . SUMMARY A. PHYLOGENETIC SYSTEMATICS In the cladograms employing taxonomic outgroups, Wyethia and Balsamorhiza always appear as p a r a p h y l e t i c taxa, but more exact r e l a t i o n s h i p s remain u n c l e a r . When f u n c t i o n a l outgroups are d e r i v e d from small c l a d e s of ingroup s p e c i e s and used f o r outgroup comparison with a l a r g e r set of morp h o l o g i c a l and chemical v a r i a b l e s , the gen e r a l p a t t e r n changes. When the Wyethia elata/W. r e t i c u l a t a outgroup i s used, Balsamorhiza appears as a monophyletic group with respect to i t s present members, but Wyethia i s p a r a p h y l e t i c with respect to i t . When s e c t i o n A r t o r h i z a i s used as the outgroup, the genera do not appear to be p a r a p h y l e t i c . More data and the mis s i n g taxa are r e q u i r e d before f u l l p h y l o g e n e t i c hypotheses can be c o n s t r u c t e d f o r Wyethia and Balsamorhiza. F l a v o n o i d chemistry may provide st r o n g evidence of synapomorphies f o r many s p e c i e s of Wyethia. Taxonomic outgroups may gi v e poor r e s u l t s because t h e i r r e l a t i o n s h i p s to the ingroup taxa are simply too d i s t a n t . T h e i r r e a l value here was f o r the i d e n t i f i c a t i o n of f u n c t i o n a l outgroups, which were used i n more d e t a i l e d a n a l y s e s . The f u n c t i o n a l outgroups and other small c l a d e s were used to form a framework f o r the second p a r t of t h i s study, which attempted to r e l a t e the p a t t e r n of s p e c i a t i o n events w i t h i n and among these 44 l i n e a g e s to p a t t e r n s d e r i v e d from an a n a l y s i s of i n t r a - s p e c i f i c v a r i a t i o n . B. EXPLORATORY DATA ANALYSIS Three d i f f e r e n t data s e t s , which c o n s i s t e d of v a r i a b l e s d e s c r i b i n g the shoots, t e r m i n a l heads and c a u l i n e l e a v e s of twe n t y - f i v e s p e c i e s , i n c l u d i n g four outgroup s p e c i e s i n a d d i t i o n to the ingroup, were examined. The ana l y s e s performed here were based on PCA and f a l l under the ge n e r a l heading of e x p l o r a t o r y data a n a l y s i s . With these techniques I was able to examine morphological data c o n s i s t i n g of measured v a r i a b l e s taken from i n d i v i d u a l p l a n t s f o r the sources and s t r u c t u r e of among and w i t h i n - i n d i v i d u a l v a r i a t i o n . The three data s e t s were s t r a t i f i e d i n t o taxonomic l e v e l s d e f i n i n g genera, s e c t i o n s , s p e c i e s , p o p u l a t i o n s and a r e s i d u a l term. The v a r i a t i o n present among shoots, heads and leaves was apportioned to each of these. I t was found that f o r heads, the m a j o r i t y of v a r i a t i o n c o u l d be a t t r i b u t e d to s p e c i e s and some to s e c t i o n s . For shoots and l e a v e s , the m a j o r i t y of the v a r i a t i o n appeared i n the term d e s c r i b i n g genera. T h i s i m p l i e s that most of the v a r i a t i o n i n the data agrees e i t h e r with l i n e a g e s of s p e c i e s (genera and s e c t i o n s ) or with the s p e c i e s themselves. P o p u l a t i o n s were always seen to account f o r l e s s v a r i a t i o n i n the data than the r e s i d u a l term. Although the p o p u l a t i o n s may have shown s t a t i s t i c a l l y s i g n i f i c a n t d i f f e r e n c e s t h e i r importance as a source of v a r i a t i o n here i s approximately e q u i v a l e n t to measurement e r r o r . I t i s p o s s i b l e that the samples 45 a n a l y z e d here were too small to adequately account f o r d i f f e r e n c e s among p o p u l a t i o n s . I t i s a l s o p o s s i b l e that the v a r i a t i o n d e s c r i b e d here i s s e l e c t i v e l y n e u t r a l and environmental f a c t o r s have had l i t t l e e f f e c t . The t h i r d p o s s i b i l i t y i s that p o p u l a t i o n s do not c o n s t i t u t e an important l e v e l of b i o l o g i c a l o r g a n i z a t i o n f o r these organisms. N i n e t y - f i v e percent c o n f i d e n c e e l l i p s e s t a t i s t i c s were c a l c u l a t e d f o r the p o i n t swarms d e s c r i b i n g each of the twenty-f i v e s p e c i e s . Graphic r e p r e s e n t a t i o n s , "dry f l i e s " , were d e r i v e d from these s t a t i s t i c s so that p a t t e r n s d e s c r i b i n g t h e t a , area and e c c e n t r i c i t y f o r the shoots, heads and l e a v e s of each s p e c i e s c o u l d be compared v i s u a l l y . These dry f l i e s were then p r o j e c t e d onto the branch t i p s of cladograms d e p i c t i n g s m a l l , monophyletic groups of s p e c i e s . T h i s was done i n order to see i f any p a t t e r n s i n the source and s t r u c t u r e of v a r i a t i o n w i t h i n a s p e c i e s c o u l d be a t t r i b u t e d to the h i s t o r y of a c l a d e . New p a t t e r n s , a p p a r e n t l y r e l a t e d to the h i s t o r y of s p e c i e s , emerge when the v a r i a t i o n w i t h i n s p e c i e s i s viewed w i t h i n the context of v a r i a t i o n among them. In a d d i t i o n , u n p r e d i c t a b l e changes in v a r i a t i o n and o r g a n i z a t i o n are always seen to be present i n the a b s t r a c t d e s c r i p t i o n of each s p e c i e s . These unique changes i n w i t h i n and among-individual v a r i a t i o n would appear to be p a r t of the m a n i f e s t a t i o n of a s p e c i a t i o n event. The emergence of new v a r i a t i o n and o r g a n i z a t i o n expressed d u r i n g ontogeny can be r e l a t e d to p h y l o g e n e t i c v a r i a t i o n , demonstrating t h a t i n c r e a s e s i n complexity appear to accompany of both of these e v o l u t i o n a r y p r o c e s s e s . 46 2. EXPLANATIONS T h i s study i l l u s t r a t e s a method of comparing aspects of w i t h i n - s p e c i e s v a r i a t i o n to s p e c i a t i o n events w i t h i n small c l a d e s . Apportionment of v a r i a t i o n to d i f f e r e n t taxonomic l e v e l s has a l s o p r o v i d e d i n s i g h t i n t o the sources of d i f f e r e n t i a t i o n f o r the v a r i o u s p l a n t s t r u c t u r e s . The use of d i f f e r e n t m u l t i v a r i a t e techniques has p e r m i t t e d analyses to be made of v a r i a t i o n at d i f f e r e n t l e v e l s of b i o l o g i c a l o r g a n i z a t i o n so that an attempt c o u l d be made to r e l a t e them. I th i n k t h i s approach w i l l prove to be worthwhile as a method with which the nature of the r e l a t i o n s h i p s between ontogeny and phylogeny may be more thoroughly e x p l o r e d . These techniques should permit the development of i n d i v i d u a l s to be t r a c k e d through time f o r d i f f e r e n t s p e c i e s , a l l o w i n g ' d e t a i l e d comparisons to be made acros s p h y l o g e n i e s . So f a r , t h i s study has p r o v i d e d an account of how p a t t e r n s of v a r i a t i o n can be analyzed at d i f f e r e n t h i e r a r c h i c a l l e v e l s u s ing d i f f e r e n t methods. The r e s u l t s have been compared in d i f f e r e n t ways and new p a t t e r n s have been r e p o r t e d . However, d e s c r i p t i o n alone i s i n s u f f i c i e n t . I b e l i e v e t h i s study has succeeded i n examining some of the v a r i a t i o n t h at i s the m a n i f e s t a t i o n of e v o l u t i o n a r y processes as expressed in these p l a n t s . I wish to provide these r e s u l t s with some c a u s a l e x p l a n a t i o n . Dobzhansky, et §_1. (1977, pp.506) o f f e r the f o l l o w i n g d e f i n i t i o n of at l e a s t a p a r t of the e v o l u t i o n a r y p r o c e s s . "The 47 theory of e v o l u t i o n by n a t u r a l s e l e c t i o n advances arguments of the f o l l o w i n g g e n e r a l form. Among a l t e r n a t i v e g e n e t i c v a r i a n t s , some r e s u l t i n f e a t u r e s that are u s e f u l to t h e i r c a r r i e r s as ad a p t a t i o n s to t h e i r environment...useful a d a p t a t i o n s become e s t a b l i s h e d i n p o p u l a t i o n s . "To e x p l a i n a p a r t i c u l a r a d a p t a t i o n , a v a l i d s e l e c t i o n i s t argument has to show (1) that n a t u r a l s e l e c t i o n i s i n v o l v e d at a l l ; and (2) that n a t u r a l s e l e c t i o n f a v o r s that p a r t i c u l a r a d a p t a t i o n . " As f a r as can be seen from these r e s u l t s , no a d a p t a t i o n s of any n o t i c e a b l e importance have become e s t a b l i s h e d i n these p o p u l a t i o n s with enough f o r c e to cause t h e i r d i f f e r e n t i a t i o n . Genera, which account f o r the m a j o r i t y of the v a r i a t i o n i n the l e a f and shoot data, occupy a tremendous d i v e r s i t y of h a b i t a t s and, presumably, of s e l e c t i o n p r e s s u r e s as w e l l . The l e v e l of s p e c i e s accounts f o r a great d e a l of the v a r i a t i o n in the head data. A s p e c i e s such as Balsamorhiza s a g i t t a t a i s sympatric with B. h o o k e r i , B. macrophylla and Wyethia a m p l e x i c a u l i s , but these d i s s i m i l a r s p e c i e s demonstrate a str o n g source of d i f f e r e n t i a t i o n . The i d e n t i f i c a t i o n of r e l e v a n t s e l e c t i v e f o r c e s , given these r e s u l t s , would be ex c e e d i n g l y d i f f i c u l t . There i s no evidence that these organisms have become e i t h e r more s i m i l a r or more d i f f e r e n t due to environmental f a c t o r s . A c c o r d i n g to the above requirements f o r a v a l i d s e l e c t i o n i s t argument, I f i n d I am unable to demonstrate l e g i t i m a t e l y that any e v o l u t i o n by n a t u r a l s e l e c t i o n has taken p l a c e here. Perhaps the sources and o r g a n i z a t i o n of the v a r i a t i o n d e s c r i b e d i n these 48 r e s u l t s are the product of e v o l u t i o n by something e l s e . There are at l e a s t two other f a c t o r s i n v o l v e d i n e v o l u t i o n a c c o r d i n g to Gould and Lewontin (1979) who would perhaps c r i t i c i z e the r e s t r i c t i v e d e f i n i t i o n c i t e d above. "An a d a p t a t i o n i s t programme has dominated e v o l u t i o n i s t thought i n England and the U.S. d u r i n g the past 40 y e a r s . I t i s based on f a i t h i n the power of n a t u r a l s e l e c t i o n as an o p t i m i z i n g agent. I t proceeds by breaking an organism i n t o u n i t a r y ' t r a i t s ' and p roposing an a d a p t i v e s t o r y f o r each c o n s i d e r e d s e p a r a t e l y . T r a d e - o f f s among competing s e l e c t i v e demands exert the only brake upon p e r f e c t i o n ; n o n - o p t i m a l i t y i s thereby rendered as a r e s u l t of a d a p t a t i o n as w e l l . We c r i t i c i z e t h i s approach and attempt to r e - a s s e r t a competing notion..„that organisms must be a nalysed as i n t e g r a t e d wholes... so c o n s t r a i n e d by p h y l e t i c h e r i t a g e , pathways of development and general a r c h i t e c t u r e that the c o n s t r a i n t s themselves become more i n t e r e s t i n g and more important in d e l i m i t i n g pathways of change than the s e l e c t i v e f o r c e that may mediate change when i t o c c u r s . " These i n s i g h t s p r o v i d e some f u r t h e r understanding of the r e s u l t s presented here. P h y l o g e n e t i c h i s t o r y does indeed seem to count as a c o n s t r a i n t on w i t h i n s p e c i e s v a r i a t i o n . A l a r g e amount of the v a r i a t i o n q u a n t i f i e d here i s a t t r i b u t a b l e to genera and s e c t i o n s , implying that shared h i s t o r i e s among s p e c i e s have acted as a strong source of d i f f e r e n t i a t i o n . In a d d i t i o n , i f shown dry f l i e s f o r any two s p e c i e s of Balsamorhiza, s e c t i o n A r t o r h i z a , I c o u l d p r e d i c t that area and e c c e n t r i c i t y f o r shoots w i l l r e f l e c t a high degree of c o n s t r a i n t 49 on t h i s source of v a r i a t i o n w i t h i n the t h i r d s p e c i e s . I c o u l d not make t h i s same p r e d i c t i o n f o r a t h i r d s p e c i e s of H e l i a n t h e l l a , given the two shown here. That these d i f f e r e n c e s are more l i k e l y to be i n f l u e n c e d by h i s t o r y than by environment i s f u r t h e r evidenced by the f a c t t h at Balsamorhiza s a g i t t a t a i s sympatric with H e l i a n t h e l l a u n i f l o r a , as i s B. d e l t o i d e a with H. c a l i f o r n i c a (see Appendix A). The h i s t o r y of a l i n e a g e seems to have an impact on the nature and source of both o r g a n i z a t i o n and v a r i a t i o n w i t h i n a s p e c i e s . That i s , h i s t o r y has p a r t l y determined where v a r i a t i o n w i l l or w i l l not be extreme among developmentally mature s t r u c t u r e s and what the nature of t h i s v a r i a t i o n w i l l be. T h i s s u p p l i e s p a r t of the e x p l a n a t i o n , but i t i s s t i l l incomplete. There i s another aspect to these r e s u l t s which I b e l i e v e to be n e i t h e r t r i v i a l nor d r i v e n by something c a l l e d "randomness" or " p r o b a b i l i t y " . Although these terms o f t e n serve to d e s c r i b e the phenomenon, they are not e q u i v a l e n t to i t s cause. Each dry f l y and each branch i n a cladogram d e s c r i b e s events which are unique and always u n p r e d i c t a b l e i n some way. That i s , I can p r e d i c t that v a r i a t i o n w i t h i n the s p e c i e s of s e c t i o n A r t o r h i z a w i l l be c o n s t r a i n e d among shoots and w i l l be manifested among heads and l e a v e s , but I w i l l never be able to p r e d i c t the exact form or source of t h i s new v a r i a t i o n . The source and nature of w i t h i n - s p e c i e s v a r i a t i o n i s shown to change i n a manner which i s p a r t l y unexpected and unique f o r each s p e c i e s . These d i f f e r e n c e s i n o r g a n i z a t i o n demonstrate i n c r e a s e s i n complexity at the h i e r a r c h i c a l l e v e l of s p e c i e s . T h i s o b s e r v a t i o n can be r e l a t e d 50 to the i n c r e a s e s in complexity expressed by the s t r u c t u r e s which make up each f l o w e r i n g shoot. That i s , where w i t h i n - s p e c i e s v a r i a t i o n i s seen to occur, i t i s i n f e r r e d that ontogenetic complexity has i n c r e a s e d . The unique v a r i a t i o n which a r i s e s d u r i n g s p e c i a t i o n i n the form of c h a r a c t e r s t a t e changes appears to be accompanied by s i m i l a r l y new and u n p r e d i c t a b l e v a r i a t i o n expressed among and w i t h i n i n d i v i d u a l s d u r i n g ontogeny. I b e l i e v e the u n p r e d i c t a b l e changes i n the source and nature of v a r i a t i o n w i t h i n and among s p e c i e s and c l a d e s demonstrates the m a n i f e s t a t i o n of emergent p r o p e r t i e s at these d i f f e r e n t l e v e l s of b i o l o g i c a l o r g a n i z a t i o n . Campbell (1982, pp.101) has d i s c u s s e d the problems a r i s i n g from the concepts of complexity, i n c r e a s i n g complexity and the nature and s i g n i f i c a n c e of emergent p r o p e r t i e s . He approaches the s u b j e c t by comparing the ideas of P r i g o g i n e , a chemist, and Chomsky, a l i n g u i s t , i n the f o l l o w i n g passages. "The c r e a t i o n of s t r u c t u r e depends on how f a r the system i s from e q u i l i b r i u m , and t h i s i n turn depends on the r a t e and amount of exchanges of matter and energy with the surroundings. The d i s t a n c e from e q u i l i b r i u m w i l l be great i f there are many exchanges, and very small or no n e x i s t e n t i f there are few. The same laws of p h y s i c s apply i n both cases, but near e q u i l i b r i u m P r i g o g i n e says the laws l e a d to 'doom or d e s t r u c t i o n ' , whereas f u r t h e r from e q u i l i b r i u m they may become processes of c o n s t r u c t i o n and o r g a n i z a t i o n . In t h i s l a t t e r case, he.adds, p r o b a b i l i t y theory breaks down. No n e q u i l i b r i u m can t h e r e f o r e be a source of order in open systems, whether these systems are chemical or 51 b i o l o g i c a l , r e v e a l i n g a fundamental k i n s h i p between l i f e and n o n l i f e . Complexity i s maintained i n both types. "The whole q u e s t i o n of complexity i s one which has proved s u p r i s i n g l y s u b t l e when i n v e s t i g a t e d by modern t h e o r i s t s . Complexity i s not j u s t a matter of a system having a l o t of p a r t s which are r e l a t e d to one another i n nonsimple ways. Instead, i t turns out to be a s p e c i a l p r o p e r t y i n i t s own r i g h t and i t makes complex systems d i f f e r e n t i n kind from simple ones, e n a b l i n g them to do t h i n g s and be t h i n g s we might not have expected. Chomsky suggests that human language competence, which must be among the most complicated s t r u c t u r e s i n the u n i v e r s e , a r i s e s u n i q u e l y i n e v o l u t i o n at a c e r t a i n stage of b i o l o g i c a l complexity. In other words, i t appears when, and only when, e v o l u t i o n has l e a d to an organism as complex as a human being. Chomsky goes on to say, 'This poses a problem f o r the b i o l o g i s t , s i n c e i f t r u e , i t i s an example of true "emergence"- the appearance of a q u a l i t a t i v e l y d i f f e r e n t phenomenon at a s p e c i f i c stage of o r g a n i z a t i o n . ' "Emergence i s not. a popular word in modern s c i e n c e . But i t i s not a b s o l u t e l y taboo, and i t i s u s u a l l y a s s o c i a t e d with complexity." I do not c o n s i d e r the word or the concept to be at a l l taboo. On the c o n t r a r y , I b e l i e v e the i n c l u s i o n of the concept of emergence i s necessary to an accurate d e s c r i p t i o n of b i o l o g i c a l change over time. The r e c o g n i t i o n of the emergent p r o p e r t i e s of a s p e c i a t i o n event or an ontogeny s u p p l i e s the l a s t m i s s i n g p i e c e r e q u i r e d f o r the understanding of these 52 r e s u l t s . The appearance of an u n p r e d i c t a b l e and " q u a l i t a t i v e l y d i f f e r e n t phenomenon" with r e s p e c t to w i t h i n - s p e c i e s v a r i a t i o n i s viewed here as the p a r t i a l m a n i f e s t a t i o n of a s p e c i a t i o n event. I t s t i l l remains to f i n d a s i n g l e and complete e x p l a n a t i o n capable of i n c l u d i n g and making i n t o a coherent whole a l l the r e s u l t s of t h i s study. Brooks and Wiley (1986) have p r o v i d e d what I c o n s i d e r to be j u s t such a necessary and comprehensive t h e o r e t i c a l e x p l a n a t i o n . T h e i r U n i f i e d Theory of B i o l o g y i s e s p e c i a l l y powerful because i t i s the f i r s t theory of e v o l u t i o n ever o f f e r e d which f i t s w i t h i n the formal d e f i n i t i o n of a deductive Covering Law Model (Hempel, 1966). Brooks and Wiley have based t h e i r theory on an expanded view of the Second Law of Thermodynamics as i t a p p l i e s to open systems. They have succeeded i n doing t h i s without reducing the processes of b i o l o g i c a l e v o l u t i o n to the m i c r o s c o p i c laws of chemistry and p h y s i c s . T h i s i s obvious from the e x p l a n a t i o n s and p r e d i c t i o n s s t a t e d i n the f o l l o w i n g passages (pp. 154). "...ontogenesis i s a n o n e q u i l i b r i u m phenomenon, that organisms are d i s s i p a t i v e s t r u c t u r e s whose temporal sequence of change i s caused by an i n t e r a c t i o n between r e l a t i v e l y determinate ( r e g u l a t o r y ) components and r e l a t i v e l y s t o c h a s t i c ( s t r u c t u r a l ) components w i t h i n the g e n e t i c and e p i g e n e t i c a r c h i t e c t u r e each organism possesses... a f u r t h e r a m p l i f i c a t i o n of t h i s phenomenon r e s u l t s i n the p r o d u c t i o n of new s p e c i e s . Thus, the emergence of new s p e c i e s f o l l o w s the same process as the emergence of new s t r u c t u r e s i n ontogenesis... 53 (pp. 350)..."By adopting an e x p l i c i t l y s t r u c t u r a l i s t approach to e v o l u t i o n , we have d i s c o v e r e d a number of s u r p r i s i n g and, i n some cases, c o u n t e r - i n t u i t i v e t h i n g s . We have been able to r e l a t e b i o l o g i c a l e v o l u t i o n to entropy without having to p o s t u l a t e any negentropic p r o p e r t i e s . T h i s i s , we d i s c o v e r e d , because the currency of e n t r o p i c behavior i n l i v i n g systems i s i n f o r m a t i o n s p e c i f y i n g s t r u c t u r e and not energy. We have c h a r a c t e r i z e d l i v i n g systems at a l l f u n c t i o n a l l e v e l s i n terms of four c o r n e r s t o n e p r i n c i p l e s . The p r i n c i p l e of i r r e v e r s i b i l i t y a s s e r t s that b i o l o g i c a l e v o l u t i o n i s one m a n i f e s t a t i o n of the more general phenomenon of c o s m o l o g i c a l e v o l u t i o n . The p r i n c i p l e of i n d i v i d u a l i t y amounts to an a s s e r t i o n that o r i g i n s are e s s e n t i a l to understanding e v o l u t i o n , and i t a l s o amounts to a r e j e c t i o n of n a t u r a l kinds i n b i o l o g y . The p r i n c i p l e of i n t r i n s i c c o n s t r a i n t s a s s e r t s that e v o l u t i o n a r y changes are non-random, o r d e r l y and c o n s t r a i n e d by the past h i s t o r y of each s p e c i e s . And the p r i n c i p l e of compensatory changes a s s e r t s that b i o l o g i c a l e v o l u t i o n has a c o n s e r v a t i v e tendency which all o w s the same dynamics to produce both determinism and new v a r i a t i o n s and o p t i o n s s i m u l t a n e o u s l y . We have shown that i n f o r m a t i o n can be produced by e n t r o p i c phenomenon. T h i s allows many aspects of e v o l u t i o n a r y h i s t o r y and e v o l u t i o n a r y processes to be t e s t e d d i r e c t l y and e x p l i c i t l y . . . "The pragmatic aspect of whether or not our theory i s c o n s i d e r e d a new view of b i o l o g i c a l e v o l u t i o n , or j u s t a r e f o r m u l a t i o n of the consensus depends on two t h i n g s . F i r s t , to what extent i s our p r e s e n t a t i o n viewed as p r o v i d i n g h e l p f u l 54 e x p l a n a t i o n f o r f i n d i n g s which would seem to be anomalous under the c u r r e n t paradigm?... "Second, we must make some p r e d i c t i o n s about the kinds of f i n d i n g s r e s e a r c h guided by our theory w i l l uncover... " B i o l o g i c a l systems are a combination of d e t e r m i n i s t i c and indeterminate f a c t o r s ; thus, i t should be p o s s i b l e to make d e t e r m i n i s t i c p r e d i c t i o n s about some aspects of e v o l u t i o n . I f we are c o r r e c t , however, the d e t e r m i n i s t i c aspects of e v o l u t i o n are the h i s t o r i c a l components..." T h i s theory of history-dependent, n o n e q u i l i b r i u m e v o l u t i o n appears necessary and s u f f i c i e n t to e x p l a i n a l l of the r e s u l t s presented i n t h i s study. F i r s t , i t e x p l a i n s why l i n e a g e s should be so important a f a c t o r i n the sources and nature of the v a r i a t i o n q u a n t i f i e d here. According to the methods of ph y l o g e n e t i c s y s t e m a t i c s , the h i s t o r i e s of s p e c i e s may be i n f e r r e d from the unique c h a r a c t e r s they share with other s p e c i e s . T h i s i m p l i e s that emergent p r o p e r t i e s are manifested at the o r i g i n s of a l i n e a g e and are r e t a i n e d i n descendants. For example, i t would have been impossible to p r e d i c t that the common ancestor of s e c t i o n A r t o r h i z a should suddenly produce something l i k e arrow-shaped b a s a l l e a v e s . The unique occurence of t h i s c h a r a c t e r i n an a n c e s t r a l s p e c i e s became an h i s t o r i c a l l y p r e s e r v e d synapomorphy which was expressed i n l a t e r s p e c i a t i o n events. I t now c o n t r i b u t e s to the u n i t y of the l i n e a g e as a developmental c o n s t r a i n t . However, each s p e c i a t i o n event i n t h i s c l a d e has a l s o been accompanied by some unique and unexpected autapomorphy of i t s own. That i s , emergent p r o p e r t i e s have 55 marked the a r i s a l of each s p e c i e s through time and these are expressed through l i n e a g e s of i n d i v i d u a l s . S i m i l a r p a t t e r n s of o r g a n i z a t i o n and v a r i a t i o n are a l s o seen among and w i t h i n i n d i v i d u a l s and appear to demonstrate the h i e r a r c h i c a l nature of b i o l o g i c a l o r g a n i z a t i o n at a f i n e r l e v e l . The use of d i f f e r e n t a n a l y t i c a l methods has p e r m i t t e d the q u a n t i f i c a t i o n of these phenomena at the l e v e l s of i n d i v i d u a l p l a n t s and the s t r u c t u r e s they produce, s p e c i e s and l i n e a g e s composed of s p e c i e s . I t has been shown t h a t w i t h i n - s p e c i e s v a r i a t i o n demonstrates the same p a t t e r n s of h i s t o r i c a l c o n s e r v a t i s m and emergence as does each s p e c i a t i o n event w i t h i n a c l a d e . Although the d e t a i l s d i f f e r , these s i m i l a r p a t t e r n s seem to e x i s t from the h i e r a r c h i c a l l e v e l of l i n e a g e s , through to the f l o w e r i n g shoots of i n d i v i d u a l p l a n t s . T h i s study has attempted to demonstrate that both history-dependent and unique v a r i a t i o n occur at d i f f e r e n t l e v e l s of b i o l o g i c a l o r g a n i z a t i o n and that these are r e l a t e d . These r e s u l t s agree w e l l with the e x p l a n a t i o n s and p r e d i c t i o n s o f f e r e d by Brooks and Wiley. I t h i n k i t i s time to take a c l o s e and s e r i o u s look at what new understanding might be gained from t h e i r theory of n o n e q u i l i b r i u m e v o l u t i o n . 56 V. EPILOG Or, can you s p e l l "deductive-nomological"? Perhaps a f i n a l aspect worthy of b r i e f d i s c u s s i o n remains with regard to the ch o i c e of c a u s a l e x p l a n a t i o n s made here. I t seems an e s s e n t i a l d i f f e r e n c e has t r a d i t i o n a l l y e x i s t e d between b i o l o g i c a l e x p l a n a t i o n s and those used i n other d i s c i p l i n e s f a l l i n g under the general heading " n a t u r a l s c i e n c e s " . T h i s d i f f e r e n c e has been p o i n t e d out r a t h e r s h a r p l y by at l e a s t one p h y s i c i s t p h i l o s o p h e r . I t has to do with the past f a i l u r e of e v o l u t i o n a r y e x p l a n a t i o n s to f i t w i t h i n the framework of a dedu c t i v e Covering Law Model (Hempel, 1966). I t seems to me that some b i o l o g i s t s and p h i l o s o p h e r s have made.fantastic and c o n t o r t e d attempts to pro v i d e e v o l u t i o n a r y theory with the r e s p e c t a b l e t i t l e of deductive e x p l a n a t i o n , which i t has so f a r not deserved. (Dobzhansky, et_ a l . , 1977; Rosenberg, 1985; F i s h e r , 1985). S i m i l a r problems may not a r i s e i n s t u d i e s which address the d e t a i l s of b i o l o g i c a l f u n c t i o n because the m i c r o s c o p i c laws of chemistry and p h y s i c s are adequate to e x p l a i n , f o r i n s t a n c e , the p r o p e r t i e s of membranes or the nature of DNA bonding. F u n c t i o n a l b i o l o g y asks "how does i t work?" which i s a very d i f f e r e n t q u e s t i o n from "how d i d i t come to be?". The f a c t that a given s t r u c t u r e performs a c e r t a i n f u n c t i o n i s i n s u f f i c i e n t to e x p l a i n why that s t r u c t u r e e x i s t s (O'Grady, 1984). That i s , s t r u c t u r e s cannot be s a i d to a r i s e i n order to perform c e r t a i n f u n c t i o n s . 57 Leaves, f o r i n s t a n c e , cannot be s a i d to have evolved i n order to conduct p h o t o s y n t h e s i s , t r a n s p o r t s o l u t e s and exchange gases. Such an a s s e r t i o n cannot be c a l l e d a l e g i t i m a t e c a u s a l e x p l a n a t i o n i n s c i e n c e . The f a c t that leaves do e x i s t and perform these f u n c t i o n s i s a c o n d i t i o n a l statement. As such, i t may be a necessary p a r t of an e x p l a n a t i o n , but i t must remain only a p a r t . The e x i s t a n c e of leaves would h a r d l y appear to p r o v i d e the f u l l e x p l a n a t i o n of an event d e s c r i b i n g the e x i s t a n c e of l e a v e s . Hempel (1966, pp. 73) q u i c k l y d i s m i s s e s b i o l o g i c a l e x p l a n a t i o n s which seek to give l i v i n g systems such autonomy that they must have t h e i r own set of "laws". "The b a s i c e n t i t i e s and processes p o s i t e d by a theory, and the laws assumed to govern them, must be s p e c i f i e d with a p p r o p r i a t e c l a r i t y and p r e c i s i o n ; otherwise, the theory cannot serve i t s s c i e n t i f i c purpose. T h i s important p o i n t i s i l l u s t r a t e d by the n e o v i t a l i s t i c c o nception of b i o l o g i c a l phenomena. L i v i n g systems, as i s w e l l known, d i s p l a y a v a r i e t y of s t r i k i n g f e a t u r e s that seem to be d i s t i n c t l y purposive or t e l e o l o g i c a l i n c h a r a c t e r . . . the remarkable c o o r d i n a t i o n of the many processes i n a d e v e l o p i n g organism which, as though f o l l o w i n g a common p l a n , l e a d to the formation of a mature i n d i v i d u a l . According to n e o v i t a l i s m , such phenomena do not occur i n n o n l i v i n g systems and cannot be e x p l a i n e d by means of the concepts and laws of p h y s i c s and chemistry alone; r a t h e r they are m a n i f e s t a t i o n s of u n d e r l y i n g t e l e o l o g i c a l a g e n c i e s . . . "This conception may w e l l seem to o f f e r us a deeper 58 understanding of the remarkable b i o l o g i c a l phenomena in question...but understanding in this sense is not what i s wanted in science, and a conceptual system that conveys insight into the phenomena in th i s i n t u i t i v e sense does not for that reason alone qu a l i f y as s c i e n t i f i c theory. The assumptions made by a theory about underlying processes must be d e f i n i t e enough to permit the derivation of s p e c i f i c implications concerning phenomena that the theory i s to explain. " T e s t a b i l i t y - i n - p r i n c i p l e and explanatory import, though c r u c i a l l y important, are nevertheless only minimal necessary conditions that a s c i e n t i f i c theory must s a t i s f y . . . " I find I must agree with Hempel and i t seems that, u n t i l recently, evolutionary explanation has been in a sorry state of a f f a i r s because of these apparently irreparable flaws. My understanding and respect for science has grown enough in the past few years that I believe Hempel's damning statements cannot be denied. The "New Synthesis" or any other variations on a neo-Darwinian theme simply cannot, by d e f i n i t i o n , be considered deductive s c i e n t i f i c explanation because they do not invoke any natural laws as part of the explanation. Uniquely b i o l o g i c a l "laws" simply don't count and never have. P r o b a b i l i t i e s or "chance" events do not count as laws, either. These may serve in a descriptive capacity, but they do not provide cause in the sense that a law does. Natural laws, unlike p r o b a b i l i t i e s , define what appear to be universal r e g u l a r i t i e s . If there are any absolutes in science, they are equivalent to these laws, which are a requisite part of causal explanation. Deductive 59 s c i e n c e does not r e a l l y seem to be such an amorphous or mutable t h i n g . Although the meaning of the term "causal e x p l a n a t i o n " goes f a r beyond the narrow notions of "hypothesis t e s t i n g " , i t s s t r u c t u r e and l i m i t s seem to me to be w e l l d e f i n e d . What, then, might c o n s t i t u t e a "good" theory of e v o l u t i o n and why should the d i f f e r e n c e be so important? Hempel (pp. 75) d e s c r i b e s some of the necessary p r o p e r t i e s i n the f o l l o w i n g passage. "In a f i e l d of i n q u i r y i n which some measure of understanding has a l r e a d y been achieved...a good theory w i l l deepen as w e l l as broaden that understanding. ...such a theory o f f e r s a s y s t e m a t i c a l l y u n i f i e d account of q u i t e d i v e r s e phenomena. I t t r a c e s a l l of them back to the same u n d e r l y i n g processes and p r e s e n t s the v a r i o u s e m p i r i c a l u n i f o r m i t i e s they e x h i b i t as m a n i f e s t a t i o n s of one common set of b a s i c laws." If "basi c laws" cannot be t r a n s l a t e d i n t o "uniquely b i o l o g i c a l laws", but are i n s t e a d d e f i n e d as "laws of nature", then I cannot see that I have any c h o i c e but to at l e a s t examine the U n i f i e d Theory of B i o l o g y presented by Brooks and Wiley (1986). They have succeeded i n p r o v i d i n g e v o l u t i o n a r y phenomena with a s i n g l e c a u s a l e x p l a n a t i o n based on the expansion of an e x i s t i n g n a t u r a l law. The charge that t h i s c o n s t i t u t e s a r e d u c t i o n of b i o l o g i c a l e n t i t i e s and processes to the m i c r o s c o p i c laws of chemistry and p h y s i c s i s immediatly i n v a l i d a t e d by Brooks and Wiley's r e c o g n i t i o n of emergence. To my mind, r e d u c t i o n i s m and emergence are mutually e x c l u s i v e concepts. Brooks and Wiley have c o n s t r u c t e d a theory of b i o l o g i c a l change which has p e r m i t t e d a f u l l e x p l a n a t i o n of the r e s u l t s 60 presented here without the need to t e l l a s i n g l e " j u s t - s o " s t o r y , and i t has done so s t r i c t l y w i t h i n the formal boundaries of d e ductive s c i e n t i f i c e x p l a n a t i o n . As w e l l , I b e l i e v e i t has p e r m i t t e d the a q u i s i t i o n of a b e t t e r understanding of how e v o l u t i o n a r y processes might be manifested by the organisms I have s t u d i e d . In a d d i t i o n , suggestions have been made reg a r d i n g new ideas f o r f u t u r e r e s e a r c h , an e x c e e d i n g l y v a l u a b l e commodity i n s c i e n c e . I f no deeper understanding can be achieved, or no new knowledge obtained, given past t h e o r i e s , then i t i s time to thank the o l d f o r what l i g h t i t c o u l d shed and look to the new. T h i s i s , I b e l i e v e , something that i s long overdue i n b i o l o g y . We have not had a r e v o l u t i o n s i n c e 1859. I t i s time. 61 VI. TAXONOMIC DESCRIPTIONS I. WYETHIA The 14 p r e s e n t l y recognized s p e c i e s of Wyethia are, i n g e n e r a l , e a s i l y d i s t i n g u i s h e d . They are d e s c r i b e d below in an a b b r e v i a t e d v e r s i o n of Weber's (1946) monograph. I had no wish to r e w r i t e t h i s e x c e l l e n t paper and my disagreements with Weber mostly concern the groupings of s p e c i e s i n t o s e c t i o n s . D i r e c t q u o t a t i o n s are used where a d e s c r i p t i o n or grouping seemed q u e s t i o n a b l e . These have been d i s c u s s e d elsewhere. Wyethia Nutt., Jour. Acad. P h i l a . 7:39. 1834 A l a r c o n i a D C , Prodr. 5:537. 1836 M e l a r h i z a K e l l o g g , Proc. C a l i f . Acad. S c i . l ( e d . 2 ) : 3 7 . 1873 Herbaceous, taprooted p e r e n n i a l s , the a e r i a l shoots mostly l e a f y and glabrous to v a r i o u s l y pubescent. Well developed b a s a l l e a v e s present i n most taxa. C a u l i n e leaves a l t e r n a t e and ranging from l i n e a r to d e l t o i d , e n t i r e to dentate. Heads s o l i t a r y to s e v e r a l , u s u a l l y r a d i a t e . P h y l l a r i e s 2 to 4 - s e r i a t e , herbaceous to c o r i a c e o u s , l i n e a r to ovate. The outermost s e r i e s in some taxa are f o l i a c e o u s and very l a r g e . Receptacle convex, paleae f i r m , c o n d u p l i c a t e and p e r s i s t i n g a f t e r the f a l l of the achenes ( c y p s e l a e ) . Disk f l o r e t s yellow, p e r f e c t , t u b u l a r -campanulate, 5-lobed. Anther appendages ovate, the bases s a g i t t a t e . S t y l e branches s l e n d e r , attenuate, h i r s u t u l o u s 62 ( p o l l e n - r e c e p t i v e ) f o r t h e i r e n t i r e l e n g t h . Ray f l o r e t s p i s t i l l a t e and f e r t i l e , mostly yellow (white i n W. h e l i a n t h o i d e s ) , o b l o n g - l a n c e o l a t e . Disk achenes quadrangulate to rhomboidal i n c r o s s - s e c t i o n , glabrous to v a r i o u s l y pubescent. Ray achenes s i m i l a r , but more compressed d o r s a l l y and t r i g o n a l . Pappus u s u a l l y an uneven crown of p e r s i s t a n t , l a c i n i a t e s c a l e s , prolonged i n t o awns at the angles of the achenes i n some taxa, e n t i r e l y l a c k i n g i n a few. The ranges of a l l s p e c i e s are r e s t r i c t e d to areas of North America west of the Rocky Mts. Type s p e c i e s : W. h e l i a n t h o i d e s Nutt. A. AGNORHIZA Agnorhiza (Jeps.)Weber. "Basal leaves absent or g r e a t l y reduced; leaves f a l s l y t r i p l e - n e r v e d by the a c c e n t u a t i o n of two of the lowermost nerves or by anastomoses of the l a t e r a l nerves... F i v e of the s i x s p e c i e s i n c l u d e d i n t h i s s e c t i o n are very r e s t r i c t e d i n t h e i r d i s t r i b u t i o n a l area, one of these, W. r e t i c u l a t a being known d e f i n i t l y from a very small area near Rescue, E l Dorado Co., C a l . They are a p p a r e n t l y s p e c i e s of long s t a n d i n g , c o n s e r v a t i v e , and d e p l e t e d of b i o t y p e s . W. scabra, i n c l u d e d i n t h i s s e c t i o n by reason of the t r i p l e - n e r v e d leaves and l a c k of b a s a l l e a v e s , does not possess the broad leaves c h a r a c t e r i s t i c of the other s p e c i e s . " 1. Wyethia ovata Torrey and Gray. 63 Stems 1-3 dm. high, much exceeded by the c a u l i n e l e a v e s . P l a n t s s i l k y - v i l l o u s , becoming g l a b r a t e . "Basal leaves reduced to s c a l e s or club-shaped rudiments." C a u l i n e leaves broadly ovate with subcordate bases, blades 7-20 cm. long, mostly mucronate at the a p i c e s , e n t i r e , c o r i a c e o u s and c o a r s l y r e t i c u l a t e , "...two of the lowermost l a t e r a l nerves u s u a l l y conspicuous, g i v i n g the l e a v e s a t r i p l e - n e r v e d appearance." P e t i o l e s w e l l developed. Heads s e v e r a l , almost s e s s i l e and c l u s t e r e d i n the a x i l s of the uppermost l e a v e s . Outer p h y l l a r i e s few, f o l i a c e o u s and broadly l i n e a r . F l o r e t s g l a b r o u s , the rays yellow and few i n number. Achenes ca. 10 mm. long, g l a b r o u s . Pappus shallow and c o r o n i f o r m with short awns at the a n g l e s . In the p a s t , p l a n t s have been c o l l e c t e d on h i l l s i d e s and mountain slopes from T u l a r e , Los Angeles, R i v e r s i d e , Orange and San Diego Cos., C a l . I t seems l i k e l y that the s p e c i e s range may have shrunken c o n s i d e r a b l y i n the l a s t c e ntury. I have only seen i t twice i n the f i e l d even a f t e r a reasonably thorough se a r c h . 2. Wyethia b o l a n d e r i (A.Gray) Weber. Stems s l e n d e r , p u r p l i s h , 1.5-3 dm. h i g h . P l a n t s g l a n d u l a r to g l a b r o u s . "Basal leaves reduced to s c a l e - l i k e b r a c t s . " C a u l i n e leaves ovate with t r u n c a t e to cordate bases, blades e n t i r e , 4-12 cm. long, c o r i a c e o u s and c o a r s l y r e t i c u l a t e with mucronate a p i c e s , " . . . p i n n a t e l y veined or tending to have a t r i p l e - n e r v e d appearance..." P e t i o l e s w ell-developed. Heads s o l i t a r y and t e r m i n a l . Outer p h y l l a r i e s few, f o l i a c e o u s , ovate. F l o r e t s glabrous, rays y e l l o w . Achenes ca. 7 mm. long, glabrous, 64 epappose. As i t flowers i n the e a r l y s p r i n g , I d i d not have the o p p o r t u n i t y to c o l l e c t these p l a n t s myself and i t was only sampled once f o r t h i s study. I t i s recorded to range through the S i e r r a Nevada f o o t h i l l s from Butte to Mariposa Co., C a l . 3. Wyethia r e t i c u l a t a Greene P l a n t s ca. 6 dm. high, g l a n d u l a r and somewhat h i s p i d , becoming scabrous. "Enlarged b a s a l leaves absent." The c a u l i n e leaves s e v e r a l , the l a r g e s t 12-15 cm. long, ovate to d e l t o i d and e n t i r e to s e r r u l a t e , with t r u n c a t e to cordate bases. P e t i o l e s w e l l - d e v e l o p e d . "Two of the lowermost l a t e r a l nerves u s u a l l y more conspicuous than the r e s t , g i v i n g the l e a f a t r i p l e - n e r v e d appearance." Heads 1-4. Outer p h y l l a r i e s unequal, l a n c e - l i n e a r . F l o r e t s s p a r s e l y g l a n d u l a r , y e l l o w . Achenes ca. 6 mm. l o n g , glabrous, with a minute, awnless, c o r o n i f o r m pappus. P l a n t s are found only i n the v i c i n i t y of Rescue/Cameron Park, E l Dorado Co., C a l . and are c o n s i d e r e d endangered. Although t h i s s p e c i e s has a very r e s t r i c t e d range i n the c h a p a r r a l - c o v e r e d low f o o t h i l l s of the S i e r r a Nevada, i t seems to be abundant and a g g r e s s i v e i n that area. 4. Wyethia e l a t a H.M.Hall P l a n t s 5-10 dm. high, g l a n d u l a r and with a dense pubescence. "Enlarged b a s a l leaves absent." C a u l i n e leaves s e v e r a l , the l a r g e s t ca. 15-20 cm. long, lance-ovate to d e l t o i d , 65 e n t i r e to dentate, the bases t r u n c a t e to c o r d a t e . P e t i o l e s w e l l -developed. "The p r i n c i p a l v e i n s prominent beneath, two of the lower l a t e r a l ones u s u a l l y conspicuous, g i v i n g the l e a f a t r i p l e - n e r v e d appearance." Heads 1-4. Outer p h y l l a r i e s unequal, l a n c e o l a t e . Disk f l o r e t s glabrous, the rays g l a n d u l a r , y e l l o w . Achenes 8-12 mm. long, mostly g l a b r o u s . Pappus c o r o n i f o r m w i i t h short awns at the achene ang l e s . Found i n open pine woods of the S i e r r a Nevada f o o t h i l l s i n Mariposa, Madera, Fresno and T u l a r e Cos., C a l . The range of t h i s s p e c i e s may have shrunken d u r i n g the l a s t century, but where i t i s p r esent, p l a n t s are abundant. 5. Wyethia invenusta (Greene) Weber P l a n t s 3-6 dm. high, p i l o s e and g l a n d u l a r . "Enlarged b a s a l leaves absent." L a r g e s t c a u l i n e leaves 15-20 cm. long, l a n c e -ovate, e n t i r e , with mostly t r u n c a t e bases. P e t i o l e s w e l l -developed, "...the p r i n c i p a l v e i n s prominant beneath, two of the lowermost l a t e r a l ones o f t e n conspicuous, g i v i n g the leaves a t r i p l e - n e r v e d appearance." Heads mostly s o l i i t a r y , " . . . d i s c o i d , or with a few short ray f l o w e r s . " Outer p h y l l a r i e s unequal, f o l i a c e o u s , l a n c e o l a t e . F l o r e t s g l a b r o u s , yellow. Achenes 7-8 mm. long, g l a b r o u s , " . . . t h e apex de v e l o p i n g k n o b - l i k e p r o j e c t i o n s on the angl e s . Pappus absent." T h i s s p e c i e s i s s a i d to be d i s t r i b u t e d through open woodlands of the s. S i e r r a Nevada, i n Fresno, T u l a r e and Kern Cos., C a l . The s p e c i e s proved extremely d i f f i c u l t to l o c a t e , p a r t l y because much of the range was i n a c c e s s a b l e due to f i r e s 66 and the e f f e c t s of l a n d s l i d e s i n the area d u r i n g the time the f i e l d work was done. My s i n g l e c o l l e c t i o n c o n s i s t e d of p l a n t s which do not completely f i t Weber's d e s c r i p t i o n of the s p e c i e s , but are even l e s s s i m i l a r to any o t h e r s . T h i s p o t e n t i a l i n c o n s i s t a n c y w i l l h o p e f u l l y be c l e a r e d up i n a f u t u r e study, but i t seemed to present no s e r i o u s problem here. 6. Wyethia scabra Hook. Stems s e v e r a l , 1.5-4 dm. high, slender and w h i t i s h . P l a n t s h i s p i d to h a r s h l y scabrous throughout. "Enlarged b a s a l leaves absent." C a u l i n e leaves numerous, the l a r g e s t 3-15 cm. long, s t i f f , l i n e a r , e n t i r e and mainly s e s s i l e , narrowed at the bases, the a p i c e s mucronate. "...the nerves very p a l e , the l a t e r a l ones c o n f l u e n t toward the margin, g i v i n g the leaves a t r i p l e - n e r v e d appearance." Heads s o l i t a r y , t e r m i n a l . Outer p h y l l a r i e s numerous, l i n e a r , a t t e n u a t e . Disk f l o r e t s g l a b rous, the rays pubescent, y e l l o w . Achenes 6-8 mm. long, g l a b r o u s . Pappus short and c o r o n i f o r m , awns l a c k i n g . P l a n t s have been c o l l e c t e d i n d e s e r t s from c e n t r a l and sw. Wyo. to ne. and c e n t r a l Utah. Weber recognized two a d d i t i o n a l v a r i e t i e s to the s p e c i e s . His d e s c r i p t i o n of these f o l l o w : "Wyethia scabra i s a p o l y t y p i c s p e c i e s with three m o r p h o l o g i c a l l y r e c o g n i z a b l e , g e o g r a p h i c a l l y d i s c o n t i n u o u s r a c e s . T h e i r m o r p h o l o g i c a l d i s c o n t i n u i t y i s manifested by d i f f e r e n c e s i n the shape and indument of the p h y l l a r i e s . The most s t r i k i n g departure from the h i s t o r i c a l type of the s p e c i e s i s found i n the race (var. attenuata) i n h a b i t i n g Kane Co., Utah 67 and adjacent Coconino Co., A r i z . In t h i s race the p h y l l a r i e s are extremely long, narrow and very l o o s e l y i m b r i c a t e , with the stout h a i r s tending to be r e s t r i c t e d to the margins. In the second race (var. canescens), occupying the adjacent c o r n e r s of A r i z . , N.M., Colo., and Utah, separated from the f i r s t by the canyons of the Colorado and Green r i v e r s , a dense o v e r - a l l pubescence of s h o r t , s t i f f h a i r s completely covers the p h y l l a r i e s , which are very c l o s e l y i m b r i c a t e , the outer having c o n s p i c u o u s l y recurved t i p s . When t h i s v a r i e t y does not winter-' k i l l , the upper p o r t i o n of the stem sometimes branches, an unusual c o n d i t i o n i n the genus. "The s p e c i e s i n the o r i g i n a l sense i s found i n Wyo. and Utah. The pubescence of the p h y l l a r i e s i s somewhat i n t e r m e d i a t e , but only a few specimens from s. c e n t . Utah, where t h i s race approaches the d i s t r i b u t i o n a l area of var. canescens, might be s a i d to e x h i b i t primary i n t e r g r a d a t i o n . " Only one c o l l e c t i o n of p l a n t s , f i t t i n g the d e s c r i p t i o n and d i s t r i b u t i o n of v ar. a t t e n u a t a , were i n c l u d e d i n t h i s study. A several-day search f o r other r e p r e s e n t a t i v e s of the s p e c i e s was u n s u c c e s s f u l . I t i s p o s s i b l e that v a r . scabra and var. canescens were past f l o w e r i n g by the time my c o l l e c t i n g had extended i n t o t h e i r ranges. I t was hoped that t h i s l a c k of specimens would not have too d e t r i m e n t a l an e f f e c t on t h i s study's a b i l i t y to respond to the main i s s u e s addressed here. B. ALARCONIA 68 A l a r c o n i a D C , Prodr. 5:537.1836, as genus; Nutt. i n Trans. Am. P h i l . Soc. 11.7:353.1840, as s e c t i o n . "Heads very l a r g e , s o l i t a r y , the outer s e r i e s of p h y l l a r i e s g r e a t l y exceeding the d i s k . Basal and c a u l i n e leaves e l l i p t i c -ovate, very l a r g e . Achenes 10-15 mm. long, the l a r g e s t i n the genus. Pappus c a l y x - l i k e , conspicuous, o f t e n prolonged i n t o s t o u t awns on the a n g l e s . Median v a s c u l a r bundles of c o r o l l a -lobes w e l l developed." 7. Wyethia g l a b r a A.Gray P l a n t s 1.5-4 dm. h i g h , g l a n d u l a r and glabrous to s p a r s l y p i l o s e . "Basal leaves very l a r g e , 3.5-4 dm. l o n g . . . p i n n a t e l y v e i n e d . . . " , oblong to e l l i p t i c . Margins e n t i r e and o f t e n undulate, bases narrowed to short p e t i o l e s . C a u l i n e leaves s i m i l a r , c a. 1.5 dm. l o n g . Heads u s u a l l y s o l i t a r y and t e r m i n a l , very l a r g e . Outer p h y l l a r i e s very l a r g e , ovate, f o l i a c e o u s . F l o r e t s s l i g h t l y g l a n d u l a r , y e l l o w . Achenes 10-12 mm. long, puberulent above. Pappus an uneven crown with s h o r t , broad awns. I d i d not have the o p p o r t u n i t y to c o l l e c t t h i s myself because of i t s e a r l y f l o w e r i n g time. Although only one c o l l e c t i o n was made f o r t h i s study, p l a n t s have been found through the Coast ranges from Mendicino to San L u i s Obispo Cos., C a l . 8. Wyethia h e l e n i o i d e s (DC.) Nutt. P l a n t s 3-6 dm. h i g h , densly tomentose, becoming g l a b r a t e 69 with age. "Basal l e a v e s very l a r g e , about 3 dm. l o n g . . . p i n n a t e l y v e i n e d . . . " , oblong to e l l i p t i c , mainly e n t i r e , bases a b r u p t l y narrowed to short p e t i o l e s . C a u l i n e leaves s i m i l a r but s m a l l e r , 1-2 dm. lo n g . Heads mostly s o l i t a r y , t e r m i n a l , very l a r g e . Outer p h y l l a r i e s very l a r g e , unequal, lance-ovate, f o l i a c e o u s . Disk f l o r e t s with pubescent l o b e s ; the rays glabrous, y e l l o w . Achenes 12-15 mm. long, pubescent above. Pappus co r o n i f o r m , uneven, with s h o r t , broad awns. The range of t h i s s p e c i e s o v e r l a p s that of the other member of the s e c t i o n i n the Coast ranges from Mendocino to San L u i s Obispo Cos., C a l . In a d d i t i o n , W. h e l e n i o i d e s occurs i n the w. f o o t h i l l s of the S i e r r a Nevada, from E l Dorado to Mariposa Cos., C a l . I t a l s o blooms i n the e a r l y s p r i n g and I have not c o l l e c t e d i t myself. C. WYETHIA (EUWYETHIA). "True Wyethia" A.Gray i n Brewer, Watson and Gray, Bot. C a l i f . 1:349.1876, as s e c t i o n . "Basal leaves p r e s e n t , much l a r g e r than the c a u l i n e l e a v e s , l a n c e o l a t e i n general o u t l i n e . P h y l l a r i e s graduate, subequal, e r e c t and appressed, not or only s l i g h t l y exceeding the d i s k . Median v a s c u l a r bundles of c o r o l l a lobes u s u a l l y l a c k i n g . Achenes 6-10 mm. long. "From a g e o l o g i c a l standpoint Euwyethia i s probably the most recent s e c t i o n of the genus. The s p e c i e s , f o r the most p a r t , occupy e x t e n s i v e a r e a s ; they have a broad range of 70 h a b i t a t s and are a p p a r e n t l y a g g r e s s i v e . " 9. Wyethia m o l l i s A. Gray P l a n t s densly tomentose, becoming g l a b r a t e i n age. "Basal leaves 2-4 dm. l o n g . . . p i n n a t e l y v e i n e d . . . " , l a n c e o l a t e to oblong, e n t i r e , bases narrowed to w e l l developed p e t i o l e s . C a u l i n e l e a v e s s i m i l a r but s m a l l e r . Heads 1-3. Outer p h y l l a r i e s few, herbaceous, la n c e - o v a t e . Disk f l o r e t s with pubescent lob e s , rays g l a b r o u s , yellow. Achenes 8-10 mm. long, pubescent above. Pappus c o r o n i f o r m with long awns. The s p e c i e s range extends from the s. Cascades of Lake Co., Ore., through the S i e r r a Nevada to Fresno Co., C a l . and adjacent Nev. 10. Wyethia h e l i a n t h o i d e s Nutt. P l a n t s 3-5 dm. high, p i l o s e . "Basal leaves l a r g e , but narrower than those of W. a m p l e x i c a u l i s . . . a b r u p t l y or g r a d u a l l y narrowed to the p e t i o l e , the l a t e r a l nerves ascending...", 1-3 dm. long, l a n c e - e l l i p t i c , u s u a l l y e n t i r e . C a u l i n e l e a v e s s m a l l e r s e s s i l e to amply p e t i o l a t e . . . " , e n t i r e . Heads s o l i t a r y , t e r m i n a l . P h y l l a r i e s numerous, t h i n , herbaceous, l a n c e - l i n e a r , with spreading h a i r s . Disk f l o r e t s g l a b rous, rays s l i g h t l y p i l o s e , white. Achenes 8-10 mm. long, pubescent. Pappus a shallow, l a c i n i a t e crown, o f t e n with s h o r t , t h i n awns. The s p e c i e s ranges from the Blue and Ochoco mts. of Ore., through s. Ida., ne. Nev., sw. Mont, and w. Wyo. i n mountain 71 meadows and c l e a r i n g s . 11. Wyethia a n g u s t i f o l i a (DC.) Nutt. "Plant 1.5-6 dm. h i g h , from almost scapose to t a l l and l e a f y , s o f t l y appressed-pubescent to h i r s u t e and scabrous. Leaves very v a r i a b l e , the b a s a l l a r g e , the c a u l i n e reduced upward,... l o n g - p e t i o l a t e to s e s s i l e , tapered at both ends or sometimes the bases of the upper ones almost c o r d a t e , e n t i r e or sometimes i r r e g u l a r l y s e r r a t e - or undulate-margined...", 15-35 cm. long, l a n c e - l i n e a r to l a n c e - o v a t e . Heads mostly 1 (to 3). P h y l l a r i e s numerous and l a n c e o l a t e , with c i l i a t e margins. Disk f l o r e t s with pubescent or g l a b r a t e l o b e s . Rays yellow with pubescent or g l a b r a t e tubes. Achenes c a . 8 mm. long, s l i g h t l y pubescent. Pappus a shallow crown with t h i n awns. The s p e c i e s ranges from sw. Wash, and Ore., w. of the C a s c a d e - S i e r r a a x i s to T u l a r e and San L u i s Obispo Cos., C a l . , from sea l e v e l to ca.2,000 m.. These p l a n t s appear to express a great d e a l of w i t h i n -s p e c i e s v a r i a t i o n , as Weber noted. "Wyethia a n g u s t i f o l i a d i s p l a y s a great range of v a r i a t i o n between l o c a l p o p u l a t i o n s , one extreme being an almost scapose, low-growing form, somewhat l o c a l i z e d i n the San F r a n c i s c o Bay a r e a , the other a t a l l , l e a f y form most w e l l marked in c o l l e c t i o n s from Ore.. However, the s p e c i e s does not appear to e x h i b i t s u f f i c i e n t m o r p h o l o g i c a l or geographic d i s c o n t i n u i t y to warrant nomenclatural r e c o g n i t i o n of the v a r i o u s forms f o r which taxonomic s t a t u s has been proposed." 72 12. Wyethia l o n q i c a u l i s A.Gray P l a n t s 3-6 dm. high with a sparse, f i n e pubescence. "Basal leaves o b l o n g - l a n c e o l a t e , 15-40 cm. l o n g g l o s s y , acute, e n t i r e or d e n t i c u l a t e , s e s s i l e or t a p e r i n g to a u s u a l l y short p e t i o l e ; l a t e r a l v e i n s ascending." C a u l i n e leaves s i m i l a r but s m a l l e r , s e s s i l e . Heads 1-4. Outer p h y l l a r i e s f o l i a c e o u s , l a n c e o l a t e . Disk and ray f l o r e t s g l a b r o u s . Achenes c a . 7 mm. long with a s h o r t , lobed c o r o n i f o r m pappus. P l a n t s are s a i d to be found on open slo p e s of the n. Coast ranges i n e. Humboldt, adjacent T r i n i t y and ne. Mendocino Cos., C a l . I was a b l e to l o c a t e i t only once i n the Mendocino Nat. For. 13. Wyethia a m p l e x i c a u l i s (Nutt.) Nutt. P l a n t s glabrous and r e s i n - v a r n i s h e d . "Basal leaves very l a r g e , o b l o n g - l a n c e o l a t e , 20-40 cm. l o n g . . . e n t i r e , sometimes d e n t i c u l a t e o r . . . s h a l l o w l y dentate, p i n n a t e l y v e i n e d . . . g r a d u a l l y narrowed to the short p e t i o l e . . . " C a u l i n e leaves s i m i l a r but s m a l l e r and s e s s i l e , u s u a l l y with c l a s p i n g bases, e n t i r e or d e n t i c u l a t e . "Heads l a r g e , s e v e r a l ( o c c a s i o n a l l y s o l i t a r y ) , the t e r m i n a l head l a r g e s t . " Outer p h y l l a r i e s subequal, f o l i a c e o u s , l a n c e - o v a t e . F l o r e t s yellow, g l a b r o u s . Achenes 8-10 mm. long, g l a b r o u s . Pappus cor o n i f o r m , sometimes with t h i n awns. P l a n t s are common in open g r a s s l a n d s and woods at mid-a l t i t u d e s i n sw. Mont and through Ida., se. Wash., and e. Ore., w. to the Columbia Gorge, s. to se. Wyo, adjacent Colo, n. Utah 73 and ne. Nev. Weber noted, "Wyethia a m p l e x i c a u l i s i s the most widely d i s t r i b u t e d s p e c i e s i n the genus, o v e r l a p p i n g the d i s t r i b u t i o n a l areas of W. h e l i a n t h o i d e s , W. a r i z o n i c a and W. sca b r a . " Wyethia a r i z o n i c a A.Gray " S i m i l a r to W. a m p l e x i c a u l i s , But densly h i r s u t e - p u b e s c e n t , becoming scabrous, or sometimes g l a b r a t e , the le a v e s narrower and s m a l l e r , with slender p e t i o l e s , only the uppermost c a u l i n e leaves s e s s i l e or somewhat c l a s p i n g . Disk flowers pubescent on the l o b e s . Ray flowers yellow, d o r s a l l y h i r s u t e . " P l a n t s are found on dry h i l l s and mountain s l o p e s , w. Colo, and adjacent Utah, A r i z , and N.M. 74 I I . BALSAMORHIZA The organisms i n c l u d e d i n Balsamorhiza Nutt. have proven d i f f i c u l t to group. Species and subspecies have been r e -o r g a n i z e d s e v e r a l times (Sharp, 1935; Weber, 1946; Ownbey and Weber, 1943; C r o n q u i s t , 1955). T h i s r e f l e c t s the c o n f u s i o n which seems to surround attempts to d e l i m i t s p e c i e s i n t h i s genus. The c l a r i f i c a t i o n of Balsamorhiza w i l l h o p e f u l l y be the s u b j e c t of a f u t u r e study. For the present, the s p e c i e s are used i n a somewhat gen e r a l sense, as d e s c r i b e d . P a r t i c u l a r d i f f i c u l t i e s and the assumptions made with respect to them are noted throughout the f o l l o w i n g taxonomic d e s c r i p t i o n s , f o l l o w i n g C r o n q u i s t ' s (1955) treatment. Heads s o l i t a r y , or s e v e r a l i n some, r a d i a t e , the rays p i s t i l l a t e and f e r t i l e , mostly yellow. P h y l l a r i e s mostly herbaceous, the outer sometimes f o l i a c e o u s . Paleae f i r m , p e r s i s t a n t . Disk f l o r e t s numerous, mostly yellow, anther bases s a g i t t a t e , s t y l e branches s l e n d e r , the submarginal s t i g m a t i c l i n e s h i s p i d u l o u s throughout, or toward a p i c e s o n l y . Pappus u s u a l l y l a c k i n g , d i s k achenes compressed-quadrangular. ". . . s c a p i f o r m p e r e n n i a l s with r o s e t t e s of l a r g e b a s a l leaves, o f t e n with one or more much reduced, g e n e r a l l y b r a c t l i k e c a u l i n e l e a v e s . " "About 12 s p e c i e s . . . a l l t a p r o o t e d p e r e n n i a l s . In the s e c t i o n A r t o r h i z a the t a p r o o t i s very l a r g e and deep-seated, and i s surrmounted by a m u l t i c i p i t a l caudex b e a r i n g . . . s e v e r a l r o s e t t e s of l e a v e s . In the s e c t i o n Eubalsamorhiza the taproot i s 75 s m a l l e r and more c a r r o t l i k e . . . the leaves of a l l the s p e c i e s are s i m i l a r on j u v e n i l e p l a n t s , being e l l i p t i c and e n t i r e . By p r o g r e s s i v e stages the a d u l t - t y p e leaves are e v e n t u a l l y produced." C r o n q u i s t d e s c r i b e s p a r t of the d i f f i c u l t y i n the assignment of p a r t i c u l a r i d e n t i t i e s to some members of Balsamorhiza i n the f o l l o w i n g . "The taxonomic d i f f i c u l t i e s of the genus a r i s e l a r g e l y from the weakness or absence of g e n e t i c b a r r i e r s , so that any two s p e c i e s w i l l h y b r i d i z e where they grow together. In most cases these h y b r i d s do not swamp the p a r e n t s , o c c u r i n g i n s t e a d only i n the immediate area of c o n t a c t , with l o c a l i n t r o g r e s s i o n seldom d e t e c t a b l e f o r more than a few hundred yards. S e v e r a l of these h y b r i d s have been mistaken in the herbarium f o r d i s t i n c t s p e c i e s , and have been named as such. B. t e r e b i n t h a c e a (Hook.) Nutt. was e v i d e n t l y based on a h y b r i d between B. d e l t o i d e a and B. hookeri ..." He goes on to l i s t other named h y b r i d s . Although s e v e r a l c o l l e c t i o n s of h y b r i d s were made durin g the f i e l d work, data from them were not used f o r any of the analyses and they are not mentioned f u r t h e r here. A couple of s p e c i e s , such as B. rosea were omitted because attempts to l o c a t e them were u n s u c c e s s f u l . The only taxa d e s c r i b e d here are those that were i n c l u d e d i n the study. A. ARTORHIZA P l a n t s with a l a r g e , woody root and a m u l t i c i p i t a l caudex. 76 Basal leaves d e l t o i d to s a g i t t a t e , e n t i r e to c r e n a t e . C a u l i n e leaves much smal l e r than the b a s a l , a l t e r n a t e , l a n c e - l i n e a r , e n t i r e . Rays yellow. 1. Balsamorhiza s a g i t t a t a (Pursh) Nutt. Basal leaves cordate to t r i a n g u l a r - h a s t a t e , up to 30 cm. l o n g . P e t i o l e s w e l l developed. P l a n t s with a f i n e , woolly pubescence. Stems 2-8 dm. h i g h . C a u l i n e leaves few, s m a l l , a l t e r n a t e and l i n e a r . Heads u s u a l l y s o l i t a r y , t e r m i n a l . Outer p h y l l a r i e s few, lanate-tomentose. Achenes gl a b r o u s , epappose. Common on open h i l l s i d e s and f l a t s up to h i g h e l e v a t i o n s , s. B.C. to s. C a l . , w. of the C a s c a d e - S i e r r a a x i s , e. to Mont., S.D. and C o l o . T h i s i s the most widespread s p e c i e s i n the genus. It i s e a s i l y r e c o g n i z a b l e and o f t e n forms a major component of sagebrush and ponderosa pine v e g e t a t i o n . 2. Balsamorhiza careyana A.Gray "...stem 2-10 decimeters t a l l , s c a p i f o r m but u s u a l l y with s e v e r a l s t r o n g l y reduced narrow l e a v e s . . . " Basal l e a v e s up to 30 cm. long, with cordate to t r i a n g u l a r - h a s t a t e bases, u s u a l l y r a t h e r s t i f f and veiny, s l i g h t l y h i r s u t e , sometimes g l a n d u l a r , on long p e t i o l e s . Heads u s u a l l y s e v e r a l , r a t h e r s m a l l . P h y l l a r i e s s l i g h t l y woolly, not much en l a r g e d . Rays tend to p e r s i s t on the achenes and become w h i t i s h as they dry. Achenes pubescent. Grows i n open, dry p l a c e s e. of the Cascades from s. B.C. 77 to c. Ore. I have only one c o l l e c t i o n of reasonable c e r t a i n i d e n t i t y . 3. Balsamorhiza d e l t o i d e a Nutt. "...stem 2-10 dm. t a l l , s c apiform, but u s u a l l y with s e v e r a l , s t r o n g l y reduced narrow l e a v e s ; c e n t r a l head l a r g e . . . l a t e r a l heads when present o b v i o u s l y s m a l l e r . . . " Basal leaves up to 30 cm. long with t r i a n g u l a r - h a s t a t e to cordate bases, u s u a l l y crenate margined, s l i g h t l y h i r s u t e and g l a n d u l a r . Outer p h y l l a r i e s r a t h e r l a r g e and f o l i a c e o u s , s l i g h t l y woolly. Rays "...soon deciduous, not becoming papery; achenes g l a b r o u s . " P l a n t s range from s. Van. I., w. of the Cascades, to s. C a l . Only one c o l l e c t i o n c o u l d be i n c l u d e d here. C r o n q u i s t a l s o r e c o g n i z e s a taxon he c a l l s B. careyana v a r . intermedia, whose d e s c r i p t i o n sounds i d e n t i c a l to that f o r B. d e l t o i d e a , except that i t s ray f l o r e t s p e r s i s t and become papery i n age. He g i v e s no j u s t i f i c a t i o n f o r the a p p a r e n t l y overwhelming importance of t h i s s i n g l e c h a r a c t e r . The s i t u a t i o n seems c o n f u s i n g and i n need of f u r t h e r study. B. BALSAMORHIZA (EUBALSAMORHIZA) 12 most of the taxa i n t h i s s e c t i o n possess a s m a l l , c a r r o t -l i k e t a p r o o t . A l l have w e l l developed b a s a l leaves with margins ranging from s e r r a t e to p i n n a t i f i d to b i p i n n a t e l y compound. C a u l i n e l e a v e s are u s u a l l y l i m i t e d to a s i n g l e , o p p o s i t e p a i r . Heads s o l i t a r y , t e r m i n a l . The c o n f u s i o n surrounding the taxonomic groupings of 78 e n t i t i e s reaches i t s z e n i t h i n t h i s s e c t i o n . Although the s e c t i o n i t s e l f seems mostly n a t u r a l , the s p e c i e s and subspecies have been s h i f t e d around s e v e r a l times and are g r e a t l y i n need of f u r t h e r study. 4. Balsamorhiza hookeri Nutt. Taproots small and crowns simple. Stems "...0.6-4 dm. t a l l , with one or more inconspicuous, l i n e a r , e n t i r e or toothed-p i n n a t i f i d b r a c t near the base." Basal leaves 1-4 dm. long, pinnate to b i p i n n a t e l y compound, the d i v i s i o n s narrow. Heads s o l i t a r y , t e r m i n a l . P h y l l a r i e s l i n e a r to ovate, pubescent at l e a s t on margins. Achenes glabrous, epappose. "Dry, rocky outcrops i n the f o o t h i l l s and lowlands; d i s t r i b u t i o n a p p a r e n t l y d i s c o n t i n u o u s ; dry p l a i n s of c. Wash., w. as f a r as Bingen; a p p a r e n t l y t o t a l l y absent from Ore.; w.c. Ida., e. to sw. Wyo. and nw. Colo., s. to C a l . and s. Utah... "Th i s s p e c i e s shows a most c o n f u s i n g p a t t e r n of i n t r a s p e c i f i c v a r i a b i l i t y , only p a r t l y c o r r e l a t e d with geography..." Cronquist r e c o g n i z e s at l e a s t 5 subspecies of B. h o o k e r i . P l a n t s resembling B. hookeri ssp. lagocephala (Sharp)Cronq. were used i n t h i s study. 5. Balsamorhiza incana Nutt. Crowns simple, stems 1.5-7 dm. t a l l . Basal l e a v e s 1-4.5 dm. long, p i n n a t e l y d i v i d e d , "...the d i v i s i o n s 1.5-6 cm. long, e n t i r e or with a few coarse t e e t h or segments, broader than i n 79 B. hookeri and B. h i r s u t a , sometimes as much as 4 cm. wide; a p a i r of reduced, but s t i l l f a i r l y w e l l developed and p i n n a t i f i d c a u l i n e leaves commonly borne j u s t above the base of the... stem..." P l a n t s covered with a long, s i l k y white tomentum. Heads s o l i t a r y , t e r m i n a l . P h y l l a r i e s woolly, "...with ovate or l a n c e o l a t e base, sometimes merely p o i n t e d , sometimes with long, r e f l e x e d , subcaudate apex..." Achenes glabrous, epappose. "Meadows and other moderately moist to moderately dry open p l a c e s , i n g e n e r a l l y more mesic h a b i t a t s than B. h o o k e r i ; moderate e l e v a t i o n s i n the mts. of ne. Ore. and se. Wash., e. through n. Ida. to n. and w.c. Wyo. and se. Mont." One c o l l e c t i o n (#8411, see Appendix A.) was made o u t s i d e t h i s range, but f i t s the s p e c i e s d e s c r i p t i o n i n other r e s p e c t s and so was in c l u d e d here. 7. Balsamorhiza s e r r a t a Nels.& Macbr. Stems 1-4 dm. t a l l , " . . . o f t e n with a p a i r of much reduced leaves near the base..." P l a n t s with a simple crown. Basal "...leaves green, scabrous and s t r o n g l y r e t i c u l a t e - v e i n y , p e t i o l a t e , the blade 4-20 cm. l o n g . . . v a r y i n g from d e l t o i d - o v a t e and merely s h a r p l y s e r r a t e , to e v i d e n t l y p i n n a t i f i d , o f t e n even on the same p l a n t , but n e a r l y a l l p l a n t s with some merely toothed l e a v e s . . . " Heads s o l i t a r y , t e r m i n a l . P h y l l a r i e s g l a b r a t e to s t r o n g l y pubescent, l a n c e - l i n e a r . Achenes glabrous, epappose. P l a n t s occur i n f r e q u e n t l y on dry, rocky h i l l s i d e s and outcrops i n Ore. and s. Wash., e. of the Cascades. 80 7. Balsamorhiza macrophylla Nutt. P l a n t s are l a r g e r than other s p e c i e s i n t h i s s e c t i o n and resemble members of A r t o r h i z a i n the p o s s e s s i o n of l a r g e t a p r o o t s and multi-branched crowns. P l a n t s g l a n d u l a r with a long, sparse pubescence. Stems 3-10 dm. high with a s i n g l e , w e l l developed p a i r of c a u l i n e l e a v e s , p i n n a t i f i d to p i n n a t e l y compound. Basal leaves 3-6 dm. long, p i n n a t e l y compound, the d i v i s i o n s e n t i r e to c o a r s l y few-toothed. Heads s o l i t a r y , t e r m i n a l , l a r g e . P h y l l a r i e s r a t h e r long and f o l i a c e o u s , l a n c e o l a t e . Achenes glabrous, epappose. P l a n t s occur i n open areas at ra t h e r h i g h e l e v a t i o n s from n. Utah, n. to se. Ida., through w. Wyo. and n. to Mont. 81 I I I . TAXONOMIC OUTGROUPS Se v e r a l other taxa were a l s o used f o r outgroup comparison and these are d e s c r i b e d i n the f o l l o w i n g s e c t i o n . A l l of these are c o n s i d e r e d c l o s e l y enough r e l a t e d to Wyethia and Balsamorhiza to at l e a s t be i n c l u d e d i n the t r i b e H e l i a n t h e a e . That i s , they are mostly p e r e n n i a l s or annuals with herbaceous p h y l l a r i e s , t a i l l e s s anther bases, and s t i g m a t i c l i n e s which are marginal and run the l e n g t h of the s t y l e branches. Paleae or pappus, when present, are composed of s c a l e s or awns. Within the Heliantheae there has been some d i s p u t e over the l i m i t s of some of the s u b t r i b e s and the i n c l u s i o n of some genera in them (Stuessy, 1976; Robinson, 1981). The attempt was made to u t i l i z e outgroup genera whose c l o s e n e s s to the study taxa i s g e n e r a l l y agreed upon. These i n c l u d e H e l i a n t h e l l a , E n c e l i a , H e l i o p s i s , F l o u r e n s i a and Rudbeckia. C o l l e c t i o n s were a l s o made f o r use i n the a n a l y s i s of w i t h i n -s p e c i e s v a r i a t i o n . Species were chosen f o r t h i s based mostly on t h e i r sympatry with Wyethia and Balsamorhiza. In other words, i f I came a c r o s s them while i n the f i e l d , I c o l l e c t e d them. The genera represented i n c l u d e H e l i a n t h e l l a , H e l i a n t h u s and Rudbeckia . A. HELIANTHELLA Torrey & Gray 8 2 The f o l l o w i n g d e s c r i p t i o n s are condensed from Weber's ( 1 9 5 2 ) monograph of the genus. He r e c o g n i z e d 8 s p e c i e s of herbaceous, ta p r o o t e d p e r e n n i a l s . "The leaves may be e i t h e r c a u l i n e or b a s a l . Basal r o s e t t e s of leaves are u s u a l l y formed by buds which do not produce a f l o w e r i n g stem duri n g the c u r r e n t season. Flowering stems r a r e l y possess l a r g e b a s a l l e a v e s , although the lower p a i r s of c a u l i n e leaves are sometimes q u i t e l a r g e . The leaves of the b a s a l r o s e t t e are s i m i l a r i n shape, t e x t u r e and indument to those of the stem. "The lowermost c a u l i n e l e a v e s are always o p p o s i t e , while the uppermost leaves may be a l t e r n a t e or o p p o s i t e . In some s p e c i e s , l e a v e s are o c c a s i o n a l l y produced three at a node. The l e a f margins are always e n t i r e . The l e a f blades are simple and range i n shape from l i n e a r to o v a t e - l a n c e o l a t e V e n a t i o n i s pinn a t e , but most leaves appear to be t r i p l e - n e r v e d because of the a c c e n t u a t i o n of two of the lowermost l a t e r a l nerves. " A l l s p e c i e s are more or l e s s h i r s u t e or s t r i g o s e . . . t h e pubescence of the f o l i a g e and stem becomes more sparse as the organs e l o n g a t e . . . i n d i v i d u a l s are scabrous or g l a b r a t e at m a t u r i t y . In most s p e c i e s , the i n f l o r e s c e n c e i s a s i n g l e , t e r m i n a l head. A few small a x i l l a r y heads are sometimes present but these o f t e n a b o r t . . . the p h y l l a r i e s range in o u t l i n e from ovate to obtuse...to l a n c e o l a t e or l a n c e - 1 i n e a r . . . I n a few s p e c i e s the outermost p h y l l a r i e s are g r e a t l y enlarged and l e a f -l i k e . P a l e a e . . . o b l a n c e o l a t e , acute, c o n d u p l i c a t e and deciduous with the mature f r u i t s or f a i l i n g . . . a f t e r a n t h e s i s . Ray f l o r e t s neuter, y e l l o w . Disk f l o r e t s p e r f e c t , yellow, p u r p l e or 83 brownish. Disk c o r o l l a s l a c k median v a s c u l a r t r a c e s . Anthers with ovate appendages and s a g i t t a t e bases. S t y l e branches s l e n d e r , t e r e t e or f l a t t e n e d , minutely pubescent from base to apex. Achenes l a t e r a l l y compressed, o f t e n s l i g h t l y wing-margined, cuneate-obovate to obcordate, pubescent to g l a b r o u s . Pappus of s l e n d e r , p e r s i s t a n t awns and a short l a c r e a t e crown, or l a c k i n g . N=15." Of the 8 s p e c i e s r e c o g n i z e d by Weber (1952), only 2 were a c t u a l l y c o l l e c t e d f o r use i n the study of w i t h i n - s p e c i e s v a r i a t i o n . These are d e s c r i b e d below. For the p h y l o g e n e t i c a n a l y s i s employing H e l i a n t h e l l a as a taxonomic outgroup, c h a r a c t e r s were taken from the g e n e r i c d e s c r i p t i o n above. The s p e c i e s d e s c r i b e d below were used in the PCA-based a n a l y s e s . 1. H e l i a n t h e l l a u n i f l o r a (Nutt.) Torrey & Gray Stems 4-12 dm. high, o f t e n somewhat h i r s u t e , becoming scabrous, simple or few-branched. Leaves mostly l a n c e - e l l i p t i c , " . . . e n l a r g e d b a s a l leaves absent, the c a u l i n e l e a v e s i n 3-6 o p p o s i t e p a i r s , the middle p a i r s l a r g e s t , 3-nerved from below the middle... s h o r t - p e t i o l e d or s e s s i l e , the blades 12-25 cm. long, the upper s e s s i l e . Heads s o l i t a r y and t e r m i n a l , . . . a few supressed l a t e r a l heads o f t e n present..." P h y l l a r i e s subequal, l a n c e o l a t e - a t t e n u a t e , " . . . t h e outermost f r e q u e n t l y elongated and l e a f - l i k e . . . " F l o r e t s y e l l o w . Achenes"... 6-7 mm. long, narrowly obovate, c i l i a t e above, appressed-pubescent on the l a t e r a l f a c e s ; pappus of two s l e n d e r awns... int e r m e d i a t e squamellae short or o b s o l e t e . " 84 P l a n t s are found through the Rocky Mts., intermountain p l a t e a u s and ranges of the Great Basin from s. Ida., se. Ore. and sw. Mont., s. to n. Nev., s. Utah and nw. C o l o . , " . . . i n a wide v a r i e t y of h a b i t a t s . . . " I n d i v i d u a l s used i n t h i s study f i t the d e s c r i p t i o n of v a r. d o u g l a s i i (T.& G.) Weber. 2. H e l i a n t h e l l a c a l i f o r n i c a A.Gray P l a n t s 1.5-6 dm. high, h i r s u t u l o u s , becoming sc a b e r u l o u s . " . . . l e a v e s o b l o n g - l a n c e o l a t e , 5-26 cm. long, t a p e r i n g to elongate p e t i o l e s , the lowermost c a u l i n e leaves o p p o s i t e ; heads u s u a l l y s o l i t a r y on long p e d u n c l e s ; . . . p h y l l a r i e s l a n c e o l a t e , densly pubescent to c i l i a t e or g l a b r a t e . . . " Achenes compressed, 6-10 mm-., glabrous, epappose. The r e p r e s e n t a t i v e used here was v ar. nevadensis (Greene) Jepson, which a p p a r e n t l y d i f f e r s from the s p e c i e s i n i t s achenes, "...which always bear two s h o r t , p e r s i s t a n t awns and u s u a l l y s e v e r a l low int e r m e d i a t e squamellae." The v a r i e t y ranges from s. Ore., to Kern Co., C a l . , through the S i e r r a Nevada, e. to Washoe Co., Nev and w. of the C e n t r a l V a l l e y to Lake Co., C a l . I t i s r e p l a c e d by the s p e c i e s on the w. s i d e of the V a l l e y . B. HELIANTHUS L. 85 Mostly p e r e n n i a l , sometimes annual herbs, i n c l u d i n g c a . 60 s p e c i e s (Hitchcock, et a_l. , 1955). The c a u l i n e leaves simple, at l e a s t the lowermost ones o p p o s i t e . Ray f l o r e t s yellow, n e u t r a l . Disk f l o r e t s p e r f e c t and f e r t i l e , the anthers with e n t i r e or s l i g h t l y s a g i t t a t e bases, the s t y l e branches f l a t t e n e d , h i s p i d u l o u s , as are t h e i r appendages. P h y l l a r i e s subequal, mostly herbaceous. Paleae p e r s i s t a n t . Achenes t h i c k , but somewhat compressed, u s u a l l y g l a b r o u s . Pappus mostly of two deciduous awns. Only one s p e c i e s was i n c l u d e d i n assessment of w i t h i n - s p e c i e s v a r i a t i o n . I t i s d e s c r i b e d below. 1 . H e l i a n t h u s c u s i c k i i A.Gray P e r e n n i a l p l a n t s 2 - 1 2 dm. t a l l , the stems a r i s i n g from the crown of a th i c k e n e d t a p r o o t , o f t e n f r e e l y branching, h i r s u t e to scabrous. Leaves l a n c e o l a t e to somewhat l i n e a r , e n t i r e , 3 - 1 2 cm. long, a p p a r e n t l y t r i p l e - n e r v e d , s u b s e s s i l e and o p p o s i t e , at l e a s t below. P h y l l a r i e s l a n c e - l i n e a r , u s u a l l y with spreading h a i r s . P l a n t s i n dry p l a i n s and f o o t h i l l s of e. Ore., sw. Ida. and ne. C a l . , n. between the Columbia R. and the Cascades to E l l e n s b u r g , Wash. C. RUDEECKIA L. P l a n t s mostly p e r e n n i a l , some annual. C a u l i n e leaves 86 a l t e r n a t e , e n t i r e to p i n n a t i f i d . Heads r a d i a t e or d i s c o i d , the rays n e u t r a l , the d i s k f l o r e t s p e r f e c t and f e r t i l e . Anthers with s a g i t t a t e or obtuse bases, s t y l e branches f l a t t e n e d and with h i r s u t e appendages. P h y l l a r i e s subequal or unequal, mostly herbaceous. Receptacle c o n i c or columnar. Achenes quadrangular or somewhat f l a t t e n e d , g l a b r o u s . Pappus a s h o r t , toothed crown, or l a c k i n g . T h i s North American genus c o n t a i n s about 24 s p e c i e s and was used here f o r outgroup comparison. M o r p h o l o g i c a l c h a r a c t e r s were taken from Hitchcock, et a l . (1955). The f o l l o w i n g s p e c i e s was a l s o used i n the e x p l o r a t o r y data a n a l y s i s p a r t of t h i s study. 1 . Rudbeckia o c c . i d e n t a l i s Nutt. P l a n t s p e r e n n i a l , 5-20 dm. t a l l , g l a b r a t e or with short pubescence. C a u l i n e l e a v e s a l t e r n a t e , up to 25 cm. long, b roadly ovate to e l l i p t i c with subcordate bases, e n t i r e to s e r r a t e , p e t i o l e s s h o r t . Heads d i s c o i d . Pappus a short crown. P l a n t s i n woodlands from sw. Mont, and nw. Wyo. to Wash, Utah and C a l . D. HELIOPSIS Pers. Herbaceous p e r e n n i a l s , r a r e l y annuals with o p p o s i t e , p e t i o l a t e c a u l i n e l e a v e s . The ray f l o r e t s y ellow, p i s t i l l a t e and f e r t i l e , t h e i r c o r o l l a s p e r s i s t i n g on the achenes and becoming 87 papery. Disk f l o r e t s p e r f e c t and f e r t i l e , the anthers mostly with e n t i r e bases, the s t y l e branches f l a t t e n e d and with s h o r t , h i r s u t e appendages. P h y l l a r i e s subequal, at l e a s t the apex herbaceous. Receptacle c o n i c . Achenes u s u a l l y quadrangular. Pappus a s h o r t , few-toothed crown, or l a c k i n g . T h i s genus c o n t a i n s about 6 s p e c i e s n a t i v e to the New World. I t was used here f o r outgroup comparison only, based on d e s c r i p t i o n s i n Hitchcock, et a l . (1955). E. ENCELIA Adans. Low, branching shrubs with a l t e r n a t e , mostly e n t i r e l e a v e s . Heads s o l i t a r y or i n p a n i c l e s , r a d i a t e or d i s c o i d . Ray f l o r e t s n e u t r a l . Disk f l o r e t s f e r t i l e . P h y l l a r i e s 2 to 3 - s e r i a t e . Receptacle convex. Paleae s c a r i c u s and deciduous with the achenes. Achenes f l a t , obovate, pubescent. Pappus of two slender awns, or none. The genus i n c l u d e s about 14 s p e c i e s n a t i v e from the sw. U.S., s. to Peru, C h i l e and the Galapagos Ids.. M o r p h o l o g i c a l i n f o r m a t i o n was taken from Munz and Keck (1968) f o r use i n outgroup comparison. F. FLOURENSIA DC. 88 Resinous shrubs with a l t e r n a t e , l a n c e o l a t e to o v a l , r e t i c u l a t e l y - v e i n e d l e a v e s . Heads s o l i t a r y or i n p a n i c l e s , r a d i a t e or d i s c o i d . Ray f l o r e t s n e u t r a l . Disk f l o r e t s p e r f e c t and f e r t i l e . P h y l l a r i e s 2 to 4 - s e r i a t e , subequal, l a n c e o l a t e to ovate, herbaceous. Paleae s c a r i o u s and deciduous with the achenes. Anthers with ovate appendages and c o r d a t e - s a g i t t a t e bases. S t y l e branches with s h o r t , h i s p i d u l o u s appendages. Achenes th i c k e n e d or somewhat compressed, s i l k y - v i l l o u s at l e a s t on margins. 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F l a v o n o i d s from Wyethia g l a b r a . Phytochem. 24(7 ) : 1614-1616. McCORMICK, S., K. ROBSON and B. BOHM. 1985c. Methylated f l a v o n o l s from Wyethia b o l a n d e r i and Balsamorhiza  macrophylla. Phytochem. 24(9):2133. MUNZ, P. and D. KECK. 1968. A C a l i f o r n i a f l o r a and supplement Univ. of C a l i f . Press, B e r k e l e y . 0'GRADY, R.T. 1984. E v o l u t i o n a r y theory and t e l e o l o g y . J . Theor B i o l . 107:563-578. OWNBEY, M. and W.A. WEBER. 1943. N a t u r a l h y b r i d i z a t i o n i n the genus Balsamorhiza. Am. J . Bot. 30:179-187. ROBINSON, H. 1981. A r e v i s i o n of the t r i b a l and s u b t r i b a l l i m i t s of the Helia n t h e a e ( A s t e r a c e a e ) . Smiths. C o n t r i b . Bot. 51. ROBSON, K.A., R.K. SCAGEL and J . MAZE. A study of morphological v a r i a t i o n w i t h i n and between p o p u l a t i o n s of Balsamorhiza  s a g i t t a t a . i n prep. 93 ROSENBERG, A. 1985. 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P r e n t i c e H a l l , Englewood C l i f f s , N.J. 95 APPENDIX A - PLANT COLLECTIONS 1. Wyethia Wyethia a m p l e x i c a u l i s (Nutt.) Nutt. Hwy. 10 between Cle Elum and Teanaway, by Camp I l l a h e e t u r n o f f . K i t t i t a s Co., Wash. Weedy roadside with g r a s s e s , Pinus ponderosa, Symphoricarpos, Lomatium, Balsamorhiza  s a q i t t a t a . 8401, N i c h o l l s and Robson, June 6, 1984. Wyethia a m p l e x i c a u l i s (Nutt.) Nutt. 1-84, 12 mi. SE of Pendleton, below viewpoint f o r S-bound l a n e s . U m a t i l l a Co., Ore. Mesic h i l l s i d e with g r a s s e s , Lupinus sulphureus, Rosa, C a s t i l l e j a , Balsamorhiza  s a g i t t a t a . 8405, N i c h o l l s and Robson, June 6, 1984. Wyethia a m p l e x i c a u l i s (Nutt.) Nutt. Hwy 89, ca. 2 mi. W of Garden C i t y , R i c h Co., Utah. Clay d e p r e s s i o n i n damp meadow with grasses, M e l i l o t u s , Geranium, A r t e m i s i a . 8433, Robson, J u l y 20, 1984. Wyethia a m p l e x i c a u l i s (Nutt.) Nutt. Road to Tony Grove Lake, 25 mi. E of Logan, Cache Nat. For., Cache Co., Utah. Meadow on dolomite s o i l with nearby P i c e a engelmanni i , Abies l a s i o c a r p a . 8434c, Barkworth and Robson, J u l y 21, 1984. Wyethia amplex i c a u l i s (Nutt.) Nutt. Hwy. 75, ca. 10 mi. N of Sun Valley/Ketchum, Sawtooth Nat. For., B l a i n e Co., Ida. On moist, sandy s o i l with Artemi s i a , Eriogonum and nearby Populus tremuloides, Pinus c o n t o r t a . 8436, Robson, J u l y 23, 1984. 96 Wyethia a m p l e x i c a u l i s (Nutt.) Nutt. U.S.F.S. Rd. 40, ca. 35 mi. S of Pomeroy, U m a t i l l a Nat. For., G a r f i e l d Co., Wash. In open areas between stands of Pinus ponderosa with grasses, H e l i a n t h e l l a u n i f l o r a . 8508, Robson, June 18, 1985. Wyethia a m p l e x i c a u l i s (Nutt.) Nutt. Ca. 3 mi. along t r a i l to J a r d i n e J u n i p e r , 12 mi. E of Logan, Cache Nat. For., Cache Co., Utah. Open montane meadow on dolomite s o i l with g r a s s e s , A r t e m i s i a , Balsamorhiza macrophylla, H e l i a n t h e l l a u n i f l o r a . 8511a, Barkworth and Robson, June 23, 1985. Wyethia a m p l e x i c a u l i s (Nutt.) Nutt. Hwy. 47, w i t h i n a 1 mi. r a d i u s of Lower Mesa F a l l s , Targhee Nat. For., Fremont Co., Ida. C l e a r i n g s among Populus  t r e m u l o i d e s , Pinus c o n t o r t a with g r a s s e s , Wyethia  h e l i a n t h o i d e s and h y b r i d s . 8513, Robson, June 24, 1985. Wyethia angust i f o l i a (DC.) Nutt. Along Sonoma Mt. Rd. N of Penngrove, Marin Co., C a l . Roadside with dry g r a s s e s , Quercus. 8414, N i c h o l l s and Robson, June 13, 1984. Wyethia a n g u s t i f o l i a (DC.) Nutt. Corner of E e l R i v e r and Van A r s d a l e Rds., between Lake P i l l s b u r y and P o t t e r V a l l e y , Mendocino Co., C a l . G r a v e l l y bank by running d i t c h with g r a s s e s , Toxicodendron, Pinus  ponderosa, Ceanothus, A r c t o s t a p h y l o s , Lotus, M e l i l o t u s , T r i f o l i u m . 8416, Robson, June 21, 1984. Wyethia angust i f o l i a (DC.) Nutt. Hwy. 36, ca. 13 mi. E of B r i d g e v i l l e , Humboldt Co., C a l . Moist rocky slope with grasses, Plantago. 8417, Robson, June 21, 1984. 97 Wyethia a n g u s t i f o l i a (DC.) Nutt. Hwy. 4, 0.7 mi. W of A r n o l d , C a l a v e r a s Co., C a l . In weedy roadside d i t c h , p l a n t s with i n s e c t damage. 8421, Robson, June 24, 1984. Wyethia a n g u s t i f o l i a (DC.) Nutt. On Hwy. 49, 16.4 mi. N of Jackson, Amador Co., C a l . B1-5, Maze, May, 1985. Wyethia a n g u s t i f o l i a (DC.) Nutt. S u t t e r Creek, Amador Co., C a l . C1-5, Maze, May, 1985. Wyethia a n g u s t i f o l i a (DC.) Nutt. P a c i f i c Angwin C o l l e g e , Napa Co., C a l . D1-5, Maze, May, 1985. Wyethia a n g u s t i f o l i a (DC.) Nutt. Jasper Ridge B i o l o g i c a l Preserve, Santa C l a r a Co., C a l . For chemistry o n l y . 1772, Bohm, "May, 1984. Wyethia a n g u s t i f o l i a (DC.) Nutt. Napa-Bothe State Park, Napa Co., C a l . I t i s i l l e g a l to take p l a n t s from a s t a t e park and i t i s not necessary. Such an act i s i n d i c a t i v e of a l a z y and arrogant c h a r a c t e r . T h i s c o l l e c t o r d i d i t twice. For chemistry o n l y . 1774, 1775, Bohm, May, 1984. Wyethia a r i z o n i c a A.Gray Hwy. A89, 12.4 mi. S of F l a g s t a f f , Coconino Co., A r i z . In g r a v e l l y , damp de p r e s s i o n by road with g r a s s e s , Pinus  ponderosa, Quercus, M e l i l o t u s . 8430, Robson, J u l y 16, 1984. 98 Wyethia a r i z o n i c a A.Gray Ca. 16 mi. along road to Warner Lake from j e t . with Hwy. 191 S of Moab, Manti-La S a l Nat. For., Grand Co., Utah. In damp, rocky c l a y with g r a s s e s , Quercus, Artemi s i a , Lupinus, Sedum. 8432, Robson, J u l y 18, 1984. Wyethia b o l a n d e r i (A.Gray) Weber Ca. 2 mi. W of Dobbins, Yuba Co., C a l . Most rays l o s t . 1786, Bohm, May 1, 1984. Wyethia e l a t a H.M.Hall Hwy. 49, 4 mi. N of Ahwahnee, Mariposa Co., C a l . Dry roadside c l a y with grasses, Pinus ponderosa, Quercus  k e l l o q g i i , A r c t o s t a p h y l o s , Lupinus, C l a r k i a , Madia• 8422, Robson, June 25, 1984. Wyethia e l a t a H.M.Hall Rd. 222, 1.5 mi. S of North Fork, Madera Co., C a l . G r a v e l l y r o adside c l a y with g r a s s e s , Toxicodendron, Ceanothus, near Pinus s a b i n i a n a , Quercus d o u g l a s i i , A r c t o s t a p h y l o s . ' 8426, Robson, J u l y 10, 1984. Wyethia g l a b r a A.Gray Solano Co., C a l . 3223, Warnock and McCormick, March 16, 1984. Wyethia h e l e n i o i d e s (DC.) Nutt. Ca. 21 mi. S of H o l l i s t e r on Panoche Rd., San Benito Co., C a l . Along roadside i n oak woodland. A1-5, Maze, May, 1985. Wyethia h e l e n i o i d e s (DC.) Nutt. Mi. 74, Hyw. 101, Mendocino Co., C a l . 1786, Bohm, A p r i l 26, 1984. 99 Wyethia h e l i a n t h o i d e s Nutt. 1.1 mi. along r d . to Red Rock Lakes B i r d Sanctuary from j e t . Hwy. 20, Fremont Co., Ida. Open meadow with g r a s s e s , t h i s p o p u l a t i o n almost out of bloom. 8514, Robson, June 25, 1985. Wyethia h e l i a n t h o i d e s Nutt. Near shores of A l t u r a s Lake, o f f Hwy. 75 between Galena Summit and O b s i d i a n , Sawtooth Nat. For., B l a i n e Co., Ida. Open areas among Pinus c o n t o r t a , with g r a s s e s , W. a m p l e x i c a u l i s and h y b r i d s A r t e m i s i a , A r n i c a , P o t e n t i l l a . 8515, Robson, June 26, 1985. Wyethia h e l i a n t h o i d e s Nutt. Along r d . to B u l l Trout Lake, c a . 35 mi. NE of Lowman, Hwy. 21 Boise Nat. For., Boise Co., Ida. In damp, rocky c l e a r i n g s among Pinus c o n t o r t a with few g r a s s e s , Ranunculus, Senecio. 8516, Robson, June 26, 1985. Wyethia invenusta (Greene) Weber Hwy"! 155 (=Evans Rd.) at j e t . of A l t a S i e r r a Rd., Sequoia Nat. For., Kern Co., C a l . Very small pop. i n sandy s o i l with g r a s s e s , Pinus ponderosa, Calocedrus decurrens, Quercus k e l l o g g i i , A s c l e p i a s . 8427, Robson, J u l y 11, 1984. Wyethia l o n g i c a u l i s A.Gray U.S.F.S. Rd. 1N02, 8 mi. S of Lake P i l l s b u r y , Mendocino Nat. For., Lake Co., C a l . Dry, g r a v e l l y s o i l with g r a s s e s , near Pinus ponderosa, Pseudotsuga m e n z i e s i i , Arbutus  menziesi i , A r c t o s t a p h y l o s . 8415, Robson, June 20, 1984. Wyethia m o l l i s A.Gray Hwy. 395, 2 mi. S of Ore. border, Goose Lake V a l l e y , Modoc Co., C a l . Roadside, grassy rangeland. 8410, N i c h o l l s and Robson, June 7, 1984. 100 Wyethia m o l l i s A.Gray Hwy. 89, ca. 5 mi. NW of G r e e n v i l l e , near W shore of Lake Almanor, Plumas co., C a l . In rocky, t h i n c l a y with grasses, Pinus ponderosa, Calocedrus decurrens, A r c t o s t a p h y l o s , Lupinus, H e l i a n t h e l l a c a l i f o r n i c a , Balsamorhiza d e l t o i d e a . 8418b, Robson, June 22, 1984. Wyethia m o l l i s A.Gray USFS Rd. 07 (=Henness Pass Rd.), ca. 6 mi. E of j e t . Hwy 89, 9 mi. S of S i e r r a v i l l e , Tahoe Nat. For., S i e r r a Co., C a l . In damp, rocky, g r a n i t i c s o i l with g r a s s e s , Pinus  ponderosa, Abies c o n c o l o r , Ribes. 8419, Robson, June 23, 1984. Wyethia m o l l i s A.Gray Hwy. 431 at j e t . of Gale S t . , near N Lake Tahoe, Washoe Co., Nev. In dry rocky, g r a n i t i c s o i l with g r a s s e s , Pinus  ponderosa, Calocedrus decurrens, A r c t o s t a p h y l o s , L o n i c e r a , P u r s h i a , Balsamorhiza s a g i t t a t a . 8425b, Robson, June 28, 1984. Wyethia ovata Torrey and Gray Hwy. 74, ca. 5 mi. S of Mtn. Center, near Lake Hemet, San Bernardino Nat. For., R i v e r s i d e Co., C a l . Very sandy g r a n i t i c s o i l with g r a s s e s , Pinus ponderosa, A r t e m i s i a , A r c t o s t a p h y l o s , Ceanothus. 8429, Robson, J u l y 15, 1984. Wyethia r e t i c u l a t a Greene J e t . of La Canada and Cameron Park Dr., Cameron Park, E l Dorado Co., C a l . In Rescue stony, sandy loam with g r a s s e s , Ceanothus, S a l v i a . 8423, Ayres and Robson, June 27, 1984. Wyethia r e t i c u l a t a Greene Deer V a l l e y Rd., .2 mi. W of Green V a l l e y , Rescue, E l Dorado Co., C a l . In Argonaut c l a y loam with grasses, Toxicodendron, Adenostema f a s c i c u l a t a , Quercus. 8424, Ayres and Robson, June 27, 1984. 101 Wyethia scabra var. a t t e n u a t a Weber 10 mi. N of Kanab on Hwy.89, Kane Co., Utah. On red sand dunes with A r t e m i s i a , P u r s h i a , Yucca, Eriogonum, J u n i p e r u s , Quercus, g r a s s e s . 8431, Robson, J u l y 17, 1984. 2. Balsamorhiza Balsamorhiza careyana A. Gray J e t . of Hwy. 395 and Hwy. 74 at Nye, U m a t i l l a Co., Ore. In rocky, exposed l a v a with v a r i o u s g r a s s e s . 8407, N i c h o l l s and Robson, June 7, 1984. Balsamorhiza d e l t o i d e a Nutt. On Hwy. 89 near W shore of Lake Almanor, c a . 5 mi. NW of G r e e n v i l l e , Plumas Co., C a l . C l e a r i n g i n rocky, t h i n , d r y i n g c l a y with Pinus ponderosa, Calocedrus decurrens, A r c t o s t a p h y l o s , Rosa, Ceanothus, Lupinus, H e l i a n t h e l l a  c a l i f o r n i c a , Wyethia m o l l i s . 8418c, Robson, June 22, 1984., Balsamorhiza hookeri var. lagocephala (Sharp) Cronq. 14 mi. W of Vantage, Grant Co., Wash. 8504, Mcintosh and Hawkes, May 17, 1985. Balsamorhiza hookeri var. lagocephala (Sharp) Cronq. Colockum Pass, S of Malaga, K i t t i t a s Co., Wash. In dry, rocky, b a s a l t i c s o i l with A r t e m i s i a t r i d e n t a t a , Agropyron  spicatum, Lithophragma p a r v i f l o r a , Eriogonum. 8501, Maze, Scagel and Robson, May 1, 1985. Balsamorhiza incana Nutt. Old Cabbage H i l l Rd., 1 mi. N of j e t . 1-84, 14 mi. SE of Pendleton, U m a t i l l a Co., Ore. D r y i n g , rocky h i l l s i d e with g r a s s e s . 8406, N i c h o l l s and Robson, June 7, 1984. 1 02 Balsamorhiza incana Nutt. Ukiah c i t y l i m i t , U m a t i l l a Co., Ore. Along roadside at margin of moist pasture with v a r i o u s g r a s s e s . 8409, N i c h o l l s and Robson, June 7, 1984. Balsamorhiza incana Nutt. Along Hwy. 395, Goose Lake V a l l e y , N of A l t u r a s , c a . 41 mi. S of Ore. border, Modoc Co., C a l . Grassy r o a d s i d e . 8411, N i c h o l l s and Robson, June 7, 1984. Balsamorhiza incana Nutt. On Hwy. 126, 14.7 mi. S of A s o t i n , A s o t i n Co., Wash. On roadside with grasses, Medicago s a t i v a . 8509, Robson, June 19,1985. Balsamorhiza macrophylla Nutt. Along road to Tony Grove Lake, Cache Nat. For., c a . 25 mi. E of Logan, Cache Co., Utah. 2 c o l l e c t i o n s made, separated by c a. 1000 f t . e l e v . In grassy c l e a r i n g s of dolomite s o i l among P i c e a enqelmanni i , Abies l a s i o c a r p a . 8434a,b, Barkworth and Robson, J u l y 21, 1984. Balsamorhiza macrophylla Nutt. T r a i l to J a r d i n e J u n i p e r , c a . 12 mi. E of Logan, Cache Co., Utah. In open montane meadow of dolomite s o i l with Artemi s i a , Wyethia a m p l e x i c a u l i s , H e l i a n t h e l l a uni f l o r a . 8511b, Barkworth and Robson, June 23, 1985. Balsamorhiza s a g i t t a t a (Pursh) Nutt. Entrance to Blue Lake Camp, Modoc Nat. For., Modoc Co., C a l . On rocky h i l l s i d e with A r t e m i s i a , Pinus ponderosa, Wyethia m o l l i s , g r a s s e s . 8412, N i c h o l l s and Robson, June 7, 1984. Balsamorhiza s a g i t t a t a (Pursh) Nutt. Hwy. 431 at j e t . of Gale St., near N Lake Tahoe, Washoe Co., Nev. In dry, rocky, g r a n i t i c s o i l with g r a s s e s , Pinus  ponderosa, Calocedrus decurrens, A r c t o s t a p h y l o s , L o n i c e r a , , P u r s h i a , Wyethia m o l l i s . 8425a, Robson, June 28, 1984. 103 Balsamorhiza s a g i t t a t a (Pursh) Nutt. 0.7 mi. S of Galena Summit on Hwy. 75, Sawtooth Nat. For., B l a i n e Co., Ida. On damp, rocky h i l l s i d e with A r t e m i s i a , G i l i a , P otenti11a, g r a s s e s . 8437, Robson, J u l y 24, 1984. Balsamorhiza s a g i t t a t a (Pursh) Nutt. 23 mi. S of N border of U m a t i l l a Nat. For., on U.S.F.S. Rd. 40, between Pomeroy and Troy, G a r f i e l d Co., Wash. On rocky c l e a r i n g s with Lupinus, H e l i a n t h e l l a u n i f l o r a , g r a s s e s . 8507a, Robson, June 18, 1985. Balsamorhiza s e r r a t a N e l s . & Macbr. On Hwy. 395 S of Nye, U m a t i l l a Co., Ore. Mesic, rocky roadside with Lupinus, Phlox, A s t r a g a l u s . 8408, N i c h o l l s and Robson, June 7, 1984. 3. Outgroups H e l i a n t h e l l a c a l i f o r n i c a v a r . nevadensis (Greene) Jepson Hwy. 89 c a . 5 mi. NW of G r e e n v i l l e , near W shore of Lake Almanor, Plumas Co., C a l . In rocky, t h i n c l a y with grasses, Pinus ponderosa, Calocedrus decurrens, A r c t o s t a p h y l o s , Lupinus, Balsamorhiza d e l t o i d e a , Wyethia m o l l i s . 8418a, Robson, June 22, 1984. H e l i a n t h e l l a c a l i f o r n i c a v a r . nevadensis (Greene) Jepson 3 mi. E of Alleghany on Co. Rd. S309, 7.8 mi. E of j e t . with Co. Rd. S180, S i e r r a Co., C a l . In sandy, dry, g r a n i t i c s o i l with grasses, Pinus ponderosa, Abies c o n c o l o r , A r c t o s t a p h y l o s , Mimulus• 8420, Robson, June 23, 1984. H e l i a n t h e l l a u n i f l o r a v a r . d o u g l a s i i (T. & G.) Weber 23 mi. S of N bo'rder of U m a t i l l a Nat. For., on U.S.F.S. Rd. 40 between Pomeroy and Troy, G a r f i e l d Co., Wash. In rocky c l e a r i n g s with g r a s s e s , Lupinus, Balsamorhiza s a g i t t a t a . 8507b, Robson, June 18, 1985. 1 04 H e l i a n t h e l l a uni f l o r a v a r . d o u g l a s i i (T. & G.) Weber T r a i l t o J a r d i n e J u n i p e r , c a . 12 mi. E of Logan, Cache Nat. For., Cache Co., Utah. In open montane meadows of dolomite s o i l with A r t e m i s i a , Wyethia a m p l e x i c a u l i s , Balsamorhiza  m a c r o p h y l l a . 8511c, Barkworth and Robson, June 23, 1985. He l i a n t h u s c u s i c k i i A. Gray Mi. 13 on Hwy. 240 through Hanford Nuke Reserve, Benton Co., Wash. Weedy roadside with g r a s s e s , S a l s o l a , B r a s s i c a . 8404, N i c h o l l s and Robson, June 7, 1984. Rudbeckia o c c i d e n t a l i s Nutt. At E. J . Harney Viewpoint on Hwy. 204, ca. 2 mi. W of T o l l g a t e , U m a t i l l a Co., Ore. In rocky, d i s t u r b e d area with g r a s s e s , P o t e n t i l l a , Penstemon. 8517, Robson, June 27, 1985. 105 Table I. L i s t of s p e c i e s and numbers of i n d i v i d u a l s from which measurements were taken. SPECIES CASES Wyethia amp!ex1can 1 1 s 38 Wyethia angust i f o l l a 35 Wyethia a n z o n l c a 10 Wyeth1 a he 1 1 a n t h o i d e s 2 1 Wyeth i a 1onq i c a u 1 i s 5 Wyeth i a mo 1 1 i s 20 Wyethia e l a t a 10 Wyeth1 a r e t 1 c u 1 a t a 10 Wyeth i a bo 1 a n g e r i 15 Wyeth1 a 1nvenus ta 5 Wyethia o v a t a 5 Wyet h i a s c a b r a 5 WyethIa he 1 e n i o i d e s 10 Wyeth i a g l a b r a 10 B a 1 s a m o r h l z a macrophy\1 a 19 B a 1 s a m o r h i z a s e r r a t a 5 Ba1samorh i za 1ncana 20 Ba1samorh i za hooker 1 1 1 B a 1 s a m o r h i z a de1 to 1dea 5 Ba1samorh i za c a r e y a n a 5 B a1 s a m o r h1 z a sag 11 t a t a 19 He 1 i a nthe 1 l a c a 1 1 f o r n t c a 10 H e l i a n t h e l l a u n l f 1 o r a 23 Rudbeck1 a o c c 1 denta 11s 7 He1 i a n t h u s c u s i c k 1 1 6 106 T a b l e 11. C h a r a c t e r s u s e d w i t h t a x o n o m i c o u t g r o u p s and i n g r o u p s p e c i e s . 05 CO • H ra cu i - l • H T3 CO CO 3 i—l • H •H re cu TO o ra o fH • H ra •a re ra CJ U-l o 3 TO M 4-1 • H 4-1 01 ra • H • H • H AJ TO I - I cu CO o re -o c X 4J C e CJ cn 3 - o 3 ra • H ra 4-1 • H ra ra 01 CO o « • H • H TO U c c TO u C 4-1 o ^, c r-l N • H oo f-l 4-1 • H ra cu 4-1 0) • H 4J cu TO cx 60 • H r-l c r-l TO 4-1 T-t > TO ra r H TO oo >—1 CJ E c U 0) o o r H 0) o c > CJ 01 i-i TO 0) ra C TO ra ra r-l e 0) l-l . o O CO X- oo CO -a u • H ra • H ra CO CO • H • H c CO ra cu U & s- cu o — JO oi o T3 r—1 CJ T—i 3 CU c Li- «: # heads/shoot 0 - 1 - 2 1 - many 2 1 2 0 1 1 1 1 0 1 2 0 1 1 1 1 2 0 0 0 0 1 2 1 1 1 leaf margin 0 - 1 - 2 entire-divided 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 2 2 2 1 0 0 1 1 0 cauline leaves 0 - 1 - 2 a l t . - opp. 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 2 2 2 1 1 0 0 2 0 basal leaves 0 - 1 abs. pres. 2 2 2 2 2 2 0 0 1 2 2 0 2 2 2 2 2 2 2 2 2 1 0 0 0 0 lea f shape 0 - 1 - 2 l i n . - ovate 1 1 1 1 1 2 2 2 2 2 2 0 1 1 0 0 0 1 0 1 1 1 1 2 2 2 p e t i o l e s 0 - 1 - 2 long - abs. 2 1 0 0 1 0 0 0 0 0 0 2 0 0 1 0 0 0 2 0 0 1 1 1 1 1 caul, venation 0 - 1 3-nerv. - r e t i c . 1 1 1 1 1 1 0 0 1 1 1 0 1 1 0 0 0 1 1 1 1 0 1 0 1 1 r a y f l r t s . 0 - 1 neut. - p i s t . 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 1 0 achenes 0 - 1 quad. - f l a t . 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 a* H cn ro • I 3 i—' cn CO O Mi <r M • i <: 01 I 3 t— O r) 3 o 3 O T3 cn 10 O K n I 3* ro M i - 3 I •o 13 N J C C f CO n o * a 3" (-< CD " "-( H * ro cn oi o o 3 a c M l I r-> ft) 3 co 3 M O D p. co 3 I C 3* N J CO •3 ro (t O H 3 t-t, I 3 Co >— >-f 00 o O J o ca al o ca ab 1 1 3" 3 c r t i c cn m 1 1 • M • 1—' co < H-ro • 1 3 1 1 to ve o ro •d 1 N J cn 3 -d N J Co f | 3* f t D> •a H ro 0 ro r t • f ( cn ro o r t H - • • < r t M . O * O 3 to ox) Co X) I T3 c cn X) f t ft) cn 0 cr Co cn 1 Co < cn o o o »—-» o o o o t—• o o o o Artorhiza I—» o t — * o k—' *—» o N J 1—' o t—• o w . amplexicaulis f - o M o i—' t—* o t—» 1—» o »—' o w . angustifolia o k—t »—» O o t—* 1—' o N J I—1 o t—« o w . arizonica o o N J o t—1 I—' o o 1—< o t—» o w . helianthoides t—» i—' o o o k—' t—' o 1 o 1—' o w . longicaulis »—' >—' 1—I N J — 1—• N J o r - l •—1 o r — ' o w . mollis o o t—• o o t—• L O o — o o »—' w . elata o o t—• o o \—* U ) o »—1 o o 1—' 1—• w . reticulata o o r — * o t—» t—» N J o o o o 3 W. bolanderi t—• o >—• t—» N J o 1—• I - i o o o w . invenus ta o o t—* o 1—" 1—' N J o N J 1—' o »—' o w . ovata •—« o o o 1—• o o o o 1—• W. scabra I—* >—» 1—• t—• t—< 1—' N J o o o »—' o w . helenioides — I—< 1—* I—» »—< *—• N J o o *—< o o W. glabra o o r — • o o »—» t—• o *—' N J o o B. incana o o o o o o o o o I—1 *—' o o B. hookeri o o — o o t—« I—' o >—• N J o o B. macrophylla o o o o >—' >—' o N J o o B. serrata 0) cn cr rti i—> c ro n •—1 rt t—i i—i o • Q) o Q) O i-l c rt- o lO rt-»-l (D O n c in n • c cn ro a rt-3" CD-CD h-' cn 0) 3 O ft N 0) tTl ro o rt-O > ft rt O >-t N CU o 108 T a b l e IV. C h a r a c t e r s u s e d w i t h t h e s p e c i e s p a i r W y e t h i a  r e t i c u l a t a / W . e l a t a as f u n c t i o n a l o u t g r o u p . basal lvs. 0 - 1 abs. - pres. pappus 0 - 1 pres. - abs. cauline arr. 0 - 1 - 2 alt. - opp. caul, venation 0 - 1 3-nrv. - retic. // heads/shoot 0 - 1 - 2 1 - few - sev. I f . marg. 0 - 1 ent. - div. caul.shape 0 - 1 - 2 - 3 delt.-ov.-lan.-lin. caul. lvs. 0 - 1 lg/many - few/sm. phyllaries 0 - 1 many - few achene pubesc. 0 - 1 - 2 glab. - pubesc. petioles 0 - 1 long - none flavanones 0 - 1 abs. - pres. r H 10 CO 01 li CO • H de to CO m ftJ r H • H cu r H 4-1 CO O to o r H • H CO T3 r H CO to CO u U-t o jr. 3 t-i u • H 4-1 cu to r H • H • H • H 4-1 to CU cn O • H TO to -a c 3 X J-J c C o cn -o 3 to • H to TO U a 4-1 4-1 • H to U cu CO 0 ro • H - H c C to u C r-l c 01 o TO 4-1 O >, • H —{ 3 N • H DO r H to CU 4-1 0) JO to u • r l 4-i cu 4-> c 60 • H r H c r H rH > to to r H TO y o o u 60 rH r-l 01 E c CU o o o c > o 01 r H c o TO <u tO 0) to to to to •C r H E • H o CO J= co • H jr, E cn CO •o a prenyls 0 - 1 abs. - pres. 3l3|3|^|S|3|3|3|S|3|3|3|S |pa|m|pa|m|cQ|-o|m 0 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 1 1 0 0 0 0 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 1 0 0 2 1 2 2 0 0 0 0 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 0 0 0 1 2 1 2 0 1 1 0 1 2 0 0 0 0 0 0 0 1 1 2 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1 0 0 0 0 2 2 2 2 2 1 1 1 1 3 2 2 2 3 2 2 3 3 3 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 0 0 0 0 0 0 1 1 1 1 0 0 1 0 0 0 0 1 1 1 01 1 0 2 0 2 0 0 0 0 1 1 0 0 0 0 0 0 2 0 1 1 0 0 1 0 0 0 0 1 0 0 0 1 0 0 1 1 1 0 1 1 1 0 1 1 0 1 0 1 1 1 0 0 0 0 0 0 0 0 1 1 0 0 1 1 0 1 0 1 1 1 0 0 0 0 0 0 0 1 09 T c i b l e V. L i s t of f l a v o n o i d s from s p e c i e s so f a r surveyed Wyethia a n g u s t i f o l i a - i s o f l a v o n e s , flavanones, prenylated flavanones, prenylated i s o f l a v o n e s , prenylated d i h y d r o f l a v o n o l , dihydrooxepin r i n g Wyethia scabra - i s o f l a v o n e s , flavanones, prenylated flavanones, methylated f l a v o n o l s Wyethia r e t i c u l a t a - i s o f l a v o n e s , methylated f l a v o n o l s Wyethia e l a t a - d i h y d r o f l a v o n o l s , methylated f l a v o n e s , methylated f l a v o n o l s Wyethia b o l a n d e r i - methylated f l a v o n o l s Wyethia ovata - i s o f l a v o n e s , methylated f l a v o n o l s Wyethia invenusta - i s o f l a v o n e s , flavanones, prenylated flavanones, methylated f l a v o n o l s Wyethia gla b r a - i s o f l a v o n e s , flavanones, prenylated flavanones, chalcone Wyethia h e l e n i o i d e s - i s o f l a v o n e s , flavanones, prenylated flavanones, methylated f l a v o n o l s Balsamorhiza s a g i t t a t a - methylated f l a v o n o l s Balsamorhiza d e l t o i d e a - methylated f l a v o n o l s Balsamorhiza careyana - methylated f l a v o n o l s Balsamorhiza macrophylla - methylated f l a v o n o l s 1 1 0 Table VI. R e s u l t s of PCA f o r three s e t s of morph o l o g i c a l v a r i a b l e s . Ax i s 1 HEAD DATA 2 Eigenvalue 5.451 1 .905 HDWD -.30188 .34683 RAY .01780 .60984 PHLN1 -.38009 .15667 PHLN2 -.37914 . 14409 PHLN3 -.37859 .15806 RECPHLN3 .38795 .14598 RECPHWD2 .38836 .15479 RECPHWD3 .38830 .14 129 PH# .15776 SHOOT DATA .61014 Eigenvalue 1.5974 1 .0689 OPP -.23763 .84803 ALT .58857 -.25366 STHT .54393 LEAF DATA .39200 E i genvalue 3.2833 1.6229 MDLN1 .42969 .34533 MDLN2 .45898 .25998 MDWD1 .47538 .09426 MDWD2 .48051 .04774 MDPT1 .31098 -.58706 MDPT2 .22608 -.67628 111 Table V I I . Sources of v a r i a t i o n by s t r a t a . A=GENUS BISECTION C=SPECIES D=POPULATION E=RESIDUAL HEAD VARIABLES PCA 1 = 60.57% PCA 2 = 21.17% A = 7.739 A = 1.892 B = 13.685 B = 2.354 C = 25.142 C = 6.761 D = 5.710 D = 4.712 E = 8.295 E = 5.514 SHOOT VARIABLES PCA 1 = 39.94% A = 16.094 B = 3.966 C = 4.355 D = 5.853 E = 9.672 LEAF PCA VARIABLES 2 = 26.72% A = 8.466 B = 1.284 C = 5.729 D = 2.813 E = 8.428 PCA 1 = 54.72% A = 22.050 B = 5.433 C = 5.967 D = 8.018 £ = 13.251 PCA 2 = 27.05% A = 8.571 B = 1 . 300 C = 5.799 D = 2.848 E = 8.531 1 1 2 F i g u r e 1. Measurements u s e d i n e x p l o r a t o r y d a t a a n a l y s i 329 i n d i v i d u a l s 1 1 3 F i g u r e 2 . C l a s s e s o f f l a v o n o i d s a n d m a j o r s u b s t i t u t i o n p a t t e f l a v o n o l methylated f l a v o n o l flavanone pre n y l a t e d flavanone i s o f l a v o n e 1 1 4 q u r e 3 . C l a d o g r a m u s i n g H e l i a n t h e l l a as t a x o n o m i c o u t g r o u p . cn nH nH rt rt r H o AJ re cu U-l rt d -o rt nH AJ re •H o AJ AJ o nH in nH CU AJ OO 3 OO M r H C M re re CU o C in CI •a r H CO # » . pa I « i m L > 1 1 5 CO 0) T J rt rt nH o AJ « O nH cn C 3 u c O rt C x> cu N AJ cu rt r H nH rt > r H CU M > c x rt o nH . • i > 1 * i S cn cu •o rt nH nH AJ o VJ rt x cu r H AJ •a 3 c rt c O rt rt u rt nH AJ nH X I r H AJ « r H rt o co r H CU o X ) t-i CU X (0 1 s ra r H r H nH rt X rt t-i A J O. CU rt o C u M rt o u U o o cu rt c x: cn 6 nH . . pa pa 1 pa pa Tree Length - 310 Consistency Index - 59% 1 15 F i g u r e 4 . C l a d o g r a m u s i n g E n c e l i a as t a x o n o m i c o u t g r o u p . 1 1 6 F i g u r e 5 . C l a d o g r a m u s i n g H e l i o p s i s a s t a x o n o m i c o u t g r o u p . 1 17 F i g u r e 6. Cladogram using F l o u r e n s i a as taxonomic outgroup. F i g u r e 7 . C l a d o g r a m u s i n g R u d b e c k i a as t a x o n o m i c o u t g r o u p . 1 1 9 F i g u r e 8. Cladogram using Balsamorhiza s e c t i o n A r t o r h i z a as f u n c t i o n a l outgroup. 1 2 0 F i g u r e 9 . Cladogram using s p e c i e s p a i r Wyethia ret iculata/W. e l a t a as f u n c t i o n a l outgroup. 121 F i g u r e 10. Dry f l i e s of s p e c i e s . eo; AO . -80 W. a m p l e x i c a u l i s 80 Uo W. a n g u s t i f o l i a W. a r i z o n i c a . -BOl.6 W. l o n e i c a u l i s 80U0 • -801.6 W. h e l i a n t h o i d e s W. h e l e n i o i d e s 1 22 F i g u r e 11. Dry f l i e s of s p e c i e s . W. r e t i c u l a t a 80140 . -801.6 W. i n v e n u s t a ' 80 AO y / s . -80 .6 W. b o l a n d e r i H e l i a n t h e l l a c a l i f o r n i c a H e l i a n t h e l l a u n i f l o r a 123 F i g u r e 12. Dry f l i e s of s p e c i e s . B. macrophylla B.. incana B.. serrata B.. hookeri F i g u r e 13. Cladograms with dry f l i e s f o r Balsamorhiza s e c t i o n A r t o r h i z a and two s p e c i e s of H e l i ? n t h e l l a . 125 F i g u r e 14. Cladograms with dry f l i e s f o r three s p e c i e s of Balsamorhiza s e c t i o n Balsamorhiza and the Wyethia r e t i c u l a t a / W . e l a t a s p e c i e s p a i r . 

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