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Generic limits and systematics of boykinia and allies (Saxiferagaceae) Gornall, R. J. 1980

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GENERIC LIMITS AND SYSTEMATICS OF BOYKINIA AND ALLIES (SAXIFRAGACEAE) by RICHARD JOHN GORNALL B . S c , The U n i v e r s i t y o f S t . Andrews, M.Sc., The U n i v e r s i t y o f Birmingham,  1975 1976  A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in THE FACULTY OF GRADUATE STUDIES (Department o f Botany)  We a c c e p t t h i s t h e s i s as c o n f o r m i n g to t h e r e q u i r e d  standard  THE UNIVERSITY OF BRITISH COLUMBIA June ©  1980  R i c h a r d John G o r n a l l ,  1980  In p r e s e n t i n g t h i s  thesis  in p a r t i a l  f u l f i l m e n t o f t h e requirements f o r  an advanced degree at the U n i v e r s i t y of B r i t i s h the I  Library shall  f u r t h e r agree  for  freely available  that permission  for  I agree  that  r e f e r e n c e and s t u d y .  f o r e x t e n s i v e copying o f  this  thesis  s c h o l a r l y purposes may be granted by the Head of my Department o r  by h i s of  make it  Columbia,  this  representatives. thesis  It  i s understood that copying o r p u b l i c a t i o n  f o r f i n a n c i a l gain s h a l l  written permission.  Department of  BOTANY  The U n i v e r s i t y o f B r i t i s h  2075 Wesbrook P l a c e V a n c o u v e r , Canada V6T 1W5  Date  2 1 s t J u l y 1980  ' Columbia  not be allowed without my  ii  ABSTRACT The  g o a l s o f t h e study were t o d e f i n e t h e l i m i t s o f t h e genus  B o y k i n i a and t o c l a r i f y i t s i n f r a g e n e r i c taxonomy.  As c i r c u m s c r i b e d by  Engler, Boykinia contained nine species i n three s e c t i o n s .  Three  segregate  genera have been d e s c r i b e d however, and a t i t s most fragmented, E n g l e r ' s B o y k i n i a would comprise: B o y k i n i a , N e o b o y k i n i a , The  P e l t o b o y k i n i a and T e l e s o n i x .  s i t u a t i o n i s c o m p l i c a t e d by o t h e r c l o s e l y r e l a t e d genera, whose l i m i t s  a l s o need c l a r i f i c a t i o n .  N a r r o w l y d e f i n e d , they i n c l u d e S u k s d o r f i a ,  Hemieva, H i e r o n y m u s i a , B o l a n d r a  and S u l l i v a n t i a .  Methods o f approach i n c l u d e d an e x a m i n a t i o n herbarium m a t e r i a l to allow morphological  comparisons o f a l l t a x a . L i v e  p l a n t s o f almost a l l s p e c i e s were m a i n t a i n e d anatomical  o f a wide range o f  i n c u l t i v a t i o n and some  o b s e r v a t i o n s were made, and an i n t r a - and i n t e r - g e n e r i c h y b r i d -  i z a t i o n programme was conducted, on t h e s e . each genus were i n v e s t i g a t e d .  The trichome  complements o f  P o l l e n g r a i n s and seeds were examined by  l i g h t and s c a n n i n g e l e c t r o n m i c r o s c o p y .  Chromosome numbers were  o r e s t a b l i s h e d f o r t h e f i r s t time f o r a l l b u t t h r e e s p e c i e s .  confirmed  A complete  f l a v o n o i d a n a l y s i s o f r e p r e s e n t a t i v e s o f a l l genera, i n c l u d i n g a l l s p e c i e s o f B o y k i n i a s . l . , was conducted.  G e n e r a l e c o l o g i c a l o b s e r v a t i o n s were  made d u r i n g f i e l d work i n A l a s k a and t h e Western C o r d i l l e r a o f Canada and the U.S.A. The  r e s u l t s o f these i n v e s t i g a t i o n s c o n f i r m e d  form a c l o s e l y r e l a t e d group.  t h a t t h e genera  However, much o f t h e e v i d e n c e combined t o  suggest t h e f o l l o w i n g c o n c l u s i o n s : 1) B o y k i n i a comprises n i n e s p e c i e s i n t h r e e s e c t i o n s ( s e c t i o n B o y k i n i a i n c l u d e s B. a c o n i t i f o l i a , B. i n t e r m e d i a , B. l y c o c t o r i i f o l i a , B. o c c i d e n t a l i s , B. r o t u n d i f o l i a and B. liia j o r ; s e c t i o n R e n i f o l i u m udes B ^ r i c l i a ^ d ^ o j r i i _ ; and a new s e c t i o n , T e l e s o n i x , i n c l u d e s  incl-  iii  B. j a m e s i i and B_. h e u c h e r i f ormis I 2) P e l t o b o y k i n i a t e l l i m o i d e s i s r e c o g n i s e d as a d i s t i n c t monotypic genus, w i t h as much a f f i n i t y w i t h S a x i f r a g a as w i t h B o y k i n i a . I t has  two s u b s p e c i e s : s s p . t e l l i m o i d e s and s s p .  watanabei.  3) S u k s d o r f i a i s expanded to i n c l u d e Hemieva and Hieronymusia,  making  a t o t a l o f t h r e e s p e c i e s : S_. r a n u n c u l i f o l i a , S. v i o l a c e a and S. alchemilloides.  The range o f v a r i a t i o n i n S u k s d o r f i a i s s i m i l a r t o  t h a t i n B o y k i n i a and the two genera a r e c o n s i d e r e d t o have e v o l v e d in  parallel.  4) B o l a n d r a i s p r o v i s i o n a l l y r e t a i n e d as a s e p a r a t e genus.  I t i s very  c l o s e t o Suksdorf i a , and f u r t h e r work on B_. c a l i f o r n i c a , which i s c l o s e s t i n terms o f gross morphology, may l e a d t o a merging o f these genera. 5) S u l l i v a n t i a i s a d i s t i n c t  genus.  G e o g r a p h i c a l d i s t r i b u t i o n s , t o g e t h e r w i t h the taxonomic d a t a , allowed some s p e c u l a t i o n on the 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 o f the genera. Boykinia i s probably a T e r t i a r y r e l i c t , between Japan, western  s i n c e i t has a d i s j u n c t  N o r t h America and e a s t e r n North America.  share a common a n c e s t o r  distribution I t may  (a S a x i f r a g a - l i k e p l a n t ) w i t h P e l t o b o y k i n i a , which i s  endemic t o Japan. The-, remaining;Hgenera*have<'probably e v o l v e d from B o y k i n i a s t o c k by a d a p t i v e r a d i a t i o n i n t o d i f f e r e n t h a b i t a t s . h a b i t a n t o f wet c l i f f  faces, probably diverged f i r s t .  S u l l i v a n t i a , an i n S u k s d o r f i a and  B o l a n d r a . c h a r a c t e r i s t i c o f open and shaded, s p r i n g - w e t , r e s p e c t i v e l y , p r o b a b l y e v o l v e d l a t e r , i n response c l i m a t e i n the l a t e P l i o c e n e .  summer-dry h a b i t a t s  t o the onset o f such a  iv  T a b l e o f Contents  Page  Abstract  •  i i  L i s t of Tables L i s t of Figures  ix and I l l u s t r a t i o n s  Acknowledgements  .  I.  INTRODUCTION  II.  RHIZOMES  .  .  .  .  .  .  x  x  i  i  v  . . . . . . . . . . .  1 5  Root p a r a s i t i s m III.  10  LEAVES AND STIPULES  .  .  .  .  .  .  .  .  .  .  .  .  .  11  Basal leaves  11  Shape and o u t l i n e  11  Texture  21  Venation  21  Stomata Cauline IV.  V.  leaves  . and s t i p u l e s  26  TRICHOMES  •.  35  Developmental r e l a t i o n s h i p s  36  Observations  43  Taxonomic i n f e r e n c e s  46  Environmental m o d i f i c a t i o n of v e s t i t u r e i n Boykinia occidentalis  49  Function  51  of the glandular  INFLORESCENCE STRUCTURE  trichomes '  Environmental m o d i f i c a t i o n of i n f l o r e s c e n c e s t r u c t u r e VI. .  25  52 .  59  THE FLOWER  63  Calyx  63  Corolla  63  P e t a l movements  71  V  T a b l e o f Contents c o n t ' d . VI.  Page  . -THE FLOWER c o n t ' d .  VII.  Androecium  71  Stamen movements  72  Gynoecium  72  F l o r a l anatomy  81  Embryology  82  Taxonomic i n f e r e n c e s  85  T e r a t o l o g i c a l phenomena  86  POLLEN  .  .  .  .  .  .  ••.  . • ;  •'•  •  •  •••  •  87  Microsporogenesis  87  Shape and s i z e  88  Apertures  89  Tectum  91  SEEDS  .  .  .  .  Observations  .  .  .  . .  .  .  .  .'.  •  .  .  96 .  .'•  . . .  . .  • .  . . . . .  .  .  .  X.  SEEDLINGS AND EARLY DEVELOPMENT  .98 .  Taxonomic i n f e r e n c e s IX.  87  M a t e r i a l s and methods  Taxonomic i n f e r e n c e s VIII.  .  98 109  •.  .  .112  M a t e r i a l s and methods  112  Observations  112  CYTOLOGY Methods  118 .  .  .  .  .  •  118  Results  119  I n f r a s p e c i f i c geographical v a r i a t i o n  126  Karyotypes  126  Discussion  126  vi Table of Contents cont'd. XI.  Page  BREEDING STUDY  130  M a t e r i a l s and methods  130  Experimental  131  pollinations  Seed g e r m i n a t i o n  .  I n t r a - s p e c i f i c cross p o l l i n a t i o n s  . .  132 .  .  132  Self pollinations  135  N a t u r e o f t h e s e l f - i n c o m p a t i b i l i t y system Inbreeding depression  .  .  .  .  .  . .  . .  .  136  Inter-specific pollinations  137  I n t e r - s p e c i f i c crosses i n Boykinia  141  H y b r i d i n v i a b i l i t y and morphology  .  .  I n t e r - s p e c i f i c crosses i n Suksdorfia  .  .  .  .  .  .  . .  . .  . .  143 .  Inter-generic crosses XII.  135  144 147  FLAVONOID CHEMISTRY  . . . . . .  Previous chemical i n v e s t i g a t i o n s  149  o f B o y k i n i a and a l l i e s  Methods  . 149 150  Plant material  150  Chromotography and s p e c t r o s c o p y  .  .  .  .  .  .  .  150  E x t r a c t i o n and i s o l a t i o n  152  I d e n t i f i c a t i o n of flavonoids  153  I d e n t i f i c a t i o n of acylated flavonoids  154  Results  156  Discussion  170  Disease  resistance  . . .  170  S t a b i l i t y of expression Taxonomic c o n c l u s i o n s  172 .  .  .  .  .  172  vii T a b l e o f Contents c o n t ' d . XII.  XIII.  Page  FLAVONOID CHEMISTRY c o n t ' d . Evolutionary interpretation  175  Parallelism  176  ECOLOGY  .  Habitats  .  .  . .  . .  . .  . . . .  .  .  . . .  . .  . .  Boykinia  .  .  .  .  .  .  .  .  . 178  .  .  178  .  .  Peltoboykinia  180  Suksdorfia  180  Bolandra  181  Sullivantia  181  Discussion  .  .  181  Seed D i s p e r s a l  183  Phenology and p o l l i n a t i o n  .  .  .  .  . 184  D i s e a s e and p r e d a t o r s XIV.  GENERIC LIMITS AND TAXONOMY  190 .  •.  .  .  .192  Taxonomic approach Synopsis o f the genera and s p e c i e s  192 .  .  .  Peltoboykinia  XV.  178  .  .  .  .  194  .  .  •  •  . 194  Boykinia  198  Suksdorfia  224  Bolandra  230  PHYTOGEOGRAPHY AND EVOLUTION  .  .  .  . 234  Methods o f approach  234  The c l a d i s t i c approach  235  Summary o f some important T e r t i a r y events i n the n o r t h e r n hemisphere .  235  E v o l u t i o n o f B o y k i n i a and a l l i e s  .' .  .  .  .  .  .  .  238  viii T a b l e o f Contents c o n t ' d . XV.  Page  PHYTOGEOGRAPHY AND EVOLUTION c o n t ' d . Boykinia  .  .  .  .  .  .  .  .  .  . 240  Peltoboykinia  248  Bolandra  248  . . . . . . .  Suksdorfia XVI.  249  ETHNOBOTANY Horticulture Medicine  . .  .  .  .  .  .  .  .  .  254 254  . . . . . .  255  References  256  Appendices  270  1. 2. 3.  D e t a i l s o f c o l l e c t i o n s and h e r b a r i u m l o c a t i o n s o f p l a n t m a t e r i a l c i t e d i n the study . . . . . . S o l v e n t systems and media used i n t h e t h i n - l a y e r chromatographic a n a l y s i s o f f l a v o n o i d compounds . UV S p e c t r a l a b s o r p t i o n maxima (nm) o f some o f . t h e f l a v o n o i d s encountered i n B o y k i n i a and a l l i e s . .  .  .  .  270 .  .  276 .277  ix L i s t of Tables Table I. II.  Page L i s t of taxa recognised body o f t h e study  and i n c l u d e d i n t h e main 2  Frequency o f v e i n l e t t e r m i n a t i o n s and a r e o l a e i n species of Boykinia, P e l t o b o y k i n i a , Suksdorfia, B o l a n d r a and S u l l i v a n t i a  25  S t o m a t a l s i z e s and f r e q u e n c y on upper s u r f a c e s o f leaves . . . . . . ,  27  IV.  L i s t o f m a t e r i a l used i n t h e t r i c h o m e , s t u d y  36  V.  D i s t r i b u t i o n o f t r i c h o m e s t r u c t u r a l types among Boykinia, P e l t o b o y k i n i a , Suksdorfia, Bolandra, S u l l i v a n t i a , P e l t i p h y l l u m and J e p s o n i a  44  Trichome s t r u c t u r e s i n i n f l o r e s c e n c e branches o f Saxifraga species  47  V a r i a t i o n i n t r i c h o m e d e n s i t y among e i g h t p o p u l a t i o n s o f B o y k i n i a o c c i d e n t a l i s grown i n a growth chamber .  50  P r o t a n d r y and n e c t a r i e s i n some s p e c i e s c u l t i v a t e d i n a growth room .  79  P a r t i a l summary o f f l o r a l anatomy i n S a x i f r a g i n a e species with a x i l e placentation  83  X.  M a t e r i a l used i n t h e p o l l e n study  88  XI.  S i z e s o f a i r - d r i e d p o l l e n g r a i n s mounted i n Hoyer's Medium  90  XII.  L i s t o f m a t e r i a l used i n t h e seed study  99  XIII.  Seed c o l o u r , shape and s i z e , and t u b e r c l e l e n g t h i n B o y k i n i a , P e l t o b o y k i n i a , Suksdorfia, Bolandra, J e p s o n i a , P e l t i p h y l l u m and S u l l i v a n t i a  100  Chromosome numbers p r e s e n t l y and p r e v i o u s l y found i n the genera B o y k i n i a , P e l t o b o y k i n i a , S u k s d o r f i a , B o l a n d r a and S u l l i v a n t i a  123  Tentative karyotypes f o r species o f Boykinia, P e l t o b o y k i n i a , B o l a n d r a and S u l l i v a n t i a . . . .  127  XVI.  L i s t of c u l t i v a t e d material  131  XVII.  Seed s e t and c o m p a t i b i l i t y from i n t r a - s p e c i f i c and c r o s s and s e l f p o l l i n a t i o n s  133  Seed s e t and g e r m i n a b i l i t y from i n t e r - s p e c i f i c and i n t e r - g e n e r i c cross p o l l i n a t i o n s  138  III.  VI. VII. VIII. IX.  XIV.  XV.  XVIII.  . . .  X  L i s t of Tables  cont'd.  Table XIX.  Page P o p u l a t i o n s sampled, and d r y w e i g h t s o f t i s s u e used, f o r f l a v o n o i d analyses  151  XX.  F l a v o n o i d b e h a v i o u r i n UV l i g h t  154  XXI.  The d i s t r i b u t i o n o f f l a v o n o i d s i n B o y k i n i a , Suksd o r f i a , P e l t o b o y k i n i a , B o l a n d r a and S u l l i v a n t i a .  157  XXII.  XXIII. XXIV. XXV. XXVI. XXVII. XXVIII. XXIX.  P a r t i a l summary o f T a b l e XXI, showing t h e t a x o nomic d i s t r i b u t i o n o f some i m p o r t a n t f l a v o n o i d s t r u c t u r a l features .  •  171  F l o w e r i n g times i n B o y k i n i a , P e l t o b o y k i n i a , S u k s d o r f i a , B o l a n d r a and S u l l i v a n t i a  185  Summary o f f l o w e r v i s i t o r s Saxifraginae  185  ( p o l l i n a t o r s ? ) of the  F l o w e r v i s i t o r s ( p o l l i n a t o r s ? ) o f some B o y k i n i a and S u k s d o r f i a s p e c i e s  188  Summary o f t h e d i s t r i b u t i o n o f some i m p o r t a n t taxonomic c h a r a c t e r s among t h e genera  195  Characters supposedly d i a g n o s t i c of taxa s e g r e g a t e d from B o y k i n i a o c c i d e n t a l i s T. & G.  .  M o r p h o l o g i c a l v a r i a t i o n between B o y k i n i a j a m e s i i and B_. h e u c h e r i f o r m i s F l o r a l differences separating Suksdorfia r a n u n c u l i f o l i a , S. a l c h e m i l l o i d e s and S_. v i o l a c e a  XXX.  Morphological  v a r i a t i o n i n Bolandra  XXXI.  P u t a t i v e l y e v o l u t i o n a r y i n t e r p r e t a t i o n o f some c h a r a c t e r s t a t e sequences, t o g e t h e r w i t h a hypothetical evolutionary tree  .  216 222 . 226 •. 233  236  xi L i s t o f F i g u r e s and  Illustrations  Figure  Page  1.  Rhizome morphology i n P e l t o b o y k i n i a and B o y k i n i a  2.  Rhizome morphology i n B o y k i n i a , B o l a n d r a  3.  B a s a l l e a f shapes and o u t l i n e s of B o y k i n i a o c c i d e n t a l i s  4.  B a s a l l e a f shapes and o u t l i n e s o f B o y k i n i a s p e c i e s  5.  B a s a l l e a f shapes and o u t l i n e s o f B o y k i n i a ma.jor  6.  B a s a l l e a f shapes and o u t l i n e s o f B o y k i n i a s p e c i e s  7.  B a s a l l e a f shapes a n d . o u t l i n e s o f B o y k i n i a s e c t i o n T e l e s o n i x , and S u k s d o r f i a  16  8-.  B a s a l l e a f shapes and o u t l i n e s o f B o l a n d r a  17  9.  B a s a l l e a f shapes and o u t l i n e s o f P e l t o b o y k i n i a tellimoides  18  L e a f v e n a t i o n v i s i b l e t o the naked eye i n s p e c i e s o f Boykinia  23  L e a f v e n a t i o n v i s i b l e t o t h e naked eye i n s p e c i e s o f P e l t o b o y k i n i a , S u k s d o r f i a , B o l a n d r a and S u l l i v a n t i a .  24  12.  S t i p u l e morphology i n B o y k i n i a s e c t i o n B o y k i n i a  29  13.  S t i p u l e morphology i n B o y k i n i a s e c t i o n s T e l e s o n i x and R e n i f o l i u m ; and i n P e l t o b o y k i n i a  31  14.  S t i p u l e morphology i n S u k s d o r f i a and B o l a n d r a  33  15.  Semi-diagrammatic i l l u s t r a t i o n s o f m u l t i c e l l u l a r g l a n d u l a r trichomes i n B o y k i n i a . . .  37  Semi-diagrammatic i l l u s t r a t i o n s o f m u l t i c e l l u l a r g l a n d u l a r trichomes i n B o y k i n i a  38  10. 11.  16.  17.  .  .  6  and S u k s d o r f i a  . .  12 .  . .  . . .  . . .  .  .  13 .14  .  . . . .  Semi-diagrammatic i l l u s t r a t i o n s o f m u l t i c e l l u l a r g l a n d u l a r trichomes i n P e l t o b o y k i n i a t e l l i m o i d e s  8  15  39  18.  Semi-diagrammatic i l l u s t r a t i o n s o f m u l t i c e l l u l a r g l a n d u l a r trichomes i n S u k s d o r f i a and B o l a n d r a . . .  40  19.  Semi-diagrammatic i l l u s t r a t i o n s o f m u l t i c e l l u l a r g l a n d u l a r trichomes i n S u l l i v a n t i a , J e p s o n i a and Peltiphyllum  41  Semi-diagrammatic i l l u s t r a t i o n s o f e g l a n d u l a r trichomes . . . . .  42  20.  xii L i s t o f F i g u r e s and I l l u s t r a t i o n s  cont'd.  Figure 21. 22.  Page I n f l o r e s c e n c e s t r u c t u r e i n some s p e c i e s o f B o y k i n i a section Boykinia  53  I n f l o r e s c e n c e s t r u c t u r e i n s p e c i e s o f B o y k i n i a and Peltoboykinia  55  23.  I n f l o r e s c e n c e s t r u c t u r e i n S u k s d o r f i a and B o l a n d r a  24.  I n f l o r e s c e n c e s o f droughted and watered c l o n e s o f Boykinia occidentalis  61  25.  P e t a l shapes i n B o y k i n i a s e c t i o n B o y k i n i a  65  26.  P e t a l shapes i n B o y k i n i a s e c t i o n s T e l e s o n i x and R e n i f o l i u m ; and i n P e l t o b o y k i n i a  67  27.  P e t a l shapes i n S u k s d o r f i a and B o l a n d r a  69  28.  L o n g i t u d i n a l s e c t i o n s o f f l o w e r s o f some s p e c i e s o f Boykinia  73  L o n g i t u d i n a l s e c t i o n s o f f l o w e r s o f some s p e c i e s o f B o y k i n i a and P e l t o b o y k i n i a  75  29. 30. 31. 32.  33.  34.  35.  36. 37.  .  L o n g i t u d i n a l s e c t i o n s o f f l o w e r s o f S u k s d o r f i a and Bolandra  .  •  Scanning e l e c t r o n m i c r o g r a p h s o f p o l l e n g r a i n s o f species of Boykinia section Boykinia . .  57  77 92  Scanning e l e c t r o n m i c r o g r a p h s o f p o l l e n g r a i n s o f s p e c i e s o f B o y k i n i a s e c t i o n s R e n i f o l i u m and T e l e s o n i x , and o f P e l t o b o y k i n i a , B o l a n d r a and S u k s d o r f i a . . .  . 9 4  Scanning e l e c t r o n m i c r o g r a p h s o f seeds o f s p e c i e s o f Boykinia section Boykinia  101  Scanning e l e c t r o n m i c r o g r a p h s o f seeds o f s p e c i e s o f B o y k i n i a s e c t i o n s R e n i f o l i u m and T e l e s o n i x , and o f S u k s d o r f i a and B o l a n d r a •  103  Scanning e l e c t r o n m i c r o g r a p h s o f seeds o f s p e c i e s o f P e l t o b o y k i n i a . S u l l i v a n t i a , Peltiphyllunu.and Jepsonia S i x week o l d s e e d l i n g s o f a) B o y k i n i a o c c i d e n t a l i s ; b) B_. i n t e r m e d i a ; c) B_. major and d) B_. r o t u n d i f o l i a S i x week o l d s e e d l i n g s o f a) B o y k i n i a h e u c h e r i f o r m i s ; b) B_. r i c h a r d s o n i i ; and c) P e l t o b o y k i n i a t e l l i m o i d e s ssp.watanabei . . . .  .  .  105  . 113  115  xiii  L i s t o f F i g u r e s and I l l u s t r a t i o n s  cont'd.  Figure  Page  38.  Chromosomes of some s p e c i e s o f B o y k i n i a and S u k s d o r f i a  39.  Somatic chromosomes o f some s p e c i e s of B o y k i n i a Peltoboykinia  40.  C r o s s i n g r e l a t i o n s h i p s between  120  .  and 121  s p e c i e s and genera,  expressed i n terms o f O v e r a l l C o m p a t i b i l i t y  142  41.  The B o y k i n i a r o t u n d i f o l i a x B_. maior h y b r i d  145  42.  B a s a l l e a f shapes o f h y b r i d s and p a r e n t s p e c i e s i n  43.  Boykinia Key to f l a v o n o i d s t r u c t u r e s and a b b r e v i a t i o n s r e f e r r e d to i n the t e x t and t a b l e s  169  V a r i a t i o n i n c h a r a c t e r s used to d i s t i n g u i s h B o y k i n i a t u r b i n a t a from B_. a c o n i t i f o l i a  211  45.  The d i s t r i b u t i o n o f some B o y k i n i a s p e c i e s  241  46.  The d i s t r i b u t i o n s of B o y k i n i a major, B_. i n t e r m e d i a Bolandra  44.  47.  48.  146  and 242  The d i s t r i b u t i o n s o f B o y k i n i a l y c o c t o n i f o l i a , P e l t o b o y k i n i a t e l l i m o i d e s and S u l l i v a n t i a The d i s t r i b u t i o n o f the genus S u k s d o r f i a  .  .  .  . • .  243 .  250  xiv  Acknowledgements I should  l i k e to thank my  v a r i o u s ways d u r i n g the course  of t h i s study: my  Bohm f o r h i s c o n t i n u a l i n t e r e s t and work; P r o f . W.B.  supervisor, Prof.  generous f i n a n c i a l support  S c h o f i e l d f o r advice  t o g e t h e r w i t h P r o f . F.R.  t h e s i s committee f o r a s s i s t i n g i n  on n o m e n c l a t u r a l  matters  Ganders, f o r r e a d i n g c r i t i c a l l y and  of  B.A. field  and,  carefully  an  e a r l i e r d r a f t of t h i s m a n u s c r i p t . I should r e p l i e d to my  a l s o l i k e to thank the f o l l o w i n g people who  l e t t e r s and p r o v i d e d me  i n f o r m a t i o n : Dr. R. U n i v e r s i t y of New  w i t h v a r i o u s b i t s and  U n i v e r s i t y of A l a s k a ; P r o f . J.G. S o l t i s , I n d i a n a U n i v e r s i t y ; and Harvard U n i v e r s i t y .  p i e c e s of  Haywood, A l p i n e Garden S o c i e t y ; P r o f . W.C.  Mexico; Dr. B. Mathew, Kew  Gardens; Dr.  kindly  D.F.  Martin, Murray,  P a c k e r , U n i v e r s i t y of A l b e r t a ; Mr. Dr.  S.A.  P l a n t m a t e r i a l was  D.E.  Spongberg, A r n o l d Arboretum of generously  provided  by Dr.  M.  Wakabayashi, Tokyo M e t r o p o l i t a n U n i v e r s i t y ; the D i r e c t o r of the J.W. U n i v e r s i t y B o t a n i c Garden, F r a n k f u r t ; and  the D i r e c t o r of the Nippon  Shinyaku I n s t i t u t e f o r B o t a n i c a l Research, Oyake.  Herbarium specimens  were k i n d l y loaned by the c u r a t o r s of the h e r b a r i a l i s t e d and  I thank P r o f . J . Maze f o r w r i t i n g the numerous l o a n I am  and Mr.  S.W.  g r a t e f u l to Dr. M. Weintraub  DeBeer f o r making a v a i l a b l e to me  microscope f a c i l i t y me  f l o w e r s , s e e d l i n g s and  i n Chapter  1,  requests.  ( D i r e c t o r ) , Ms. the scanning  at A g r i c u l t u r e Canada, Vancouver, and  i n i t s operation.  Goethe  B.  Schroeder  electron  for instructing  Thanks a l s o go to L e s l i e Bohm f o r her drawings of stipules.  F i n a l l y , f i n a n c i a l support  throughout t h i s study was  provided  by an Izaak Walton K i l l a m F e l l o w s h i p , which i s g r a t e f u l l y acknowledged.  1  I.  INTRODUCTION  The p r e s e n t work f o c u s e s on the l i m i t s and d i v e r s i t y of the genus B o y k i n i a N u t t .  As t r e a t e d by E n g l e r (1928), the genus c o n t a i n s  n i n e s p e c i e s i n N o r t h America and Japan and b e l o n g s Saxifraginae.  to the s u b - t r i b e  Both s e c t i o n a l and g e n e r i c b o u n d a r i e s are p r o b l e m a t i c  and much d e t a i l e d i n f o r m a t i o n i s needed to determine g e n e i t y of the genus (Spongberg 1972).  The problem  the l i m i t s and homoconcerning  stems from the e x i s t e n c e of two s m a l l s e g r e g a t e genera  limits  (Neob o y k i n i a and  P e 1 t o b o y k i n i a ) and seven o t h e r c l o s e l y r e l a t e d ones ( B o l a n d r a , Hemieva, Hieronymusia,  S a x i f r a g a , S u k s d o r f i a , S u l l i v a n t i a and T e l e s o n i x ) . The  taxa  r e c o g n i s e d and i n c l u d e d i n the main body of the s t u d y are shown i n T a b l e I . S u l l i v a n t i a i s b e i n g monographed by Mr. D. S o l t i s of I n d i a n a U n i v e r s i t y , and i n o r d e r t o a v o i d d u p l i c a t i o n o f e f f o r t , t h i s genus i s r e p r e s e n t e d m a i n l y by S_. oregana and o c c a s i o n a l l y by S_. hapemanii  i n the p r e s e n t s t u d y .  Most o f the s p e c i e s t r e a t e d here were o r i g i n a l l y a s s i g n e d t o the genus S a x i f r a g a L.  However, as more and more members of the S a x i f r a g -  i n a e were d i s c o v e r e d , a g r e a t e r a p p r e c i a t i o n of the p a t t e r n s of v a r i a b i l i t y r e q u i r e d t h a t the b r o a d l y c o n c e i v e d genus S a x i f r a g a be broken up i n t o more coherent groups.  There was no immediate agreement on the l i m i t s of the  r e s u l t a n t s e g r e g a t e genera and many of the s p e c i e s have been a s s i g n e d to d i f f e r e n t genera a t one time o r a n o t h e r . alignments i s given i n Table I .  A h i s t o r y of these d i v e r s e  R e g i o n a l f l o r a s show no  consensus  r e g a r d i n g taxonomic t r e a t m e n t . There are s e v e r a l s o u r c e s of the confused s i t u a t i o n from T a b l e I .  apparent  F i r s t , m o r p h o l o g i c a l p a r a l l e l i s m and the g e n e r a l p a u c i t y of  b i o l o g i c a l i n f o r m a t i o n i n e v i t a b l y l e d the many f l o r a w r i t e r s to i n c o m p l e t e and/or c o n t r a d i c t o r y views c o n c e r n i n g the r e l a t i o n s h i p s of the v a r i o u s t a x a . Second, a t about the t u r n of the c e n t u r y , N o r t h American taxonomy s u f f e r e d  2  TABLE I . L i s t o f t a x a r e c o g n i s e d study.  and i n c l u d e d i n t h e main body o f the  A b r i e f h i s t o r y of t h e i r various generic alignments i s g i v e n ; only  the combining a u t h o r s a r e r e f e r e n c e d . Taxon  A s s i g n e d Genus  BOYKINIA NUTT. (nom. conserv.) section Boykinia aconitifolia intermedia  Nutt.  B o y M n i a ( N u t t a l l 1834) ; S a x i f r a g a ( F i e l d i n g & Gardner 1844).  ( P i p e r ) G.N. Jones  lycoctonifolia  (Maxim.) E n g l .  B o y k i n i a ( P i p e r 1899). S a x i f r a g a (Maximowicz 1886); B o y k i n i a ( E n g l e r 1891); N e o b o y k i n i a (Hara 1937).  major A. Gray  B o y k i n i a (Gray 1876).  O c c i d e n t a l i s T o r r . & Gray  S a x i f r a g a ( N u t t a l l i n T o r r e y & Gray 1840); B o y k i n i a ( T o r r e y & Gray 1840).  r o t u n d i f o l i a P a r r y i n Gray  B o y k i n i a ( P a r r y i n Gray 1878).  section Renifolium  Rosendahl  r i ch ards o n i i (Hook.) R o t h r o c k  S a x i f r a g a (Hooker 1832); Hemieva ( R a f i n e s q u e 1838); B o y k i n i a ( R o t h r o c k 1868).  s e c t i o n T e l e s o n i x (Raf.) G o r n a l l j a m e s i i (Torr.) E n g l .  heucheriformis  (Rydb.)  S a x i f r a g a (Hooker 1832); T e l e s o n i x ( R a f i n e s q u e 1838) ; B o y k i n i a ( E n g l e r 1891) . Rosend.  B o y k i n i a (Rydberg 1897); T e l e s o n i x ( R a f i n e s q u e 1838); S a x i f r a g a (Jones 1910).  PELTOBOYKINIA (ENGL.) HARA t e l l i m o i d e s (Maxim.) E a r a ssp. t e l l i m o i d e s  s s p . w a t a n a b e i (Yatabe) G o r n a l l  S a x i f r a g a (Maximowicz 1871); Boyk i n i a ( E n g l e r 1919); P e l t o b o y k i n i a (Hara 1937). S a x i f r a g a (Yatabe 1892); B o y k i n i a ( E n g l e r 1919) ; P e l t o b o y k i n i a (Hara 1937).  SUKSDORFIA A. GRAY (nom. conserv.) ranunculifolia  (Hook.) E n g l .  S a x i f r a g a (Hooker 1832); Hemieva ( R a f i n e s q u e 1838); S u k s d o r f i a ( E n g l e r 1891); B o y k i n i a (Greene 1891).  3  TABLE I c o n t ' d . Taxon  Assigned  Genus  v i o l a c e a A. Gray  S u k s d o r f i a (Gray 1879); Hemieva (Wheelock 1896).  a l c h e m i l l o i d e s (Griseb.) Engl.  S a x i f r a g a ( G r i s e b a c h 1879); Suksd o r f i a ( E n g l e r 1891) ; H i e r o n y m u s i a ( E n g l e r 1918)  BOLANDRA A. GRAY  c a l i f o r n i c a A. Gray  Bolandra  (Gray 1867).  oregana Wats.  Bolandra  (Watson 1879a).  SULLIVANTIA TORR. & GRAY IN GRAY*  * c u r r e n t l y b e i n g monographed by Mr.  2 - 6 spp. a s s i g n e d o c c a s i o n a l l y to S a x i f r a g a ( T o r r e y & Gray 1840; Fedde 1906); Heuchera ( C o u l t e r & F i s h e r 1892); B o y k i n i a (Brandegee 1899); but m a i n l y t o S u l l i v a n t i a (Gray 1842; Watson 1879b; C o u l t e r 1892; S m a l l 1905; Rosendahl 1927). D. S o l t i s o f I n d i a n a U n i v e r s i t y .  a p e r i o d o f e x c e s s i v e " s p l i t t i n g " . Many i n f r a s p e c i f i c t a x a were e l e v a t e d to s p e c i f i c s t a t u s and the numbers o f genera were p r o p o r t i o n a l l y i n c r e a s e d . T h i s l e g a c y o f an i n f l a t e d number o f genera and s p e c i e s has o f t e n tended to o b s c u r e p a t t e r n s o f v a r i a b i l i t y and has h i n d e r e d T h i r d , a p a r t from E n g l e r (1891, 1928)  taxonomic s y n t h e s i s .  and Dandy (1927),  t h e r e has been no  attempt t o d e a l w i t h the v a r i o u s genera i n a w o r l d - w i d e p e r s p e c t i v e .  A  c o n s i s t e n t taxonomy w i t h i n the f a m i l y depends on a d o p t i n g b r o a d e r h o r i z o n s than those u s u a l l y shown by r e g i o n a l f l o r a w r i t e r s . I t i s a g a i n s t t h i s background t h a t the g a t h e r i n g o f s o r e l y needed i n f o r m a t i o n on the genus B o y k i n i a and i t s a l l i e s was c o n s i d e r e d highly desirable.  By u s i n g e v i d e n c e from anatomy, morphology,  cytology, breeding behaviour,  chemistry, ecology  palynology,  and phytogeography the  v a r i o u s genera a r e r e d e f i n e d and t h e i r 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 commented on.  The i n f r a g e n e r i c c l a s s i f i c a t i o n o f B o y k i n i a i s a l s o r e v i s e d . The  s t u d y i s based on l i v i n g m a t e r i a l and on p r e s s e d  specimens  4  from t h e f o l l o w i n g h e r b a r i a ( a b b r e v i a t i o n s as i n Holmgren and Keuken 1974):  ALA, ALTA, AUA, BM, CAN, CM, COLO, DAO, F, GA, GH, I D , K, MIN, MO,  MONTU, NCU, ND, NDG, NY.OSC, P, RM, RSA, SD, TENN, T I , UBC, UNA, UTC, US, UVIC, V, WILLU, WS, WTU.  5  II.  RHIZOMES  Among t h e t a x a s t u d i e d t h e r e i s some v a r i a t i o n i n t h e n a t u r e of t h e i r underground  s t o r a g e organs.  B o y k i n i a and P e l t o b o y k i n i a have  t h i c k , s c a l y rhizomes r a n g i n g from 5 - 20mm i n diameter ( F i g s . 1, 2 ) . B o y k i n i a i n t e r m e d i a i s o f p a r t i c u l a r i n t e r e s t because l a r g e p a r t s o f i t s rhizome f r e q u e n t l y can be found c r e e p i n g over t h e s u r f a c e o f t h e ground. These above-ground segments a r e s t r o n g l y pigmented w i t h a n t h o c y a n i n s , q u i t e f l e s h y and n o t a t a l l s c a l y .  Roots and l e a v e s emerge from each node  ( F i g . l d ) . I t seems r e a s o n a b l e t o i n t e r p r e t t h e s e organs as s t o l o n s , a l t h o u g h the i n t e r n o d e s a r e q u i t e s h o r t .  These organs a r e u s u a l l y m i s s i n g  from h e r b a r i u m specimens and the taxon has never been r e c o r d e d as b e i n g stoloniferous.  Some h e r b a r i u m specimens o f the Japanese B. l y c o c t o n i f o l i a  a l s o show s i g n s o f b e i n g s t o l o n i f e r o u s ( F i g . I f ) , and i n d e e d , Maximowicz (1886) o r i g i n a l l y d e s c r i b e d i t as s u c h , a l t h o u g h t h i s f e a t u r e seems to have been i g n o r e d s u b s e q u e n t l y . c r e e p i n g rhizome  I n d i c a t i o n s o f t e n d e n c i e s toward a s u r f a c e -  can a l s o be found i n B. major and B_. a c o n i t i f o l i a  (Figs.  I c , e) . Both S u k s d o r f i a and B o l a n d r a have b u l b i f e r o u s rhizomes ( F i g . 2).  Each b u l b i l i s a s i m p l e reduced shoot a x i s and can v a r y from w h i t e ,  p a l e y e l l o w , p i n k t o brown o r b l a c k . phragma ( T a y l o r 1965).  S i m i l a r b u l b i l s a r e found i n L i t h o -  E n g l e r (1891, 1918) r e p o r t e d t h a t S. a l c h e m i l l o i d e s  l a c k e d a b u l b i f e r o u s rhizome, and t h i s f e a t u r e may have i n f l u e n c e d him i n the c r e a t i o n o f the s e g r e g a t e genus H i e r o n y m u s i a species.  ( E n g l e r 1918) f o r t h i s  However, a l l h e r b a r i u m specimens examined i n t h i s s t u d y d e f i n i t e -  l y have b u l b i l s ( F i g . 2 f ) , b u t t h e s e a r e u s u a l l y obscured by s o i l  caked  around the r o o t s t o c k s . G r i s e b a c h (1879), i n h i s p r o t o l o g u e , d e s c r i b e d the rhizome o f S. a l c h e m i l l o i d e s as " t u b e r i f o r m e " . S u b t e r r a n e a n b u l b i l s a r e g e n e r a l l y r e c o g n i s e d t o be a d a p t a t i o n s  6  FIGURE 1 . Rhizome morphology i n P e l t o b o y k i n i a and B o y k i n i a . A) P_. t e l l i m o i d e s s s p . t e l l i m o i d e s ; B) Underground p o r t i o n of the rhizome o f B_. major; C) Above-ground p o r t i o n o f the rhizome o f B_. ma.jor; D) S t o l o n p r o d u c t i o n i n B_. i n t e r m e d i a ; E) Tendency toward s t o l o n p r o d u c t i o n i n B_. a c o n i t i f o l i a : F) Suggestion s t o l o n p r o d u c t i o n i n a herbarium specimen o f B_.  of  lycoctonifolia.  7  8  FIGURE 2. Rhizome morphology A)'- B o y k i n i a o c c i d e n t a l i s ;  i n B o y k i n i a , B o l a n d r a and  Suksdorfia.  B) B o y k i n i a r o t u n d i f o l i a ; C) B o y k i n i a  r i c h a r d s o n i i : D) B o y k i n i a h e u c h e r i f o r m i s ; of S u k s d o r f i a r a n u n c u l i f o l i a ;  E) B u l b i f e r o u s rhizome,  F) B u l b i f e r o u s rhizome o f S u k s d o r f i a  a l c h e m i l l o i d e s : G) B u l b i f e r o u s rhizome o f B o l a n d r a oregana.  SCALE  BAR  A - D  2 cm  E - G  0-5 c m  10  f o r r e p r o d u c t i o n d u r i n g adverse e n v i r o n m e n t a l from s e x u a l s t e r i l i t y ( D a r l i n g t o n 1939). correlated with xeric habitats. c a l l y i n spring-wet,  snow-melt, and then d i e back  I n the present  Both Bolandra  summer-dry h a b i t a t s .  c o n d i t i o n s o r as an escape case they a r e  and S u k s d o r f i a grow t y p i -  They f l o w e r d u r i n g t h e s p r i n g  completely.  L i k e B o y k i n i a , S u l l i v a n t i a a l s o produces r h i z o m e s , b u t they are much more s l e n d e r .  Only i n S. oregana does v e g e t a t i v e  o c c u r , b u t t h i s i s i n t h e form of s l e n d e r s t o l o n s w i t h l o n g  reproduction internodes,  which are q u i t e d i f f e r e n t i n appearance from those o f B_. i n t e r m e d i a . Root P a r a s i t i s m Orobanche u n i f l o r a L. p a r a s i t i z e s the r o o t s o f the N o r t h American species of Suksdorfia.  The a s s o c i a t i o n of 0_^ u r i i f l o r a w i t h t h e Sax-  i f ragaceae has been r e c o r d e d by Munz (1959) , F e r r i s (1960) and T a y l o r (1965).  11 III.  LEAVES AND  STIPULES  L e a f morphology more than any o t h e r s i n g l e f a c t o r c o n t r i b u t e s to the o v e r a l l appearance of these p l a n t s , and assignment of g e n e r i c l i m i t s and omy  o f B o y k i n i a ( E n g l e r 1891;  t h i s has i n f l u e n c e d  the  the arrangement of the i n f r a g e n e r i c t a x o n -  Rosendahl 1905).  morphology of b o t h l i v e and p r e s s e d  I n the p r e s e n t  study,  the  l e a v e s has been examined, i n c l u d i n g  the use of c l e a r e d specimens. The spiral.  Two  l e a v e s of a l l genera:are produced i n a 3/8  c a t e g o r i e s of l e a f may  be i d e n t i f i e d :  phyllotactic  b a s a l and  cauline.  B a s a l l e a v e s are produced i n a r o s e t t e a t the base of a p l a n t , and  cauline  l e a v e s , v a r i a b l e i n number, s i z e and shape, form an a c r o p e t a l s e r i e s showing progressive morphological  r e d u c t i o n from the b a s a l l e a v e s , becoming  minute b r a c t s i n the upper branches of the i n f l o r e s c e n c e .  Those on  stem i t s e l f are a s s o c i a t e d w i t h v a r i o u s k i n d s of s t i p u l e s . category  the  Each l e a f  i s discussed i n turn.  BASAL LEAVES B o y k i n i a t h e b a s a l l e a v e s are e v e r g r e e n w i t h two B. r i c h a r d s o n i i and B_^ r o t u n d i f o l i a .  These two  exceptions:  species, together  with  P e l t o b o y k i n i a , S u k s d o r f i a and B o l a n d r a are r h i z o c a r p i c .  In nature,  b a s a l l e a v e s o f S u k s d o r f i a and B o l a n d r a o f t e n d i e b e f o r e  anthesis.  Shape and  the  Outline L e a f shapes and o u t l i n e s are v a r i a b l e w i t h i n i n d i v i d u a l  a r e p r e s e n t a t i v e a r r a y f o r each i s shown i n F i g s . 3 - 9 .  species,  At l e a s t p a r t o f  the v a r i a t i o n w i t h i n many of the s p e c i e s r e s u l t s from h e t e r o b l a s t y .  The  phenomenon i s most n o t i c e a b l e i n B o y k i n i a s e c t i o n B o y k i n i a where the e a r l y l e a v e s i n the b a s a l r o s e t t e tend to be more r e n i f o r m than l a t e r  leaves,  and t h e i r t e e t h tend t o be rounded-ovate, r a t h e r than acute or l a n c e o l a t e , as i n the l a t e r l e a v e s .  T h i s s i t u a t i o n i s i l l u s t r a t e d f o r B. o c c i d e n t a l i s  12  FIGURE 3.  B a s a l l e a f shapes and o u t l i n e s of B o y k i n i a o c c i d e n t a l i s (xJg).  a - h) V a r i a t i o n between p o p u l a t i o n s . i  - 1) H e t e r o b l a s t y ,  ( i ) b e i n g the f i r s t  l e a f and  (1) b e i n g the  last.  13  FIGURE 4. B a s a l l e a f shapes and  o u t l i n e s of B o y k i n i a s p e c i e s  a - c) B. i n t e r m e d i a ; d - e) B^  lycoctonifolia;  folia.  (xlg) .  f - j)B_. aconiti-  15  FIGURE 6 . B a s a l l e a f shapes and  o u t l i n e s of Boykinia  a - c) B_. r o t u n d i f o l i a ; d - f ) B_.  richardsonii.  species  16  FIGURE 7. B a s a l l e a f shapes and o u t l i n e s o f B o y k i n i a s e c t i o n T e l e s o n i x and S u k s d o r f i a  (x3/4).  a - d) B_. j a m e s i i and B_. h e u c h e r i f o r m i s  (both s p e c i e s show s i m i l a r  v a r i a t i o n ) ; e - j ) S_. r a n u n c u l i f o l i a ; k - o) S_. v i o l a c e a ; p - r ) S. a l c h e m i l l o i d e s .  ^ww!^ h  i  ^f^3 j  k  18  F I G U R E 9. B a s a l l e a f shapes and o u t l i n e s  of  tellimoides  e - h) s s p .  a,e)  ( x % ) . a - d) s s p . w a t a n a b e i ;  mature f i r s t l e a f ; b , f )  t h i r d l e a f ; d,h)  mature t h i r d  Peltoboykinia  mature second l e a f ; c,g) leaf.  tellimoides. young  19 i n F i g . 3, b u t i t a l s o o c c u r s i n many o t h e r s p e c i e s , i n c l u d i n g B^. a c o n i t ifolia.  A s e g r e g a t e s p e c i e s has been d e s c r i b e d from t h e l a t t e r ( B . turfa-  i n a t a Rydberg) p a r t l y on t h e b a s i s o f i t s h a v i n g acute t e e t h .  rounded-ovate r a t h e r than  However, i n view o f the h e t e r o b l a s t y i n v o l v i n g t h i s  character  and  the f a c t t h a t t h e r e i s i n t e r g r a d a t i o n between t h e two extremes ( F i g .  4),  i t cannot be used t o s u p p o r t f o r m a l taxonomic s e p a r a t i o n . L e a f shapes i n B o y k i n i a s e c t i o n B o y k i n i a a r e r e l a t i v e l y homo-  genous, w i t h t h e e x c e p t i o n o f B. r o t u h d i f o l i a , a l t h o u g h B. o c c i d e n t a l i s i s somewhat t r a n s i t i o n a l between t h i s s p e c i e s and the r e s t o f t h e s e c t i o n . Boykinia Occidentalis i s also t r a n s i t i o n a l to section t e l e s o n i x i n i t s a b i l i t y t o produce doubly c r e n a t e  leaves.  Boykinia r i c h a r d s b n i i of section  R e n i f o l i u m c o u l d be l i n k e d t o s e c t i o n B o y k i n i a by way o f B_j_ r o t u n d i f o l i a as suggested by Spongberg (1972) i h i g h l y - t o o t h e d l e a v e s , although (Fig.  n  v i e w o f the s i m i l a r  shallowly-lobed,  p a r a l l e l i s m i s t h e more l i k e l y  explanation  6 ) . Hooker (1832) and R o s e n d a h l (1905) based t h e i r s u g g e s t i o n o f an  a l l i a n c e between B_^ r i c h a r d s o n l j and s e c t i o n T e l e s o n i x p a r t l y on l e a f shape a l t h o u g h , two  as w i l l be seen l a t e r , t h e r e i s good r e a s o n t o m a i n t a i n  s e c t i o n s f o r these t a x a . A s i m i l a r s i t u a t i o n t o t h a t i n B. a c o n i t i f o l i a e x i s t s i n  Bolandra.  H e r e , p a r t o f the h i s t o r i c a l d i s t i n c t i o n between B o r e g a n a and  B. c a l i f o r i i i c a has been made on t h e b a s i s o f l e a f shape and o u t l i n e . B o l a n d r a oregana was c o n s i d e r e d  t o be 9 - 13 l o b e d w i t h a c u t e t e e t h , and  B. c a l i f o r n i c a was d e s c r i b e d as b e i n g 5 - 7 mucronate t e e t h ( B a c i g a l u p i 1944).  l o b e d w i t h rounded-ovate,  Y e t , i f a range o f m a t e r i a l i s examined,  e s p e c i a l l y i n c l u d i n g p l a n t s from e a s t e r n Oregon and Idaho, w h i c h seem t o be e s p e c i a l l y v a r i a b l e , i t i s seen t h a t t h e r e i s an o v e r l a p between t h e C a l i f o r n i a n and the more n o r t h e r l y m a t e r i a l , t h e l a t t e r i n t e r g r a d i n g w i t h the former ( F i g . 8 ) . The c h a r a c t e r o f l e a f shape and o u t l i n e cannot be  20  used t o s u p p o r t a s p e c i f i c d i s t i n c t i o n between t h e two t a x a . In Pe11oboykinia  the i n i t i a l m e t a p h y l l s o f b o t h s u b s p e c i e s a r e  q u i t e p r o m i n e n t l y l o b e d and d e n t a t e d u r i n g the e a r l y p a r t o f t h e i r d e v e l opment, b u t as they age and e n l a r g e t h e l o b e s broaden l a t e r a l l y t o r e a c h the mature o r b i c u l a t e , v e r y s h a l l o w l y l o b e d c o n d i t i o n ( F i g . 9 ) . I n s s p . t e l l i m o i d e s t h e r e i s l i t t l e o r no h e t e r o b l a s t y , a l l b a s a l l e a v e s as j u s t d e s c r i b e d .  However, i n s s p . w a t a n a b e i ,  behaving  although the e a r l i e s t  b a s a l l e a v e s a r e o r b i c u l a t e and s h a l l o w l y l o b e d when mature, the l a t e r b a s a l l e a v e s do n o t undergo t h e l a t e r a l growth i n t h e l o b e s t o such an e x t e n t as seen i n t h e e a r l y l e a v e s . prominently 5 - 9 than w i d e .  R a t h e r , the l a t e r l e a v e s adopt a  l o b e d c o n d i t i o n , each l o b e e l o n g a t i n g and becoming l o n g e r  T h i s h e t e r o b l a s t y i n s s p . w a t a n a b e i i s n o t apparent  b a r i u m specimens, s i n c e b a s a l l e a v e s a r e e a r l y deciduous one o r two o f t h e youngest a r e p r e s e n t a t f l o w e r i n g .  from her-  and u s u a l l y o n l y  The f a c t t h a t l e a f  morphology p r e s e n t s the o n l y d i s c o n t i n u i t y t h a t e x i s t s between t h e two t a x a , and t h a t the e a r l y l e a v e s i n b o t h a r e so s i m i l a r , s u p p o r t s  their  c o n s p e c i f i c s t a t u s (Makino 1901) . ^  n  S u k s d o r f i a , S. v i o l a c e a and  i n l e a f shape and o u t l i n e .  a l c h e m i l l o i d e s are s i m i l a r  Suksdorfia ranunculifolia i s d i s t i n c t i v e i n  the genus i n h a v i n g t e r n a t e l y p a r t e d b a s a l l e a v e s , a l t h o u g h t h i s c o n d i t i o n can be approached i n some S_» v i o l a c e a p o p u l a t i o n s ( F i g . 7 ) . I n S u l l i v a r i t i a t h e r e i s the same k i n d o f v a r i a t i o n t h a t i s found i n B o y k i n i a and B o l a n d r a , and i t i s l i k e l y t h a t some o f the s i x s p e c i e s r e c o g n i s e d by Rosendahl (1927) c o u l d be reduced The  t o synonymy.  s e r i e s o f l e a f shapes, from c r e n a t e t o d e e p l y - p a r t e d , can  be found i n a l l t h e genera under study and i t i s l i k e l y t h a t t h i s phenomenon has i t s b a s i s i n homologous v a r i a t i o n .  The a d a p t i v e s i g n i f i c a n c e o f  p a r t e d l e a v e s has been l i n k e d e x p e r i m e n t a l l y t o more e f f e c t i v e  temperature  21  c o n t r o l under c o n d i t i o n s of drought and h i g h l i g h t i n t e n s i t y (Lewis  1969;  W i l k i n s and L e w i s 1969) . Texture Leaves o f P e l t o b o y k i n i a and B o y k i n i a s e c t i o n s B o y k i n i a R e n i f o l i u m are c a n a l i c u l a t e on t h e i r upper s u r f a c e s and lower surfaces.  and  c o s t a t e on  their  I n s e c t i o n T e l e s o n i x and S u k s d o r f i a the upper l e a f  face i s canaliculate-smooth  and  the l o w e r s u r f a c e i s somewhat r i b b e d .  B o l a n d r a i s t r a n s i t i o n a l between these two shown by B o y k i n i a o c c i d e n t a l i s .  sur-  conditions, a condition also  S u l l i v a n t i a l e a v e s are smooth on b o t h  sides. Leaves i n P e l t o b o y k i n i a , B o y k i n i a , S u k s d o r f i a and  Bolandra  bear g l a n d u l a r t r i c h o m e s (Ch. IV) and t h e i r e g l a n d u l a r d e r i v a t i v e s .  They  o c c u r i n v a r y i n g d e n s i t i e s s c a t t e r e d over the l a m i n a e , o f t e n a s s o c i a t e d w i t h the v e i n r e t i c u l u m and a l s o on the l e a f m a r g i n s . i s e s p e c i a l l y pronounced i n B o y k i n i a s e c t i o n T e l e s o n i x . l e a v e s are s u b - g l a b r a t e  This l a t t e r condition Sullivantia  to g l a b r a t e and not a t a l l v i s c i d .  Leaves of P e l t o b o y k i n i a are s t r i k i n g l y l u s t r o u s on l o w e r s u r f a c e s ( c . f . Ohwi (1965) who  their  s t a t e d , t h a t the upper s u r f a c e i s  l u s t r o u s ) , a c o n d i t i o n not found i n the o t h e r genera, a l t h o u g h o c c i d e n t a l i s does approach i t sometimes.  Boykinia  Leaves i n B o y k i n i a s e c t i o n T e l e -  s o n i x and i n S u k s d o r f i a are q u i t e s u c c u l e n t , a c o n d i t i o n p r o b a b l y  related  to t h e i r x e r i c h a b i t a t s . Venation Study of l e a f v e n a t i o n i n a l l t a x a was  based on c l e a r e d b a s a l  l e a v e s , f o l l o w i n g the method of Payne (1969), w h i c h i n v o l v e s b o i l i n g  the  m a t e r i a l i n methanol to e x t r a c t the c h l o r o p h y l l and then s o a k i n g i n 10% KOH  f o r one  t o t h r e e days at about 30°C.  I n most cases f r e s h m a t e r i a l  used and supplemented w i t h h e r b a r i u m m a t e r i a l .  The  terminology  i s that  was  22  of Hickey  (1973).  I l l u s t r a t i o n s of the v a r i o u s v e n a t i o n p a t t e r n s are  shown i n F i g s . 10 and 11. A l l s p e c i e s examined a r e v e r y s i m i l a r , h a v i n g  actinodromous  v e n a t i o n w i t h the p r i m a r y v e i n s a r i s i n g from the b a s a l p o s i t i o n .  The  p r i m a r y v e i n s are branched and f a i r l y s t r a i g h t , o r somewhat s i n u o u s , i n a l l species. 80°),  Secondaries  a r i s e w i t h an acute angle o f d i v e r g e a n c e  and b r a n c h from a s t r a i g h t o r somewhat s i n u o u s  course.  (45 -  Tertiary  v e n a t i o n i n B o y k i n i a , S u k s d o r f i a , S u l l i v a n t i a and B o l a n d r a i s random r e t i c u l a t e b u t tends to be o r t h o g o n a l r e t i c u l a t e i n P e l t o b o y k i n i a . Q u a t e r n a r y v e n a t i o n i n a l l genera i s o r t h o g o n a l r e t i c u l a t e .  Higher  order  v e n a t i o n forms a r e t i c u l u m i n w h i c h the i n d i v i d u a l l e v e l s cannot be distinguished.  I n B o l a n d r a and B o y k i n i a s e c t i o n s B o y k i n i a and R e n i f o l i u m  t h i s corresponds  t o l e v e l 5 o r 6.  In section Telesonix, Suksdorfia,  S u l l i v a n t i a and P e l t o b o y k i n i a i t i s l e v e l 4 o r 5. a t i o n i s incomplete l e a f margin.  M a r g i n a l u l t i m a t e ven-  i n a l l t a x a , v e i n l e t s ending f r e e l y a d j a c e n t t o the  L a r g e r v e i n s ( p r i m a r y and secondary) converge a t t h e t i p o f  each l a r g e l e a f l o b e t o produce a hydathode, s t r u c t u r e s common i n t h e S a x i f r a g i n a e ( M e t c a l f e and Chalk  1950).  The h i g h e r o r d e r v e n a t i o n forms a network o f i m p e r f e c t a r e o l a e , variable i n size. oriented.  The a r e o l a e a r e i r r e g u l a r l y p o l y g o n a l and randomly  D e t a i l s o f the d e n s i t y o f a r e o l a e and v e i n l e t t e r m i n a t i o n s a r e  g i v e n i n Table I I .  S t r i k i n g d i f f e r e n c e s among t a x a o c c u r i n t h e prominence  o f t h e v e n a t i o n p a t t e r n s v i s i b l e t o the naked eye.  This i s a r e f l e c t i o n  o f l e a f t e x t u r e and t h e s i z e and number o f v e i n o r d e r s p r e s e n t .  Bolandra  and B o y k i n i a s e c t i o n s B o y k i n i a and R e n i f o l i u m have more l e v e l s o f v e n a t i o n than the o t h e r t a x a and c o r r e s p o n d i n g l y the r e t i c u l u m appears more c o m p l i cated.  D i f f e r e n c e s between s p e c i e s e x i s t i n t h e d e n s i t y o f a r e o l a e and  of v e i n l e t t e r m i n a t i o n s .  I n p a r t i c u l a r , P e l t o b o y k i n i a i s unique i n t h e  23  FIGURE 1 0 . Leaf venation v i s i b l e to the naked eye i n species of Boykinia. A l l drawings prepared from cleared specimens a) B_. intermedia; b) B_. major; c) B_. a c o n i t i f o l i a ; e) B_. heucheriformis;  (x%).  d) B_. o c c i d e n t a l i s ;  f) B_. r i c h a r d s o n i i ; g) B_. r o t u n d i f o l i a .  24  FIGURE 1 1 . L e a f v e n a t i o n Peltoboykinia,  v i s i b l e t o the naked eye i n s p e c i e s o f  Suksdorfia,  B o l a n d r a and S u l l i v a n t i a . A l l  drawings p r e p a r e d from c l e a r e d specimens, xJg except where n o t e d , a) P_. t e l l i m o i d e s s s p . t e l l i m o i d e s ; b) S u k s d o r f i a c) S^ a l c h e m i l l o i d e s f) S u l l i v a n t i a  ( x l ) ; d) S_. v i o l a c e a  oregana.  ranunculif o l i a ;  ( x l ) ; e) B^ oregana;  25 TABLE I I . Boykinia.  Frequency o f v e i n l e t t e r m i n a t i o n s and a r e o l a e i n s p e c i e s o f Peltoboykinia,  Suksdorfia, Mean No.  Mean No. v e i n l e t  Taxon  areolae/mm  Boykinia a c o n i t i f o l i a B. i n t e r m e d i a B. major B. o c c i d e n t a l i s B. r o t u n d i f o l i a B. r i c h a r d s o n i i B. h e u c h e r i f o r m i s  3.0 3.3 4.6 2.4 2.5 3.0 6.1  Suksdorfia violacea S. a l c h e m i l l o i d e s S. r a n u n c u l i f o l i a  + + + + + + +  B o l j n j r a and S u l l i v a n t i a .  2  endings/areola  0.2* 0.3 0.3 0.5 0.5 0.4 0.1  + + + + + + +  0.2 0.1 0.1 0.2 0.4 0.1 0.1  3.0 2.8 5.9 1.9 3.5 3.3 5.9  5.8 + 0.3 1.4 + 0.3 3.5 + 0.4  1.3 + 0.1 0.9 + 0.2 2.8 + 0.4  7.7 1.2 9.9  B o l a n d r a oregana  1.3 + 0.1  1.6  ±  0.1  2.0  S u l l i v a n t i a oregana  1.4  0.5  0.5 + 0.1  Peltoboykinia t e l l i m oides ssp. t e l l i m o i d e s  0.5 + 0.3  1.3 + 0.5  * standard  ±  1.0 0.9 1.3 0.8 1.5 1.2 1.0  Mean No. v e i n l e t 2 endings/mm  + + + + + + +  0.6 0.5 0.6 .0.6 0.4 0.6 0.4 0.6 0.3 1,0 0.2  0.6  ± ± ± ±  0.5  ±  0.2  0.1  deviation.  large s i z e of i t s areolae.  O t h e r w i s e no d e f e n s i b l e  taxonomic g r o u p i n g s  are suggested on t h e b a s i s o f h i g h e r o r d e r v e n a t i o n , n o r a r e t h e r e any clear ecological interpretations  t o be made from t h e d a t a .  Stomata The  stomata o f the S a x i f r a g i n a e  have been t h e s u b j e c t o f  s t u d i e s by Moreau (1971, 1976), G o r e n f l o t and Moreau (1970) and G o r e n f l o t (1971).  S t p m a t a l c h a r a c t e r i s t i c s were c l a i m e d t o be o f taxonomic s i g n i f -  i c a n c e a l t h o u g h i t i s n o t c l e a r , from t h e s e s t u d i e s , q u i t e what t h i s stitutes.  S t o m a t a l development was i n v e s t i g a t e d  including Boykinia  Boykinia  i n a wide range o f g e n e r a ,  o c c i d e n t a l i s and P e l t o b o y k i n i a  Terminology i s t h a t o f R a d f o r d e t a l . (1974).  con-  t e l l i m o i d e s (Moreau 1976).  Moreau (1976) r e p o r t e d  that  o c c i d e n t a l i s had b o t h anomocytic and a n i s o c y t i c , mesoperigenous  stomata as w e l l as c y c l o c y t i c p e r i g e n o u s types ("anomocytique p e r i g e n e avec couronnes de c e l l u l e s compag.").  Peltoboykinia  t e l l i m o i d e s was r e p o r t e d  26 t o p o s s e s s b o t h mesoperigenous and p e r i g e n o u s anomocytic stomata. b o t h s p e c i e s , s t o m a t a l development o c c u r s n o n - s y n c h r o n o u s l y over  In the  whole l e a f ("type melange"). I n the p r e s e n t  study no attempt was  made to observe  stomatal  development, r a t h e r the d i s t r i b u t i o n o f mature s t r u c t u r e s on b o t h upper lower s u r f a c e s o f the b a s a l l e a v e s was  examined.  Epidermal  and  peels of f r e s h  l e a v e s were used whenever p o s s i b l e , and when n o t , h e r b a r i u m m a t e r i a l  was  substituted. A l l s p e c i e s s t u d i e d had anomocytic stomata s c a t t e r e d i n a dense a r r a y o v e r the lower s u r f a c e o f t h e i r b a s a l l e a v e s . one o r two a d j a c e n t e p i d e r m a l  Occasionally  c e l l s were somewhat s m a l l e r and i n these  cases the stomata c o u l d be d e s c r i b e d as a n i s o c y t i c . No  c y c l o c y t i c stomata  were o b s e r v e d . V a r i a t i o n between s p e c i e s o c c u r s i n the presence o r absence of stomata on the upper s u r f a c e s o f the b a s a l l e a v e s ( T a b l e I I I ) .  In  S u k s d o r f i a and B o y k i n i a s e c t i o n s R e n i f o l i u m and T e l e s o n i x anomocytic s t o mata can be found commonly on the upper b a s a l l e a f s u r f a c e .  In  Peltoboy-  k i n i a , S u l l i v a n t i a and some s p e c i e s of s e c t i o n B o y k i n i a such stomata are v e r y i n f r e q u e n t and u s u a l l y absent.  I n the r e m a i n i n g  species of s e c t i o n  B o y k i n i a ( o c c i d e n t a l i s , r o t u n d i f o l i a and a c o n i t i f o l i a ) and i n B o l a n d r a they a r e p r e s e n t  i n low numbers.  S p e c i e s w i t h s m a l l l e a v e s tend to have  stomata common on b o t h l e a f s u r f a c e s , and l a c k stomata on the upper s u r f a c e . however, e.g.  those w i t h l a r g e l e a v e s tend to  There are s e v e r a l e x c e p t i o n s  S u l l i v a n t i a oregana and B o y k i n i a r i c h a r d s o n i i .  to t h i s ,  Stomatal  s i z e s are a l s o g i v e n i n T a b l e I I I . There i s no c l e a r c o r r e l a t i o n w i t h chromosome number ( c f . Ch. CAULINE LEAVES AND  X).  STIPULES  C a u l i n e l e a v e s form a r e d u c t i o n s e r i e s , b e g i n n i n g  from the  27 TABLE I I I . S t o m a t a l s i z e s and f r e q u e n c y on upper s u r f a c e s o f l e a v e s . Taxon  S t o m a t a l f r e q u e n c y on upper l e a f s u r f a c e .  Mean + SD S t o m a t a l length,^urn.  Boykinia B.  intermedia  Absent/v. r a r e  95 + 11  B.  aconitifolia  Occasional  86 + 11  B. l y c o c t o n i f o l i a  Absent/v. r a r e  122 + 22  B.  Moderate  106 +  B. maj or  Absent/v. r a r e  114 + 21  B. r o t u n d i f o l i a  Moderate  109 +  B.  richardsonii  Common  153  B.  heucheriformis  Common  121 +  Common  127  Absent/v. r a r e  144 + 15  Absent/v. r a r e  123 + 19  S. r a n u n c u l i f o l i a  Common  134  S. v i o l a c e a  Common  110 + 12  S. a l c h e m i l l o i d e s  Common  119 + 13  Occasional  133 + 13  Absent/v. r a r e  128 + 10  B.  occidentalis  jamesii  ± ±  7 9 15 7 14  Peltoboykinia P.  tellimoides ssp. t e l l i m o i d e s  P. t . s s p . w a t a n a b e i Suksdorfia  ±  17  Bolandra B.  oregana  Sullivantia S.  oregana  l a r g e r b a s a l l e a v e s , and becoming s m a l l b r a c t s i n the h i g h e r l e v e l s o f the i n f l o r e s c e n c e .  Lower l e a v e s are p e t i o l a t e and the upper ones become  increasingly sessile. I n a l l s p e c i e s , s t i p u l e s subtend t h e l e a v e s . u l e s a r e composed o f w i n g - l i k e e x t e n s i o n s  o f the base o f t h e p e t i o l e .  Genera v a r y i n the morphology o f t h e upper s t i p u l e s . B o l a r i d r a , B o y k i n i a major and B_j_ i n t e r m e d i a  The l o w e r s t i p -  In Suksdorfia,  t h e upper s t i p u l e s a r e  28  f o l i a c e o u s ( F i g s . 12, 14) and o f t e n become adnate t o t h e c a u l i n e l e a v e s , obscuring  the l e a f - s t i p u l e d i s t i n c t i o n .  I n a l l other species of B o y k i n i a ,  S u l l i v a n t i a and P e l t o b o y k i n i a t h e upper s t i p u l e s a r e s i m i l a r t o t h e l o w e r s t r u c t u r e s , or are represented  by s m a l l f l a p s ( F i g s . 12, 1 3 ) . I n B o y k i n i a  s e c t i o n B o y k i n i a the s t i p u l e s are f r i n g e d w i t h brown b r i s t l e s .  29  FIGURE 12. S t i p u l e morphology i n B o y k i n i a s e c t i o n B o y k i n i a  (allx  a - c) _B_. major, i n b a s i p e t a l sequence; d - f ) B_. i n t e r m e d i a , i n b a s i p e t a l sequence; g) _B_. l y c o c t o n i f o l i a ; h) B_. a c o n i t i f o l i a r o t u n d i f o l i a s i m i l a r ) ; i ) B_. o c c i d e n t a l i s .  (B^.  30  31  FIGURE 13. S t i p u l e morphology i n B o y k i n i a s e c t i o n s T e l e s o n i x  and  R e n i f o l i u m ; and i n P e l t o b o y k i n i a . a)  heucheriformis  ssp. watanabei ( x l ) .  ( x l % ) ; b) _B_ r i c h a r d s o n i i ( x l ) ; c) P_. t e l l i m o i d e s L  32  33  FIGURE 14. S t i p u l e morphology i n S u k s d o r f i a and B o l a n d r a ( a l l a,b) S. r a n u n c u l i f o l i a , i n b a s i p e t a l sequence; c) S. v i o l a c e a d) S. a l c h e m i l l o i d e s ; e , f ) B. oregana, i n b a s i p e t a l sequence.  34  35 IV. The Engler  TRICHOMES  t r i c h o m e s i n t h e S a x i f r a g i n a e were o r i g i n a l l y s t u d i e d by  (1891, 1916-1919) and Rosendahl (1905) .  f o u r main c a t e g o r i e s o f t r i c h o m e :  These r e s e a r c h e r s  a) m u l t i c e l l u l a r , m u l t i s e r i a t e g l a n d u l a r ;  b) m u l t i c e l l u l a r , u n i s e r i a t e g l a n d u l a r ; c) m u l t i c e l l u l a r , s e s s i l e and d) u n i c e l l u l a r , e g l a n d u l a r . (1963) a t t a c h e d  observed  Engler  glandular;  (1916), Rosendahl (1905) and Huber  c o n s i d e r a b l e importance t o h a i r types i n t h e i n f r a g e n e r i c  taxomony o f S a x i f r a g a , a c l a s s i f i c a t i o n w h i c h h a s been s u p p o r t e d e s s e n t i a l l y by p o l l e n e x i n e s t u d i e s (Ferguson & Webb 1970).  Rosendahl (1905), i n  a s t u d y o f N o r t h American genera, observed a s i m i l a r i t y i n t r i c h o m e s t r u c t u r e between M i t e 1 1 a , Heuchera, T e l l i m a , T i a r e 1 1 a  and T o l m i e a , between  B o l a n d r a and S u k s d o r f i a , and between B o y k i n i a and S u l l i v a n t i a .  These a r e  a l l groups w h i c h a r e r e c o g n i z a b l e i n terms o f gross morphology, so the .evidence i n d i c a t e s t h a t the t r i c h o m e complement may p r o v i d e a r e l i a b l e guide t o taxonomic r e l a t i o n s h i p s . Moreau (1976) examined the t r i c h o m e s i n a number o f herbaceous Saxifragaceae,  i n c l u d i n g P e l t o b o y k i n i a and B o y k i n i a o c c i d e n t a l i s , b u t h i s  study d i d n o t encompass a l l organs o f the p l a n t and t h e r e s u l t s were c l e a r l y incomplete.  I n s t u d i e s o f p l a n t h a i r s i t i s important  the e n t i r e p l a n t ( C a r l q u i s t 1961);.  t o examine  M a t e r i a l used i n the p r e s e n t  study i s  l i s t e d i n T a b l e I V . Trichome s t r u c t u r e s from a l l organs were o b s e r v e d . S i x c a t e g o r i e s o f trichome were i d e n t i f i e d :  a) m u l t i c e l l u l a r , m u l t i s e r -  i a t e g l a n d u l a r ; b) m u l t i c e l l u l a r , m u l t i s e r i a t e e g l a n d u l a r w i t h many brown, dead c e l l s (brown c h a f f ) ; c) m u l t i c e l l u l a r , u n i s e r i a t e g l a n d u l a r ; d) m u l t i c e l l u l a r , u n i s e r i a t e e g l a n d u l a r w i t h many brown, dead c e l l s  (brown  c h a f f ) ; e) m u l t i c e l l u l a r , u n i s e r i a t e e g l a n d u l a r ; and f ) u n i c e l l u l a r , eglandular.  Trichome s t r u c t u r e i n h e r b a r i u m m a t e r i a l was compared w i t h  t h a t i n c u l t i v a t e d m a t e r i a l , and no d i f f e r e n c e was found.  Illustrations  36 TABLE IV.  L i s t of m a t e r i a l used i n the t r i c h o m e s t u d y .  d e t a i l s and hebarium l o c a t i o n s are g i v e n i n Appendix 1. G o r n a l l are p r e f i x e d by  Full collection Collections  by  'G'.  Boykinia a c o n i t i f o l i a : G342; H a r v i l l and Segars 11, K r a i 35423. B. i n t e r m e d i a : G23; G24; Jones 8411. B. l y c o c t o n i f o l i a : Sato s.n.; Ohba e t a l . 73099; K-anai s.n. B. major: G45; G335; G337. B. o c c i d e n t a l i s : G2; G8; G22; G86; G216; G255; G256; S t r a l e y 1753. B. r o t u n d i f o l i a : G98; G101; G105. B. h e u c h e r i f o r m i s : G115; G194; G344. B. j a m e s i i : Payson 1564; G i l l e t t & Mosquin 12114; Jones 955. B. r i c h a r d s o n i i : G272; G302; G306. S u k s d o r f i a r a n u n c u l i f o l i a : G15; G245; G332. S. v i o l a c e a : G248; G330; Bohm 1112. S. a l c h e m i l l o i d e s : F i e b r i g 3161; V e n t u r i 3296. Peltoboykinia tellimoides ssp. t e l l i m o i d e s : G328; Maekawa P. t . ssp. w a t a n a b e i : G343; Yamazaki B o l a n d r a oregana:  s.n. s.n.  G260; G262; G340.  S u l l i v a n t i a oregana: G261. S. h a p e m a n i i : Hapeman s.n. J e p s o n i a h e t e r a n d r a : Ornduff 5072. J . p a r r y i : Cromwell 709; M u l r o y s.n. P e l t i p h y l l u m peltatum:  Bohm 1201;  G66;  G67;  G80;  Guppy s.n.;  Steward 7438.  of the t r i c h o m e complement i n each s p e c i e s are shown i n F i g s . Developmental  Relationships  O b s e r v a t i o n s of young p l a n t s o f B o y k i n i a Peltoboykinia  occidentalis  t e l l i m o i d e s ssp. watanabei i n d i c a t e t h a t the brown  t r i c h o m e s are d e r i v e d glandular  15-20.  d e v e l o p m e n t a l l y from the m u l t i s e r i a t e and  t y p e s of the r e s p e c t i v e  u l a r head f o l l o w e d  species.  t u r n brown, c u r l and  twist.  chaffy uniseriate  There i s a l o s s of the  by p r o g r e s s i v e b a s i p e t a l c e l l d e a t h .  The  and  gland-  dead c e l l s  T h i s p r o c e s s i s l e s s marked i n B^ r o t u n d i f o _ l i a ,  where o n l y a few m u l t i s e r i a t e g l a n d s n o r m a l l y degenerate l e a v i n g the viscid-hirsute. of brown c h a f f y  stem  Some h e r b a r i u m specimens, however, possess q u i t e a number hairs.  37  FIGURE 15. Semi-diagrammatic i l l u s t r a t i o n s o f  multicellular  g l a n d u l a r t r i c h o m e s i n B o y k i n i a . A l l drawings x l 8 5 . a,b) B_. l y c o c t o n i f o l i a ; e) J^. o c c i d e n t a l i s ;  c) B_. a c o n i t i f o l i a ; d) B_. i n t e r m e d i a ;  f - h) B_. m a j o r .  38  FIGURE 16. Semi-diagrammatic i l l u s t r a t i o n s o f m u l t i c e l l u l a r g l a n d u l a r t r i c h o m e s i n B o y k i n i a . Drawings a - e x l 8 5 , f x70. a>b) B_. h e u c h e r i f o r m i s ;  c,d) B_. r i c h a r d s o n i i ; e , f ) B_. r o t u n d i f o l i a .  39  FIGURE 17.  Semi-diagrammatic i l l u s t r a t i o n s of m u l t i c e l l u l a r glandular  trichomes i n Peltoboykinia tellimoides. Drawings a - c xl85, d x70, e - g x490. a,b) ssp. watanabei; c,d) ssp. t e l l i m o i d e s ; e,f) tellimoides; g) gland head of ssp.  watanabei.  gland heads of ssp.  40  FIGURE 18. Semi-diagrammatic i l l u s t r a t i o n s of m u l t i c e l l u l a r glandular trichomes i n Suksdorfia and Bolandra. A l l drawings a - d) S. r a n u n c u l i f o l i a ; e - g) S_. violacea; h) S. i,j)  B_. oregana.  xl85. alchemilloides;  41  FIGURE 19. Semi-diagrammatic i l l u s t r a t i o n s of m u l t i c e l l u l a r glandular trichomes i n S u l l i v a n t i a , Jepsonia, and P e l t i p h y l l u m . A l l drawings x l 8 5 . a) J3_. hapemanii; b) S_. oregana; c - e) J_. heterandra; f ) J_. p a r r y i ; g - i ) P_. peltatum.  42  FIGURE 20. Semi-diagrammatic i l l u s t r a t i o n s o f e g l a n d u l a r t r i c h o m e s . a,h)  Brown, c h a f f y , m u l t i s e r i a t e t r i c h o m e  b - g) U n i c e l l u l a r t r i c h o m e s  (x70).  (xl85).  i ) U n i s e r i a t e t r i c h o m e , found o n l y on a n t h e r s and s t i g m a t a o f S u k s d o r f i a alchemilloides  (x750).  43  Observations The and has  v a r i a t i o n i n t r i c h o m e s i z e among the s p e c i e s i s  a c o n s i d e r a b l e i n t r a - i n d i v i d u a l component.  onomic s i g n i f i c a n c e can be a t t a c h e d important i n Table  to i t .  and w h i c h w i l l be emphasized.  Therefore  continuous,  little  tax-  I t i s the s t r u c t u r e s w h i c h are  A summary of the r e s u l t s i s g i v e n  V. Boy^kinia s e c t i o n B o y k i n i a i s c h a r a c t e r i z e d by a t r i c h o m e com-  plement of m u l t i s e r i a t e g l a n d s , brown c h a f f , and u n i c e l l u l a r hairs.  S e c t i o n s R e n i f o l i u m and  normally  eglandular  T e l e s o n i x are s i m i l a r e x c e p t t h a t  (but not always) l a c k the brown c h a f f .  The  they  brown c h a f f i s most  common a t the base o f the stem, b a s a l l e a f p e t i o l e s , and around the l o w e r stem nodes.  T h i s t r i c h o m e type i s e s p e c i a l l y common i n B. o c c i d e n t a l i s  and some specimens can l o o k q u i t e v i l l o u s .  On the upper p a r t of the stem  toward the i n f l o r e s c e n c e , the m u l t i c e l l u l a r , m u l t i s e r i a t e g l a n d s become i n c r e a s i n g l y frequent. and hypanthium.  The  They r e a c h t h e i r maximum d e n s i t y on the p e d i c e l s  m u l t i c e l l u l a r g l a n d head i s u s u a l l y s p h e r i c a l o r  o v a l , but o c c a s i o n a l l y has p r o t r u d i n g p a p i l l a e (as i n BoJLandra and dorfia) .  The  u n i c e l l u l a r eglandular  trichomes  occur densely  f l o r e s c e n c e b r a n c h e s j p e d i c e l s and upper b r a c t s . to the m u l t i s e r i a t e g l a n d u l a r  Suks-  on the i n -  They form an  understory  hairs.  On the l e a v e s , trichomes  o f t e n are a s s o c i a t e d w i t h the  veins  and a l s o o c c u r a l o n g the l e a f m a r g i n s , b e i n g e s p e c i a l l y prominent h e r e i n section Telesonix.  The  l a m i n a e are g e n e r a l l y l e s s d e n s e l y pubescent than  are the stem o r p e t i o l e s . i s e r i a t e glands and more f r e q u e n t  A l l t h r e e h a i r types are p r e s e n t , w i t h the m u l t -  c h a f f b e i n g the commonest.  on the l e a v e s n e a r e r  U n i c e l l u l a r h a i r s become  t o the stem apex.  I n B.  rotundifolia,  an o c c a s i o n a l l e a f a l s o p o s s e s s e d a v e r y s m a l l number o f u n i s e r i a t e glandular  trichomes.  44  TABLE V.  D i s t r i b u t i o n o f trichome s t r u c t u r a l types among B o y k i n i a ,  boykinia,  Suksdorfia,  Bolandra, S u l l i v a n t i a , Peltiphyllum  Unicellular  and J e p s o n i a .  Multicellular Uniseriate  Eglandular Boykinia occidentalis intermedia major aconitifolia lycoctonifolia rotundifolia richardsonii jamesii heucheriformis  Gland.  + + + + + + + + +  -  Bolandra oregana Sullivantia oregana hapemanii  -  (+) + +  -  -  -  (+)  Peltiphyllum peltatum  <-  Jepsonia heterandra parryi  -  Chaff  Multicellular Egland.•  —  _  -  —  —  Gland.  + + + + + + + + +  _  —  — —•  Peltoboykinia tellimoides <ssp. t e l l i m o i d e s ssp. watanabei Suksdorfia ranunculifolia violacea alchemilloides  Pelto-  —  _  _  -  -  -  -  +*  (+) (+)  —  _  -  -  -  -  +  +* +* +  -  -  +*  —  —  -  -  +*  -  -  + +  -  —  (+)* a  + + + (+) - (+) - (+) - (+)  C+)*  _  -  +  _  —  (+)* (+)*•  Chaff  —  - .  + +-  _  +' +  *  ;  trichomes o c c u r which are t r a n s i t i o n a l between m u l t i - and u n i s e r i a t e .  a  :  some o f these may be s u b - s e s s i l e .  ( ) :  Only e n t r y b r a c k e t e d = always o c c u r r i n g b u t i n v e r y low numbers. A l t e r n a t i v e e n t r y b r a c k e t e d = a r a r e c o n d i t i o n , n o t always encountered.  The trichomes.  genus P e l t o b o y k i n i a  has m u l t i c e l l u l a r , u n i s e r i a t e  glandular  Some g l a n d s , however, a r e m u l t i s e r i a t e below and u n i s e r i a t e  above, and thus may be regarded as t r a n s i t i o n a l t y p e s .  The t r i c h o m e s  45  o c c u r most p r o m i n e n t l y on the stem, p e t i o l e s , i n f l o r e s c e n c e branches p e t a l s , but are a l s o p r e s e n t on the l e a v e s .  I n o l d e r p l a n t s , the  nodes bear brown, c h a f f y , u n i s e r i a t e , e g l a n d u l a r  glandular,  and  lower  trichomes.  I n B o l a n d r a , the t r i c h o m e s are m u l t i c e l l u l a r , m u l t i s e r i a t e  and  a l t h o u g h t r a n s i t i o n s to a u n i s e r i a t e c o n d i t i o n o f t e n o c c u r .  In  t r a n s i t i o n a l c a s e s , the t r i c h o m e i s m u l t i s e r i a t e below but  uniseriate  above; much v a r i a t i o n o c c u r s and  almost a l l s t a g e s between the m u l t i -  u n i s e r i a t e s t a t e s can be found.  The  a s s o c i a t e d w i t h the v e i n s , and branches.  The  g l a n d s o c c u r on the l e a v e s ,  on the p e t i o l e s , stem and  hypanthium, however, i s u s u a l l y g l a b r o u s .  g l a n d heads u s u a l l y have from one  and  often  inflorescence The m u l t i c e l l u l a r  to t h r e e prominent p a p i l l a e .  S u k s d o r f i a possesses m u l t i c e l l u l a r , m u l t i s e r i a t e glands, t o g e t h e r w i t h types t r a n s i t i o n a l to the u n i s e r i a t e c o n d i t i o n .  The  multi-  c e l l u l a r g l a n d heads have from one  t o t h r e e p a p i l l a e , a l t h o u g h these are  l e s s marked i n S_. a l c h e m j l l o i d e s .  U n i c e l l u l a r h a i r s are d e n s e l y  on the p e d u n c l e s and p e d i c e l s of S_. v j o l a c e a and  arrayed  S_j_ a l c h e m j l l o i d e s .  They  are v e r y r a r e , and u s u a l l y absent from S^ r a n u n c u l i f o l i a . The d i s t r i b u t i o n of a l l these h a i r types o v e r the p l a n t f o l l o w s the p a t t e r n found i n Boykinia .  Suksdorfia  a l c h e m j l l o i d e s i s a unique s p e c i e s  i n having  i s t i c red, slender, m u l t i c e l l u l a r , u n i s e r i a t e eglandular o c c u r on the a n t h e r s and I n the two  hairs.  characterThey  sometimes among the s t i g m a t i c p a p i l l a e .  S u l l i v a n t i a s p e c i e s examined, the l e a v e s  and  pet-  i o l e s are v e r y s p a r i n g l y p u b e r u l e n t w i t h m u l t i c e l l u l a r , m u l t i s e r i a t e u l a r trichomes. r e g i o n , but  They become i n c r e a s i n g l y common i n the  are r a t h e r s p a r s e on the hypanthium.  The  gland-  inflorescence  m u l t i c e l l u l a r gland  heads are s p h e r i c a l or o y a l as i n B o y k i n i a , w i t h no d i s t i n c t p a p i l l a e .  In  S. hapemanii the peduncles and p e d i c e l s a d d i t i o n a l l y have a v e r y s p a r s e complement of u n i c e l l u l a r , e g l a n d u l a r  trichomes.  These seem t o be  absent  46  f rom _S_. oregana, a t l e a s t i n t h e m a t e r i a l examined (Rosendahl 1927). The  l e a v e s and p e t i o l e s o f J e p s o n i a bear m u l t i c e l l u l a r , mult*-  i s e r i a t e glandular trichomes,  a g a i n a s s o c i a t e d w i t h v e i n s i n t h e laminae.  Toward the t o p o f the stem, u n i s e r i a t e glands become i n c r e a s i n g l y common. They comprise t h e o n l y h a i r type i n t h e i n f l o r e s c e n c e r e g i o n .  The g l a n d  heads a r e o v a l . P e l t i p h y l l u m has a mixed a r r a y o f b o t h u n i s e r i a t e and m u l t i s e r i a t e glandular trichomes, i n c l u d i n g t r a n s i t i o n a l types.  A l l occur on  the l e a v e s , the stem, i n f l o r e s c e n c e branches and h y p a n t h i a .  Many o f the  trichomes The  have v e r y s h o r t s t a l k s and some may be d e s c r i b e d as s u b - s e s s i l e .  g l a n d heads a r e s p h e r i c a l o r o v a l .  Engler  (1928) and Moreau (1976)  d i d n o t r e p o r t t h e p r e s e n c e o f m u l t i s e r i a t e glands i n t h i s genus, b u t they were p r e s e n t i n a l l c o l l e c t i o n s examined i n t h e p r e s e n t  study.  Taxonomic I n f e r e n c e s Trichome complements c l e a r l y i n d i c a t e a v e r y c l o s e r e l a t i o n s h i p between B o y k i n i a , S u k s d o r f i a , S u l l i v a n t i a and B o l a n d r a . be c h a r a c t e r i z e d by the r e g u l a r o c c u r r e n c e eglandular trichomes,  B o y k i n i a can  o f a dense a r r a y o f u n i c e l l u l a r ,  together w i t h m u l t i c e l l u l a r , m u l t i s e r i a t e trichomes  w i t h s p h e r i c a l o r o v a l g l a n d u l a r heads, and i n s e c t i o n B o y k i n i a , by t h e brown, c h a f f y d e r i v a t i v e s . I n t h e p a s t , s p e c i e s o f s e c t i o n T e l e s o n i x have been p l a c e d i n the genus S a x i f r a g a ( e . g . Jones 1910; H a r r i n g t o n However, the t r i c h o m e  1954).  complement argues a g a i n s t such a placement, s i n c e  S a x i f r a g a has n e v e r been r e p o r t e d t o p o s s e s s the u n i c e l l u l a r  trichomes,  and a survey o f 26 s p e c i e s from t e n s e c t i o n s ( T a b l e V I ) d i d n o t r e v e a l any. F u r t h e r m o r e , the o c c u r r e n c e supports  o f the brown, c h a f f y h a i r ( a l t h o u g h r a r e ) a l s o  an a l l i a n c e w i t h B o y k i n i a r a t h e r than S a x i f r a g a . B o y k i n i a l y c o c t o n i f o l i a i s o b v i o u s l y a member o f s e c t i o n Boy-  k i n i a (Table V) on t h e b a s i s o f i t s t r i c h o m e s , w h i c h do n o t s u p p o r t i t s  47  TABLE V I . species.  Trichome  s t r u c t u r e s i n i n f l o r e s c e n c e branches o f S a x i f r a g a  C o l l e c t i o n s by G o r n a l l a r e p r e f i x e d by  I n f r a g e n e r i c category  Section Micranthes Grex P u n c t a t a e p u n c t a t a (G298, 318) m e r t e n s i a n a ( M i l l e r 764)* Grex D a v u r i c a e l y a l l i i (G294, 297) Grex N i v a l i - v i r g i n i e n s i s r e f l e x a (G287) h i e r a c i f o l i a (G292) c a l i f o r n i c a ( M i l l e r 766) o c c i d e n t a l i s (G334) Grex S t e l l a r e s f e r r u g i n e a (G324) b r y o p h o r a ( S h a r s m i t h 4556A) Grex I n t e r m e d i a e t o l m i e i (Bohm 1372) Section Hirculus Grex Sediformes c h r y s a n t h a (Pennard s.n.) Grex H e m i s p h a e r i c a e e s c h s c h o l t z i i (G291) Section T r i d a c t y l i t e s n u t t a l l i i (Bohm 1289) adscendens (Pojair 1/8/75) S e c t i o n Nephrophyllum Grex S i b i r i c a e cernua (G288, 319) Section Dactyloides Grex C a e s p i t o s a e c a e s p i t o s a (G325) Section Trachyphyllum t r i c u s p i d a t a . ( G 2 8 6 , 316) b r o n c h i a l i s (G289) Section Xanthizoon a i z o i d e s (Beamish e t a l . 730369) Section Euaizoonia Grex P e r a i z o o n i a e a i z o o n ( G a r t o n 15349) Grex C o t y l e d o n i a e c o t y l e d o n ( A l a v a e t a l . 9517) Section Kabschia Grex Squarrosae c a e s i a ( F r e y s.n.)  'G'.  Trichome S t r u c t u r e ( w i t h m u l t i c e l l u l a r g l a n d u l a r heads except where n o t e d ) .  Uniseriate Multiseriate Uniseriate Uniseriate Uniseriate Uniseriate Uniseriate  + brown  chaff  Multiseriate Multiseriate Multiseriate  Multiseriate Glabrous Multiseriate, transitional to u n i s e r i a t e Uniseriate Uniseriate Uniseriate Multiseriate, transitional to u n i s e r i a t e Multiseriate U n i s e r i a t e , u n i c e l l u l a r head  Multiseriate Multiseriate, transitional to u n i s e r i a t e  U n i s e r i a t e , some w i t h u n i c e l l u l a r heads  48  TABLE V I , c o n t ' d . Trichome S t r u c t u r e Cwith m u l t i c e l l u l a r g l a n d u l a r heads except where n o t e d ) .  I n f r a g e n e r i c category  Section Porphyrion Grex O p p o s i t i f o l i a e o p p o s i t i f o l i a (G284)  Multiseriate  * M u l t i s e r i a t e t r i c h o m e s i n S_. m e r t e n s i a n a s t r o n g l y s u p p o r t i t s t r a n s f e r to g r e x S t e l l a r e s , a move advocated by S a v i l e (1975) on t h e b a s i s o f f l o r a l morphology.  s e g r e g a t i o n as the monotypic  N e o b o y k i n i a (Hara 1937) .  S u k s d o r f i a and B o l a n d r a a r e v e r y s i m i l a r i n t h e s t r u c t u r e o f t h e i r m u l t i c e l l u l a r glands.  I n b o t h , t r a n s i t i o n a l types between a u n i s e r -  i a t e and a m u l t i s e r i a t e c o n d i t i o n o c c u r , and the g l a n d u l a r heads u s u a l l y possess from one t o t h r e e prominent  papillae.  Two S u k s d o r f i a s p e c i e s ,  S. v i o l a c e a and S_. a l c h e m i l l o i d e s , possess u n i c e l l u l a r h a i r s i n t h e i n f l o r escence.  T h i s f e a t u r e s u p p o r t s the abandonment o f the monotypic  Hierony-  musia t o which S_. a l c h e m i l l o i d e s had been e l e v a t e d ( E n g l e r 1918) a l t h o u g h the p e c u l i a r u n i s e r i a t e , e g l a n d u l a r t r i c h o m e s on i t s a n t h e r s and s t i g m a t a clearly distinguish i t .  The u s u a l absence o f u n i c e l l u l a r h a i r s  from  S. r a n u n c u l i f o l i a ( f o r m e r l y i n t h e monotypic Hemieva) does n o t j e o p a r d i z e i t s p o s i t i o n because somewhat s i m i l a r v a r i a t i o n e x i s t s i n S u l l i v a n t i a w i t h r e s p e c t t o t h i s trichome t y p e .  The presence o f these u n i c e l l u l a r  h a i r s i n b o t h S u k s d o r f i a and S u l l i v a n t i a i n d i c a t e s a v e r y c l o s e r e l a t i o n ship with B o y k i n i a . The r e c o g n i t i o n o f P e l t o b o y k i n i a a t a g e n e r i c l e v e l 1937)  i s s u p p o r t e d by i t s t r i c h o m e complement (Moreau 1976).  (Hara  I t lacks  b o t h the u n i c e l l u l a r and t h e m u l t i s e r i a t e t r i c h o m e s o f B o y k i n i a , h a v i n g instead u n i s e r i a t e types.  I t i s p o s s i b l e , however, t h a t t h e r a r e o c c u r r e n c e  i n B_. r o t u n d i f o l i a o f u n i s e r i a t e g l a n d s may i n d i c a t e some r e l a t i o n s h i p . P e l t o b o y k i n i a i s a l s o the o n l y genus i n the group t h a t p o s s e s s e s g l a n d u l a r  49 petals.  The u n i s e r i a t e g l a n d s suggest an a f f i n i t y w i t h  similarly-  endowed S a x i f r a g a s p e c i e s , o r w i t h P e l t i p h y l l u m ( E n g l e r 1891) and i n d e e d , Jepsonia. The presence i n J e p s o n i a o f b o t h m u l t i - and u n i s e r i a t e g l a n d s i m p l i e s an a l l i a n c e w i t h P e l t i p h y l l u m , a l t h o u g h s e s s i l e types.  the l a t t e r a l s o has sub-  S e s s i l e glands o c c u r i n B e r g e n i a , Mukenia and the p e l t a t e -  l e a v e d A s t i l b o j d e s (Moreau 1976). number 2n = 34 (Hamel 1953).  A l l f o u r genera share t h e chromosome  T h i s i s an u n u s u a l chromosome number i n t h e  S a x i f r a g i n a e and thus b o t h t r i c h o m e and chromosome f e a t u r e s suggest a close r e l a t i o n s h i p , notwithstanding  the f a c t t h a t Engler. (1928) p l a c e d  A s t i l b o i d e s i n s u b - t r i b e A s t i l b i n a e r a t h e r than S a x i f r a g i n a e . Environmental M o d i f i c a t i o n of V e s t i t u r e i n Boykinia o c c i d e n t a l i s B o y k i n i a o c c i d e n t a l i s i s a v a r i a b l e species w i t h segregate v a r i e t i e s and even s p e c i e s h a v i n g been d e s c r i b e d on t h e b a s i s o f v a r i a t i o n i n t r i c h o m e d e n s i t y (Rydberg 1905; Rosendahl 1905).  A s e r i e s of observa-  t i o n s was made on t h e amount o f pubescence on e i g h t p o p u l a t i o n s o f p l a n t s , sampled from Vancouver I s l a n d t o C a l i f o r n i a , and grown under u n i f o r m e n v i r onmental c o n d i t i o n s (17-20°C, 16hr p h o t o p e r i o d ) (Table V I I ) . A f t e r the p l a n t s had been c u l t i v a t e d f o r two y e a r s , which time they f l o w e r e d  during  a t l e a s t t w i c e , a s e r i e s o f measurements was made:  1) t r i c h o m e d e n s i t y on t h e p e t i o l e s o f two b a s a l l e a v e s  (one young and one  o l d ) o f each p l a n t ; 2) l e n g t h o f t h r e e r e p r e s e n t a t i v e t r i c h o m e s on e a c h p e t i o l e examined i n ( 1 ) ; 3) trichome d e n s i t y on t h e hypanthium.  Since the  i m p r e s s i o n o f h a i r i n e s s on t h e p e t i o l e s owes much t p the t a n g l e d n a t u r e o f the t r i c h o m e s ,  a t h i r d parameter, p e t i o l e h a i r i n e s s i n d e x , was c a l c u l a t e d  by m u l t i p l y i n g (1) by the mean o f ( 2 ) .  The r e s u l t s a r e p r e s e n t e d i n  Table V I I . The d a t a show f i r s t t h a t a l l measures o f t r i c h o m e d e n s i t y have  TABLE V I I . V a r i a t i o n i n t r i c h o m e d e n s i t y among e i g h t p o p u l a t i o n s o f B o y k i n i a o c c i d e n t a l i s grown i n a growth chamber. Mean P e t i o l e Trichome D e n s i t y -2 mm  Mean P e t i o l e Trichome L e n g t h  P o p u l a t i o n //  Young  Old  Young  Old  Young  Old  G8 ( 3 ) ; Vancouver I s .  10.3 (1.2)*  4.7 (2.3)  2.3 (0.1)  1.5 (0.3)  23.5 (3.2)  7.4 (4.4)  35.0 + 12.3  G2 ( 4 ) ; Vancouver, B.C.  14.8  C2.2)  7.0 (0.8)  2.2 (l.D  1.9 (1.1)  30.5 (11.2)  13.6 (9.8)  28.0 + 11.8  G22 ( 2 ) ; J e f f e r s o n Co., Wash.  16.0 (2.8)  8.5 (3.5)  2.9 (0.1)  1.9 (0.1)  46.2 (5.9)  15.6 (5.9)  19.0 +  9.9  G216 ( 2 ) ; K i n g Co., Wash.  14.5 (6.4)  6.0 (2.8)  1.9 (0.2)  1.2 (0.2)  26.2 (8.7)  6.6 (2.0)  29.0 +  1.4  G255 ( 2 ) ; L i n c o l n Co., Oreg.  12.5 (3.5)  4.0 (1.4)  3.1 (0.9)  1.6 (0.9)  39.8 (22.3)  7.0 (5.7)  25.7  ±  2.5  G256 ( 3 ) ; C u r r y Co., Oreg.  13.3 (4.0)  10.3 (2.1)  3.5 (0.4)  3.2 (1.7)  46.8 (13.2)  33.2 (21.6)  11.0 +  2.7  S1753 ( 5 ) ; D e l Norte Co., C a l . 17.8 (5.5)  8.2 (1.6)  3.5 (1.2)  2.5 (0.7)  61.2 (22.6)  21.1 (9.2)  47.6 + 21.8  G86 ( 8 ) ; Humboldt Co., C a l .  3.9 (1.3)  2.4 (0.8)  1.7 (0.4)  24.5 (14.1)  6.7 (2.8)  12.8 +  9.8 (3.5)  Mean P e t i o l e H a i r i n e s s Index  mm  Mean Hypanthium Trichome D e n s i t y -2 mm  8.1  #  C o l l e c t i o n numbers a r e those o f G o r n a l l (G) o r S t r a l e y ( S ) . The number o f p l a n t s i s g i v e n i n b r a c k e t s .  *  Standard  Deviations are given i n brackets.  51  completely  c o n t i n u o u s d i s t r i b u t i o n s , and  correlation with this variation. than o l d p e t i o l e s .  t h a t t h e r e i s no  geographical  Second, young p e t i o l e s are more pubescent  This demonstration of glabrescence  v i n d i c a t e s Torrey  and Gray's (1840) s u s p i c i o n t h a t the " c h a f f y h a i r s " , w h i c h were o r i g i n a l l y thought to d i s t i n g u i s h N u t t a l l ' s S a x i f r a g a e l a t a from t h e i r own o c c i d e n t a l i s , were d e c i d u o u s . s i n g l e species  The  r e s u l t s s u p p o r t the r e c o g n i t i o n o f a  only.  F u n c t i o n of the G l a n d u l a r The  Trichomes  g l a n d u l a r p o r t i o n of the h a i r s c o n s i s t s of a m u l t i c e l l u l a r  g l o b u l a r head w h i c h o f t e n c o n t a i n s a n t h o c y a n i n s , and J e p s o n i a .  Boykinia  and at l e a s t i n B o y k i n i a  the s p i c y , a r o m a t i c p r i n c i p l e s (somewhat l i k e  w h i c h h e l p c h a r a c t e r i z e these genera,  ge1toboykinia.  and S u l l i v a n t i a have no odour d e t e c t a b l e by humans.  coriander)  Suksdorfia. Bolandra The  spicy smell  may  s e r v e to a t t r a c t p o l l i n a t o r s . The  s t i c k y g l a n d s a l s o have the p r o p e r t y of t r a p p i n g i n s e c t s ,  p a r t i c u l a r l y s m a l l d i p t e r a , and a p r o t e c t i v e r o l e a g a i n s t f e e d i n g and/or egg l a y i n g i s l i k e l y ( L e v i n 1973). f l o r e s c e n c e r e g i o n a l s o has been shed from the c a p s u l e s .  The  dense a r r a y of g l a n d s i n the i n -  the e f f e c t of t r a p p i n g seeds a f t e r they have T h i s o b v i o u s l y i n h i b i t s d i s p e r s a l to a l a r g e  e x t e n t and presumably a f f e c t s the g e n e t i c a l s t r u c t u r e of p o p u l a t i o n s , the l o n g - t e r m s i g n i f i c a n c e i s o b s c u r e .  but  52  V.  INFLORESCENCE STRUCTURE  The genera B o y k i n i a . P e l t o b o y k i n i a , S u k s d o r f i a , B o l a n d r a and S u l l i v a n t i a a l l have m o n o t e l i c (cymose) i n f l o r e s c e n c e s , w i t h a t e r m i n a l f l o w e r and b a s i p e t a l development o f s u c c e s s i v e p a r a c l a d i a .  I n a l l taxa  the i n f l o r e s c e n c e i s i n i t i a l l y h i g h l y congested and i t i s o n l y a f t e r anthesis  and e s p e c i a l l y d u r i n g f r u i t i n g , f o l l o w i n g peduncle growth, t h a t i t s  s t r u c t u r e becomes r e a d i l y apparent. t h i s stage.  The f o l l o w i n g d e s c r i p t i o n s a p p l y t o  Diagrammatic r e p r e s e n t a t i o n s  are shown i n F i g s . 21-23. cauline leaf.  of t y p i c a l b r a n c h i n g  Each p a r a c l a d i u m i s subtended by a b r a c t o r  I n many cases t h e i n f l o r e s c e n c e c o n s i s t s o f f i r s t  p a r a c l a d i a only, although i n well-developed c l a d i a may be p r e s e n t ,  patterns  order  i n d i v i d u a l s second-order  para-  a g a i n subtended by b r a c t s .  A l l s p e c i e s o f B o y k i n i a s e c t i o n B o y k i n i a have s i m i l a r i n f l o r escence shapes and b r a n c h i n g p a t t e r n s ,  They have l a x p a n i c l e s w i t h each  p a r a c l a d i u m c o n s i s t i n g o f a t e r m i n a l f l o w e r borne a t t h e j u n c t i o n o f a dichotomous b r a n c h p a i r , each b r a n c h of which b e a r s s e v e r a l f l o w e r s i n a t y p i c a l l y h e l i c o i d arrangement. one  The t e r m i n a l f l o w e r i s o f t e n d i s p l a c e d t o  s i d e o f t h e b r a n c h j u n c t i o n , and o c c a s i o n a l l y i t i s a b s e n t .  Boykinia  major i s unique i n t h e s e c t i o n i n h a v i n g a corymbiform i n f l o r e s c e n c e , a l t h o u g h B. i n t e r m e d i a  can approach t h i s c o n d i t i o n .  Boykinia richardsonii  of s e c t i o n R e n i f o l i u m has a s i m i l a r b a s i c s t r u c t u r e t o t h a t i n s e c t i o n B o y k i n i a ; however i t r a r e l y , i f e v e r , b e a r s second-order p a r a c l a d i a and i t has  a d i f f e r e n t i n f l o r e s c e n c e shape.  c l a d i a are t i g h t l y organized  I n s t e a d o f a l a x p a n i c l e , the p a r a -  i n t o a contracted  p a r a c l a d i u m comprised o f a t h r e e - f l o w e r e d  t h r y s o i d s t r u c t u r e , each  d i c h a s i a l cyme.  Species of sec-  t i o n T e l e s o n i x have a s i m i l a r i n f l o r e s c e n c e shape t o t h a t o f B^ r i c h a r d s o n i i but t h e i r p a r a c l a d i a are represented i c a l l y o n l y one l a t e r a l  flower.  by a t e r m i n a l f l o w e r w i t h  typ-  53  FIGURE 21.  I n f l o r e s c e n c e s t r u c t u r e i n some s p e c i e s o f B o y k i n i a  section Boykinia. a) B_. l y c o c t o n i f o l i a ; b) B_. o c c i d e n t a l i s ; c) B_. i n t e r m e d i a ; d) B_. major . O Flower  • Terminal flower  C  Bract or  leaf  55  FIGURE 22. I n f l o r e s c e n c e s t r u c t u r e i n s p e c i e s o f B o y k i n i a  and  Peltoboykinia. a) B_. r o t u n d i f o l i a ; b) B_. r i c h a r d s o n i i ; c) B_. h e u c h e r i f o r m i s ; d) P_^ t e l l i m o i d e s (both s u b s p e c i e s ) . O  Flower  • Terminal  flower  /"  Bract or l e a f  56  57  FIGURE 23. I n f l o r e s c e n c e s t r u c t u r e i n S u k s d o r f i a and  Bolandra.  a) S_. r a n u n c u l i f o l i a ; b) j3_. v i o l a c e a ; c) S_. a l c h e m i l l o i d e s ; d) B_. oregana. O Flower  • Terminal  flower  /*"  B r a c t or l e a f  59  A s e r i e s s i m i l a r to t h a t i n B o y k i n i a i s found i n S u k s d o r f i a . Thus the i n f l o r e s c e n c e s t r u c t u r e of S_. r a n u n c u l i f o l i a i s v e r y s i m i l a r to t h a t of B o y k i n i a s e c t i o n B o y k i n i a . s e c o n d - o r d e r p a r a c l a d i a and  fewer  fewer f l o w e r s p e r b r a n c h so t h a t i t s i n f l o r -  escence s t r u c t u r e i s i n t e r m e d i a t e and  S u k s d o r f i a a l c h e m j l l o i d e s has  between t h a t of B o y k i n i a r i c h a r d s o n i i  that of Boykinia s e c t i o n Telesonix.  The  s i m p l e s t c o n d i t i o n i s seen  i n S u k s d o r f i a v i o l a c e a whose shoots have v e r y few p a r a c l a d i a , each of w h i c h c o n s i s t s of a t e r m i n a l f l o w e r w i t h a s i n g l e l a t e r a l bloom. p l a n t s have but a s i n g l e t e r m i n a l  Some  flower.  I n B o l a n d r a and P e l t o b o y k i n i a , the i n f l o r e s c e n c e s t r u c t u r e i s as i n B o y k i n i a s e c t i o n B o y k i n i a , a l t h o u g h the number of f l o w e r s per c l a d i u m i s fewer.  T h i s i s e s p e c i a l l y t r u e o f B o l a n d r a where the v e r y l a x ,  open p a n i c l e t y p i c a l l y has p a r a c l a d i a w i t h a t e r m i n a l and f l o w e r s o n l y , b u t second-order p a r a c l a d i a are q u i t e The  para-  i n f l o r e s c e n c e of S u l l i v a n t i a was  by Rosendahl (1927).  two  lateral  frequent.  described  and  illustrated  I t i s a l s o v e r y s i m i l a r to the p a t t e r n found i n  Boykinia section Boykinia.  However i t does d i f f e r i n t h a t the  paracladia  are o f t e n composed of " t r i c h a s i a l cymes", i . e . t h r e e , many-flowered b r a n c h e s , the c e n t r a l one of w h i c h b e a r s the t e r m i n a l f l o w e r a t i t s base. E n v i r o n m e n t a l M o d i f i c a t i o n of I n f l o r e s c e n c e  Structure  Boykinia occidentalis i s especially variable i n i t s i n f l o r escence s t r u c t u r e . basis;  A s e g r e g a t e s p e c i e s has even been d e s c r i b e d on  this  Therophon ( B o y k i n i a ) c i n c i n n a t u m Rosendahl e t Rydberg (1905).  These i n d i v i d u a l s have p a r a c l a d i a formed as i n B_. r o t u n d i f o l i a ( F i g . 2 2 ) , i . e . comprised of f l o w e r s w i t h curved p e d i c e l s borne i n h i g h l y h e l i c o i d cymes.  I n o t h e r i n d i v i d u a l s of B. o c c i d e n t a l i s , the  s t r u c t u r e i s l e s s r e g u l a r and straight pedicels.  regular inflorescence  the f l o w e r s are l e s s secund, some w i t h  O b s e r v a t i o n has  shown t h a t the r e g u l a r i t y of  an  60 i n f l o r e s c e n c e i n c r e a s e s w i t h age. I n d i v i d u a l s from d i f f e r e n t p o p u l a t i o n s grown i n the growth chamber were a l l c i n c i n n a t e by t h e time t h e i r had  ripened.  capsules  E x a m i n a t i o n o f h e r b a r i u m specimens showed t h a t t h e v a r i a t i o n  i s continuous,  w i t h no apparent g e o g r a p h i c a l  correlation.  Indeed, on  s i n g l e p l a n t s , some p a r a c l a d i a had f l o w e r s on s t r a i g h t p e d i c e l s and o t h e r s b o r e secund f l o w e r s .  P i p e r (1906) concluded t h a t T_. c i n c i n n a t u m was o n l y  " f e e b l y d i s t i n g u i s h a b l e by a l a r g e r i n f l o r e s c e n c e w i t h the branches more o r l e s s r a c e m i f o r m and curved p e d i c e l s " . I n the p r e s e n t study some attempts were made t o a l t e r e x p e r i m e n t a l l y b o t h i n f l o r e s c e n c e s t r u c t u r e and p e d i c e l c u r v a t u r e . e x p e r i m e n t , c l o n e d i n d i v i d u a l s o f each o f t h r e e p o p u l a t i o n s 256,  S t r a l e y 1753) were d e l i b e r a t e l y underwatered.  eloped very small b a s a l leaves  I n the f i r s t ( G o r n a l l 22,  I n each case they dev-  and produced a h i g h l y depauperate and i r r e g -  u l a r shoot system when compared w i t h t h e i r w e l l - w a t e r e d  counterparts  with  i d e n t i c a l genotypes ( F i g . 2 4 ) . I n t h e second e x p e r i m e n t , one i n d i v i d u a l o f p o p u l a t i o n 86 had i t s d e v e l o p i n g  Gornall  shoot system t i e d w i t h c o t t o n i n such a way t h a t one  f l o w e r i n g stem was o r i e n t e d h o r i z o n t a l l y and t h e o t h e r v e r t i c a l l y . opment o f the f l o w e r s from bud t o f r u i t was then o b s e r v e d .  Devel-  I n the v e r t i c a l  stem, where t h e p a r a c l a d i a developed a t about 60° t o the h o r i z o n t a l , the f l o w e r s began w i t h s t r a i g h t p e d i c e l s .  As they aged and s e t seed, many o f  them, e s p e c i a l l y those a t t h e p r o x i m a l ends o f the p a r a c l a d i a , developed somewhat curved p e d i c e l s ; those n e a r e r the d i s t a l ends u s u a l l y r e t a i n e d their straight pedicels.  I n t h e h o r i z o n t a l stem, w i t h t h e p a r a c l a d i a  o r i e n t e d e i t h e r upward o r downward, a l l t h e f l o w e r s on t h e i n v e r t e d c l a d i a developed curved p e d i c e l s v e r y r a p i d l y , some becoming  para-  completely  reflexed.  The u p r i g h t p a r a c l a d i a developed f l o w e r s w i t h o n l y s t r a i g h t  pedicels.  I t was concluded t h a t p e d i c e l c u r v a t u r e  i n v o l v e s t h e photo-  61  FIGURE 24. Inflorescences of droughted ( l e f t ) and watered clones of Boykinia occidentalis  (x3/5).  (right)  62  and/or g e o t r o p i c  responses of; the  Since s t r u c t u r e i n B.  of  environmental  served The  v a r i a t i o n i n both p e d i c e l curvature  o c c i d e n t a l i s has  these c h a r a c t e r s  plant.  a considerable  and  inflorescence  e n v i r o n m e n t a l component,  cannot be used r e l i a b l y f o r taxonomic purposes,. m o d i f i c a t i o n of i n f l o r e s c e n c e s t r u c t u r e was  i n i n d i v i d u a l s of B_. major t h a t had  s u f f e r e d a t t a c k by  This kind also  ob-  weevils.  p l a n t s developed g r e a t l y reduced shoot systems compared w i t h those  produced by unmolested specimens.  I n n a t u r e i t was  p l a n t s of a l l taxa, when growing i n a p p a r e n t l y produced shoots w i t h v e r y  simple or no  also noticed  unfavourable  branching.  that  conditions,  63  VI.  The  THE FLOWER  flowers of Boykinia, Peltoboykinia, Suksdorfia,  Sullivantia  and B o l a n d r a a r e r e g u l a r and a r e composed o f f o u r o r f i v e w h o r l s o f p a r t s , v i z . c a l y x , c o r o l l a , a one o r two-whorled androecium, and a b i c a r p e l l a t e gynoecium.  The f l o w e r s a r e t y p i c a l l y a c t i n o m o r p h i c ,  i s o c c a s i o n a l l y seen.  although  some asymmetry  V a r i a t i o n i n t h i s g e n e r a l morphology does o c c u r and  i s d i s c u s s e d i n a s p e c i a l s e c t i o n on t e r a t o l o g i c a l phenomena. Calyx I n a l l s p e c i e s the c a l y x i s composed o f f i v e s e p a l s w h i c h a r e f u s e d w i t h t h e b a s a l p o r t i o n s o f t h e p e t a l s and stamens t o form t h e f r e e hypanthium.  The a p i c e s o f the s e p a l s a r e t r i a n g u l a r and f r e e .  The hypan-  thium i s c l o t h e d i n g l a n d u l a r h a i r s , and sometimes, s m a l l e r u n i c e l l u l a r t r i c h o m e s as w e l l (Ch. iv) i n a l l genera except B o l a n d r a , where i t i s g l a brous .  I n S u l l i v a n t i a and B o y k i n i a l y c o c t o n i f o l i a t h e s e p a l s a r e i m b r i c a t e  at t h e i r b a s e s .  The s e p a l s i n S u l l i v a n t i a a l s o have w h i t i s h m a r g i n s , a  c o n d i t i o n n o t found i n the o t h e r  genera.  Corolla Most s p e c i e s have w h i t e p e t a l s , b u t some have o t h e r e.g.  yellow-green  colours,  i n B o y k i n i a l y c o c t o n i f o l i a , white w i t h rose veins i n  B. r i c h a r d s o n i i , c r i m s o n - p u r p l e  i n B_. j a m e s i i , v i o l e t o r almost w h i t e i n  S u k s d o r f i a v i o l a c e a , r o s e o r w h i t e i n S_. a l c h e m j l l o i d e s . p e t a l colour i n both subspecies  i sidentical,  In Peltoboykinia  c . f . Ohwi (1965):  newly-  emergent p e t a l s a r e a cream c o l o u r w h i c h darkens t o a p a l e y e l l o w a t anthes i s , and s u b s e q u e n t l y fades back t o a cream c o l o u r . somewhat v a r i a b l e i n p e t a l c o l o u r : i n some they can be y e l l o w - g r e e n yellow-green.  Bolandra i s also  most p o p u l a t i o n s have r e d p e t a l s , b u t  w i t h r e d margins and o c c a s i o n a l l y j u s t  Some p o p u l a t i o n s o f B_j_ major have f l o w e r s w i t h  purple  64  b l o t c h e s a t the base of the p e t a l s (e.g. K r u c k e b e r g 3722, from the T r i n i t y A l p s , N.  California). P e t a l margins are u s u a l l y e n t i r e , except i n P e l t o b o y k i n i a  which produces toothed p e t a l s .  However, i n some p o p u l a t i o n s of  some i n d i v i d u a l s have t o o t h e d or s e r r a t e d p e t a l s . where t h i s phenomenon was  major  I n the s i n g l e case  encountered i n n a t u r e , almost a l l the p l a n t s  had p e t a l s w i t h e i t h e r s e r r a t e d margins or w i t h e n t i r e m a r g i n s . "Very few i n d i v i d u a l s had morphism i s unknown.  f l o w e r s of b o t h k i n d s .  Only a  The b a s i s f o r t h i s p o l y -  I t o c c u r s i n b o t h the B i t t e r r o o t Mountains and  the  Oregon Cascades. A s i m i l a r polymorphism a p p a r e n t l y e x i s t s i n S u k s d o r f i a V i o l a c e a , a l t h o u g h I have never observed i t ( S u k s d o r f , i n l e t t e r w i t h h o l o type; Rydberg 1905). P e t a l shape v a r i e s among the s p e c i e s s t u d i e d and the t i o n s i n F i g s . 25 - 27 s h o u l d be c o n s u l t e d .  illustra-  Some s p e c i e s show a pronounced  claw on the p e t a l , i n o t h e r s i t i s reduced and i n s t i l l o t h e r s i t i s absent.  P e l t o b o y k i n i a i s unique i n the group i n i t s p o s s e s s i o n of  p e t a l s , and B o l a n d r a  i s d i s t i n c t with i t s subulate p e t a l s .  glandular  Sullivantia  i s a l s o d i f f e r e n t , i n t h a t i t s p e t a l s are c h a r a c t e r i s t i c a l l y u n d u l a t e ,  a  c o n d i t i o n approached o n l y s l i g h t l y by B o y k i n i a major. A n a t o m i c a l l y , each p e t a l p o s s e s s e s a s i n g l e v a s c u l a r b u n d l e i n the f r e e p a r t o f i t s b a s a l p o r t i o n .  V a r i o u s numbers of t r a c e s  from t h i s p r i n c i p a l bundle i n d i f f e r e n t s p e c i e s ( F i g s . 25  diverge  -27).  I n n a t u r e , marcescent p e t a l s occur i n S u l l i v a n t i a and lycoctonifolia.  They occur a l s o i n some growth-room-maintained  t i o n s of B o y k i n i a o c c i d e n t a l i s , B. i n t e r m e d i a and B_. major.  Boykinia popula-  In this  pro-  t e c t e d environment, the a b s c i s s i o n l a y e r a t the base of the p e t a l s f a i l s to develop f u l l y , and  the p e t a l s p e r s i s t on the f l o w e r s , r e s i s t i n g even  vigorous disturbance.  T h i s i s i m p o r t a n t because the c h a r a c t e r has been  65  FIGURE 25. P e t a l shapes i n B o y k i n i a s e c t i o n B o y k i n i a ( a l l x l O ) . a,b) B^ i n t e r m e d i a ; c - e) B_. a c o n i t i f o l i a ; f,g) B_. major; h , i ) B^ o c c i d e n t a l i s ; j ,k) B_. r o t u n d i f o l i a ; 1) B^ l y c o c t o n i f o l i a .  67  FIGURE 26. P e t a l shapes i n B o y k i n i a Renifolium;  and  sections Telesonix  and  i n Peltoboykinia.  a) B. j a m e s i i ( x l 3 ) ; b,c)  B_. h e u c h e r i f o r m i s  ( x l 3 ) ; d)  r i c h a r d s o n i i ( x l O ) ; e , f ) P^ t e l l i m o i d e s (both ssp.)  B.  (x3).  69  FIGURE 27. P e t a l shapes i n S u k s d o r f i a  and B o l a n d r a ( a l l x l O ) .  a - c) _S_. r a n u n c u l i f o l i a ; d) S_. a l c h e m j l l o i d e s ; e , f ) S_. v i o l a c e a ; g) B_. oregana.  70  71  considered  t a x o n o m i c a l l y v a l u a b l e i n the p a s t (Hara 1937).  P e t a l s are  deciduous i n growth-room m a t e r i a l o f P e l t o b o y k i n i a . P e t a l Movements I n B o y k i n i a ma.1 or and B_. i n t e r m e d i a , the p e t a l s c l o s e around the s t y l e s as the f l o w e r ages, f o r m i n g a tube, a t the c e n t r e o f one of w h i c h s i t the s t i g m a t a .  The  stamens are no l o n g e r v i s i b l e ,  s h i e l d e d from v i e w by the i m b r i c a t e p e t a l laminae.  pals,  e.g.  being  These p e t a l movements  i n o l d , p o s s i b l y u n r e c e p t i v e or a l r e a d y f e r t i l i z e d f l o w e r s , may p o l l i n a t o r s to keep away.  end  signal  I n o t h e r s p e c i e s w i t h p e t a l s l o n g e r than the se<-<  B o y k i n i a o c c i d e n t a l i s , B. r i c h a r d s o n i i , B.  l f o l i a . S u l l i v a n t i a oregana, B o l a n d r a  j a m e s i i , B. a c o n i t -  oregana, S u k s d o r f i a and P e l t o b o y k i n i a .  the p e t a l s become i n c r e a s i n g l y r e f l e x e d as the f l o w e r opens and  ages.  S p e c i e s w i t h s h o r t , or i n c l u d e d p e t a l s ( B o y k i n i a r o t u n d i f o l i a . B. i f o r m i s ) show l i t t l e or no movement of these  heucher-  organs.  Androecium A l l stamens a r e f r e e and a r e i n two w h o r l s o f f i v e each i n P e l t o b o y k i n i a and B o y k i n i a s e c t i o n T e l e s o n i x , and i n a s i n g l e w h o r l f i v e i n a l l the o t h e r s p e c i e s .  I n the diplostemonous s p e c i e s the  of  outer  w h o r l i s a n t i - s e p a l o u s and d e h i s c e s e a r l i e r than the a n t i - p e t a l o u s i n n e r whorl.  I n the haplostemonous s p e c i e s the stamens are a n t i - s e p a l o u s .  stamen w h o r l s are adnate t o and a r i s e from the hypanthium.  The  A single  v e i n e d , c o n i c a l f i l a m e n t b e a r s the anther w h i c h i s c o r d a t e at the base and i n B o y k i n i a s e c t i o n B o y k i n i a , P e l t o b o y k i n i a and S u k s d o r f i a a l c h e m i l l o i d es , p o s s e s s e s a s m a l l a p i c a l awn. young u n d e h i s c e d a n t h e r s . the a n t h e r ages. microsporangia.  T h i s awn  i s u s u a l l y most marked i n  I t becomes l e s s d i s t i n c t and may  Each anther has  disappear  two t h e c a e , each of w h i c h c o n t a i n s  D e h i s c e n c e i s by l a t e r a l l o n g i t u d i n a l s l i t s ,  p o r t i o n s of each theca becoming c o m p l e t e l y  r e f l e x e d so  the  as two  lateral  as to p r e s e n t  the  72  p o l l e n b o t h i n t r o r s e l y and e x t r o r s e l y , almost In a 360° a r c .  In many  c a s e s , p o l l e n p r e s e n t a t i o n i s v e r y c l o s e t o the s t i g m a t a ( B o y k i n i a o c c i d e n t a l i s . B. r o t u n d i f o l i a , B. l y c o c t o n i f o l i a , S u k s d o r f i a v i o l a c e a and jT. a l c h e m j l l o i d e s ' ) .  Most s p e c i e s examined a r e p r o t a n d r o u s ,  e x t e n t v a r i e s from s p e c i e s t o s p e c i e s (Table V I I I ) .  although the  The a n t h e r s o f P e l t o -  b o y k i n i a change from p a l e cream t o b l a c k j u s t b e f o r e d e h i s c e n c e .  In a l l  o t h e r s p e c i e s the a n t h e r s remain y e l l o w . Stamen Movements I n i t i a l l y the a n t h e r s A  s  form a c l o s e r i n g around the young s t y l e s .  the f l o w e r ages and the ovary and s t y l e s e l o n g a t e , t h e p e t a l s and  s e p a l s spread o u t , p u l l i n g the stamens away from t h e s t y l e s .  Anther de-  h i s c e n c e u s u a l l y o c c u r s b e f o r e f l o w e r reaches i t s maximum s p r e a d .  There  are no independent stamen movements as a r e found i n S a x i f r a g a (Spongberg 19 72), a l l p o s i t i o n a l changes b e i n g dependent on hypanthium shape a l t e r a t i o n s mediated by ovary  growth.  Gynoecium A l l genera s t u d i e d have a x i l e p l a c e n t a t i o n , u s u a l l y w i t h two c a r p e l s f o r m i n g a b i l o c u l a r ovary.  C a r p e l f u s i o n v a r i e s among t h e s p e c i e s  from p a r t i a l t o complete ( F i g s . 28 - 30).  I n those s p e c i e s w i t h t h e upper  p a r t s of t h e i r o v a r i e s f r e e , the p l a c e n t a t i o n i n t h i s r e g i o n i s c o r r e c t l y termed m a r g i n a l .  I n s p e c i e s w i t h almost complete c a r p e l f u s i o n , t h e s t y l e s  are j o i n e d by a d e l i c a t e membrane w h i c h o f t e n r u p t u r e s a l o n g some o f i t s length a l l o w i n g the s t y l e s to separate.  As a taxonomic c h a r a c t e r i t i s  thus p o t e n t i a l l y v a r i a b l e and s h o u l d be used w i t h c a r e , e s p e c i a l l y i n h e r b a r i u m specimens.  Ovary p o s i t i o n v a r i e s between t h e s p e c i e s from  nearly i n f e r i o r to nearly superior.  The v a r i a t i o n has a d e v e l o p m e n t a l  component because as a f l o w e r ages the ovary e l o n g a t e s , becoming l e s s inferior.  The l e n g t h o f the f r e e hypanthium a l s o v a r i e s between s p e c i e s ,  73  FIGURE 28. L o n g i t u d i n a l s e c t i o n s o f f l o w e r s of some s p e c i e s of a) B_. o c c i d e n t a l i s ( x l O ) ; b) J3_. i n t e r m e d i a aconitifolia  (x8).  Boykinia.  ( x 8 ) ; c) B_. major ( x 7 ) ; d)  B_.  74  75  FIGURE 29. L o n g i t u d i n a l s e c t i o n s of f l o w e r s of some s p e c i e s o f  Boykinia  and P e l t o b o y k i n i a . a) B_. r o t u n d i f o l i a  ( x l 4 ) ; b) B_. l y c o c t o n i f o l i a  ( x l 4 ) ; c) B.  heucheri-  f o r m i s ( x 8 ) ; d) P_. t e l l i m o i d e s s s p . w a t a n a b e i ( x 5 ) ; e) B_;_ r i c h a r d s o n i i (x5) .  76  77  FIGURE 30. L o n g i t u d i n a l s e c t i o n s o f f l o w e r s of S u k s d o r f i a and a) S_. a l c h e m i l l o i d e s ( x l 3 ) ; b) S_. r a n u n c u l i f o l i a (x8) ; d) B_. oregana (x8) .  Bolandra.  ( x 8 ) ; c) S_^ v i o l a c e a  78  79  TABLE V I I I .  P r o t a n d r y and n e c t a r i e s i n some s p e c i e s c u l t i v a t e d i n a  growth room. Taxon  Approx. degree of protandry, h r s .  Nectary  tissue  Boykinia occidentalis  96  Insignificant; YellowishGreen  B. i n t e r m e d i a  96  Prominent;  Yellow  B. major  96  Prominent;  Yellow  B. r o t u n d i f o l i a  48  Absent  Suksdorfia ranunculifolia  108  Prominent;  Yellow  S. v i o l a c e a  12  Absent  Peltoboykinia tellimoides*  72  I n s i g n i f i c a n t ; Green  S u l l i v a n t i a oregana  96  I n s i g n i f i c a n t ; Green  *  both  subspecies  a l t h o u g h t h i s c h a r a c t e r i s more o r l e s s c o n s t a n t throughout  the l i f e o f a  f l o w e r ; i t i s thus of v a l u e t a x o n o m i c a l l y , e s p e c i a l l y i n s p e c i e s d e l i m i t ation.  Ovary p o s i t i o n to a l a r g e e x t e n t d i c t a t e s hypanthium shape:  t u r b i n a t e t o campanulate i n B o y k i n i a , S u k s d o r f i a , and S u l l i v a n t i a ; t u b u l a r campanulate i n P e l t o b o y k i n i a and B o l a n d r a .  As seeds r i p e n the ovary s w e l l s  and the c a p s u l e becomes u r c e o l a t e i n a l l t a x a except S u l l i v a n t i a , where i t remains campanulate (Rosendahl  1927), and i n some p o p u l a t i o n s o f B o y k i n i a  a c o n i t i f o l i a whose c a p s u l e s remain  turbinate.  I n many s p e c i e s a n e c t a r y o c c u r s i n a band around the s t y l e base where the f r e e hypanthium and the ovary w a l l meet ( T a b l e V I I I ) . band i s d i s c - l i k e i n S u k s d o r f i a r a n u n c u l i f o l i a .  The  The n e c t a r y i s composed of  y e l l o w , green, o r y e l l o w - g r e e n g l a n d u l a r t i s s u e and s e c r e t e s a sugary s o l t i o n s e v e r a l hours p r i o r to the s t i g m a t a becoming r e c e p t i v e . F o l l o w i n g p o l l i n a t i o n , as the f l o w e r ages, the s t y l e s , s t i g m a t a and p e t a l s become flushed w i t h anthocyanins.  The amount i s v a r i a b l e and u s u a l l y n o t  apparent i n t h e genus P e l t o b o y k i n i a .  I f the p o l l i n a t i o n i s s u c c e s s f u l ,  80  the c a r p e l s e l o n g a t e and expand, an event n o t i c e a b l e a f t e r about two weeks. D e v e l o p i n g o v u l e s remain e n c l o s e d i n t h e ovary i n a l l genera. seeds r i p e n , t h e s t y l e s i n s p e c i e s w i t h l i t t l e  As t h e  c a r p e l f u s i o n become i n -  c r e a s i n g l y d i v e r g e n t ( B o y k i n i a . P e l t o b o y k i n i a and S u k s d o r f i a ) .  I n other  s p e c i e s t h e s t y l e s r e m a i n c l o s e t o g e t h e r u n t i l d e h i s c e n c e when they r e f l e x ( S u l 1 i v a n t i a and B o l a n d r a ) .  I n a l l s p e c i e s the c a p s u l e s d e h i s c e by a  l o n g i t u d i n a l s l i t between t h e s t y l e s . The s t i g m a t a a r e c a p i t a t e w i t h u n i c e l l u l a r p a p i l l a e i n B o y k i n i a , P e l t o b o y k j n i a , B o l a n d r a and i n S u k s d o r f i a r a n u n c u l i f o l i a , b u t t r u n c a t e i n S u k s d o r f i a v i o l a c e a . S. a l c h e m i l l o i d e s , and S u l l i v a n t i a .  Recent c y t o -  c h e m i c a l and p h y s i o l o g i c a l i n v e s t i g a t i o n s o f the angiosperm s t i g m a have r e s u l t e d i n i t s c l a s s i f i c a t i o n i n t o two main t y p e s : ffarrison  1975) .  wet and d r y ( H e s l o p -  Wet types show a d i s t i n c t s u r f a c e s e c r e t i o n d u r i n g t h e i r  r e c e p t i v e p e r i o d , whereas d r y types do n o t .  W i t h i n each type f u r t h e r sub-  d i v i s i o n s have been made based on the morphology o f t h e r e c e p t i v e s u r f a c e . A b r o a d s u r v e y o f the angiosperms has shown t h a t many f a m i l i e s a r e homogeneous i n s t i g m a t y p e , b u t some ( i n c l u d i n g S a x i f r a g a c e a e s . s . ) , a r e n o t . Thus wet (group I I I ) genera i n c l u d e Mukdenia, B e r g e n i a , P e l t i p h y l l u m , Heuchera, T e l l i m a and T i a r e l l a .  Dry (group I I B ) genera i n c l u d e A s t i l b e ,  L i t h o p h r a g m a , and R o d g e r s i a ( H e s l o p - H a r r i s o n and Shivanna  1977) .  (1965) however, d e s c r i b e d L i t h o p h r a g m a as h a v i n g wet s t i g m a t a . i s unique i n p o s s e s s i n g b o t h t y p e s :  Saxifraga  14 s p e c i e s had d r y s t i g m a t a and o n l y  one had wet s t i g m a t a ( H e s l o p - H a r r i s o n and Shivanna suggested  Taylor  1977).  These a u t h o r s  t h a t f u r t h e r heterogeneous genera w i l l be found as s t u d i e s p r o -  ceed b u t t h a t " t h o s e encountered emphasize t h e remarkable characteristic".  i n the s u r v e y so f a r s e r v e m a i n l y t o  c o n s i s t e n c y o f s t i g m a s u r f a c e t y p e as a g e n e r i c  I n the p r e s e n t s t u d y , wet (group I I I ) s t i g m a t a were ob-  s e r v e d i n B o y k i n i a , P e l t o b o y k i n i a , S u k s d o r f i a , S u l l i v a n t i a and B o l a n d r a .  81  Soltis (in l i t t . ) The exception  reported  t h a t a l l S u l l i v a n t i a s p e c i e s have wet s t i g m a t a .  s t y l e s i n a l l s p e c i e s s t u d i e d have a c a n a l w h i c h , w i t h the  o f S u l l i v a n t i a oregana, i s l i n e d w i t h t r a n s m i t t i n g t i s s u e . I n  the case o f P e l t o b o y k i n i a . b o t h s u b s p e c i e s have c a n a l s c o m p l e t e l y with this tisue.  filled  I t i s probable that a l l species of S u l l i v a n t i a lack t r a n s -  m i t t i n g t i s s u e s i n c e B e n s e l and P a l s e r (1975) a l s o r e p o r t i t s absence from S. r e n i f o l i a .  S u l l i v a n t i a i s a l s o r a t h e r d i f f e r e n t from the o t h e r  i n the l a r g e s i z e o f i t s s t y l a r c a n a l . described  genera  I t s s t y l e s c o u l d more a c c u r a t e l y be  as h o l l o w .  F l o r a l Anatomy On the b a s i s o f a d e t a i l e d study o f 26 s p e c i e s o f E n g l e r ' s Saxifragoideae,  i n c l u d i n g R i b e s , B e n s e l and P a l s e r (1975) c o n c l u d e d t h a t  the s u b - f a m i l y  was remarkably homogeneous i n terms o f b o t h f l o r a l anatomy  and morphology. They p r o v i d e d "The  the f o l l o w i n g d e s c r i p t i o n :  vascular c y l i n d e r i n the p e d i c e l generally consists of  s e v e r a l t o many d i s c r e t e bundles from which d i v e r g e t e n compound t r a c e s at the base o f the r e c e p t a c l e , l e a v i n g an i n n e r c y l i n d e r o f v a s c u l a r strands  that coalesce  a t a h i g h e r l e v e l i n t o e i t h e r as many v e n t r a l bun-  d l e s as c a r p e l s o r t w i c e t h a t number. bundle c o n s i s t s o f one-half  I n the former c a s e , each v e n t r a l  of the v a s c u l a r supply  p e l and s e p a r a t e s i n t o i n d i v i d u a l v e n t r a l s t r a n d s the o v a r y .  t o each a d j a c e n t  car-  i n the d i s t a l h a l f of  The v e n t r a l bundles p r o v i d e v a s c u l a r t r a c e s t o t h e o v u l e s and,  a l o n g w i t h the d o r s a l s , e x t e n d up the s t y l e t o the s t i g m a .  Each t r a c e  d i v e r g i n g i n a s e p a l p l a n e t y p i c a l l y s u p p l i e s one o r more c a r p e l - w a l l b u n d l e s , a median s e p a l b u n d l e , and a stamen b u n d l e . trace usually provides  Each p e t a l p l a n e  one o r more c a r p e l - w a l l b u n d l e s , a l a t e r a l  trace  t o each a d j a c e n t s e p a l , a p e t a l b u n d l e and, i n f l o w e r s w i t h t e n stamens, a stamen b u n d l e .  D o r s a l c a r p e l bundles a r e u s u a l l y r e c o g n i z e a b l e and  82 may  o r i g i n a t e from t r a c e s i n e i t h e r p e r i a n t h A p a r t i a l summary of B e n s e l  plane",  and F a l s e r ' s (1975) a n a t o m i c a l  ob-  s e r v a t i o n s on herbaceous t a x a w i t h a x i l e p l a c e n t a t i o n ( t h e group of  inter-  e s t here) i s g i v e n i n T a b l e IX.  Boykin-  D i f f e r e n c e s between the s p e c i e s of  ia,, P e l t o b o y k i n i a and S u l l i v a n t i a s t u d i e d i n v o l v e m a i n l y the v a s c u l a r i z a t i o n of the c a r p e l s .  I n v e s t i g a t i o n of o t h e r s p e c i e s i n these genera and  S u k s d o r f i a and B o l a n d r a may  in  r e v e a l c o n s i s t e n t taxonomic d i f f e r e n c e s of  value i n generic d e l i m i t a t i o n i Embryology According Saxifragoideae late.  to D a v i s (1966) the u s u a l o v u l e c o n d i t i o n i n E n g l e r ' s  ( i n c l u d i n g R i b e s ) i s a n a t r o p o u s , b i t e g m i c and c r a s s i n u c e l -  P e l t i p h y l l u m and S a x i f r a g a s e c t i o n M i c r a n t h e s ,  mic o v u l e s .  The  (1966) r e p o r t e d  however, have u n i t e g -  embryo sac i s u n i f o r m l y of the Polygonum type and the embryogeny as conforming to the Soland t y p e .  Davis While  t h i s i s t r u e o f P e l t i p h y l l u m (Lebegue 1952), most S a x i f r a g o i d e a e have the Caryophyllad  type of embryo development (Johansen 1950;  K a p l a n 1976).  The  sub-family  Lebegue  i s thus heterogeneous and  i n v e s t i g a t i o n of the genera a t hand may  1952;  embryological  prove u s e f u l i n t h e i r d e l i m i t a t i o n .  F u r t h e r v a r i a t i o n between the genera o c c u r s w i t h r e s p e c t endosperm f o r m a t i o n .  D a h l g r e n (1930) has  shown t h a t endosperm  to  formation  i s ab i n i t i o C e l l u l a r i n A s t i l b e (2 s p p . ) , Heuchera (2 s p p . ) , P e l t i p h y l l u m , T e l l i m a . M i t e l l a p e n t a n d r a and B o y k i n i a o c c i d e n t a l i s ; and H e l o b i a l i n C h r y s o s p l e n i u m (2 s p p . ) , S a x i f r a g a (6 spp.,  that i t i s  1 h y b r i d and  d o u b t f u l sp.) , M i t e l l a d i p h y l l a and P e l t o b o y k i n i a t e l l i m o i d e s . (1976) a l s o r e p o r t e d H e l o b i a l endosperm i n S a x i f r a g a . are s p a r s e ,  k i n i a from B o y k i n i a .  Kaplan  A l t h o u g h the  they p r o v i d e f u r t h e r s u p p o r t f o r the s e p a r a t i o n of  1  data  Peltoboy-  A l t h o u g h the genus M i t e l l a i s a l s o heterogeneous i n  endosperm t y p e , t h i s i s not s u f f i c i e n t e v i d e n c e by i t s e l f f o r the  recog-  TABLE I X .  P a r t i a l summary o f f l o r a l anatomy i n S  i f r a g i n a e s p e c i e s w i t h a x i l e p l a c e n t a t i o n , (from B e n s e l  and F a l s e r (1975)) .  tn •rl  cd  rH cd u e cu x)  •H -A fl  •rl  A;  <->  oO  o w  Anatomical Feature* No, v a s c u l a r b u n d l e s i n p e d i c e l  cd  CO  •A x i  C •H o f3 •H  •r( O 43  o •H cu 4-1  cd  0)  rH  rH CU 4-1  S-t fl cd  XI  Cd - r l  •A  rH  O fl M-l  4-1  Cd  'A  > fl  •rl  rH rH  CU U  cd u • r l cu Pi 4-1 CU cn eu cu  o  rC  3  many small  2-3  1  cont cyl  1  No, compound p e r i p h e r a l b u n d l e s  9-10  9-10  8-12  8-10  No. b u n d l e s a t base o f f r e e  3,5  3  1-3  s  s+p  (inner  e„ e P)  s rH rH !>,  4-1 Cd  .fl  4J  rH Pu  P. CU •rt 4J  rH CU Pu  •H rH cd H cd. •Xi  60  cd u M-l •rj  O  fl  o  S-t  4=  cd cn  CD •rl CO  C CU •H C  •H  60  U •rl"  !>  •  in  cd •rl rH  o  <4-4 Cd  vi  •A X I C cu S-t CU  60  U  oO  pq  cd  •A rH O  4-1  'rl cn cn cd S-4  CJ  *  pq  c/i  9-11  Y"entral v a s c u l a r s u p p l y r i n g of bundles)  cd  cont cyl 1  4-6  ^ cont cyl  5-6  1  3  10-12  9-10  3-  cont cyl  cont cyl  8-10  8-10  8-10  10  9-10  3  3  3  2-3  3  3  s  s+p  s+p  s+p  s+p  s+p  s+p  8-20  4-5  6-10  16-20  3-11  5-8  10-15  9-10  -  -  +  +  +..  +  +  V e n t r a l b u n d l e arrangement  same cpd  adj cpd  adj cmp  adj cmp  simp  adj cmp  Level of separation v e n t r a l bundles  mid ovary  O r i g i n o f stamen bundles T o t a l No.  bundles/carpel  D o r s a l bundle d i s t i n c t  of cmp.  V e n t r a l bundle branch i n carpel w a l l No. i n t e r m e d i a t e bundles/carpel  carpel-wall  sepal  13-15  dist h of ov  +  +  11-13  18-20  dist h of ov  top o f loc  -  dist h of ov  simp  3-7  13-17  simp  + 0-9  + simp  -  + 1-2  -  2-5  8-13  + 6-7  TABLE I X , c o n t ' d .  to  •rl rH ca 4-1 C ca •H  iH  s oO •H o  Anatomical  Feature*  Q  pof  No, i n t e r m e d i a t e c a r p e l - w a l l b u n d l e s i n s t y l e base *  a d j .= a d j a c e n t cont c y l = c o n t i n u o u s cpd = compound  cylinder  1-11  ca  CO cu  C •H  "r-C O  >» O ,0 O 4J r-l  Ti rH -H CJ  4->  ca ca Ti 4-1  fi - K  ca  CD  CM.  4-1  > fi  •rl rH rH  CJ  M  3  u  ca -i-c iH O  fi ca ca t-i  u  a)  fi 4J o cn ,e CJ  &.  CJ  0  3 3g 4->  ca rfi 4-> PH r H Ti CJ 4-1 O, rH CJ  CU  C/3  16-20  e rH rH >,  0  9-15  P = p e t a l plane  dist = distal  s = s e p a l plane  ov = ovary  simp = simple  loc = locule  m Ti rH CS •rl U X oo o ca u o 4-4 5-1  xi  TI  K  fi  .pi CO fi Q) i-l fi Ti oo  u  fi  >  0  ca Ti -ri TJ u CJ O  60 o u  CJ  •  M  0  4-4  i-l  CB  W  ca -ni rH O  M  ca -rl rH O 4-|  iH co co ca U  a •  pq  5-9  4-7  85  n i t i o n of s e g r e g a t e s .  I t may,  however, l e a d t o a n a t u r a l i n f r a g e n e r i c  taxonomy. Taxonomic  Inferences As n o t e d by B e n s e l and P a l s e r (1975) f l o r a l anatomy and morph-  ology  i n the S a x i f r a g i o d e a e  are r e l a t i v e l y homogeneous and  consequently  they are o f l i m i t e d a s s i s t a n c e i n the d e l i m i t a t i o n of genera. v a r i a t i o n e x i s t s i t appears to be t a x o n o m i c a l l y of s p e c i e s .  Although  u s e f u l o n l y a t the  level  However, a s p e c t s of f l o r a l morphology do c o n t r i b u t e t o  the  o v e r a l l f a c i e s of some g e n e r a , and i n these cases i t i s q u i t e v a l u a b l e . Thus the c h a r a c t e r i s t i c t u b u l a r - c a m p a n u l a t e f l o w e r s of B o l a n d r a w i t h unique s u b u l a t e  red p e t a l s and g l a b r o u s h y p a n t h i a s e t the genus  from i t s c l o s e r e l a t i v e s , i n c l u d i n g B o y k i n i a and S u k s d o r f i a .  apart  Similarly,  P e l t o b o y k i n i a i s d i s t i n c t i n i t s tubular-campanulate flowers w i t h u l a r , toothed p e t a l s .  their  gland-  I n t h i s c a s e , i t s H e l o b i a l r a t h e r than C e l l u l a r  endosperm a l s o adds f u r t h e r s u p p o r t f o r i t s r e c o g n i t i o n as a s e p a r a t e - g e n u s . The  r e m a i n i n g genera i n t h i s s t u d y , B o y k i n i a , S u k s d o r f i a , and  a l l show s i m i l a r v a r i a t i o n i n t h e i r f l o r a l morphology.  Sullivantia  Sullivantia  may  be d i s t i n g u i s h e d however, by i t s l a c k of s t y l a r t r a n s m i t t i n g t i s s u e . f e a t u r e would appear to be one  of the few r e l i a b l e g e n e r i c  a v a i l a b l e i n t h i s c l o s e l y - r e l a t e d group of genera.  This  characters  B o y k i n i a and  Suksdorfia  cannot be s e p a r a t e d by f l o r a l morphology, b o t h showing s i m i l a r t r e n d s f l o r a l v a r i a t i o n , v i z . l a r g e f l o w e r s w i t h prominent n e c t a r i e s to  in  small  f l o w e r s w i t h no n e c t a r i e s ; h a l f s u p e r i o r to n e a r l y i n f e r i o r o v a r i e s ; f r e e v s . u n i t e d s t y l e s ; awned v s . unawned a n t h e r s ;  polymorphism f o r t o o t h e d  e n t i r e p e t a l s ; c r i m s o n or p i n k to w h i t e p e t a l c o l o u r ; stamens w i t h f i l a m e n t s to those w i t h s h o r t f i l a m e n t s .  long  Some of t h i s p a r a l l e l v a r i a t i o n  p r o b a b l y can be a t t r i b u t e d to changes i n the b r e e d i n g system from o u t B r e e d i n g to  inbreeding.  vs.  86  T e r a t o l o g i c a l Phenomena A b o r t i o n o f p a r t o r a l l o f the androecium was observed i n growth-room p o p u l a t i o n s some f l o w e r s  o f B o y k i n i a o c c i d e n t a l i s and B_. major so t h a t  (but n o t a l l ) on a p l a n t were e f f e c t i v e l y male s t e r i l e .  A b e r r a n t v a r i a t i o n i n p e t a l , stamen and c a r p e l number, u s u a l l y an i n c r e a s e of one, was observed o c c a s i o n a l l y i n a l l B o y k i n i a s p e c i e s and much more commonly i n S u k s d o r f i a . N u t t a l l (1834) n o t e d the tendency o f B o y k i n i a a c o n i t i f o l i a t o produce t h r e e r a t h e r than two c a r p e l s i n some f l o w e r s . S u k s d o r f i a e s p e c i a l l y showed weak c o n t r o l o v e r t h e numbers o f i t s f l o r a l p a r t s , f l o w e r s w i t h 4 t o 7 p e t a l s and 2 t o 4 c a r p e l s b e i n g n o t uncommon. I n some f l o w e r s o f S_. v i o l a c e a  one p a i r o f p e t a l s a r e i m b r i c a t e  the o t h e r s a r e s e p a r a t e , b e s t o w i n g a degree o f zygomorphism.  while  The p o s s i b l e  e f f e c t o f t h i s on p o l l i n a t o r fauna i s obscure s i n c e f i e l d  observations  i n d i c a t e d t h a t i n s e c t v i s i t o r s were e q u a l l y r a r e on b o t h  actinomorphic  and zygomorphic f l o w e r s .  On one o c c a s i o n , an i n d i v i d u a l o f B o y k i n i a  o c c i d e n t a l i s from Oregon ( G o r n a l l 256) developed one o r two s m a l l , y e l l o w orange s p o t s on the p e t a l s o f some o f i t s f l o w e r s , n o t u n l i k e those found i n some S a x i f r a g a s p e c i e s . these s p o t s .  S u b s e q u e n t l y , f l o w e r s on t h i s p l a n t  lacked  87  VTI>  The p o l l e n morphology  POLLEN  of the S a x i f r a g a c e a e has Been examined  many times (AgaBaByan 1961; WakaBayashi and Ferguson 1976).  1970; P a s t r e and Pons 1973; H i d e u x  The p r e s e n t s t u d y o f B o y k i n i a , B o l a n d r a and S u k s d o r f i a  was- u n d e r t a k e n t o complement the B r o a d e r s u r v e y conducted By H i d e u x and Ferguson (1976) who  examined B o y k i n i a a c o n i t i f o l i a . B. o c c i d e n t a l i s , B  m a j o r , B. l y c o c t o n i f o l i a , B. r i c h a r d s o n i i . P e l t o B o v k i n i a  iT  tellimoides.  S u k s d o r f i a r a n u n c u l i f o l i a , S. a l c h e m i l l o i d e s , B o l a n d r a oregana and  Sullivan-  t i a o h i o n i s . as w e l l as r e p r e s e n t a t i v e s o f a l l o t h e r genera i n the S a x i f r a g inae.  These w o r k e r s oBserved v a r i a t i o n i n tectum and a p e r t u r e s t r u c t u r e s  w h i c h has p o t e n t i a l taxonomic s i g n i f i c a n c e . M a t e r i a l s and Methods D e t a i l s o f specimens s t u d i e d are g i v e n i n TaBle X. examined By B o t h l i g h t and s c a n n i n g e l e c t r o n m i c r o s c o p y . scopy, the p o l l e n was Hbyer's Medium.  a c e t o l y s e d f o l l o w i n g Erdtman  Pollen  was  For l i g h t m i c r o -  (1960) and mounted i n  F o r e l e c t r o n m i c r o s c o p y , d r y a n t h e r s were soaked i n d i s -  t i l l e d w a t e r w i t h a w e t t i n g agent f o r 48 hours and then t r e a t e d w i t h a 1;1 m i x t u r e o f 2% OsO^J.lM phosphate b u f f e r , pH 7.2, f o r one h o u r .  They  were d r i e d down through an e t h a n o l s e r i e s and i n f i l t r a t e d w i t h amyl a c e t a t e p r i o r t o c r i t i c a l p o i n t d r y i n g w i t h C02«  A f t e r d r y i n g , they were mounted  on s t u b s and the p o l l e n g r a i n s d i s s e c t e d o u t .  P o l l e n was g o l d - c o a t e d and  viewed on a H i t a c h S-500 s c a n n i n g e l e c t r o n m i c r o s c i p e .  A l l s i z e measure-  ments were made w i t h a l i g h t m i c r o s c o p e on a i r - d r i e d g r a i n s mounted i n Hoyer's Medium.  M i c r o s p o r o g e n e s l s was s t u d i e d a c c o r d i n g t o the method  o u t l i n e d i n Chapter X, Microsporogenesls I n t h o s e B o y k i n i a and S u k s d o r f j a s p e c i e s examined  (B. O c c i d e n t -  88  TABLE X.  M a t e r i a l used i n the p o l l e n s t u d y .  g i v e n i n A p p e n d i x 1. Boykinia  F u l l c o l l e c t i o n d e t a i l s are  C o l l e c t i o n s b y G o r n a l l a r e p r e f i x e d b y 'G'.  aconitifolia: G342; H a r v i l l & Segars 25; i b i d .48; "Massey 3522; R a d f o r d 449.52; Shanks e t a l . 4348.  B. i n t e r m e d i a :  G23; O t i s 1317; W i g g i n s 9453.  B:, l y c o c t o n i f o l i a : N a g a i s.n.; Ohba e t a l . 73099; Sato s.n. ;R. m a j o r :  D e t l i n g 3469; E v e r e t t & B a l l s 22043; G45; G337; H i t c h c o c k & M u h l i c k 1539.1; P a r k s 24191; W i l s o n 116.  B_, o c c i d e n t a l i s :  G86; G256; Howe s.n.; Meyer 1000; S z c z a w i n s k i s.n.  B,, r o t u n d i f o l i a :  G98; G105; P a r i s h 7147; Sprout s.n.; T h o m e e t a l . 40871.  B,, h e u c h e r i f o r m i s : Clokey 7963; G198; H i t c h c o c k 18016; M a c g u i r e 14046; M c C a l l a 7084; T a y l o r & G i l l e t t 4726. B.. j a m e s i i ;  G i l l e t t & Mosquin 12114; Jones 955; Payson 1564.  B:, r i c h a r d s o n i i ( A l a s k a Range):' Anderson & Brown 10120; G306; Greene 333; P o r s i l d & P o r s i l d 1367; Spetzman 5038. B.. r i c h a r d s o n i i (Brooks Range): Argus & Chunys 5195; P a c k e r 2099; Parmelee 2819; Spetzman 689; Spetzman 2739. Suksdorfia  alchemilloides: Sleumer 3533.  F i e b r i g 3161; V e n t u r i  3296; S p a r r e 9613;  S\ r a n u n c u l i f o l i a : Beamish & Vrugtman 60358; Beamish & Vrugtman 60827; Bohm 1092; Bohm 1096; G15; G245; G253; H a i n a u l t 8008; L u i t j e n s & Campbell 8; Nagy & B l a a s 943a. S?„ v i o l a c e a ;  Bohm 1112; Bohm 1118; Bohm 1122; C a l d e r & S a v i l e 11306; C".. :der & S a v i l e 8732; C a l d e r & S a v i l e 7682; Eastham 32; G248; G252; M c C a l l a 6535; T a y l o r s.n.  Bolandra c a l i f o r n i c a : B. oregana:  S h a r s m i t h 3676.  G340; N e l s o n 2552; Ownbey & Keck 3153; Ownbey & Ownbey 2752.  Peltoboykinia tellimoides s s p . t e l l i m o i d e s : G328; I t o s.n.; Maekawa s.n. ssp. w a t a n a b e i ; alls.  G343; Yamazaki s.n.; Shimozono s.n.  B. r o t u n d i f o l i a , B. m a j o r , B. h e u c h e r i f o r m i s ,  B. r i c h a r d s o n i i , S.  r a n u n c u l i f o l i a and S. v i o l a c e a ) , c y t o k i n e s i s o f t h e p o l l e n mother c e l l s i s simultaneous.  I n species  o f a l l genera s t u d i e d , t h e p o l l e n g r a i n s a r e b i -  n u c l e a t e and a r e d i s s e m i n a t e d as monads. Shape and S i z e P o l l e n g r a i n s a r e i s o p o l a r w i t h a r a d i a l o r d e r t h r e e symmetry  89  throughout the S a x i f r a g i n a e , i n c l u d i n g the genera s t u d i e d h e r e ,  In polar  view the g r a i n s of the s p e c i e s v a r y somewhat i n the amount of i n f l u e n c e the a p e r t u r e s have on p o l l e n shape. and  G e n e r a l l y the a p e r t u r a l a n g l e i s obtuse  the i n t e r - a p e r t u r a l zone convex, a l t h o u g h  such t h a t some p o p u l a t i o n s  infraspecific variation exists  of S-uksdorfia r a n u n c u l i f o l i a , S. y i o l a c e a ,  B o l a n d r a oregana and B o y k i n i a r i c h a r d s o n l i may w h i c h i s more l p b a t e than convex. depending on the t r e a t m e n t .  have an i n t e r - a p e r t u r a l zone  I n e q u a t o r i a l v i e w , p o l l e n shape v a r i e s  Thus, when u n t r e a t e d  g r a i n s are mounted i n  E o y e r ' s Medium they appear o b l a t e s p h e r o i d a l i n a l l s p e c i e s .  Acetolysed  g r a i n s mounted i n Hoyer's Medium however, v a r y from s p h e r o i d a l to subprol a t e , the shapes r e p o r t e d by I k u s e (1956) and H i d e u x and Ferguson I n m a t e r i a l prepared f o r e l e c t r o n microscopy,.Peltoboykinia,  (1976).  Bolandra  and  a l l B o y k i n i a s p e c i e s save one, have p r o l a t e g r a i n s i n e q u a t o r i a l v i e w ; The  e x c e p t i o n i s B o y k i n i a l y c o c t o n i f o l i a w h i c h , l i k e S u k s d o r f i a . has  oidal grains. relation  P o l l e n s i z e s are shown i n T a b l e X I .  w i t h chromosome number.  spher-  There i s no c l e a r c o r -  A l t h o u g h p o l l e n of the 1 4 - p l o i d cytodeme  of B o y k i n i a r i c h a r d s o n l i has a s i g n i f i c a n t l y l a r g e r mean e q u a t o r i a l d i a m e t e r than t h a t of the h e x a p l o i d  cytodeme, the range of v a r i a t i o n o b s c u r e s  any  useful distinction. Apertures I n t h i s and the n e x t s e c t i o n , s e l e c t e d r e s u l t s from H i d e u x and Ferguson (1976) w i l l be i n c o r p o r a t e d s i n c e these have y e t t o be i n t e r p r e t e d i n terms of g e n e r i c r e l a t i o n s h i p s w i t h i n the s m a l l group a t hand. The p o l l e n g r a i n s of a l l s p e c i e s i n S u k s d o r f i a , B o l a n d r a B o y k i n i a s e c t i o n B o y k i n i a are 3 - c o l p o r a t e .  The  r e p o r t by I k u s e (1956) of  colporoidate p o l l e n i n B o y k i n i a l y c o c t o n i f o l i a i s erroneous. s p e c i e s have 3 - c o l p o r a t e each colpus  and  Sullivantia  p o l l e n but with, two or t h r e e orae a s s o c i a t e d w i t h  (Agababyan 19.61; H l d e n x and Ferguson 1976).  Species  of  90  TABLE X I .  S i z e s o f a i r ^ d r i e d p o l l e n g r a i n s mounted i n Rover's Medium.  Ten  r e p r e s e n t a t i v e g r a i n s p e r c o l l e c t i o n were measured, r e s u l t s B e i n g e x p r e s s e d as the mean + one s t a n d a r d  deviation.  Taxon  N  Boykinia aconitifolia intermedia lycoctonifolia major occidentalis rotundifolia richardsonli* A l a s k a Range Brooks Range heucheriformis jamesii Peltoboykinia tellimoides ssp. t e l l i m o i d e s s s p . watanabei Suksdorfia alchemilloides ranunculifolia violacea Bolandra californica oregana  E yum  P /<m #  50 30 30 50 50 50  14.8 17.2 18.0 17.9 14.9 16.1  1.3 1.0 0.7 1.5 0.8 1.0  16.2 18.4 19.2 20.2 16.5 18.1  ± + + + + +  1.3 1.2 1.2 •1.5 0.9 0.9  50 50 50 20  27.0 + 1.3 27.4 + 1.2 21.0 + 1.0 20.3 ± 1.6  29.0 30.9 24.2 22.8  + + + +  2.0 1.5 1.0 1.4  30 30  22.7 + 1.2 22.0 ± 1.0  23.9 + 1.3 24.4 + 1.1  20 100 100  17.1 + 1.2 16.9 + 1.2 16.6 +'1.2  14.5 + 1.1 17.1 + 1.5 18.0 + 1.3  10 30  23.4 + 1.6 23.0 + 1 . 8  26.8 + 1 . 5 25.4 + 2.1  ± + + + + +  P =. p o l a r d i a m e t e r ; E = e q u a t o r i a l d i a m e t e r . There i s no s i g n i f i c a n t d i f f e r e n c e i n mean p o l a r d i a m e t e r between c o l l e c t i o n s from the A l a s k a Range (2n = 36) and those from t h e Brooks Range ( 2 n = 8 4 ) : F = 2.56, P > 5 % . x, y o Q  Mean e q u a t o r i a l d i a m e t e r i s F  l f  9  8  = 28.88,  s i g n i f i c a n t l y l a r g e r i n p l a n t s from t h e Brooks Range P < 1%. P e l t o b o y k i n i a and B o y k i n i a s e c t i o n s T e l e s o n i x and R e n i f o l i u m have broidate p o l l e n .  3-colpor-  H i d e u x and Ferguson (1976) r e p o r t e d t h a t i n a l l the gen-  e r a c o n s i d e r e d h e r e , the endexine i s t h i c k e n e d i n the a r e a around each colpus.  91  Tectum A l l genera have a homogeneous tectum but i t s s t r u c t u r e v a r i e s between the s p e c i e s . and 32.  Scanning e l e c t r o n m i c r o g r a p h s are shown i n F i g s . 31  I n a l l t h r e e S u k s d o r f i a s p e c i e s , B o l a n d r a oregana and  richardsonli  ("Fig. 3 2 ) , the tectum i s p a r t i a l l y c o n t i n u o u s and  reticulate.  The  coarsely  luminae v a r y i n shape and s i z e , a measure o f the l a t t e r  b e i n g the r a t i o of m u r i and l u m i n a w i d t h s . than one  Boykinia  This r a t i o  i s usually less  ( i . e . l a r g e luminae) at the e q u a t o r but i n c r e a s e s at the  where the tectum becomes p e r f o r a t e r a t h e r than r e t i c u l a t e . luminae s c u l p t u r a l elements may  be seen.  The  columellae  poles  Within  the  i n Suksdorfia  a l c h e m l l l o i d e s are l o n g e r , r e l a t i v e t o tectum t h i c k n e s s , than they are i n the o t h e r s p e c i e s  (Hideux and Ferguson 1976).  I n B o y k i n i a s e c t i o n B o y k i n i a ( F i g . 31) , B_. a c o n i t i f o l i a . i n t e r m e d i a , B. l y c o c t o n i f o l i a and B_. o c c i d e n t a l i s are a l l v e r y w i t h a densely  luminae w i d t h s i s about one, but as b e f o r e  increase i n this ratio,  towards the p o l e s .  from the p r e c e d i n g  s i g n s of b e i n g d e n s e l y occluded  similar,  r e t i c u l a t e , p a r t i a l l y c o n t i n u o u s ( f o v e o l a t e ) tectum.  r a t i o of m u r i and  different  B.  t h e r e i s an  Boykinia rotundifolia i s  species i n t h a t , although  reticulate  i t s tectum shows  to p e r f o r a t e , most of the luminae are  such t h a t the tectum i n c o n t i n u o u s over much of the g r a i n .  k i n i a ma.jor i s i n t e r m e d i a t e  The  i n t h i s r e s p e c t between B_. r o t u n d i f o l i a  Boyand  the r e s t of s e c t i o n B o y k i n i a . I n B o y k i n i a s e c t i o n T e l e s o n i x , B_. h e u c h e r i f o r m i s  has  pollen  v e r y s i m i l a r to those s p e c i e s of s e c t i o n B o y k i n i a w i t h a f o v e o l a t e tectum. There i s some i n f r a s p e c i f i c v a r i a t i o n ( F i g . 32g)  i n t h a t the sample from A l b e r t a  had a t y p i c a l e q u a t o r i a l m u r l / l u m i n a w i d t h r a t i o of l e s s than  one, whereas i n the sample from Montana ( F i g s , than one."  32h,  i ) t h i s r a t i o was  more  B o y k i n i a .Iamesji has p o l l e n s i m i l a r t o t h a t of Bj_ r o t u n d i f o l i a .  92  FIGURE 31. Scanning e l e c t r o n m i c r o g r a p h s o f p o l l e n g r a i n s o f s p e c i e s of B o y k i n i a s e c t i o n B o y k i n i a . a - c) B_. a c o n i t i f o l i a , G o r n a l l 342. d - f ) _B_. l y c o c t o n i f o l i a , g - i ) B_. i n t e r m e d i a ,  Sato s.n.  G o r n a l l 23.  j - 1) B_i major, G o r n a l l 337. m - o.) B_. r o t u n d i f o l i a ,  m: G o r n a l l 98; n,o: G o r n a l l 105.  p - r ) B_. o c c i d e n t a l i s , G o r n a l l 86. S c a l e b a r = 5yum i n a,b,d,e,g,h,j,k,m,n,p,q. = 1^m  in c,f,i,l,o,r.  93  94  :  FIGURE 32. Scanning electron micrographs of pollen grains of species of Boykinia sections Renifolium and Telesonix, and of Peltoboykinia, Bolandra and  Suksdorfia.  a - c) Boykinia r i c h a r d s o n i i . Gornall  306.  d - f) B_. jamesii, G i l l e t t & Mosquiri 12114. g - i ) B_. heucheriformis,  g: Gornall 344; h , i : Gornall  j - 1) J?_. tellimoides ssp. watanabei, Gornall m - o) Bolandra oregana, Gornall p - r) _S_. violacea, Bohm  198.  343.  340.  1122.  s - u) S_. ranunculif o l i a , s: Gornall 245;  t,u: Nagy & Blaas 943a.  v) JS_. alchemjlloides. (from Hideux and Ferguson (1976) who Sleumer 3533 and Sparre 9613). Scale bar = 5 yum i n a,b,d,e,g,h,j,k,m,n,p,s. = 1 yum i n c , f , i , l , o , q , r , t , u , v .  cited  96  i n t h a t t h e luminae o f i t s d e n s e l y the tectum i s  r e t i c u l a t e tectum a r e o c c l u d e d  so t h a t  continuous.  I n both subspecies  o f P e l t o b o y k i n i a t e l l i m o i d e s t h e tectum i s  p a r t i a l l y c o n t i n u o u s and p e r f o r a t e , s p a r i n g l y so i n s s p . w a t a n a b e i . muri/lumina width  The  r a t i o i s v e r y l a r g e , i . e . t h e p e r f o r a t i o n s a r e minute.  R e g a r d i n g e x i n e t h i c k n e s s , H i d e u x and Ferguson (1976) r e p o r t e d v a l u e s o f about one m i c r o n i n a l l the genera s t u d i e d h e r e , a l t h o u g h  i t can  Be as much as two m i c r o n s i n S u k s d o r f i a a l c h e m i l l o i d e s . Taxonomic  Inferences A l t h o u g h many o f the s p e c i e s i n the p r e s e n t  study, e s p e c i a l l y  those o f B o y k i n i a s e c t i o n T e l e s o n i x , have been c o n s i d e r e d  as b e l o n g i n g t o  S a x i f r a g a , p o l l e n morphology p r o v i d e s s t r o n g s u p p o r t f o r t h e i r e x c l u s i o n . Thus a l l S a x i f r a g a s p e c i e s so f a r s t u d i e d have c o l p a t e g r a i n s , l a c k i n g e n d o a p e r t u r e s (Hideux and Ferguson 1976). endoapertures are uniformly  T h i s i s n o t t h e case h e r e , where  present.  S u l l i v a n t i a i s w e l l separated  from t h e o t h e r genera by i t s  p e c u l i a r a p e r t u r e s w i t h two o r t h r e e orae p e r c o l p u s . v a r i a t i o n w i t h b o t h c o l p o r a t e and c o l p o r o i d a t e p o l l e n .  B o y k i n i a shows some The l a t t e r  occurs  i n s e c t i o n s R e n i f o l i u m and T e l e s o n i x , s u p p o r t i n g t h e i r p o s i t i o n o u t s i d e the core o f the genus.  P e l t o b o y k i n i a a l s o has c o l p o r o i d a t e p o l l e n , a  c o n d i t i o n s h a r e d by Mukdenia, O r e s i t r o p h e ,  J e p s o n i a and P e l t i p h y l l u m , o f  the genera w i t h a x i l e p l a c e n t a t i o n (Hideux and Ferguson 1976) . p o l l e n i s uniformly present  Colporate  i n s e c t i o n B o y k i n i a and t h i s s u p p o r t s the  i n c l u s i o n o f Ji,. l y c o c t o n i f o l i a i n t h e genus.  Suksdorfia species are a l l  c o l p o r a t e and thus no s u p p o r t i s g i v e n t o t h e r e c o g n i t i o n o f s e g r e g a t e genera.  B o l a n d r a has c o l p o r a t e p o l l e n , s u p p o r t i n g i t s c l o s e a s s o c i a t i o n  w i t h S u k s d o r f i a and B o y k i n i a .  Of t h e S a x i f r a g i n a e w i t h a x i l e p l a c e n t a t i o n ,  colporate p o l l e n a l s o occurs i n Bergenia  and S a x j f r a g o d e s  ( H i d e u x and  97  Ferguson 1976). I n terms o f tectum s t r u c t u r e , S u k s d o r f i a i s a g a i n a homogeneous  group, and shows an a f f i n i t y w i t h B o l a n d r a and, more s u r p r i s i n g l y , w i t h  Boykinia rlchardsonji. perforate  tectum.  Peltoboykinia i s also very d i s t i n c t i v e with i t s  Within section Boykinia,  t h e i n t e r m e d i a c y o f B^ major  between B. r o t - u n d i f o l i a and t h e r e s t o f t h e s e c t i o n p r o v i d e s s u p p o r t f o r the h y p o t h e s i s t h a t one o f the genome donors of B. major was a B_. r o t u n d i f o i l a-like plant.  The s i m i l a r i t y o f B^ l y c o c t o n i f o l i a , B. i n t e r m e d i a  B. a c o n l t l f o l i a f u r t h e r s u p p o r t s the former's i n c l u s i o n i n B o y k i n i a .  and The  d i f f e r e n c e s i n tectum s t r u c t u r e between B_. j a m e s i i and B_. h e u c h e r i f o r a i l s s u p p o r t t h e i r r e c o g n i t i o n as s e p a r a t e  species.  98  VIII. Engler  (1891:, 1928)  SEEDS and Rosendahl (1905) n o t e d the v a r i e t y of .  s i z e s and shapes of seeds i n the s u b - t r i b e S a x i f r a g i n a e , and made some p r e l i m i n a r y d e s c r i p t i o n s of the t e s t a s u r f a c e s i n d i c a t i n g t h a t some s p e c i e s were t u b e f c u l a t e and o t h e r s smooth. have confirmed K a p l a n 1976;  More r e c e n t s t u d i e s of the seed  the d i v e r s i t y i n t e s t a o r n a m e n t a t i o n ( C o n o l l y 1972,  K r a c h 1976;  M i l l e r and Thompson 1979).  Perhaps the  coat 1976;  greatest  V a r i a t i o n i s found i n the genus S a x i f r a g a where the seeds p o s s e s s o r l a c k f o l d s i n the t e s t a , and the s u r f a c e can be made up of s t r o n g l y b u l g i n g o u t e r w a l l s of seed coat c e l l s , or e l a b o r a t e d i n t o l o n g t u b e r c l e s or i n t o s h o r t , s h a r p l y - d e f i n e d p a p i l l a e . The may  be mono- or p l u r i p a p i l l o s e .  o u t e r w a l l s of the e p i d e r m a l c e l l s  C o n f i g u r a t i o n of the mature seed  f r e q u e n t l y d i f f e r s c o n s i s t e n t l y between s p e c i e s although 1976) .  coat  ( M i l l e r and Thompson 1979) ,  a c e r t a i n amount of i n f r a - s p e c i f i c v a r i a t i o n o c c u r s  (Conolly  1972,  Other genera a l s o show some i n t e r - s p e c i f i c v a r i a t i o n i n t h e i r t e s t a  o r n a m e n t a t i o n , thus i n L i t h o p h r a g m a . ( T a y l o r 1965), C h r y s o s p l e n i u m (Hara 1957)  and Heuchera (Rosendahl e t a l . 1936)  the s p e c i e s v a r y from  l a t e t o smooth, each p a t t e r n o f t e n c h a r a c t e r i z i n g a s p e c i e s or group.  tubercu-  species  D i f f e r e n c e s between genera seem t o l i e i n the d i s t r i b u t i o n  patterns  of the t u b e r c l e s , or o r i e n t a t i o n and shape of the t e s t a e p i d e r m a l c e l l s . I n the p r e s e n t  s t u d y , a s u r v e y of the morphology o f . t h e seeds  o f B o y k i n i a and r e l a t e d genera was variation.  made i n o r d e r t o document the range o f  C o l l e c t i o n s used i n the study are g i v e n i n T a b l e X I I .  Observations Seeds of the s p e c i e s s t u d i e d v a r y i n shape and s i z e , which a p p a r e n t l y the o v a r y .  features  can be i n f l u e n c e d somewhat by the degree of crowding i n  I n B o y k i n i a . P e l t o b o y k i n i a . S u l l i v a n t i a and S u k s d o r f i a  the  seeds range i n shape from n e a r l y s p h e r i c a l t o e l l i p s o i d ; i n B o l a n d r a they  99  TABLE X I I .  L i s t o f m a t e r i a l used i n t h e seed s t u d y .  Full collection  d e t a i l s and h e r b a r i u m l o c a t i o n s a r e g i v e n i n Appendix 1.  C o l l e c t i o n s by  G o r n a l l a r e p r e f i x e d By G'. !  Boykinia  aconitifolia;  B a r k s d a l e e t a l , 3172; C l o y d s.n.; K a r v i l l e t a l . 11; K r a i 35423; M c F a r l a n d s.n.  B. l y c o c t o n i d o l i a ; B. major:  K a n a i s.n.  G45; H o w e l l 983; P i p e r s.n.  B. o c c i d e n t a l i s :  G I ; G22; G86; G256; S t r a l e y 1753.  B. r o t u n d i f o l i a :  G98; G101; G105.  B. h e u c h e r i f o r m i a : B. r i c h a r d s o n l i : Suksdorfia  G115; G193; G344. G272; G302; Raup 12796.  ranunculifolia:  S. v i o l a c e a :  Bohm 1112; G248.  S, a l c h e m i l l o i d e s : Peltoboykinia ssp.  Bohm 1096; Bohm 1132.  F i e b r i g 3161.  tellimoides  tellimoides:  ssp, watanabei:  G343; I t o s.n.; Mizushima s.n. G328; Nippon S h i n y a k u B o t Gard.; Yamazaki s.n.  B o l a n d r a oregana:  G340; Henderson s.n.; P i p e r 5721.  S u l l i v a n t i a halmlcola: Jepsonia p a r r y l :  S o l t i s e t a l . 1029.  Cromwell 709; M u l r o y s.n.  P e l t i p h y l l u m peltatum:  G66; G67; G80.  are n a r r o w l y e l l i p s o i d , w h i l e  i n P e l t i p h y l l u m t h e y a r e somewhat r e c t a n g u -  l a r , and i n J e p s o n i a they v a r y from e l l i p s o i d t o n a r r o w l y t r i a n g u l a r . raphe i s prominent i n P e l t o b o y k i n i a  and S u k s d o r f i a  genera.  T a b l e X I I I shows t h e seed  s i z e s and shapes o f t h e d i f f e r e n t  species,  together with  b u t l e s s so i n t h e o t h e r  t e s t a c o l o u r and c o n f i g u r a t i o n .  A f e a t u r e most i m p o r t a n t i n s e p a r a t i n g groups i s the t e s t a s u r f a c e , been d i r e c t e d .  species  scanning electrom microscope.  and s p e c i e s  and i t i s t h i s t o which most a t t e n t i o n has  G o l d - c o a t e d seeds were examined u s i n g  V a r i a t i o n was seen.  a H i t a c h i S-500  I n a l l cases, very l i t t l e i n f r a - s p e c i f i c  Photomicrographs o f seeds o f t h e v a r i o u s  shown i n F i g s . 33 r- 35.  A  species are  TABLE XIII.  Seed colour, shape and s i z e , and tubercle length In Boykinia. Peltoboykinia. Suksdorfia,  Bolandra, Jepsonia. Peltiphyllum and S u l l i v a n t i a .  A l l measurements are i n microns and are expressed as the  mean + one standard deviation, with a sample size of 24.  Taxon Boykinia a c o n i t i f o l i a B. intermedia B. l y c o c t o n i f o l i a B. major B. occidentalis B. r o t u n d i f o l i a B. heucheriformis B. r i c h a r d s o n i i  Length (L) 524 552 520 643 567 514 1121 1493  + 57  + 51  + 34 + 56  + 48 + 41 + 213  ± •112 746 ± 60  S u l l i v a n t i a data are from Rosendahl (1927)..  Width (W)  Shape (L/W)  Tubercle Length  + + + + + + + +  31 33 15 33 47 34 94 99  1.69 + .22 .21 1.59 .16 1.93 1.84 + .16 1.52 + .21 1.72 + .22 2.08 + .34 2.10 ± .29  13 + 3 11 + 2 11 + 1 13 + 4 8 + 2 10 + 2 smooth 4 + .3  Black or Dk. Black Black or Dk. Black or Dk. Black Black Dk. Brown Dk. Brown  446 + 51  1.68 + .17  51 + 15  Dk. Brown  ±  45 + 18  Dk. Brown  311 350 271 351 378 301 543 720  ± ;  Testa Colour  Peltoboykinia tellimoides ssp. tellimoides ssp. watanabei  685 + 34  458 + 18  1.50  Suksdorfia alchemilloides S. r a n u n c u l i f o l i a S. violacea  429 + 27 771 + 100 462 + 60  250 + 10 343 + 41 234 + 40  1.72 + .12 2.24 + .42 1/95 + .15  10 + 2 4 + 2 9 + 2  Dk. Brown Dk. Brown Dk. Brown  Bolandra oregana  719 + 50  245 + 33  2.98 + • .43  8 + 2  Dk. Brown  Jepsonia p a r r y i  893 + 111  465 + 36  1.92 + .18  smooth  Brown  1090 + 120  1.76 + .25  21 + 3  Brown  1.64  smooth  Dk. Brown  1.67  smooth  Dk. Brown  1.46  smooth  Dk. Brown  1.51  smooth  Dk. Brown  1.38  smooth  Dk. Brown  Peltiphyllum peltatum Sullivantia sullivantii S. r e n i f o l i a S. hapemanii ' S. halmicola S. oregana  1900 + 240 590 (570-620) 600 (530-650) 700 (560-770) 860 (800-960)  360 (320-400) 360 (330-380) 480 (400-520) 570 (480-680)  650 (580-720)  470 (430-530)  .09  101  FIGURE 33. Scanning e l e c t r o n m i c r o g r a p h s o f seeds o f s p e c i e s o f Boykinia section  Boykinia.  a - c) B^ a c o n i t i f o l i a , C l o y d s.n. d - f ) J3_. l y c o c t o n i f o l i a , K a n a i s.n. g - 1) B_. i n t e r m e d i a ,  G o r n a l l 23.  j - 1) B_. maj o r , G o r n a l l 45. m - o) B^, r o t u n d i f o l i a ,  m,o: G o r n a l l 105; n: G o r n a l l 101.  p - r ) B_. o c c i d e n t a l i s , p: G o r n a l l 256; q: G o r n a l l 22; r : S t r a l e y 1753. S c a l e b a r = 50ya.ni i n a,b,d,e,g,h, j ,k,m,n,p,q. = 5y»t.m i n c , f , i , l , o , r .  102  103  FIGURE 34. Scanning e l e c t r o n micrographs o f seeds o f s p e c i e s o f B o y k i n i a s e c t i o n s R e n i f o l i u m and T e l e s o n i x , and o f S u k s d o r f i a and Bolandra. a - c) B o y k i n i a r i c h a r d s o n l i , G o r n a l l 302. d - f ) B_. h e u c h e r i f o r m i s ,  d , f : G o r n a l l 193; e: G o r n a l l 344.  g - i ) S_^ r a n u n c u l i f o l i a , Bohm 1132. j - 1) S_. v i o l a c e a , Bohm 1112. m - o) S_. a l c h e m j l l o i d e s , F i e b r i g 3161. p - r ) B o l a n d r a oregana, G o r n a l l 340. S c a l e b a r = 500yttm i n a,d. = 50yum i n b,e,g,h,j,k,m,n,p,q. = 5 urn i n c , f , i , l , o , r .  104  105  FIGURE 35. Scanning e l e c t r o n micrographs o f seeds o f s p e c i e s o f Peltoboykinia,  S u l l i v a n t i a . P e l t i p h y l l u m and  a-;- c) P e l t o b o y k i n i a  Jepsonia.  t e l l i m o i d e s s s p . t e l l i m o i d e s , Mizushima s.n.  d - f ) P^ t e l l i m o i d e s s s p . watanabei, G o r n a l l 343. g - i ) S± h a l m i c o l a , j - 1 )  S o l t i s & Hammond-Soltis 1029.  P e l t i p h y l l u m p e l t a t u m , G o r n a l l 80.  m - o) J \ p a r r y i , Cromwell  709.•  S c a l e b a r = 500juxa. i n a,d,g,j,m. = 50 jum i n b , c , e , f , h , i , k , l , n . = 5 yum i n o.  106  107  In  B o y k i n i a s e c t i o n B o y k i n i a ( F i g . 33)  l y u n i f o r m i n t h e i r t e s t a ornamentation u s u a l l y b l a c k , has no spaced  and  the seeds are  colour.  The  striking-  t e s t a , which i s  creases or f o l d s , and has v e r y prominent, e v e n l y -  t u b e r c l e s arranged  l o n g i t u d i n a l l y , more o r l e s s i n rows.  The  h e i g h t and s p a t i a l arrangement of these t u b e r c l e s show v e r y l i t t l e t i o n between the s p e c i e s .  The e p i d e r m a l  c e l l s are p o l y g o n a l and  w a l l of each forms the t u b e r c l e , one per c e l l . i s i n v o l v e d i n t h i s way,  varia-  the o u t e r  Much of the o u t e r c e l l w a l l  so t h a t o n l y the s m a l l a r e a around i t s edge i s  v i s i b l e a t the s u r f a c e , the c e n t r a l p o r t i o n b e i n g r a i s e d i n t o a d i s t i n c t dome. Boykinia richardsonii degree.  I t s dark brown t e s t a i s o f t e n creased o r f o l d e d but i s a l s o  covered i n t u b e r c l e s . trudes  ( F i g . 34) d i f f e r s but mostly o n l y i n  only a l i t t l e  They are s m a l l e r than i n s e c t i o n B o y k i n i a , and above the s u r r o u n d i n g s u r f a c e .  f a c e area of the o u t e r c e l l w a l l i s devoted  Moreover, l e s s s u r -  to t u b e r c l e f o r m a t i o n ,  hence the p o l y g o n a l c e l l p e r i m e t e r i s c l e a r l y  and  visible.  B o y k i n i a h e u c h e r i f o r m i s ( F i g . 34) d i f f e r s somewhat from previous taxa. of  are oblong r a t h e r than p o l y g o n a l and I t may  I t s two  uncreased  These e p i d e r m a l  the t e s t a s u r f a c e thus appears  finely  shows another k i n d of t e s t a ornament-  s u b s p e c i e s are v e r y s i m i l a r .  The  dark brown t e s t a i s  and most of i t s t u b e r c l e s are v e r y l o n g , s p i n e - l i k e  Over much o f the seed they are arranged  formed i n the same way  as b e f o r e except  t r a n s v e r s e l y oblong.  The  structures.  i n 14 - 20, h i g h l y d i s t i n c t ,  i t u d i n a l rows, w i t h a d i s t i n c t gap between each row.  are n a r r o w l y  cells  a l s o be r a t h e r c r e a s e d or f o l d e d .  P e l t o b o y k i n i a ( F i g . 35) ation.  the  I t l a c k s t u b e r c l e s of any k i n d but i n s t e a d the o u t e r w a l l s  the dark brown seed coat c e l l s bulge s l i g h t l y .  striate.  pro-  The  long-  t u b e r c l e s are  t h a t the component e p i d e r m a l  cells  outer c e l l w a l l s are d i s t e n d e d i n t o  108  t a l l spines face l e v e l .  i n t h e i r centres, T h e r e can Be  l e a v i n g l a r g e areas on e i t h e r s i d e at sur^-  some v a r i a t i o n i n t u b e r c l e l e n g t h and  ment w i t h i n i n d i v i d u a l seeds. are s h o r t e r and  The  a f f e c t e d a r e a has  for Boykinia  section  or f o l d e d , u s u a l l y w i t h three although sometimes there  The  i n rows and  Surface  seed b e i n g low is  can be more.  Tubercles  are arranged  a r i s e from i r r e g u l a r l y p o l y g o n a l  s i m i l a r to  creased  seed, longitud-  epidermal  cells.  than i n the o t h e r  v a r i a t i o n a l s o can i n v o l v e p a r t s of the t e s t a on  composed of an e n t i r e outer  two any  I n S_. r a n u n c u l i f o l i a the t u b e r c l e s  are  d i f f e r i n g i n t h a t each  c e l l w a l l , making the t u b e r c l e s more  I n S_. v i o l a c e a the' t u b e r c l e s are v a r i a b l e i n s i z e , even  on a s i n g l e seed, from low The  than  the dark brown t e s t a i s o f t e n  domes, s i m i l a r to those i n B_. r i c h a r d s o n l i but  ances.  rather  r i d g e s r u n n i n g the l e n g t h of the  more or l e s s smooth.  t i g h t l y packed.  reflects  an appearance v e r y  rows are more regimented i n S_. a l c h e m j l l o i d e s  species.  the l a t t e r  tubercles  Boykinia.  ( F i g . 34)  In Suksdorfia  inally  on a seed the  component c e l l s h e r e , which are p o l y g o n a l  t r a n s v e r s e l y oblong. that described  Thus i n some p l a c e s  they l o s e t h e i r regimented p a t t e r n ;  the shape of the  arrange^  1  taller  domes to much t a l l e r , p a p i l l a - l i k e  protruber-  t u b e r c l e s are o f t e n a s s o c i a t e d w i t h the s i d e s of  the  r i d g e s , whereas the domes tend to occur on the seed f a c e between the The  o c c u r r e n c e of i n t e r m e d i a t e s  obscures t h i s p a t t e r n .  B o y k i n i a , much of the s u r f a c e o f each o u t e r of the low surface.  domes, l e a v i n g o n l y I n the  The  case of the t a l l e r  arrangement and  i n section  c e l l i s used i n the  formation  the area around the edges v i s i b l e on  w a l l of each c e l l i s used and v e r y ible.  As  t u b e r c l e s , however, the e n t i r e little  section  In B o l a n d r a o r e g a n a • ( F i g .  34)  the outer  of the seed s u r f a c e remains v i s -  s i z e of the t u b e r c l e s i n S.  s i m i l a r to the p a t t e r n i n B o y k i n i a  ridges.  alchemjlloides  are  Boykinia. the t u b e r c l e s are a l s o arranged  109  as i n B o y k i n i a s e c t i o n B o y k i n i a , although, t h e y are somewhat s m a l l e r and t h e i r p a r e n t c e l l s are more e l o n g a t e d . S u l l i v a n t i a ( F i g . 35) i s unique i n t h a t the dark brown t e s t a i s extended l a t e r a l l y to form "wings",  The e p i d e r m a l c e l l s are p o l y g o n a l  to t r a p e z o i d and t h e i r o u t e r w a l l s B u l g e o n l y s l i g h t l y .  The  overall  appearance i s thus one o f smoothness. J e p s o n i a ( F i g . 35) has a smooth mid-brown t e s t a w h i c h i s c r e a s e d , o f t e n w i t h about t h r e e prominent  ridges.  The e p i d e r m a l  are p o l y g o n a l w i t h o n l y s l i g h t l y b u l g i n g o u t e r w a l l s .  cells  Peltiphyllum (Fig.  35) i s somewhat s i m i l a r a l t h o u g h i t s mid-brown seeds are much l a r g e r . The seeds have t h r e e or f o u r r i d g e s and u s u a l l y have c h a r a c t e r i s t i c squared ends.  The e p i d e r m a l c e l l s are p o l y g o n a l and the o u t e r w a l l s b u l g e  v e r y s t r o n g l y to g i v e the seed a v e r r u c a t e . appearance. Taxonomic I n f e r e n c e s I n v i e w of the v a r i a t i o n found i n S a x i f r a g a , C h r y s o s p l e n i u m and L i t h o p h r a g m a , genera. grounds.  i t i s i m p o s s i b l e to use seed c h a r a c t e r s alone t o d e l i m i t  However, they can be used to s u p p o r t groupings made on o t h e r Thus B o y k i n i a s e c t i o n B o y k i n i a i s a homogeneous u n i t c h a r a c t e r -  i z e d by seeds o f f a i r l y u n i f o r m s i z e , shape and c o l o u r , w i t h uncreased seed c o a t s and a remarkably  constant t e s t a ornamentation.  Boykinia lycoc-  t o n i f o l i a thus f i t s w e l l i n the core s e c t i o n o f the genus and no i s g i v e n t o i t s r e c o g n i t i o n as N e o b o y k i n i a (Hara 1937).  support  Boykinia richard-  s o n l i d i f f e r s s l i g h t l y i n i t s l a r g e r , dark brown seeds w i t h c r e a s e d coats and much s m a l l e r t u b e r c l e s .  T h i s sequence i s c o n t i n u e d i n B. h e u c h e r i f o r m i s  which a l s o has l a r g e , dark brown, c r e a s e d seeds, but l a c k s any  tubercles.  The seed coat c e l l s are a l s o o b l o n g , r a t h e r t h a n p o l y g o n a l , as i n o t h e r Boykinia species.  These d i f f e r e n c e s s u p p o r t the e x c l u s i o n of these  s p e c i e s from s e c t i o n B o y k i n i a and t h e i r assignment  two  t o two s e p a r a t e s e c t i o n s ,  110  R e n i f o l i u m and T e l e s o n i x , A l t h o u g h B o y k i n i a i s heterogeneous i n i t s seed c h a r a c t e r s , the v a r i a t i o n can be understood i n terms of an e v o l u t i o n a r y t r e n d .  jMany o t h e r  genera i n the S a x i f r a g i n a e show the t r a n s i t i o n from smooth t o t u b e r c u l a t e seeds, e.g. Chrysosplenl-um, Llthophragma, Heuchera and S a x i f r a g a .  Hara  (1957) suggested t h a t i n Chrysosplenium the smooth c o n d i t i o n was p r i m i t i v e and  tubercles derived, a proposal  trends i n Heuchera (Wells 1977).  a l s o compatible  with other  I n B o y k i n i a there i s no reason  t h a t the d i r e c t i o n o f e v o l u t i o n has been d i f f e r e n t , indeed seeded B_j_ h e u c h e r j f o r m i s  evolutionary to suppose  the smooth-  r e t a i n s other p r i m i t i v e f e a t u r e s such as t e n s t a -  mens and c o l p o r o i d a t e p o l l e n . The  long, s p i n e - l i k e , highly-regimented  tubercles with  trans-  v e r s e l y oblong component c e l l s i n P e l t o b o y k i n i a h e l p d i s t i n g u i s h i t as a separate  genus.  However, the occurrence  on some seeds o f p o l y g o n a l  com-  ponent c e l l s w i t h much s h o r t e r t u b e r c l e s , i n d i c a t e s a p o s s i b l e r e l a t i o n ship with  Boykinia. The  helps  c h a r a c t e r i s t i c a l l y narrow e l l i p s o i d seed shape i n B o l a n d r a  d e f i n e i t as a genus, although  i t s t e s t a s u r f a c e p a t t e r n i s remark-  a b l y s i m i l a r to that i n B o y k i n i a s e c t i o n B o y k i n i a . Although there are some d i s t i n c t d i f f e r e n c e s i n t e s t a s u r f a c e pattern between the t h r e e s p e c i e s o f S u k s d o r f i a . they i n v o l v e m o s t l y c l e s i z e and i t may be t h a t the c o n d i t i o n s i n S u k s d o r f i a r e p r e s e n t stages above.  of d i f f e r e n t i a t i o n according  t o the e v o l u t i o n a r y t r e n d  I n the p a s t the t h r e e s p e c i e s have been a s s i g n e d  t y p i c genera.  tuber-  early  discussed  t o s e p a r a t e mono-  S i m i l a r i t i e s i n many r e s p e c t s tp the v a r i a t i o n i n t e s t a s u r -  face p a t t e r n i n B o y k i n i a are q u i t e marked, and i t i s l i k e l y  t h a t the two  genera r e p r e s e n t p a r a l l e l l i n e s o f e v o l u t i o n . Winged seeds w i t h smooth t e s t a s d i s t i n g u i s h S u l l i v a n t i a as a  Ill  d i s t i n c t genus.  I t might be n o t e d however, t h a t winged seeds- a l s o o c c u r  i n some S a x i f r a g a s p e c i e s , E n g l e r  (1928) has suggested t h a t t h e c o n d i t i o n  i s m e r e l y an extreme form o f t e s t a f o l d i n g , a phenomenon most r e a d i l y apparent i n S u k s d o r f i a among t h e genera s t u d i e d h e r e .  S a v i l e (1975) l i n k e d  the wings t o a chasmophilous h a b i t and a e r i a l d i s p e r s a l , s i n c e  Sullivantia  t y p i c a l l y grows on c l i f f s . The  verrucate  t e s t a i n P e l t i p h y l l u m , together w i t h i t s . very  l a r g e seeds, c o n t r i b u t e s f u r t h e r t o t h e d i s t i n g u i s h i n g c h a r a c t e r s o f the genus.  I t does n o t h i n g  t o suggest an a l l i a n c e w i t h P e l t o b o y k i n i a as has  been proposed i n t h e p a s t ( E n g l e r 1891).  J e p s o n i a has u n s p e c i a l i z e d smooth  seeds and i t s r e l a t i o n s h i p s a r e ambiguous i n t h i s  respect.  112  IX.  SEEDLINGS AND EARLY DEVELOPMENT  M a t e r i a l s and Methods Germination ing  species:  and e a r l y development were s t u d i e d i n t h e f o l l o w -  B o y k i n i a o c c i d e n t a l i s , B. i n t e r m e d i a . B. m a j o r , B. r o t u n d -  i f o l i a . B. r i c h a r d s o n i i , B. h e u c h e r i f o r m i s  and P e l t o b o y k i n i a t e l l i m o i d e s .  Seeds were germinated i n pots o f s o i l , o r on beds o f agar e n r i c h e d w i t h Hoagland's s o l u t i o n , a t 17°C under a 16hr p h o t o p e r i o d .  Germination  occur-  r e d t y p i c a l l y a f t e r about t h r e e weeks i n b o t h , i f f r e s h seed were used. The  germination  success  r a t e was 70 - 90%.  not a s i g n i f i c a n t f a c t o r i n g e r m i n a t i o n .  Light i s therefore  probably  A s i m i l a r c o n c l u s i o n was reached  by T a y l o r (1965) f o r L i t h o p h r a g m a , where temperature was found t o be a controlling variable. Observations Germination  i s e p i g e a l w i t h the r a d i c l e emerging f i r s t  by t h e h y p o c t y l and t h e two, r a r e l y t h r e e , c o t y l e d o n s . Is pigmented r e d w i t h a n t h o c y a n i n s ,  followed  The r a d i c l e t i p  a c o n d i t i o n found i n a l l a c t i v e l y  growing r o o t s , whether i n t h e l i g h t o r dark.  I n Boykinia sections Boykinia  and T e l e s o n i x and i n P e l t o b o y k i n i a t h e c o t y l e d o n s  a r e o v a t e , whereas i n  B. r i c h a r d s o n i i o f s e c t i o n R e n i f o l i u m they a r e n a r r o w l y  ovate.  They may  be up t o 3mm l o n g a t m a t u r i t y , t h e e n t i r e s e e d l i n g b e i n g about 3mm  tall.  I n B o y k i n i a t h e shoot apex produces the f i r s t e o p h y l l w h i c h i s e n t i r e i n t h e b a s a l r e g i o n b u t s e r r a t e t o v a r y i n g degrees toward i t s apex. T h i s i s q u i c k l y f o l l o w e d by a second e o p h y l l , a l t e r n a t e t o t h e f i r s t , and also s e r r a t e i n the a p i c a l region.  Subsequent f o l i a g e l e a v e s a r e produced  i n a 3/8 p h y l l o t a c t i c s p i r a l t o form a r o s e t t e from w h i c h t h e f l o r a l a x i s later arises.  I n s p e c i e s whose m e t a p h y l l s  are lobed o r otherwise d i v i d e d ,  s u c c e s s i v e l e a v e s show a sequence from t h e i n i t i a l l y e n t i r e o r p a r t i a l l y s e r r a t e c o n d i t i o n t o the t y p i c a l d i v i d e d c o n d i t i o n ( F i g s . 36, 3 7 ) .  113  FIGURE 36. S i x week o l d s e e d l i n g s o f a) B o y k i n i a o c c i d e n t a l i s ; i n t e r m e d i a ; c) B_. major; and d) B_. r o t u n d i f o l i a ;  ( a l l x2) .  b)  114  115  FIGURE 37.  S i x week o l d s e e d l i n g s of a) B o y k i n i a h e u c h e r i f o r m i s ;  richardsonli;  and  b) JB  c) P e l t o b o y k i n i a t e l l i m o i d e s ssp. watanabei; ( a l l x  116  117  In P e l t o b o y k i n i a  the e o p h y l l s  are not p e l t a t e , but show the  o r b i c u l a r l a m i n a w i t h b a s a l p e t i o l e t y p i c a l of a l l the o t h e r species  ( F i g . 37).  From the f o u r t h o r f i f t h  n a t u r e becomes i n c r e a s i n g l y apparent. o f t e n young, s p r i n g l e a v e s (Jepson 1936).  In P e l t o b o y k i n i a  the p e l t a t e  I t i s i n t e r e s t i n g that eophylls  and  syndrome  the f i r s t m e t a p h y l l s have l o b e s which are These l o b e s  s u b s e q u e n t l y undergo  l a t e r a l growth to r e a c h the t y p i c a l , b r o a d l y  Development o f l a t e r b a s a l l e a v e s Ch. I ' l l .  l e a f on however,  of P e l t i p h y l l u m a l s o l a c k the p e l t a t e  almost l a c i n i a t e i n t h e i r young s t a g e . extensive  Boykinia  triangular condition.  i n the two s u b s p e c i e s i s d i s c u s s e d i n  P h y l l o t a x i s and l e a f arrangement are as i n B o y k i n i a .  118  X.  CYTOLOGY  Chromosome numbers of some of the t a x a i n t h i s s t u d y have been previously reported these  counts.  I n the p r e s e n t  a geographical first  (Hamel 1953), although  c o n f u s i o n surrounds some of  study p o p u l a t i o n s were examined from as wide  range as p o s s i b l e .  Chromosome numbers are r e p o r t e d f o r the  time f o r B o y k i n i a i n t e r m e d i a , B. major, P e l t o b o y k i n i a t e l l i m o i d e s .  ssp, watanabei and S u k s d o r f i a r a n u n c u l i f o l i a ; and  counts are confirmed  o t h e r s p e c i e s i n B o y k i n i a , P e l t o b o y k i n i a and S u k s d o r f i a . t o n l f o l i a and S u k s d o r f i a a l c h e m i l l o i d e s were not S u l l i v a n t i a spp. have been counted by Mr.  for  Boykinia lycoc-  available for  study.  D. S o l t i s of I n d i a n a U n i v e r s i t y .  Methods Both m e i o t i c and m i t o t i c events were observed. buds were f i x e d i n 3:1 24 h o u r s .  For  meiosis,  a b s o l u t e a l c o h o l : g l a c i a l a c e t i c a c i d f o r at l e a s t  P o l l e n mother c e l l s were d i s s e c t e d out, s t a i n e d and  i n 1% acetocarmine ( D a r l i n g t o n and LaCour 1976) .  M i t o s i s was  squashed observed i n  root t i p s .  A c t i v e l y growing r o o t t i p s were e x c i s e d i n t o v i a l s of c o l d  water.  drop of eC-monobromonaphthalene was  was  One  then kept at 5°C  formation  and  f o r e i g h t hours.  T h i s pretreatment  c o n t r a c t e d the chromosomes.  i n water and h y d r o l y z e d  squashed i n 45%  The  inhibited  After rinsing  leuco-basic fuchsin  to t h r e e hours i n the dark at 20°C.  acetic acid.  spindle  r o o t t i p s were then r i n s e d  f o r ten minutes at 60°C i n IN HC1.  i n water a g a i n , the r o o t s were s t a i n e d w i t h S o l u t i o n ) f o r two  added to each v i a l which  (Feulgen's  Root t i p s were then  The method i s t h a t of D a r l i n g t o n and L a Cour  (1976). Drawings or photographs were made of s u i t a b l e p r e p a r a t i o n s . The  chromosomes of B o y k i n i a and S u k s d o r f i a are s m a l l , and  occurred  at m i t o t i c metaphase, making counts d i f f i c u l t  Hamel (1953) a l s o r e p o r t e d  this d i f f i c u l t y .  clumping commonly  to i n t e r p r e t .  However, at l e a s t t h r e e  cells  119  per c o l l e c t i o n were  counted.  Results I l l u s t r a t i o n s of the chromosomes of the v a r i o u s s p e c i e s are shown i n F i g s . 38 and 39, and a l l counts, b o t h p r e v i o u s l y and p r e s e n t l y made, are g i v e n i n TaBle XIV. Boykinia:  6 and 7.  There are two B a s i c chromosome numbers i n  Section Boykinia contains both, s e c t i o n Renifolium  has x = 6, and s e c t i o n T e l e s o n i x has x = 7.  The genera S u k s d o r f i a .  Bolandra  and S u l l i v a n t i a a l l have x = 7, and i n P e l t o b o y k i n i a x = 11. The c o n f u s i o n surrounding  the counts made by Hamel (1953) f o r  B o y k i n i a r o t u n d i f o l i a and B_. e l at a v a r . c i n c i n n a t a (B. o c c i d e n t a l i s ) r e s u l t s from the f a c t t h a t h i s drawings and t e x t a s s i g n e d  2n = 14 to B. r o -  t u n d i f o l i a and 2n = 26 to B_. e l a t a , w h i l e i n h i s summary t a b l e .these numbers were transposed. have a p p a r e n t l y  A l l subsequent works q u o t i n g  counts f o r B o y k i n i a  used the summary t a b l e as t h e i r source.  Thus Munz and  Keck (1959) r e p o r t e d 2n = 26 f o r B_. r o t u n d i f o l i a and 2n = 14 f o r B_j_ e l a t a . Hitchcock if  ironic,  e t a l . (1961) a l s o claimed n = 7 f o r Bj, e l a t a .  I t i s fortunate,  t h a t B_. o c c i d e n t a l i s ( e l a t a ) does indeed have 2n = 14 i n a l l  c o l l e c t i o n s counted.  B o y k i n i a r o t u n d i f o l i a however most d e f i n i t e l y i s n o t  2n = 2 6  (1953) t a b l e , but r a t h e r 2n = 1 4  as i n Hamel's  illustration. 26 count.  This leaves  I t seems l i k e l y  as i n h i s t e x t and  the problem of where Hamel (1953) got h i s 2n = t h a t he m i s i d e n t i f i e d B. major, whose  inflor-  escence i s o f t e n more c l e a r l y c i n c i n n a t e than i s t h a t of B^ o c c i d e n t a l i s , as B_. e l a t a v a r . c i n c i n n a t a . 2n  A l l populations  of B^ major counted have  =26. B o t h s p e c i e s of B o y k i n i a s e c t i o n T e l e s o n i x haye 2n R 14  mosomes.  chro-  The r e p o r t o f 2n = 28 f o r B_j. j a m e s j i (Spongberg 1972) i s an  erroneous c i t a t i o n of Hamel's  (1953) count o f 2n = 14 (Spongberg i n l i t t . ) .  120  FIGURE 38. Chromosomes of some species of Boykinia and Suksdorfia.  5,1  a> Boykinia o c c i d e n t a l i s (Bohm 1293X, p o l l e n mother c e l l (PMC), n = 7. b) B o y k i n i a intermedia (Gornall 23), root t i p c e l l , 2n = 14. c) B o y k i n i a r i c h a r d s o n i i ( G o r n a l l 271), PMC, n = 18 d) B o y k i n i a heucheriformis (Gornall 193), PMC, n = 1 e) Suksdorfia r a n u n c u l i f o l i a ( G o r n a l l 332), PMC, n = 7. f) Suksdorfia v i o l a c e a ( G o r n a l l 329), PMC, n = 7.  121  FIGURE 39. Somatic chromosomes o f some s p e c i e s o f B o y k i n i a and Peltoboykinia. A) B o y k i n i a r o t u n d i f o l i a  (G105); B) B o y k i n i a a c o n i t i f o l i a  (G342);  C). B o y k i n i a major (G45); D) P e l t o b o y k i n i a t e l l i m o i d e s s s p . watanabei (G343).  Scale bar = 5 microns.  122  123  TABLE XIV.  Chromosome numbers p r e s e n t l y and p r e v i o u s l y found i n t h e  genera B o y k i n i a , P e l t o b o y k i n i a , S u k s d o r f i a . B o l a n d r a and S u l l i v a n t i a .  :  BOYKINIA SECTION BOYKINIA ... X R 6, 7 B. ACONITIFOLIA ... 2N = 1 2 Hamel 0-953) ex R o y a l B o t a n i c Garden, E d i n b u t g h , 2n = 12; G o r n a l l 342, Chatooga R i v e r , Macon Co., N o r t h C a r o l i n a (UBC), 2n = 12. B„ INTERMEDIA ... 2N = 1 4 G o r n a l l 23, Humptulips, Grays Harbor Co., Washington (UBC), 2n = 14. B, OCCIDENTALIS ... 2N = 1 4 Hamel (1953) ex Copenhagen B o t a n i c Garden, 2n = 26*; R. L. T a y l o r (unpub.) T a y l o r 64C-17, ex Rancho Santa Ana B o t a n i c a l Garden, ia  (DAO), 2n = 14; R, L. T a y l o r  Californ-  (unpub.) T a y l o r 63C-176, ex B o t a n i c  Garden H a u n i e n s i s , Denmark (DAO), 2n = 14; G o r n a l l 1, C a p i l a n o  Canyon,  North Vancouver, B.C. (UBC), n = 7; G o r n a l l 216, Snoqualmie F a l l s ,  King  Co., Washington (UBC), 2n = 7 ^ ; Bohm 1293, Saddle Mt., C l a t s o p Co., Oregon (UBC), 2n = 7 ^ ; R. L. T a y l o r  (unpub.) T a y l o r & G l l l e t t  4658,  Glenwood,  T i l l a m o o k Co., Oregon (DAO), 2n = 14; T a y l o r  (unpub.)  Taylor  & Gillett  4669, Diamond Creek, Douglas Co., Oregon (DAO), 2n = 14;  S t r a l e y 1753, Gasquet, D e l N o r t e Co., C a l i f o r n i a (UBC), n = 7; G o r n a l l 65, Happy Camp,. S i s k i y o u C o . , . C a l i f o r n i a (UBC), 2n = 7 ^ ; R. L . T a y l o r (unpub.) T a y l o r , Baker e t a l . 4449, 46 mi SW o f Happy Camp, S i s k i y o u Co., C a l i f o r n i a (DAO), 2n = 14. B. ROTUNDIFOLIA ... 2N = 14 Hamel (1953) ex M. de V i l m o r i n , V e r r i e r e s - l e - B u i s s o n , 2n = 14; R. L. Taylor  (unpub.) T a y l o r 63C-175, ex U n i v e r s i t y o f U p p s a l a B o t a n i c Garden,  Sweden (DAO), 2n = 14; G o r n a l l 98, C r y s t a l Lake, Los Angeles Co., C a l i f o r n i a (UBC), 2n = 14; G o r n a l l 101, hwy 39 b r i d g e over N. Fork San G a b r i e l R., L o s Angeles C o . , , C a l i f o r n i a (UBC), 2n = 1 4 ; G o r n a l l 105, Icehouse Canyon, San B e r n a r d i n o Co., C a l i f o r n i a (UBC), 2n = 1 4 . B. MAJOR ... 2N R 26 G o r n a l l 335, 337, R a c k l i f f Creek by Selway R., Idaho Co., Idaho  (UBC),  2n = i ! ! ? G o r n a l l 202, L o l o Pass, M i s s o u l a Co., Montana (UBC), 2n = 3  26; G o r n a l l "38, upper reaches o f Rogue R., Douglas Co., Oregon  (UBC),  2n = 26; G o r n a l l 45, Rogue R., Jackson Co., Oregon (UBC), 2n = 26;  124  TABLE XIV,  cont'd.  B. MAJOR ... 2N =• 26 G o r n a l l 92, K y b u r z , E l d o r a d o  Co., C a l i f o r n i a  (UBC), 2n . = 26,  BOYKINIA SECTION RENIFOLIUM ... X = 6 B. RICHARDSONII ... 2N =. 36, 84 Knaben  (1968) G j a e r v o l l 1407, Rainbow Mt., A l a s k a Range, A l a s k a (TRH),  2n = 36; G o r n a l l 271, Mt. W r i g h t , M c K i n l e y N a t i o n a l Park, A l a s k a (UBC), 2n =. 1 8 ; G o r n a l l 302, m i l e 63,7 D e n a l i Highway, A l a s k a (UBC), 2n = I T  1 8  ir  Johnson & Packer (1968) Packer 2099, Ogoturuk Creek, n o r t h w e s t e r n  Alaska  (ALTA) , 2n = 84; Packer & McPherson (1974) McPherson 72-411, Meade River, Alaska BOYKINIA SECTION  (ALTA). 2n = c, 84. TELESONIX  ... X = 7  B. HEUCHERIFORMIS ... 2N = 14 R. L. T a y l o r (unpub.) C a l d e r & S p i c e r 33811, M i e t t e Hot S p r i n g s ,  Jasper  N a t i o n a l Park, A l b e r t a (DAO), 2n - 14; Packer (1968) Packer 2725, M i e t t e Hot S p r i n g s , J a s p e r N a t i o n a l Park, A l b e r t a (ALTA), 2n = 14; G o r n a l l 193, P i n e Creek Lake, Park Co., Montana  (UBC), 2n =  G o r n a l l 132, Mt. Magog, Cache Co.., Utah (UBC), 2n = 7^; (unpub.) T a y l o r & G i l l e t t boundary, on hwy  R. L. T a y l o r  4726, 1.2mi NE o f Shoshone N a t i o n a l F o r e s t  28, Fremont Co., Wyoming (DAO), 2n = 1 4 ;  C h a r l e s t o n Peak, C l a r k Co., Nevada (UBC), 2n = 7 B. JAMESII  7^;  G o r n a l l 115,  .  ... 2N = 14  Hamel (1953) ex Ingwersen N u r s e r i e s , 2n = 14.  PELTOBOYKINIA ... X = 11 P, TELLIMOIDES Skovsted  SSP. TELLIMOIDES  ... 2N = 22  (1934) ex Copenhagen B o t a n i c Garden,.2n ='11  ; Hamel (1948)  ex J a r d i n A l p i n du Museum N a t i o n a l d ' H i s t o i r e N a t u r e l l e , P a r i s , 2n = 22, 2n = HJ-J-J Hamel (1953) ex Copenhagen B o t a n i c Garden, Denmark, 2n = 22, 2n p  G o r n a l l 343, ex J . W.  G o e t h e ' U n i v e r s i t y B o t a n i c Garden,  F r a n k f u r t , F.D.R. (UBC), 2n =. 22. P. TELLIMOIDES  SSP. WATANABEI ... 2N = 22  G o r n a l l 328, ex J . W,  Goethe U n i v e r s i t y B o t a n i c Garden, F r a n k f u r t , F.D.R.  125  TABLE XIV, cont'd. P. TELLIMOIDES SSP. WATANABEI ... 2N = 22 (UBC) , 2n = 22.  SUKSDORFIA ... X = 7 . S, VIOLACEA ... 2N = 14 Bohm  1112, Crawford Bay, Kootenay Lake, B.C. (UBC), 2n = 7  R, L. T a y l o r  T T  _ . 7  (unpub.) T a y l o r & Sherk 4878, C o r r a L i n n Dam, Kootenay  R., B.C. (DAO), 2n = 14; G o r n a l l 329, 22mi W o f Sheep Lake, R o s s l a n d , B.C.  (UBC), n = 7; G o r n a l l 330, 19mi W of Sheep Lake, n r . R o s s l a n d ,  B.C.  (UBC), n = 7; R. L, T a y l o r  B.C.  (DAO), 2n' = 14; Packer (1968) Packer 2814, B l a k i s t o n Mt., Water-  (unpub.) C a l d e r & S p i c e r 32855, Yahk,  ton Lakes N a t i o n a l Park, A l b e r t a (ATLA), 2n = 14; R. L . T a y l o r T a y l o r & G i l l e t t 4605, Bad Rock Canyon, F l a t h e a d  (unpub.)  R., F l a t h e a d Co.,  Montana (DAO), 2n = 14, 2n ~ 7-j.^. S. RANUNCULIFOLIA ... 2N = 14 G o r n a l l 332, 6mi N o f F a u q u i e r , N end Lower Arrow Lake, B.C. (UBC), n =• 7; G o r n a l l 331, 19mi W o f Sheep Lake, n r . R o s s l a n d , B.C. (UBC), n = 7; Bohm 1132, Saddle Rock, n r . Y a l e , F r a s e r Canyon, B.C. (UBC), n = 7; Bohm 1144, B l a c k w a l l Mt., Manning P r o v i n c i a l 2n =  IJJ.  Park, B.C. (UBC),  , n = 7.  BOLANDRA ... X = 7 B. OREGANA ... 2N i= 14 Hamel (1958) ex " E n g l i s h Nursey", 2n = 14; Ornduff (1962) O r n d u f f 6240, Elowah F a l l s , McCord Creek, Multnomah Co., Oregon (UC), n = 7; R. L . Taylor  (unpub.) T a y l o r & G i l l e t t 4625, B u f o r d Creek Canyon, A s o t i n Co.,  Washington (DAO), 2n - 14; R. L. T a y l o r Klickitat, Klickitat  SULLIVANTIA  (unpub.) T a y l o r & G i l l e t t 4640,  Co,, Washington (DAO), 2n - 7  ... X = 7.  A l l s p e c i e s a r e 2N F 14 (Hamel 1953; S o l t i s *Hamel's  .  in litt.)  (1953) count o f 2n = 26 f o r B o y k i n i a o c c i d e n t a l i s was almost  c e r t a i n l y based on m a t e r i a l o f B_. -major.  126  I n f r a - s p e c i f i c Geographical V a r i a t i o n With the e x c e p t i o n o f B o y k i n i a r i c h a r d s o n i i . a l l s p e c i e s have constant  chromosome numbers, a t l e a s t i n the m a t e r i a l s t u d i e d so f a r .  B o y k i n i a r i c h a r d s o n i i possesses  2n = 6x = 36 i n c o l l e c t i o n s from the  A l a s k a Range, b u t 2n = 14x F 84 i n c o l l e c t i o n s from the a r c t i c s l o p e o f the Brooks Range.  T h i s s i t u a t i o n deserves  f u r t h e r study,  a t i o n o f specimens from both areas r e v e a l e d no obvious ences.  although  examin-  morphological  differ-  P o l l e n s i z e i s v e r y s i m i l a r between the two cytodemes (Ch. V I I ) .  No f o r m a l taxonomic r e c o g n i t i o n i s warranted. B_. r i c h a r d s o n i i i s a T e r t i a r y r e l i c t . s u r v i v e d the i c e ages.  According  to Hulten  (1945),  T h i s r a i s e s the q u e s t i o n o f where i t  Both the a r c t i c s l o p e and p a r t s of the A l a s k a  Range remained u n g l a c i a t e d d u r i n g the l a s t i c e age ( H u l t e n 1968), and the two  cytodemes c o n c e i v a b l y c o u l d have e v o l v e d  independently  i n their  t i v e r e f u g i a , r a t h e r than one b e i n g d e r i v e d from t h e o t h e r . interesting  I t would be  to e s t a b l i s h the chromosome numbers of p o p u l a t i o n s  Yukon T e r r i t o r y s i n c e i c e - f r e e r e f u g i a a l s o e x i s t e d t h e r e  respec-  from the  ( H u l t e n 1968).  Karyotypes I n most cases, m i t o t i c p r e p a r a t i o n s were inadequate karyotype  analysis.  The chromosomes a r e s m a l l i n B o l a n d r a ,  for detailed  Sullivantia  (Hamel 1958, 1953), B o y k i n i a and S u k s d o r f i a ( 1 - 3 / i m ) . However, by exami n i n g a l a r g e number of p r e p a r a t i o n s , t e n t a t i v e karyotypes duced f o r many o f the s p e c i e s ( T a b l e XV). are s u b - m e t a c e n t r i c  have been p r o -  G e n e r a l l y the l o n g e r chromosomes  and the s h o r t e r ones a r e m e t a c e n t r i c .  Discussion In the context  o f the S a x i f r a g i n a e as a whole, the base number  x = 7 i s v e r y common and p r o b a b l y Base numbers o f 6 and 11 a r e found  ancestral  (Skovsted  1934; Hamel 1953).  o n l y i n S a x i f r a g a and Chrysosplenium.  A h a p l o i d number o f t h i r t e e n is- r e a s o n a b l y  common b u t c o n f i n e d to t h e  TABLE XV.  Tentative karyotypes f o r species  of B o y k i n i a ,  P e l t o b o y k i n i a , B o l a n d r a and S u l l i v a n t i a .  Chromosome P a i r s  Taxon Metacentric  Sub-metacentric  Acrocentric  Telocentric  5  2  0  0  B. i n t e r m e d i a  5  2  0  0  B. a c o n i t i f o l i a  4  2  0  0  B. major  9  4  0  0  B. j a m e s i i *  5 (1 SAT)  2  0  0  Peltoboykinia tellimoides ssp. t e l l i m o i d e s  4  4 (1 SAT)  2  1  4 (1 SAT)  2  1  3  0  0  2  0  0  Boykinia  rotundifolia •  s s p . watanabei  4  B o l a n d r a oregana*  4 (ISAT)  Sullivantia sullivantii*  5  * +  d a t a from Hamel (1958,  1953).  SAT = s a t e l l i t e d p a i r . using  the nomenclature o f Levan e t a l . (1965).  128  genus S a x i f r a g a . There i s no c o r r o b o r a t i n g e v i d e n c e to support the grouping of B o y k i n i a s p e c i e s a c c o r d i n g to chromosome base number, v i z . two x = 6 and  7 respectively,  Chrysosplenium  Rather, B d y k j n l a shares w i t h S a x i f r a g a and  the p r o p e r t y of a n e u p l o i d v a r i a t i o n .  The numbers 6 and  have p r o b a b l y been d e r i v e d from 7 and 14, r e s p e c t i v e l y . suggested  questionable. and  13  Hamel (1953)  a p o s s i b l e h e t e r o p l o i d o r i g i n f o r the n = 13 c o n d i t i o n i n B o y k i n i a ,  although he r e c o g n i z e d that the g e o g r a p h i c a l assumptions  12)  groups w i t h  i n v o l v e d made i t  Taxa w i t h x = 6 are the A p p a l a c h i a n B_;_ a c o n i t i f o l i a  the a r c t i c B^ r i c h a r d s o n j i  (2n =  (2n = 36, 84); those w i t h x = 7 are  the western North American B_. o c c i d e n t a l i s , B. i n t e r m e d i a , B.  rotundifolia,  B. h e u c h e r i f o r m i s and B_. j a m e s i i (2n = 14). On m o r p h o l o g i c a l , chemical and p a l y n o l o g i c a l grounds IT - VIT, X I I ) the two  (Chs.  genome c o n t r i b u t o r s were p r o b a b l y 1) a B_. i n t e r m e d i a -  l i k e p l a n t and 2) a p l a n t s i m i l a r t o B_. r o t u n d i f o l i a and B^  occidentalis  ( t h e i r common a n c e s t o r ? ) which on p h y t o g e o g r a p h i c a l grounds (Ch. XV) i t s e l f have been s i m i l a r to B_. i n t e r m e d i a .  A segmental  may  allopolyploid  o r i g i n i s t h e r e f o r e e n v i s a g e d from an x = 7 by x = 7 h y b r i d i z a t i o n f o l l o w e d by a n e u p l o i d y to g i v e 2n = Karyotypes ploid origin for  26.  c o u l d support the h y p o t h e s i s o f a d i b a s i c  allotetra-  major, a l t h o u g h they are e q u a l l y compatible w i t h a  simple a l l o t e t r a p l o i d o r i g i n f o l l o w e d by the l o s s of a m e t a c e n t r i c p a i r , a l o s s i n c i d e n t a l l y which seems a l s o to have o c c u r r e d i n B_. a c o n i t i f o l i a . The base number o f x.=  11 i n P e l t o b o y k i n i a t o g e t h e r w i t h i t s  d i s t i n c t i v e asymmetrical k a r y o t y p e supports i t s s e p a r a t i o n from B o y k i n i a . The o r i g i n of t h i s number j s obscure, and r e l a t i o n s h i p s may  possibly l i e  i n the d i r e c t i o n of S a x i f r a g a where some s e c t i o n s (MiscopetaTum, T r i d a c t y l i t e s , Nephybphyllum, D a c t y l o i d e s and D i p t e r a ) have x = 11 as a common,  129  i f n o t the o n l y , base number.  L i k e P e l t o b o y k i n i a , many of these a l s o haye  u n i s e r i a t e glandular trichomes.  Other p e l t a t e - l e a v e d genera,  and A s t i l b o i d e s , have 2n = 34 (Earner 1948, Sullivantia,  1953).  the two S u k s d o r f i a s p e c i e s counted, and B o l a n d r a  possess 2n = 14 chromosomes.  The l a t t e r two  t i v e l y v i a t h e i r b u l b i f e r o u s rhizomes. c o n t r a s t s w i t h Lithophragma, - y e g e t a t i v e l y by b u l b i l s  Peltiphyllum  The  genera can reproduce apparent  cytological  a s a x i f r a g a c e o u s genus which a l s o  vegetaconstancy  reproduces  and grows i n s i m i l a r summer-dry h a b i t a t s .  I t shows  e x t e n s i v e p o l y p l o i d y , i n c l u d i n g t r i p l o i d y , p e n t a p l o i d y and the presecence of  'B' chromosomes ( T a y l o r  1965).  The g r e a t s i m i l a r i t y i n the k a r y o t y p e s of B o l a n d r a , B o y k i n i a and S u l l i v a n t i a supports a c l o s e r e l a t i o n s h i p between these genera 1958).  Indeed,  the s a t e l l i t e d p a i r of chromosomes i n b o t h B o l a n d r a  and B o y k i n i a h e u c h e r i f o r m j s might be homologous.  (Hamel oregana  130  XI',  BREEDING STUDY  M a t e r i a l s and methods A list  o f a l l c u l t i v a t e d m a t e r i a l i s g i v e n i n T a b l e XVI.  P l a n t s were e i t h e r t r a n s p l a n t e d from the w i l d o r grown from seed, i n a growth chamber w i t h a 16'hour d a y l e n g t h  a t 15 - 2 0 C . P  and kept  Problems were  encountered i n g e t t i n g some of the s p e c i e s to f l o w e r , and a l s o i n e n s u r i n g t h a t d i f f e r e n t s p e c i e s f l o w e r e d synchronously.  These d i f f i c u l t i e s  account f o r the s m a l l number of c r o s s p o l l i n a t i o n s of each type  largely  attempted.  G e n e r a l l y , B o y k i n i a o c c i d e n t a l i s , S u l l i v a n t i a oregana and to some e x t e n t B o y k j n l a r o t u n d i f o l i a , were w e l l d i s p o s e d m a t e l y 25 week c y c l e .  However a f t e r two s u c c e s s i v e f l o w e r i n g p e r i o d s , the  p l a n t s became, and remained, v e g e t a t i v e . Intermedia  toward f l o w e r i n g on an a p p r o x i -  Of the B o y k i n i a major and B.  p l a n t s i n c u l t i v a t i o n , o n l y a few i n d i v i d u a l s f l o w e r e d .  a t e l y many f l o w e r s are produced on each p l a n t . formis or Bj, a c o n i t i f o l i a f l o w e r e d to o v e r w i n t e r  No p l a n t s of B_. h e u c h e r i -  i n c u l t i v a t i o n , d e s p i t e a l l o w i n g them  o u t s i d e d u r i n g 1978 <-' 1979.  Both t r a n s p l a n t e d and grown from seed,  Boykinia richardsonij material,  a l s o remained v e g e t a t i v e .  a 24 hour d a y l e n g t h may have promoted f l o w e r i n g i n t h i s s p e c i e s i t was not t r i e d .  Fortun-  Perhaps although  P o l l i n a t i o n s i n v o l v i n g B_. r j c h a r d s o n i i and B_. h e u c h e r i -  f o r m i s used f r e s h m a t e r i a l c o l l e c t e d from the w i l d . T h i s r e l a t i v e l a c k of success  a t coaxing p l a n t s i n t o  however, m i r r o r e d a s i m i l a r s i t u a t i o n i n the w i l d . r e c a l c i t r a n t western s p e c i e s , i t was observed the i n d i v i d u a l s a c t u a l l y f l o w e r i n a g i v e n S u k s d o r f i a presented was,not p o s s i b l e to m a i n t a i n  I n p o p u l a t i o n s o f the  t h a t o n l y about a t h i r d of  year.  the worst problems f o r c u l t i v a t i o n .  It  th.e two N o r t h American s p e c i e s a l i v e under  growth chamber c o n d i t i o n s f o r more than one g e n e r a t i o n . had  flower,  B e s t success  i n t r a n s p l a n t i n g i n d i v i d u a l s i n t o c l a y pots and k e e p i n g  them i n an  was  131  TABLE XVI:. herbarium  L i s t o f c u l t i v a t e d m a t e r i a l . F u l l ^ c o l l e c t i o n d e t a i l s and l o c a t i o n s are g i v e n i n Appendix 1.  C o l l e c t i o n s by G o r n a l l a r e '  p r e f i x e d by 'G'.  Roykinia a c o n i t i f o l i a ; G342. R, i n t e r m e d i a : G23; G24, B, major: G38; G45; G92; G202; G335; G337. R, o c c i d e n t a l i s : G l ; G8; G22; G25; G27; G29; G54; G65; G86; G'216; G227; G255; G256; S t r a l e y 1753. R. r o t u n d i f o l i a : G98; 610.1; G105. H. h e u c h e r i f o r m i s : '6115; 6132; 6193; 6194; 6198; G344. B'v r i c h a r d s o n i i : 6271; G272; G302; G306. S/uksdorfia r a n u n c u l i f o l i a : S. v i o l a c e a : G329; 6330.  Bohm 1096; Bohm, 1132;  G332.  Peltoboykinia tellimoides ssp, t e l l i m o i d e s : G328. ssp. watanabei: G343, R o l a n d r a oregana:  G260; G340.  S u l l i v a n t i a oregana;  G261.  Jepsonia heterandra: Ornduff 5072. J, p a r r y i : Cromwell 709; Mulroy s.n. Mitella diphylla: Radford e t a l . 9085. ;M, t r i f i d a : Bohm 1129.  outdoor  c o l d frame, where a few p l a n t s once and then remained v e g e t a t i v e  or d i e d . Experimental P o l l i n a t i o n s A l l s p e c i e s are protandrous  to v a r y i n g degrees  (Ch. V I ) .  Flowers were emasculated b e f o r e any anthers had d e h i s c e d , and p o l l i n a t i o n s were made by rubbing an e x c i s e d anther o f one d e s i r e d p a r e n t on the r e c e p t i v e s t i g m a t a o f the o t h e r .  Stigmata were judged r e c e p t i v e when t h e i r  p a p i l l a e appeared h y a l i n e under a x l O hand l e n s and t h e i r s u r f a c e s become covered i n a wet  exudate.  C o n t r o l f l o w e r s , emasculated any s e t o f e x p e r i m e n t a l p o l l i n a t i o n s .  But n o t p o l l i n a t e d , were run. w i t h  Initially,  s m a l l m u s l i n hoods w i t h  cottorn d r a w s t r i n g s were p l a c e d over e v e r y e x p e r i m e n t a l f l o w e r to prevent  132  c o n t a m i n a t i o n , and under these c o n d i t i o n s no seed was e v e r s e t i n the emasculated  control flowers.  Autonomous agamospermy i s t h e r e f o r e  absent  from t h e s p e c i e s under study; i n d e e d , i t has n o t been r e p o r t e d i n the Saxifraginae.  The use o f muslin hoods was time consuming, the numerous  f l o w e r s on some i n f l o r e s c e n c e s b e i n g congested. the  practice  I t was d e c i d e d to abandon  and subsequent work showed t h a t l e s s than one i n twenty  c o n t r o l f l o w e r s was e v e r contaminated w i t h s e l f p o l l e n . sibility  of contaminant  s e l f i n g s i n subsequent  To check the pos-  i n t r a - and i n t e r - s p e c i f i c  p o l l i n a t i o n s , seeds o f every-mating were sown and grown up to determine their identity.  Those n o t c o r r e s p o n d i n g t o t h e i r i n t e n d e d parentage were  e x c l u d e d from the r e s u l t s . A s e r i e s of i n t r a - s p e c i f i c c r o s s p o l l i n a t i o n s was r u n f o r each s p e c i e s t o e s t a b l i s h a s e t o f c o n t r o l d a t a w i t h which t o compare b o t h s,eifings and i n t e r - s p e c i f i c m a t i n g s . Seed  Germination In  those experiments  i n v e s t i g a t i n g the g e r m i n a b l i t y of seeds  from any g i v e n c r o s s , t h r e e batches o f f i f t y seeds each (when p o s s i b l e ) were sown e v e n l y i n p e t r i d i s h e s c o n t a i n i n g s t e r i l e agar e n r i c h e d w i t h Hoagland's  solution.  Germination was s c o r e d a f t e r s i x weeks.  seeds were s c a t t e r e d on the s u r f a c e of s o i l  Otherwise,  i n p o t s and k e p t m o i s t .  I n t r a - s p e c i f i c Cross P o l l i n a t i o n s In  o n l y B o y k i n i a o c c i d e n t a l i s and B_. r o t u n d j f o l i a was i t  p o s s i b l e t o compare i n t r a - p o p u l a t i o n w i t h i n t e r - p o p u l a t i o n tions.  There was l i t t l e  regimes  (Table X V I I ) .  weeks from sowing.  cross-pollina-  d i f f e r e n c e i n seed s e t between the two c r o s s i n g  The g e r m i n a t i o n o f F l seed was about 80% a f t e r s i x  Pollen fertilities  o f b o t h p a r e n t a l s t o c k and F l gener-  a t i o n s 'averaged Between 80 - 97%, as judged from s t a i n a B i l i t y i n 1% a c e t o carmine.  TABLE X V I I .  Seed s e t and c o m p a t i b i l i t y from i n t r a - s p e c i f i c c r o s s - and s e l f - p o l l i n a t i o n s .  The G e n e t i c  C o m p a t i b i l i t y Index (C.I.) i s the mean seed s e t from a given p o l l i n a t i o n expressed as a percentage o f the mean seed s e t o b t a i n e d i n i n t r a - p o p u l a t i o n i s an analogous q u o t i e n t Genetic Compatibility  crosses  u s i n g mean g e r m i n a b i l i t y .  and Seed V i a b i l i t y  of the mother s p e c i e s .  The Seed V i a b i l i t y Index  The O v e r a l l C o m p a t i b i l i t y  Index i s a product o f the  Indices.  Pollination  Seed Set Mean + SE  Genetic c.i.%  % Germ. Mean + SE  57 + 14 62 + 12 3 3 + 5  100 109 62  81 83 58  Seed V,l.%  Overall C.I.%  + 4 + 3 + 8  100 .102 70  100 111 43  Boykinia o c c i d e n t a l i s i n t r a - p o p l n . outx. ^nterr-popln, outx. self  5 10 58  B, r o t u n d i f o l i a intra-^popln, outx. I n t e r - p o p l n . outx. self  2 3 21  296 293 291  + 46 + 70 + 16  100 99 98  89 86 90  + 2 + 5 + 3  100 97 101  100 96 99  B. i n t e r m e d i a i n t r a - p o p l n . outx. self  5 14  167 + 37 0 + -  100 0  82  + 4  100  100 0  B., major intra-popln. self  5 21  97 + 46 5 + 2  100 5  78 5  + 8 + 5  100 6  100 0  P e l t o b o y k i n i a t e l l i m o i d e s s s p . watanabei i n t r a - p o p l n . outx. self  3 7  235 + 30 143 + 69  100 61  76 78  + 9 + 11  100 103  100 63  Suksdorfia r a n u n c u l i f o l i a i n t r a - p o p l n . outx. self  1 7  93 + 49 + 13  100 53  -—  -—  outx.  (V.I.)  -  * *  -  TABLE XVII, cont'd. Pollination  N  Seed Set Mean + SE  Genetic C.I.%  % Germ Mean + SE  Seed V.I.%  Overall C.I.%  100 92  100 84  Suksdorfia violacea self  (autogamy)  S u l l i y a n t j a oregana i n t r a - p o p l n . outx. self B o l a n d r a oregana i n t r a - p o p l n . outx. self Jepsonia p a r r y i i n t r a - p o p l n . outx. self Number o f p o l l i n a t i o n s attempted.  55+7  *  34 + 31 + 4  100 91  1 2  52 + 44 + 1 7  100 85  * *  -  -  3 2  73+4 8 + 3-  100 11  72+9 26 + 1  100 36  100 4  3 1 5  * 71 + 65 + 8  135  Self Pollinations R e s u l t s of s e l f p o l l i n a t i o n s are shown i n T a b l e X V I I . c o m p a t i b i l i t y i n d e x was  c a l c u l a t e d by e x p r e s s i n g  A  self-  average s e l f seed s e t  as  a percentage o f the average i n t r a r - s p e c i f i c c r o s s p o l l i n a t e d seed s e t . T a b l e XVII shows t h a t the d i p l o i d B o y k i n i a i n t e r m e d i a and  the  B, major are b o t h s t r o n g l y s e l f - i n c o m p a t i b l e ( S I ) , whereas the B, r o t u n d i f o l i a , S u k s d o r f i a v i o l a c e a and compatible  CSC).  watanabei and  self  B o y k i n i a o c c i d e n t a l i s , P e l t o b o y k i n i a t e l l i m o i d e s ssp.  S u k s d o r f i a r a n u n c u l i f o l i a were found to be p a r t i a l l y  CPSC).  the,nature  o f i t s response to i n c o m p a t i b l e no  diploid  S u l l i v a n t i a oregana are f u l l y  compatible  producing  tetraploid  P e l t o b o y k i n i a was  seeds and  self  unique among the genera s t u d i e d i n pollinations.  Instead  s l i g h t l y w i t h e r i n g , a f t e r about two  f l o w e r would a b s c i s s , the p e d i c e l b r e a k i n g  2 - 3mm  of merely  weeks the e n t i r e  below the base of  the  flower. Genetic  s e l f i n c o m p a t i b i l i t y i n the S a x i f r a g i n a e occurs  the d i p l o i d l e v e l i n Heuchera (Wells  1980), Lithophragma ( T a y l o r 1965)  J e p s o n i a where i t i s accompanied by d i s t y l y  (Ornduff  p l o i d l e v e l , SI i s found i n Tolmiea (Correns 1965)  and B e r g e n i a  (Yeo  1966).  1969^).. At the  and  poly-  1928), Lithophragma ( T a y l o r  Although few S a x i f r a g a s p e c i e s have been  i n v e s t i g a t e d f o r t h e i r breeding  system, no  a l l b e i n g p a r t i a l l y or f u l l y SC  (Spongberg 1972).  In some s p e c i e s , e.g.  at  case of SI has been r e p o r t e d , Dioecy, however,  occurs  S_. e s c h s c h o l t z i i (Chambers 1964).  Many s p e c i e s i n o t h e r genera of the s u b - t r i b e have been found to be SC,  i n c l u d i n g T e l l i m a (Ornduff, p e r s . comm.), T j a r e l l a spp.  1966) , Heuchera spp.  (Wells 1980)  and ^ M i t e l l a ( p e r s o n a l  (Kern  observation),  Nature of the S e l f I n c o m p a t i b i l i t y System SI systems can be d i v i d e d i n t o those and  those  controlled sporophytically.  c o n t r o l l e d gametophytically  A l l known d i s t y l o u s systems are  136  governed s p o r o p h y t i c a l l y and tems i t i s i m p o s s i b l e t o t e l l without  conducting  this includes Jepsonia.  c o n c l u s i v e l y which type o f c o n t r o l e x i s t s  a s e r i e s of d i a l l e l  f;amilies and p a r e n t s .  I n homomorphic s y s -  c r o s s e s and b a c k c r o s s e s  i n FI h y b r i d  However t h e r e are a number of c o r r e l a t i o n s which  ha,ye Been observed between the type of c o n t r o l and  certain cytological  p h y s i o l o g i c a l f e a t u r e s , i n c l u d i n g number.of p o l l e n n u c l e i , mode of  and  PMC  c y t o k i n e s i s , the s i t e o f male gametophyte i n h i b i t i o n i n the c a r p e l , and the type of s t i g m a t i c s u r f a c e  (Keslop-Harrison  1975).  I n the p r e s e n t  B o y k i n j a , P e l t o B o y k l n i a , S u k s d o r f i a , S u l l i v a n t i a and B o l a n d r a nucleate p o l l e n with Stigmata. evidence  The  a l l have b i -  simultaneous c y t o k i n e s i s of PMC's and wet,  s i t e of p o l l e n i n h i b i t i o n was  not  case,  investigated.  group I I I The  scanty  a v a i l a b l e t h e r e f o r e i n d i c a t e s t h a t gametophytic SI i s o p e r a t i n g .  T a y l o r (1965) p o s t u l a t e d gametophytic SI f o r Lithophragma.  The  SI does not break down i n the t e t r a p l o i d B o y k i n i a major, nor  fact  that  in polyploid  s p e c i e s i n some o t h e r genera of the s u b - t r i b e , suggests t h a t a m u l t i - l o c u s system i s o p e r a t i n g i n the S a x i f r a g i n a e ( L u n d q v i s t i s f u r t h e r supported  by  1975).  This  hypothesis  the h i g h l e v e l of c r o s s - c o m p a t i b i l i t y w i t h i n F l  s i b s h i p s of T o l m i e a m e n z i e s i i (Correns  1928;  Lundqvist  1975).  I t would be i n t e r e s t i n g to d i s c o v e r the number of l o c i and.to f i n d out how Jepsonia arose.  the s p o r o p h y t i c , s i n g l e - l o c u s , d i s t y l i c c o n d i t i o n i n  I t i s however, i n t r i g u i n g t h a t i n the genus S a x i f r a g a ,  almost a l l of whose s p e c i e s are p o l y p l o i d , no  case of SI has  T h i s would suggest e i t h e r t h a t SI has never been p r e s e n t , l o c u s system once operated Inbreeding  involved  i n this  been r e p o r t e d .  or t h a t a s i n g l e  genus.  Depression E v i d e n c e of i n b r e e d i n g d e p r e s s i o n was  t i o n of B o y k i n i a major and B^ o c c i d e n t a l i s .  found i n the SI genera-  I t was  manifested  on i n the form of a.reduced seed g e r m i n a b l l i t y (Table X V I I ) .  very e a r l y In s u r v i v i n g  137  p l a n t s , a b e r r a n t l e a f development  The SI g e n e r a t i o n from B. ma j or d i d n o t f l o w e r ,  l e s s so i n B_. o c c i d e n t a l i s . b u t t h a t from \B_.  was q u i t e common i n B. major but much  o c c i d e n t a l i s d i d and showed normal p o l l e n f e r t i l i t y ,  95% p o l l e n s t a i n a b l e i n 1% acetocarmine.  82 -  B o y k i n i a r o t u n d j f o l i a showed no  evidence o f i n b r e e d i n g d e p r e s s i o n i n i t s SI g e n e r a t i o n , e i t h e r i n seed g e r m i n a b i l i t y , development  or p o l l e n f e r t i l i t y .  The r e s u l t s agree w i t h  what would be expected based on the b r e e d i n g systems Inter-specific The  o f the s p e c i e s .  Pollinations c r o s s i n g programme was concerned m a i n l y w i t h B o y k i n i a  s p e c i e s , b o t h t h e i r r e l a t i o n s h i p s to each o t h e r and to s p e c i e s i n o t h e r genera.  Where p o s s i b l e , s p e c i e s were c r o s s e d r e c i p r o c a l l y .  shown i n T a b l e X V I I I .  Seed  set i s  A g e n e t i c c r o s s a b i l i t y i n d e x was c a l c u l a t e d by  e x p r e s s i n g the seed s e t from a g i v e n h y b r i d c r o s s as a percentage of the mother s p e c i e s . In  the f i r s t  Three k i n d s of s i t u a t i o n arose from the h y b r i d  case,' no seed was s e t and the c a p s u l e s c o n t a i n e d many, l i g h t  brown, u n f e r t i l i z e d o v u l e s .  I n the second case, seed was s e t and they  appeared o f normal s i z e and shape. seeds were s e t , they appeared shrivelled.  crosses.  I n the t h i r d  case however, a l t h o u g h  s m a l l e r than u s u a l and f r e q u e n t l y  somewhat  D i s s e c t i o n of these seeds showed t h a t both endosperm and  embryo were shrunken, and o c c a s i o n a l l y the embryo was e n t i r e l y There was sometimes a complete  lacking.  range o f v a r i a t i o n w i t h i n a s i n g l e c a p s u l e ,  from u n f e r t i l i z e d o v u l e s , to v a r y i n g s i z e s of s m a l l seeds, to good seeds. In  cases where the o v u l e s had c l e a r l y developed to some e x t e n t , . t h e y were  counted as seeds.  Germination t e s t s were conducted on a l l h y b r i d  batches to t e s t f o r good seeds and to check f o r contaminant l a t t e r were e x c l u d e d from the d a t a .  seed  selfings.  The  Germination r e s u l t s are shown i n T a b l e  XVII'I, where a seed v i a b i l i t y i n d e x was c a l c u l a t e d i n the same manner as for  the g e n e t i c c o m p a t i b i l i t y i n d e x , b u t u s i n g g e r m i n a b i l i t y r a t h e r  than  TABLE X V I I I .  Seed s e t and g e r m i n a b i l i t y  from i n t e r - s p e c i f i c and i n t e r - g e n e r i c c r o s s p o l l i n a t i o n s .  G e n e t i c C r o s s a b i l i t y Index (C.I.%) i s the mean seed s e t from a given h y b r i d of the mean seed s e t o b t a i n e d i n i n t r a - s p e c i f i c crosses (V.I.) i s an analogous q u o t i e n t  the G e n e t i c C r o s s a b i l i t y and Seed V i a b i l i t y Pollination  (?xo*)  Boykinia o c c i d e n t a l i s x B. i n t e r m e d i a x B. r o t u n d i f o l i a x B. major x B. h e u c h e r i f o r m i s x B, x i c h a r d s o n i i x S u l l i v a n t i a oregana x Suksdorfia r a n u n c u l i f o l i a x S, v i o l a c e a x B o l a n d r a oregana x Peltoboykinia tellimoides ssp. watanabei x Jepsonia p a r r y i x J . heterandra x Mitella diphylla x M. t r i f i d a rotundifolia x~B. intermedia x B. o c c i d e n t a l i s x B, maj or x S u l l i v a n t i a oregana  N*  The Seed V i a b i l i t y Index  The O v e r a l l C o m p a t i b i l i t y  Index i s a product o f  Indices.  Seed Set jMeari + SE  11 12 11 7 1 14 10 2 1 8  54 28 53 2 0 1 5 10 27 0  5 5 4 4  0 1 0 0  6 12 2 2  cross expressed as a percentage  o f the mother s p e c i e s .  u s i n g mean g e r m i n a b i l i t y .  The  + 8 + •7 + 9  ± -* 2  + 0.3* + 3*  ± * 6  -  -  ±  1  -  184 + 24* 164 + 30* 169 £ 158* 0  Genetic C.I.% 95 49 93 4 0 2 9 18 47 0 0 2 0 0 62 55 57 0  % Germ. Mean + SE 81 56 53 23 0 0 0 0  0  1 16 23  + 6 + 16 + 17  ±-  -  -  23  -  + 1 + 12  ±  -  11  Seed V.I.%  O v e r a l l C.I.%  100 69 65 28  95 34 60 1 0 0 0 0 0 0  0 0 0 0  -  -0 -  0 0 0 0  1 18 26  1 10 15 0  -  -  TABLE X V I I I , Pollination  cont'd. ( ? x $)  iN  #  Seed S e t Mean + SE  B. i n t e r m e d i a X B. o c c i d e n t a l i s X B. r o t u n d i f o l i a X B. major X B. h e u c h e r i f o r m i s X Suksdorfia r a n u n c u l i f o l i a X Jepsonia p a r r y i J . heterandra X  10 10  161 85 170 17 0 0 0  B.. major X B. o c c i d e n t a l i s X B. r o t u n d i f o l i a X B, i n t e r m e d i a X S u l l i y a n t i a oregana Suksdorfia r a n u n c u l i f o l i a X B o l a n d r a oregana X  12 6 6 3 4 7  81,+ 102 + 88 + 24 0 0 0  Peltoboykinia tellimoides ssp. t e l l i m o i d e s x Boykinia occidentalis x S u l l i y a n t i a oregana S o i l l i y a n t i a oregana x Boykinia occidentalis x E. intermedia x B. r o t u n d i f o l i a x Peltoboykinia tellimoides ssp. watanabei Suksdorfia r a n u n c u l i f o l i a x S. v i o l a c e a x Boykinia occidentalis  10 3 10 8  1  + 12 ± * + 14 + 9* 1 8  Genetic C.I.% 96 51 102 10 0 0 0 84 105 91 0 0 0  '.  % Germ. Mean +'SE  Seed V.I.%  79 + 1 0 65 + 10 0  80 +  3  9 + 4 65 + 1 3 -  O v e r a l l C.I.%  96 0 79 0 -  92 0 81 0 0 0 0  103 1  2  87 1  79 -  3  72 0 0 0  6 2  0 0  -  0 0  -  0 0  3 1 4 2  4 £ 0 0 0  3* -  0  -  0  0 0 0  -  0 0 0  3 3  0 0  '-  0 0  0 0  TABLE X V I I I , Pollination  cont'd. ( £ x c?)  Genetic C.I.%  Suksdorfia violacea x S, r a n u n c u l i f o l i a  1  0  B o l a n d r a oregana x Boykinia occidentalis x B. maj or  3 4  0 0  Jepsonia p a r r y i x Boykinia occidentalis x B. i n t e r m e d i a  10 10  0 3  J, 1  N  heterandra x Boykinia occidentalis x B, i n t e r m e d i a = number o f p o l l i n a t i o n s  10 10 attempted.  * I n c l u d e s some s m a l l o r s h r i v e l l e d seeds  % Germ. Mean + SE  -  0  Seed V.I.%  O v e r a l l C.I.%  -  0  -  0 0  0  0 0  141  seed  set.  product  An o v e r a l l c o m p a t i b i l i t y index was  of the g e n e t i c and  I n t e r - s p e c i f i c Crosses The icant la,  :  results  seed  then c a l c u l a t e d as  compatibilities  the  CTable X V I I I and F i g . 40).  i n Boykinia (Table XVIII and F i g , 40)  show t h e r e i s a s i g n i f -  l e v e l of o v e r a l l c o m p a t i b i l i t y between the s p e c i e s of s e c t i o n Boykin-;  c o n f i r m i n g the group as a n a t u r a l u n i t .  B o y k i n i a major, B_. o c c i d e n t a l -  Is-, and B_. i n t e r m e d i a a l l show a h i g h l e v e l of p o t e n t i a l gene exchange  and  rform a w e l l - d e f i n e d group , to which B. . a c o n j t i f o l i a and B_. l y c o c t o n i f o l i a a l s o belong two  on m o r p h o l o g i c a l  grounds.  s p e c i e s w i l l a l s o prove to be  I t may  be p r e d i c t e d t h a t these  c r o s s compatible  and w i t h t h e i r P a c i f i c n o r t h w e s t e r n  latter  both w i t h each o t h e r ,  couterparts.  B o y k i n i a r o t u n d i f o l i a a l s o shows a degree of c o m p a t i b i l i t y w i t h the o t h e r members of s e c t i o n B o y k i n i a , although  the b a r r i e r s to  b r i d i z a t i o n are somewhat s t r o n g e r than between the o t h e r s . with;B_. major there i s some e v i d e n c e although  In  hy-  crosses  of SI x SC u n i l a t e r a l i n c o m p a t i b i l i t y  t h i s i s o n l y r e v e a l e d i n h y b r i d seed  g e r m i n a b i l i t y experiments.  B o y k i n i a r o t u n d i f o l i a i s almost c r o s s - i n c o m p a t i b l e w i t h B_. i n t e r m e d i a again t h e r e i s evidence p o l l i n a t i o n and  and  of SI x SC u n i l a t e r a l i n c o m p a t i b i l i t y , both i n  i n h y b r i d seed g e r m i n a t i o n .  I n f a c t no h y b r i d s w i t h  B.  i n t e r m e d i a as female parent were produced, and o n l y a v e r y few were made i n the r e c i p r o c a l d i r e c t i o n .  C r o s s e s of B_. r o t u n d i f o l i a w i t h B_. o c c i d e n t 1  a l i s were more s u c c e s s f u l w i t h the l a t t e r as female p a r e n t . seeds were produced w i t h a female B. r o t u n d i f o l i a , lower than i n the r e c i p r o c a l combination. l e s s e x t e n s i v e between K. than- between B.  ALthough more  t h e i r g e r m i n a b i l i t y was  Overall, crossing barriers  are  r o t u n d i f o l i a and B. o c c i d e n t a l i s or B_. major  r o t u n d l f o l i a and "Jl. i n t e r m e d i a .  The  Increased  genetic  l a t i o n of B_. r o t u n d l f o l i a from the o t h e r s p e c i e s of s e c t i o n B o y k i n i a i s r e f l e c t e d i n i t s d i s t i n c t i v e morphology.  iso-  142  FIGURE 40. C r o s s i n g  r e l a t i o n s h i p s between s p e c i e s  terms o f O v e r a l l C o m p a t i b i l i t y and  hybrid  and genera, expressed i n  (the product o f i n t e r s p e c i f i c c r o s s a b i l i t y  seed v i a b i l i t y ) . A l l c r o s s e s a r e r e c i p r o c a l except where n o t e d .  Arrows i n d i c a t e the d i r e c t i o n o f p o l l e n  transfer.  71 - 100 Overall Compatibility  — —  31-70  1-30 O  ov /o  143  Only a s i n g l e cross w i t h B_. r l c h a r d s o n i i was I n v o l v e d B.  o c c i d e n t a l i s as  is difficult resulted  to assess the  from one  being rejected  the  female p a r e n t .  I t was  made and  this  unsuccessful.  s i g n i f i c a n c e of t h i s because f a i l u r e may  o r more o f the  following  factors:  1) p o l y p l o i d  (e.g. Rajhathy and  incompatibility;  pollen  Thomas 1974); 2) i n t e r - s p e c i f i c c r o s s -  or 3) e n v i r o n m e n t a l e f f e c t s .  In s e c t i o n T e l e s o n i x , o n l y p o l l e n of B_. h e u c h e r i f ormis  media. small,  T h i s was  dusted on  the  s t i g m a t a of  occidentalis  In b o t h cases a c e r t a i n percentage of the somewhat underdeveloped seeds.  f a i l e d , to germinate, but i f ormis c r o s s e s d i d . F l hybrids. formis and  This  The  other i n t e r g e n e r i c  seedlings  grew up  to be v e g e t a t i v e l y  supports the  i n Boykinia rather  B.  rather  intermedia  vigorous  i n c l u s i o n of B_.  than i n S a x i f r a g a  heucheri-  or the  segregate  f a i l u r e at some stage of a l l  Morphology  Seed i n c o m p a t i b i l i t y , i s a well-known phenomenon and 1953)  an e a r l y e x p r e s s i o n of h y b r i d  has  inviability,  been r e p o r t e d i n many genera,  e.g.  where.the symptoms of a b e r r a n t endosperm and  b r y o development are v e r y s i m i l a r to those found h e r e . the p h y s i o l o g i c a l b a s i s  Valentine  f o r seed i n c o m p a t i b i l i t y was  the  it  arose as  to t h e i r -mutual d e g e n e r a t i o n ,  a correlated  em-  suggested  l a c k of  chronous exchange of hormones and m e t a b o l i t e s between the embryo and sperm, l e a d i n g  inter-  hybridizations.  H y b r i d I n v i a b i l i t y and  Primula (Valentine  crosses y i e l d e d  was  some of those from the B_^ o c c i d e n t a l i s x heucher-  result strongly  B_. j a m e s i i  and  A l l those produced by B_.  genus T e l e s o n i x . e s p e c i a l l y i n view of the  that  have  on a d i p l o i d s t y l e ; t h i s i s a r e l a t i v e l y common phenomenon  i n such c r o s s e s  available.  It  Eaton (1973) c o n s i d e r e d  synendothat  response to s e l e c t i o n f o r d i f f e r i n g p h y s i o l o g i c a l  a t t r i b u t e s i n d i f f e r e n t species', as- a postT-zygotic B a r r i e r to  rather  than B e i n g s e l e c t e d  hybridization.  for  specifically  144  The those  o n l y h y b r i d s to f l o w e r d u r i n g the course  d e r i v e d from the B.  r o t u n d i f o l i a x maj or c r o s s e s .  p o l l e n s t a i n a b i l i t y i n 1% acetocarmine of 1.3%, no p o l l e n at a l l . of the t r i p l o i d  of the study were, They had  w i t h some p l a n t s  a mean producing  T h i s h i g h l e v e l of s t e r i l i t y i s p a r t l y or w h o l l y  s t a t u s of these p l a n t s  h y b r i d i s shown i n F i g . 41.  (2n = 20).  morphology i s v e r y  An i l l u s t r a t i o n of  I t i s i n t e r e s t i n g t o note t h a t l a r g e  s i z e Is dominant over s m a l l f l o w e r s i z e i n t h i s c l o s e to t h a t o f B. major.  a result  flower  c r o s s , as the h y b r i d I n some cases  this  floral  developmental  anomalies were observed i n the b a s a l l e a f complement of some p l a n t s .  In  ohe  to  of the  r o t u n d i f o l i a x maj or h y b r i d s , the f l o w e r i n g stem f a i l e d  develop c a u l i n e l e a v e s p r o p e r l y ( F i g . 41). A comparison of b a s a l l e a f shapes of the h y b r i d s w i t h respective parents' hybrids  i s shown i n F i g . 42.  d i f f e r e n c e s were observed i n  r e s u l t i n g from r e c i p r o c a l p o l l i n a t i o n s , and  morphologies were i n t e r m e d i a t e No wild.  No  The  between those  their  a l l hybrid  of t h e i r  vegetative  parents.  i n s t a n c e of n a t u r a l h y b r i d i z a t i o n has been found i n the  g e o g r a p h i c a l p r o x i m i t y of the P a c i f i c Northwestern  species  could a l l o w c r o s s - p o l l i n a t i o n i n some l o c a l i t i e s , so the absence of n a t u r a l h y b r i d s may  be  a t t r i b u t e d to any  or a combination of the f o l l o w i n g f a c t o r s :  s u b t l e e c o l o g i c a l d i f f e r e n c e s between the p a r e n t s , p r e - z y g o t i c i s o l a t i o n , seed i n c o m p a t i b i l i t y , and  i n some cases  genetic  to h y b r i d developmental  instability. I n t e r - s p e c i f i c Crosses A v e r y few  i n Suksdorfia c r o s s - p o l l i n a t i o n s were attempted between the  N o r t h American s p e c i e s , S u k s d o r f i a r a n u n c u l i f o l i a and _S\_ - v i o l a c e a . crosses f a i l e d .  two All  T h i s c r o s s ^ i n c o m p a t i b i l i t y i s i n t e r e s t i n g because the  s p e c i e s have Been p l a c e d i n separate d o r f i a , r e s p e c t i v e l y (Ch. X I V ) .  genera i n the p a s t , Hemjeya and  However, r a t h e r than r e f l e c t i n g  two  Suks-  generic  145  FIGURE 41. The Boykinia  rotundifolia  x B_. major hybrid. a) Hybrid i n v i a b i l i t y . Occasional hybrids showed developmental abnormalities. In t h i s case the cauline leaves have f a i l e d to develop  properly, although the  plant went on to flower.  b) Longitudinal section of a flower. F l o r a l morphology tends strongly toward that of Boykinia major.  c) Upper s t i p u l e and cauline l e a f . Stipule morphology i s much more l i k e that of Boykinia  rotundifolia  than B_. major.  147  s t a t u s f o r each, the c r o s s - i n c o m p a t i b i l i t y may ence of two ranges,  B i o l o g i c a l species.  They are sympatric  over most of  have v e r y s i m i l a r e c o l o g i c a l p r e f e r e n c e s , and  p o s s i B l e to ifind them growing i n adjacent Although  p e r e n n i a l s , they are proBaBly  rhizome system i s not s u b s t a n t i a l .  I n mixed or contiguous  exist-  their  i n many p l a c e s i t i s  or even s l i g h t l y mixed p o p u l a t i o n s .  s h o r t - l i v e d as t h e i r B u l B i f e r o u s  I n these  circumstances,  B a r r i e r s between r e l a t e d s p e c i e s are common and  ing  merely i n d i c a t e the  strong  to be expected  (Grant  p o p u l a t i o n s , S_. v i o l a c e a begins  about two weeks e a r l i e r than S_. r a n u n c u l i f o l i a ,  although  breeding 1971).  flower-  t h e r e i s ob-  v i o u s l y a g r e a t d e a l of o v e r l a p once the b u l k of both p o p u l a t i o n s are i n flower.  No  i n s t a n c e s of n a t u r a l h y b r i d i z a t i o n have been d i s c o v e r e d .  f e r e n c e s i n p o l l i n a t o r fauna may to be  exist  (Ch. XIII) but  the d a t a are too  Crosses  The  r e s u l t s of the i n t e r - g e n e r i c c r o s s e s are g i v e n i n T a b l e  XVIII and F i g . 40. t i a , Bolandra,  Genera t e s t e d i n c l u d e d S u k s d o r f i a , B o y k i n i a ,  J e p s o n i a , P e l t o b o y k i n i a and M i t e l l a .  h y b r i d i z a t i o n s succeed.  I n no  case d i d the  (both s p p . ) , S u l l i v a n t i a oregana and J e p s o n i a h e t e r a n d r a  Suksdorfia  (Table XVIII).  a l s o o c c u r r e d i n c r o s s e s of B o y k i n i a i n t e r m e d i a w i t h J e p s o n i a  parryi.  r e p o r t e d the same phenomenon i n c r o s s e s between B o y k i n i a  a c o n l t i f o l j a and S u l l i v a n t i a spp. s t u d i e d supports  into  T h i s seed i n c o m p a t i b i l -  i t y o c c u r r e d as a r e s u l t of c r o s s e s of B o y k i n i a o c c i d e n t a l i s w i t h  (in litt.)  Sullivan-  I n some cases, however, the ovules developed  s m a l l , sometimes s h r i v e l l e d , non-germinable seeds.  Soltis  few  certain.  Inter-generic  It  Dif-  t h e i r separate  The  c r o s s ^ i n c o m p a t i b i l i t y of the genera  i n t e g r i t i e s , e s p e c i a l l y i n v i e w of the  fact  t h a t i n t r a - g e n e r i c , i n t e r s p e c i f i c h y b r i d s are e a s i l y formed i n many of the S a x i f r a g i n a e , i n c l u d i n g B'oyklnla, J e p s o n i a (Soltis i n l i t t . ) ,  Bergenia  (Yeo  (Ornduff 1969a), S u l l i v a n t i a  1966) , Lithophragma ( T a y l o r 1965)  and  Heuchera (Wells 1980).  149  XII. Previous  FLAVONOID CHEMISTRY  Chemical I n v e s t i g a t i o n s o f B o y k i n i a and A l l i e s The  Saxifragaceae  s . l . were the s u b j e c t o f e x t e n s i v e  polyphenol  surveys by P l o u v i e r 0-965) , J a y and L e b r e t o n (1965) and J a y (1969).  The  f a m i l y was a l s o i n c l u d e d i n the d i c o t y l e d o n survey by Bate-Smith (1962) who found q u e r c e t i n and kaempferol i n B o y k i n i a  aconitifolia.  J a y (1969) l i s t e d  the presence o f l e u c o c y a n i d i n , q u e r c e t i n and kaempferol i n B. a c o n i t i f o l i a , ;B"... o c c i d e n t a l i s and B, r o t u n d i f o l i a . ported ered  F o r P e l t o b o y k i n i a t e l l i m o i d e s he r e -  l e u c o c y a n i d i n , q u e r c e t i n , kaempferol and e l l a g i c a c i d .  the o c c u r r e n c e o f e l l a g i c a c i d i n support  He c o n s i d -  Hara's (1937) r e c o g n i t i o n  of P e l t o b o y k i n i a a t the g e n e r i c l e v e l . - J a y (1969) a l s o s t u d i e d  Sullivantia  s u l l l v a n t i j which he found to c o n t a i n l e u c o d e l p h i n i d i n , l e u c o c y a n i d i n , q u e r c e t i n , kaempferol and e l l a g i c a c i d .  Recently,  Soltis  (1980) i n v e s t i -  gated a l l s i x s p e c i e s o f S u l l i v a n t i a u s i n g paper chromatography and r e p o r t e d quercetin-3-0-mono-, d i - , and t r i g l y c o s i d e s (some o f which  contained  g l u c u r o n i c a c i d ) , pedalitin-6-0-mono-.and d i g l y c o s i d e s , and a t r a c e o f luteolin. On the b a s i s o f t h e i r r e s u l t s , Jay (1969) and J a y e t a l . (1970) proposed t h r e e taxonomic groups: a " c e r c l e e l l a g i q u e " composed o f Mukdenia, A g t i l b o i d e s , Saxifraga sect. Micranthes,  B e r g e n i a and P e l t o b o y k i n i a ; a  " c e r c l e Heuchera" which i n c l u d e d T e l l i m a , T o l m i e a , T i a r e l l a ,  Mitella,  Heuchera, S u l l i v a n t i a and p o s s i b l y P e l t i p h y l l u m and B e n s o n i e l l a ; and a tentative "cercle Saxjfraga  M  comprising  S a x i f r a g a , R o d g e r s j a , A s t j l b e and  Boykinia.  The conclusions-must be regarded w i t h s c e p t i c i s m because the  techniques  used and the philosophy- behind the i n t e r p r e t a t i o n o f t h e r e s u l t s  were p r i m i t i v e . Regarding o t h e r p h y t o c h e m i c a l s t u d i e s on these genera, Izawa e t a l . (1973) r e p o r t e d o i - p e l t o b o y k i n o l i c a c i d and p - p e l t o b o y k i n o l i c a c i d  150  ( t r i t e r p e n i c a c i d s ) and b e r g e n i n Ca C - g l y c o s y l a t e d from B o y k i n i a B.  g a l l i c acid deriyatiye)  t e l l i m o i d e s . fi - P e l t o b o y k i n o l i c a c i d alone was found i n  lycoctonifolia.  They concluded t h a t these r e s u l t s supported the r e c o g -  n i t i o n o f section Peltoboykinia  a t the g e n e r i c  level  (Kara 1937).  I t may  be n o t e d , however, t h a t fi - p e l t o b o y k i n o l i c a c i d , i t s monoacetate and a s t i l b i c a c i d (6-fi- 0 R 7 r > ^ — p e l t o b o y k i n o l i c a c i d ) have a l s o been i s o l a t e d from As t i l b e r i v u l a r i s Ham. ( S a s t r y and Rao 1977). The tographic Tellima  family i s g r a d u a l l y being  and s p e c t r o s c o p i c methods.  reworked, u s i n g modern chroma-  Genera s t u d i e d so f a r i n c l u d e  ( C o l l i n s and Bohm 1974; C o l l i n s e t a l . 1975), Heuchera CWilkins and  Bohm 1976; Bohm and Wllkins' 1978a), Elmera CBohm and W l l k i n s  1978b),  T d l m l e a CBohm 1979), Chrysosplenjum CBohm and C o l l i n s 1979), P e l t i p h y l l u m CBohm and W l l k i n s  1976), J e p s o n i a  (Bohm and Ornduff 1978), L e p t a r r h e n a  CMiller and Bohm 1979) and S a x i f r a g a C M i l l e r and Bohm 1980). i n v e s t i g a t i o n forms p a r t of t h i s r e v i s i o n , and r e p o r t s Boykinia,  Peltoboykinia, Suksdorfia,  The p r e s e n t  f l a v o n o i d d a t a on  S u l l i v a n t i a and B o l a n d r a .  METHODS Wj;th some m o d i f i c a t i o n s , t h i s study f o l l o w W i l k i n s Plant  the p h y t o c h e m i c a l methods used i n  & BohmC1976).  Material D e t a i l s o f c o l l e c t i o n s and amounts o f t i s s u e used a r e g i v e n i n  T a b l e XIX.  M a t e r i a l o f almost a l l the c o l l e c t i o n s was a l s o grown under  uniform conditions daylength).  i n an e x p e r i m e n t a l growth room Cat 15 - 20°C and 16hr  Apart from some q u a n t i t a t i v e d i f f e r e n c e s , the f l a v o n o i d p r o -  f i l e s of flowering,  growth-^room m a t e r i a l were q u a l i t a t i v e l y s i m i l a r t o  those found i n m a t e r i a l  c o l l e c t e d from n a t u r e .  Chromatography and S p e c t r o s c o p y '  Analytical t h i n ^ l a y e r chromatography CTLC) was c a r r i e d out  151  TABLE XIX.  Populations  onoid a n a l y s e s .  sampled and d r y weights o f t i s s u e used f o r f l a y -  C o l l e c t i o n s By G o r n a l l are p r e f i x e d By G'. ?  Full  details  are g i v e n i n Appendix 1,  B o y k i n i a a c o n i t i f o l i a ( u r c e o l a t e c a p s u l e s ) : G342j' 19.9g; S o l t i s 1046, 25. 8g. B. a c o n i t i f o l i a C t u r B i n a t e c a p s u l e s ) : H a r v i l l & Segars 143, O . l g . B. i n t e r m e d i a : G23, 77.7g; G24, 49.1g. B. l y c o c t o n i f o l i a : OhBa e t a l . 73099, O . l g ; Sugawara s.n. 18.7g. B. major: G38, 27.2g; G45, 28.3g; G55, 15.6g; G60, 49.6g; G92, 38.5g; G202, 117.1g; G207, 60.2g; G212, 49.2g; G335, 25'.Ig. B. o c c i d e n t a l i s : Bohm 1102, 46.Og; Bohm and S c h o f l e l d 1293, 1.6g; G l , 4.5g; G8, . 27.3g; G22, 22,6g; G25, 7.5g; G27, 25.7g; G29, 31.8g; G54, 6.0g; G65, 7.0g; G86, 28.3g; G216, 25.1g; G227, 307.8g; G255, 12.5g; G256, 5.2g; S t r a l e y 1 7 5 3 , 11.2g. B. r o t u n d i f o l i a : G98, 184.8g; G101, 102.4g; G105, 66.6g. B. r i c h a r d s o n i i : G271, 9.8g; G272, 13.6g; G302, 19.4g; G303, 18.2g; G306, 19.Og; Guppy s.n., 68.4g. B.. h e u c h e r i f o r m i s : G115, 22.2g; G132, 27.2g; G193, 40.Og; G194, 2.6g; G198, 77.2g; G344, 25.4g. B. j a m e s i i : G i l l e t t & Mosquin 12114, O . l g . S u k s d o r f i a a l c h e m i l l o i d e s : F i e B r i g 3161, O . l g . S. r a n u n c u l i f o l i a : Bohm 1092, 3.4g; Bohm 1096, 3.3g; Bohm 1132, 2.2g; Bohm 1144, 1.3g; G15, 4.1g; G245, 4.8g; G253, 8.8g. S, v i o l a c e a : Bohm 1112, l.Og; Bohm 1118, 0.3g; Bohm 1122, l.Og; G248, 1.4g; G252, 0.4g. Kolandra  oregana:  G260, 0.5g; G262, 2,0g; G340, 8.5g.  S u l l i v a n t i a oregana:  G261, 4.0g.  PeltoBoykinia tellimoides s s p . t e l l i m o i d e s : G328, l . O g . ssp. watanabei: G343, 1.8g.  w i t h "Polygram" 0.25mm s i l i c a g e l and "Polygram" 0.1mm c e l l u l o s e MN300 commercially  prepared  amide DC.6 p l a t e s .  s h e e t s , and w i t h l a b o r a t o r y prepared  .33mm MN P o l y -  F o r p r e p a r a t i v e TLC, three types of p l a t e were used:  1) 0.5mm A v i c e l m i c r o c r y s t a l l i n e c e l l u l o s e ; 2) 0.33mm M N - s i l i c a g e l G; and 3) 0.33mm MN Polyamide DC 6,6. Two standard e l u e n t s f o r polyamide p l a t e s were r o u t i n e l y used;.  1) "aqueous"  Cwater :  Nr^Butanol  ; acetone :  dloxane i n the r a t i o 70:15:1055), and 2) " o r g a n i c " ' C l , 2 - d i c h l o r o e t h a n e :. methanol ; butanone : water i n the r a t i o 50:25:21:4). comparative Rf data a r e d e s c r i b e d i n Appendix 2.  S p e c i a l eluents f o r  Ultra-violet  spectra  152  were r e c o r d e d  i n methanol.  E x t r a c t i o n and  Isolation  From each p o p u l a t i o n of each taxon ( T a b l e XIX) e r a l whole p l a n t s were blended w i t h 500ml methanol and hour.  The  s l u r r y was  filtered  and  ness.  About 5g C e l i t e A n a l y t i c a l F i l t e r A i d and  samples of sev-  them warmed f o r  the methanol e x t r a c t evaporated to  water were added to d i s s o l v e the e x t r a c t .  and  The  combined b u t a n o l  the r e s i d u e was  e x t r a c t was  T h i s was  then f i l t e r e d  and  the  evaporated to dryness i n vacuo  been p r e v i o u s l y s w o l l e n .  e l u t e d w i t h s u c c e s s i v e 400ml a l i q u o t s of 20%,  30%,  and  The  methanol, and  during e l u t i o n with  finally  100%  an u l t r a v i o l e t  acetone. (UV)  40%,  " o r g a n i c " s o l v e n t systems.  The  50%,  column was  lamp (360nm).  200ml were c o l l e c t e d and monitored on polyamide TLC, and  n-  taken up i n 20% methanol ( i n water) f o r l o a d i n g onto a  Sephadex LH-20 column, which had  100%  dry-  50 - 100ml d i s t i l l e d  f i l t r a t e e x t r a c t e d f o u r times w i t h 125ml p o r t i o n s of w a t e r - s a t u r a t e d butanol.  one  column  60%,  was  80%  examined  F r a c t i o n s of 100  -  u s i n g the "aqueous"  S i m i l a r f r a c t i o n s were combined.  F i n a l p u r i f i c a t i o n of i n d i v i d u a l f l a v o n o i d s was  achieved  by  banding the r e l e v a n t column f r a c t i o n s onto TLC p l a t e s , u s i n g a s u i t a b l e medium and e l u e n t system.  I f the q u a n t i t y of p o l y p h e n o l  great f o r t h i s , and p u r i f i c a t i o n was Polyamide SC-6  column was  fi - a m i n o - e t h y l d i p h e n y l b o r i n a t e UV  light.  a s u i t a b l e eluent  system.  on TLC p l a t e s by s p r a y i n g w i t h  ( A l d r i c h Chem. Co.)  and  0.1%  examining under  Dark brown, green, y e l l o w or orange c o l o u r s i n d i c a t e d f l a v o n -  oids.' Blues  i n d i c a t e d other poly-phenols.  r e a c t i o n s on polyamide DC  6.6  TLC  The  p l a t e s was  s e n s i t i v i t y of the  found to be  i n the next s e c t i o n ,  colour  particularly  h e l p f u l as a p r e l i m i n a r y i n d i c a t o r of f l a v o n o i d s t r u c t u r e . cussed  too  p o s s i b l e u s i n g polyamide, then a 9g  run, e l u t i n g i t w i t h  F l a v o n o i d s were d e t e c t e d  m a t e r i a l was  This i s d i s -  15 3  I d e n t i f i c a t i o n of Flavonoids Mabry e t a l . (1970) d e s c r i b e d the r e l a t i o n s h i p s between onoid s t r u c t u r e and appearance on paper,  flav-  i n UV l i g h t and i n t h e presence o f  ammonia. Some u s e f u l i n f e r e n c e s can be made when v i e w i n g f l a v o n o i d s on Polyamide DC-6.6 i n UV l i g h t , b e f o r e and a f t e r s p r a y i n g w i t h 0.1% e t h y l diphenyl borinate.  A b r i e f o u t l i n e o f some o f t h e c o l o u r r e a c t i o n s  was  g i v e n by W l l k i n s and Bohm (1976) who d e a l t w i t h t h e common  and  luteolin  (a flavone).  and spray r e a c t i o n s of the 144 f l a v o n o i d s  study.  UV s p e c t r a l d a t a u s i n g s h i f t of the aglycone al.  (Appendix  allowed  3).  (Mabry e t  I n f e r e n c e s about s t r u c t u r e s p r o v i d e d by  H y d r o l y s e s were conducted  o f d i g l y c o s i d e s and t r i g l y c o s i d e s ,  the h y d r o l y s a t e s were  T h i s allowed  During  hydros  monitored  c o n c l u s i o n s to be reached  as to the n a t u r e o f the i n n e r sugar and a l s o i t s p o s i t i o n . complete (1.5 - 2 h r s ) , the h y d r o l y s a t e was evaporated  vacuo and taken up i n e t h y l a c e t a t e and water. f o r sugar a n a l y s i s .  data  i n about 2ml of 20% methanol i n  o f t r i f l u o r a c e t i c a c i d a t 80 - 100°C.  by TLC a t 10 minute i n t e r v a l s .  was  identification  and b o r i n a t e s p r a y r e a c t i o n s were confirmed by t h e s p e c t r a l  water, u s i n g s i x drops lysis  reagents  and the p o s i t i o n o f the attachment o f the sugars  1970; Mears and Mabry 1972).  UV l i g h t  flavonols  T a b l e XX shows the s t r u c t u r a l i n f o r m a t i o n which  c o u l d be drawn from UV behaviour i s o l a t e d i n the present  P -amino-  both  When h y d r o l y s i s to dryness i n  The o r g a n i c phase was.used  The aqueous e x t r a c t was evaporated  t o dryness  and the  r e s i d u e taken up i n one or two drops- o f water and s p o t t e d along w i t h  stand-  ards on s t r i p s of c e l l u l o s e TLC s h e e t .  using  two  Chromatography was conducted  developments w i t h e t h y l a c e t a t e : p y r i d i n e : water (10;3.2;2),  The  sugars were l o c a t e d By s p r a y i n g w i t h p-r-anisidine p h t h a l a t e and h e a t i n g t h e s t r i p s a t 110°C f o r about f i v e  minutes.  154  TABLE XX.  F l a v o n o i d behaviour i n UV l i g h t , b e f o r e and a f t e r s p r a y i n g w i t h  0.1% >8-aminoethyl d l p h e n y l B o r i n a t e .  Flavonoid  S t r u c t u r a l Inference  Spot'Colour  UV L i g h t  E o r i n a t e Spray + UV L i g h t  CR f- H; X F anything)  Dark p u r p l e o r  Y e l l o w to dark B r i c k r e d  3'4'-COH) ;  3-OR o r  2  3-H or 3,6-COH) .  dark Brown  2  Dark p u r p l e o r  Not  Green  3'4'-COH) ; 2  3-OR;  dark Brown  6-H;  or 3-H  Dark p u r p l e o r  Dark p u r p l e o r dark  Not  3'4'-C0H) ;  dark brown  Brown  3-H or 3-OR  2  6-OX;  o r 3,6-  C0H) . 2  D u l l yellow or  Y e l l o w to dark B r i c k r e d  3'4'-(OH) ;  Green  Not  2  3-0H.  p a l e Brown D u l l yellow or  3'4'-C0H) ;  6-H o r  p a l e Brown  2  3-0H;  6-OR.  In many cases, t r i g l y c o s i d e s were h y d r o l y z e d under t h r e e r e gimes:  at room temperture  total hydrolysis  f o r 5 minutes,  (1.5 to 2hr a t 1 0 0 ° C ) .  a t 80°C f o r 5 minutes,  and a  The sugars from each of these  procedures were converted to t h e i r a l d i t o l a c e t a t e s and a n a l y z e d g a s - c h r o m a t o g r a p h i c a l l y f o l l o w i n g the method of Jones and A l B e r s h i e m C1972) .  This  method allowed n o t o n l y the sequence of sugars t o Be determined, But a l s o t h e i r r e l a t i v e amounts.  A s t a n d a r d mixture of d e r i v a t i z e d  araBinose,  x y l o s e , g a l a c t o s e , g l u c o s e , rhamnose and mannose i n e q u a l amounts was run for  comparison.  I d e n t i f i c a t i o n of A c y l a t e d FlayonoldsThree k i n d s of f l a v o n o i d p r e s e n t e d i d e n t i f i c a t i o n The  f i r s t k i n d showed Rf v a l u e s of n i l ,  proBlems.  or s t r e a k i n g from the o r i g i n , i n  a l l n e u t r a l s o l v e n t systems But Behaved as- normal  flavonoid  i n a c i d i f i e d - s o l v e n t 'systems., UV s p e c t r a " i h d i c a t e d  -monoglycosides  the 'normal  flavonoid  155  monoglycosides, and a c i d h y d r o l y s i s gave the common f l a v o n o l s quercetin, appearing.  Ckaempferol,  o r isorhamnetin) w i t h no B l u e - f l u o r e s c i n g , p h e n o l i c  acids  The sugar chromatograms showed o n l y g l u c o s e o r g a l a c t o s e .  A l k a l i n e h y d r o l y s i s o f p a r e n t compounds, u s i n g  1 drop o f cone. NK^OH,  l l B e r a t e d the c o r r e s p o n d i n g f l a v o n o l m o n o g l y c o s i d e s .  I n j e c t i o n o f the  r e a c t i o n m i x t u r e i n t o an HPLC (C-18 a c y l a t e d s i l i c a g e l , r e v e r s e revealed  o n l y two peaks:  the p a r e n t a c y l a t e d  ing  monoglycoside.  not  glucuronic a c i d , nor a phenolic  phase)  compound and i t s c o r r e s p o n d -  These r e s u l t s suggest t h a t the charged a c y l group i s a c i d , nor a sulphate  group.  I t must  t h e r e f o r e Be an a l i p h a t i c a c i d . The  second k i n d o f a c y l a t e d f l a v o n o i d was t h a t which, i n t h e  o r g a n i c s o l v e n t system, .had an  value higher  than was e x p e c t e d from an  i d e n t i t y Based on UV s p e c t r a l and a c i d h y d r o l y s i s data. i n d i c a t e d t h e common f l a v o n o l monoglycosides. only k a e m p f e r o l , q u e r c e t i n o r i s o r h a m n e t i n .  UV s p e c t r a l d a t a  Acid hydrolysis yielded No B l u e - f l u o r e s c i n g  compounds  appeared and the sugar chromatograms showed o n l y g l u c o s e o r g a l a c t o s e . I n t h e aqueous s o l v e n t systems the compounds Behaved as t h e common f l a v o n o l monoglycosides.  HPLC Behaviour confirmed t h i s o B s e r v a t i o n .  As B e f o r e ,  a l k a l i n e h y d r o l y s i s l l B e r a t e d the c o r r e s p o n d i n g f l a v o n o l monoglycosides. These r e s u l t s suggest that there  i s an a c y l group a t t a c h e d  moiety i n t h e parent g l y c o s i d e .  Unlike  the f i r s t  t o t h e sugar  case, the a c y l f u n c t i o n  i s n o t charged.  The e v i d e n c e i n d i c a t e s t h a t i t i s a l i p h a t i c r a t h e r  aromatic.  i t s e f f e c t i s to i n c r e a s e R^'s i n the o r g a n i c  Since  l i k i h o o d i s t h a t the a c y l m o i e t y i s an a c e t a t e the kaempferol v a r i a n t w i t h a confirmed t h i s The In Both o r g a n i c  group.  than  system, the  Chromatography of  standard kaempferol-3<-0-acetylglucoside  hypothesis, t h i r d k i n d o f a c y l a t e d f l a v o n o i d was.that whose R^ v a l u e s and aqueous- s o l v e n t  systems was r o u g h l y h a l f t h e expected  156  v a l u e based kaempferol  on i d e n t i t i e s i n f e r r e d from UV s p e c t r a l d a t a which i n d i c a t e d and q u e r c e t i n - 3 - 0 - g l y c o s i d e s .  A c i d h y d r o l y s i s y i e l d e d kaempr-  f e r o l o r q u e r c e t i n as the o n l y p h e n o l i c p r o d u c t .  The sugar a n a l y s e s y i e l d e d  g l u c o s e and a d u l l g r e y ^ c o l o u r e d compound w i t h h i g h R^ ( c a . .95). T h i s was concluded to be the a c y l group;  i t s n a t u r e was not determined.  Results The  r e s u l t s o f the f l a v o n o i d survey are g i v e n i n T a b l e XXI where  the d i s t r i b u t i o n o f 144 f l a v o n o i d s among 17 taxa i s shown.  Apart  from  S u k s d o r f i a . the d i f f e r e n t p o p u l a t i o n s o f each taxon showed, q u a l i t a t i v e l y , remarkably  similar profiles.  The f l a v o n o i d  complement o f f l o w e r i n g , growth  room-grown m a t e r i a l was the same as t h a t found i n m a t e r i a l c o l l e c t e d the f i e l d .  Furthermore,  from  d i f f e r e n c e s i n the stage o f p l a n t development had  only q u a n t i t a t i v e e f f e c t s .  With the e x c e p t i o n o f S u k s d o r f i a , p o p u l a t i o n s  sampled from n a t u r e i n s u c c e s s i v e y e a r s a l s o showed q u a l i t a t i v e l y  invariant  profiles. Only i n S u k s d o r f i a d i d i n t r a - s p e c i f i c q u a l i t a t i v e occur.  Some S_. r a n u n c u l i f o l i a p o p u l a t i o n s possessed  f l a v o n o l g l y c o s i d e s ; others d i d not.  In  variation  the charged a c y l a t e d  v i o l a c e a t h e r e was some i n t e r -  p o p u l a t i o n a l v a r i a t i o n i n the t r i g l y c o s i d e complement where two o f the three q u e r c e t i n - 3 - 0 - r h a m n o s y l g l u c o s i d e - 7 - 0 - g l u c o s i d e isomers were n o t u n i formly present.  I t i s u n f o r t u n a t e t h a t t h i s genus proved  t o be m o s t . d i f f i -  c u l t t o m a i n t a i n i n c u l t i v a t i o n and so f l a v o n o i d i n f o r m a t i o n from t i o n s grown under u n i f o r m c o n d i t i o n s was not a v a i l a b l e .  popula-  The•inter-popula-  t i o n a l v a r i a t i o n may be a r e s u l t o f p h e n o t y p i c p l a s t i c i t y r a t h e r than the r e s u l t o f any g e n e t i c d i f f e r e n c e s because i n S_. r a n u n c u l i f o l i a , a second sampling  two y e a r s l a t e r o f one o f t h e p o p u l a t i o n s (Bohm 1092 and G o r n a l l  245) showed t h a t the charged the se cond ' time.  compounds-, u n d e t e c t e d  a t f i r s t , were p r e s e n t  eCD  LO  ri O  fD rt  — 1  I O.  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M ^  •<! H  O CD HCu fD  i  I  Co 1  o I  1  11 4  4  o  CO V!  M  CN CU  Co ]  8= o1 .O  CO  JO I  rs  P  P  o  CO HCu fD  1— cu o rt o  X!  ho CO HCu fD  1  1  vi  1  O  o]  1-f  P  Cu fD  H  CD  oI1  00  cf  HP o  CD HCu fD  o1 3  00  H  1  o1  1  1 00  1  oo  00  o o  n o  CO HCu fD 1  vi  1  n o  1  oc o  CD HCu fD  c  M 0  h-  c  CD HCu fD  o1 3  O  O  H P n o  o1  vi  1  o1  Co  1  VI  CD HCu fD  1  CO  1  1  1  1  1  Co  CO hCU n rt O CD HCu fD  4  CU p o CO HCU fD  SL  00  CO  1  o1  n o  CO  CO  1  vj  CO H" Cu fD  1  1  CO  1 00  M P  «o JO JO 1 1 I  .O  i  o  CO  00  cu M Co o rt O CD H4 S fD  1  o  CD  OO M 0 O o  CO Pr CU fD Cl  JO JO 1 1 JO JO Co  Co 1 1  CO J J  o1 oI 3 -si  Cu  1  1 1-i  CO i•  O CD  H-  1  00  M 0  O o  cn H* Cu fD td  J1  n  cn h-  1  h0 D O CO HCu fD  1  >  CO H« Cu fD  o p  o  4  H0 O CD h-  1  00  H  0  H-  p?  §•  O  00  o  r-> CO  < o  Cu CO  o  n  o  CD M 00 CO I— Cu o rt O CD HCu fD-  0  1  0  O  o  CD HCu fD  Boykinia i  i  I  i  + + + 1 + + +  1  +  1 1  + 1  1  1  1  1  1  1  1  1  1  1  1  1 1  1  +  +  +  +  +  1  elata  +  major  1  +  intermedia  1  +  1  1  +  1  1  1  1  1  +  aconitifolia';.  (ureceolate)  +  1 1  +  1  1  1  1  1  +  aconitifolia  (turbinate)  1  1  1  1  1  1  lycoctonifolia  1  1  1  1  1  1  1  rotundifolia  1  1  +  1  1  1  1  richardsonii . ..a, b jamesxi  1  1  1  1 1  + 1  1  I  1  1  )  1  1  1  +  1  I  I  1  t  i  l  l  I  +  1  1  1  1  heucheriformis Peltoboykinia  1  1  1  1  1  1  1  I  1  1  tellimoides  1  1  1  1  1  1  1  1  1  1  watanabei  tellimoides  Suksdorfia 1 1 1  1  1  1  1 1  1  1 1  1  1  1  1  1  1  1  ranunculifolia  1  1  1  1  1  1  violacea  1  1  1  1  1  +  3  alchemilloides Bolandra  1  1  1  1  1  +  1  1  +  +  oregana Sullivantia  1  1  1  1  1  1  1  1  1  1  e9i  oregana  3  7  Lo 1  O  I 00 P  o o  co  H» CU  fD I •vj I  o J  oo e  Lo I . O  I  P  4  1  1 Lo  LO  O  O  1  I  A J HO  ?  Lo  J  cn  O  cn h-  1  3 O  cn  H> Cu  fD  O O  X I—  cn  I  O CD  1  00  Co M Co  n  rt O CD HCu fD  00  00  C  c o o  o o  cn Cu  fD  oL Co rt  1  O  7  Lo  CD  eu fD  &  LO  Lo  )  O  O  O  H4  H4  M4  H4  Co  Co  Co  CD  c  o  p o  HCu fD  P  o  h-  CD  h-> H  C  o o  CD  >*• Cu  cn  LO  O CD  00  1 J  Co t—' Co  o  rt O CD rCU fD 1  I  1  P  P  O. CD  O  h-  00  00  00  C  e o o  P  M  cn  H  o o  H» Cu fD  fD  LO  O  o  I I  1  oo  n o  n o  C  cn  r-  1  C  CO  oo oo C c a o o o  co  HCu  ro  ro w  15  M CO < O  1  OO H  Cu  ro  W  Lo  CD  CO H* Cu  cn  Cu  cn  COM  1  o o  I i-i  P  cn  1  00  1  1  tn  HCu  Lo 1 O  ) o  Cr H-  P  7.  LO  1  o  ff 1  JO  P  o  HCu CD O  O O  I  P  LO  1  o  1 00 H  s o o  CO hOO M  1  0  fD  n o  co  H' Cu fD  Boykinia )  ]  1  1  1  1  +  1  1  1  +  1  +.  1  +  1 1 1  +  1  +  1  + 1  +  +  1  1  1  elata  +  1  1  1  maj or  1  +  +  +  1  intermedia  1  +  1  1  1  aconitifolia  (ureceolate) (turbinate)  +  1  1  +  1  1  1  1  1  +  J  +  1  1  1  1  1  +  1 1 1  +  1  1  1  aconitifolia  1  1  1  1  1  1  1 1 1  +  1  1  +  lycoctonifolia  1  1  1  1  1  ]  1  1 1 1  +  1  1  1  rotundifolia  1  1  1  1  1  1  1  1  +  +  1  1  1  1  )  1  1  +  1  I  1  1  1  +  1  1  richardsonli . ..a, b jamesii  1  1  1  1  +  1  1  l  l  +  1  +  +  1 l  l  heucheriformis Peltoboykinia tellimoides  1  1  1  1  1  1  1  1  ]  1  1  1  '  1  1  l  l  1  1 l  l  1  1  1  tellimoides  1  1  1  watanabei Suksdorfia  1  1  1  1  1  1  1  1  1  1  1  1  1  1  1  1  >  +  1 1 1  +  1  1  1  ranunculifolia  1 1 1  +  1  1  1  violacea  1 1 1  +  1  1  1  alchemilloides  3  Bolandra 1  1  1  1  +  Hh  +  1 1 1  +  1  1  1  oregana Sullivantia  i  i  i  i  i  i  i  i  i  i  +791  oregana  SL  •I LO )  o  1 OP h0 o o CO 1  M OP H P  O O CO H* CU ro i  T 1  o  ) OP M  c n o CD ro  OP M P  LO  I o 3 OP ro M i  M O CO  g  o o CD ro  OP Co H  I  i-f  D rt  I OP M  PJ  o  c o  p O CD ro ^—'  r-  1  SB  O CD HCu ro 1  1  o  o o CO HCu ro  X)  1 LO 1 O 1 l-i P*  p O CO M OP CO 1—  1  Co  o rt  O CD HCu ro J  e  .O  1 Lo  V  r -  p O CD  O CD  cj p  hOQ he  Cu ro  rt  1  P  n  o CD HCu ro 1  o  J O 1  rt rl  1  I LO 1  1 l-i  o 1 o Lo CO 1 r - O Cu 1 ro OQ Co hCo n 1  JO  1  O CO H> Cu ro CO  p rt H»  Hi H> ro Cu ^ '  J o 1  P  4  Co  rt P*  Co p O CD HCU ro i i o 1 OQ M 0 O o CD  1 LO 1  I LO 1  «<3 1 LO 1  JO  i Lo 1  1 Lo 1  J  J  1  1 H  1 H  Co  Co  3 o CD  p O CD  O  X  V! M O CO H  1  O  M o CD r-  1  OQ Co  OQ  Co O  O o CO H» Cu ro i  rt  O CD HCu ro i ~j i o i OQ M  e  o o CD  >*  P  O  % ho CO 1  hOP h-  1  1  c o  o CO H> Cu ro i  ~J  i o i OQ  i o 1 OP  P n o CD  o  Cu ro  Cu ro  w  >  M  n.  M  P  o CD  O  P*  hOQ Co H Co O  M  O  <COo  P*  o  H<  P  H*  Cu CO  O  o  p rt  1  rt  O CO H»  Cu ro 1 1 o 1 OP — I 1  p  o  o CO  de  ro  >  JO JO  OP H C o o CO H-  CL.  ro 1  1 o 1 OP M  p  o o CO  ri-  de  1 o  1 o 3 OQ ro M 1  de  1  P  Lo  id  o CO H' Cu ro  p*  LO  1 O 3 ~0P ro i ; cX5 o ] . o LO .'CO 1 O CU 1 ro %  a  Boykinia  +  +  I  1  elata  +  I  major  +  I  intermedia  1  +  aconitifolia  (ureceolate)  I  +  aconitifolia  (turbinate)  I  +  lycoctonifolia  1  1  1  1  1  +  I  I  rotundifolia  +  richardsonii . .a, b jamesxx heucheriformis  I  1  Peltoboykinia  I  I  tellimoides  tellimoides  watanabei I  1  I  I  +  1  I  +  I  I  I  1  I  +  I  I  I  I  Suksdorfia ranunculifolia violacea  3  alchemilloides I  I  I  +  I  Bolandra  I  oregana I  I  I  Sullivantia  I  oregana  C  9T  3  Ox  Ox  1  1  o  OQ  g. fD 1  00  i  P o  \-*JD e rt  o o  LO  CO J HO Cu 1 fD  O  M  M O CO  M  OO H C  o o  CO r' Cu fD  s. fD 1  rt  1  LO  to 1 H- O 'CU 1 'fD  P  1  Lo  O  O  1  1  X* H O CO  X  cu hcu O rt O CO HCu fD 1  3 P O CO  X  h-> 00 h1  P  1  o  VI  O  vl  CO rCu ft)  1  1  P  1  o  1  1  O I  o  1  VI  o  1  1  1  X>i  M O ' CO  ' K00 M  1  oo  4  7  LO  1 1  00 r-  1  o  o  HCu fD 1  vj 1  o1  00 HC n  1  o  CO  H-  Cu fD  •i  ?1  )  1  Lo  Lo  LO  Lo  o J  O  O  O  O  O  1  1  1  1  p  P  P  P  I-i  4  CU S3 p  o.  co  h00 cu hfu  1  O  »-f  cu  cu  4  p O CD  00 M C  vi  O  O  1  00  M p o o  CD Cu fD  CO H" Cu fD 1 1  CO  o  1  p o o  o  P o o  CO HCu fD 1  vi  I  00 H  P o o  CD HCU fD  CO H> Cu fD  a  n  »i  P  4  g p  O CD  1  )  OO CU M cu o rt  o  oo  H 00  CD  c  P  I— 00 h-  o  n  h-  1  1  X  1  1  00 1—  4  CD  O  )  i-i  Cu p  h-  o o  1  p  oo  ~J  1  4  M  rt O CO HCu fD  I  J  i-i  1  1  co  J  LO  1  M  o o  CD HCU fD 1  vi 1  O  J  00 M  P o o  CD HCU fD W  1  1  c  O  CD H* Cu fD  o  1  CO HCu fD  1  vi  1  O  vj O  1  1  00 M  00  o oO C  P o o  P  r' CU fD 1  1  CD H>  de  CO HCu fD  P  ?1  Lo  >  1  M  O  <O  00  O  Lo  1  r->  p o  o  CO  tr 1  P  HCu CO  o  o  o p  M  00 h-  1  p o  o  CO  H-  Cu fD  I  vl  1  O  1  00 h-  1  P  O  o  CO  H-  Cu fD  td  Boykinia 1  +  +  +  + + +  1  -+  elata  1  major  I  1  1  1  +  I  1  1  i  +•  -J  intermedia  1  1  1  1  1  1  aconitifolia  (ureceolate)  1  1  1  .  1  '1  aconitifolia  (turbinate)  1  1  1  1  1  J  lycoctonifolia  1  1  1  1  1  1  rotundifolia  I  1  1  1  I  1  1  1  1  1  1  1  richardsonii . ..a, b jamesn  1  I  >  I  1  1  heucheriformis Peltoboykinia tellimoides  i  1  .  1  1  •1  1  tellimoides  i  1  1  1  )  1  1  watanabei Suksdorfia  i  1  1  1  i  +  1  1  i  1  1  1  1  1  ranunculifolia  +  1  1  violacea  -4-  1  1  alchemilloides  1  3 .  Bolandra 1  1  1  1  I  I  oregana Sullivantia  1  1  1  1  ]  I  991  oregana  cl  1  LO  1 o 1  H  pr  CO  p o  CO  JO  1  OJ  M  1 o 1  H  pr  CD  p o cn  ^M  P  O O CO H*  fD  fD  PL  )  ~j 1 o 1  1  P  O O  PL  1  1  O  1  OQ h-  1  P  O O CO  OQ  1—  1  P O  o CD FPL  ro i i o I  1  P O  H-  F-  fD  fD  fD  PL  o  PL  W  •O 1  1 o 1  1 o 1  OJ  OQ., Co  PL  >  OJ  OQ  M P  It  i  OJ  1 o ]  rOQ M  M OQ  F> P o  o o  F-  CO  PL  fD  fD  /  o F-  F-  N  P P  PL  fD  P  •^J  ~J  O  Frt) F-  o  OQ  I  OQ h-  PL  1 1 1  F  1  P  O O CO  PL  F-  1  O CO  P  CO  h-  P  t—*  fD  V  fD  rt  fD  PL  1  P  OQ h-  1  P n  OJ  1 o  o  CO  fD  g  fD  -o  1  ON  1 o  —  )  o g  fD  1  1  Q  i  it  1  ON  1 o 1  ON  1  P  CD  h-  Pi p  fD  F-  OQ M  P  O O CD  FPL  fD  PL  1 o  g  ro -1  —  I o  fD  1  JO it  J Lo  fD  OJ  O ] OQ M  O g  1  1  -O rt  1  I o 1 H  OQ M  1 o 1  o o  OQ  OQ H C  CO  CO  o o CD FPL  fD  ro  1—  o o  PL  pr  1  o  M  o  O  OQ H  fD  PL  CO  CL  1  O  1  1  P  o o  J  cn.  1  HPL  P O  it  fD 1  OQ h-  CD  O O P  PL  P  ro 1  P  F-  co F-  ~J  OQ  tr  cn  o  F"  O CO  O  o F'  Cp  1  -J  o F-  P  n  <! o p o  1  O  p  Pi  I— OQ M 1  J  n o CO F-  1  P  r Co  LO  It  1  pr  1  pr  ** •O  ON  ON  OJ  P  o o  OJ  V  o  1  OQ I-  w  1-f  OJ  C7>  1  O CD  O O CD  o  CO  CD  O  OQ  CO  o CO FPL n> I  1 o 1  p.  n  F-  1  OJ  H H  OQ  PL  O CD  <o 1  It M  F-  O O CD  I  OQ h-  o CD F-  cn  1  Co O  CO  1  o cn F-  PJ  o  1  P  pr P  h-  O  OQ h-  OJ  1 o 1  hOQ  M OQ  JO  I  fD  o CD FCL  fD  CL  fD  fD  Boykinia I  1  +  1  +  +  +  elata major intermedia aconitifolia  (ureceolate)  aconitifolia  (turbinate)  lycoctonifolia rotundifolia richardsonli . . a, b  •jamesii  + + 1  1  I  I  heucheriformis Peltoboykinia tellimoides tellimoides watanabei  1  I  I  +  +  +  +  I  I  I  I  +  +  I  I  I  I  I  1  I  I  Suksdorfia ranunculifolia violacea  3  alchemilloides 1  Bolandra  I  oregana I  I  I  I  +  I  +  I  Sullivantia  I  oregana  L91  a  a  1  TABLE XXI, cont'd.  a  Not grown i n growth chamber.  b  F l a v o n o i d s i d e n t i f i e d by co-chromatography o n l y , owing t o the s m a l l amount o f m a t e r i a l ; the l i s t o f compounds may t h e r e f o r e be incomplete,  c  A b b r e v i a t i o n s and s t r u c t u r e s are g i v e n i n F i g . 43.  d  '-' = n o t d e t e c t e d ;  e  .Charged a l i p h a t i c a c y l a t i n g f u n c t i o n .  f  A c y l group appears as a grey spot  h  Absent from c o l l e c t i o n s Bohm 1092, 1096, 1132, 1144.  j  Absent from c o l l e c t i o n s Bohm 1112, 1118, 1122.  '+' = p r e s e n t ;  'g' = found o n l y i n g l y c o s i d i c form,  (Rf = .95) on sugar  chromatograms.  00  169  FIGURE 43. Key t o f l a v o n o i d s t r u c t u r e s and a b b r e v i a t i o n s r e f e r r e d t o i n the t e x t and T a b l e s .  M = M y r i c e t i n ( R ^ R ^ R ^ O H ; R =H)  Gt = G a l e t i n (R =R^=OH; R ^ R ^ H )  Q = Quercetin  L = Luteolin  (Rj=OH; R =R =R^=H)  A = Apigenin  (R^R^R^R^H)  4  3  ( R ^ R ^ O H ; R =R =H) 2  4  K = Kaempferol (R =OH; R ^ R ^ R ^ H ) 3  Qt = Q u e r c e t a g e t i n  (R =R =R^=OH; R =H) 1  3  2  2  3  Sc = S c u t e l l a r e i n (R^OH; R  1  = R  2  = R  3  = H )  OMe = methyl e t h e r Examples: 3'-0Me-Q = q u e r c e t i n - 3 ' - m e t h y l 3,7-dimethyl e t h e r .  e t h e r ; 3,7-OMe-Q = q u e r c e t i n -  170  A s e t o f poly-Orvmethylated f l a y o n o l s and f l a y o n e s was found i n many t a x a . But  They are noteworthy n o t o n l y f o r t h e i r taxonomic s i g n i f i c a n c e  a l s o Because they were o f t e n n o t r e p r e s e n t e d  flavonoid p r o f i l e , u n l i k e the'other  aglycones.  as g l y c o s i d e s i n the Experience  with  glycosides  of s i m i l a r compounds from Chrysosplenlum (Bohm and C o l l i n s 1979) has shown t h a t h y d r o l y s i s does n o t occur d u r i n g the e x t r a c t i o n and p u r i f i c a t i o n cedures.  I t seems t h e r e f o r e , t h a t these  compounds o f t e n occur  pro-  free i n  the p l a n t . In order  to s i m p l i f y T a B l e XXI f o r taxonomic purposes,  import-  ant s t r u c t u r a l f e a t u r e s have Been s e l e c t e d and t h e i r d i s t r i B u t i o n shown i n TaBle X X I I .  S e v e r a l p o i n t s are worth m e n t i o n i n g .  6-0xygenation i s aBsent  from S u k s d o r f i a , P e l t o B o y k i n i a , B o y k i n i a i n t e r m e d i a , 6^0xygenated f l a y o n e s occur The  Acetylated f l a v o n o i d s occur  and/ S u l l i v a n t i a oregana. arabinose  Bolandra,  one e x c e p t i o n ,  r e s t r i c t e d to  only i n Boykinia ( s p o r a d i c a l l y )  G l y c o s y l a t i o n a t the d i g l y c o s i d e l e v e l , i n v o l v i n g  and/or x y l o s e , occurs  P o l y g l y c o s y l a t i o n occurs  oid  i n Both B o y k i n i a r o t u n d i f o l i a and S u l l i v a n t i a .  charged a c y l a t e d f l a v o n o i d s a r e , w i t h  Boykinia.  and p o s s i B l y B_. j a m e s i i .  o n l y i n B o y k i n i a and B o l a n d r a .  3,7-0-  throughout B o y k i n i a , S u k s d o r f i a , and p o s s i B l y  But i t i s aBsent from S u l l i v a n t i a and P e l t o B o y k i n i a .  syndrome i n P e l t o B o y k i n i a i s remarkaBly  The f l a v o n -  simple.  DISCUSSION Disease  Resistance The  flavones  I n many  occurrence  of unglycosylated polymethylated  o f the s p e c i e s  I sInteresting  f l a v o n o l s and  s i n c e t h e i r presence i n t h i s  form makes them good c a n d i d a t e s : f o r a r o l e i n the p r o t e c t i o n o f the p l a n t a g a i n s t c h r o n i c d i s e a s e (McClure 1975). notaBle  As- c i r c u m s t a n t i a l  evidence,  i t is  that the o n l y B o y k i n i a s p e c i e s s u b j e c t t o i n f e s t a t i o n By r e d - s p i d e r  I  o I  rt i-i HO  ft rT o1 I  0  0  cn  cn  O  ro  cn H" Cu H fD O cn cn fD  O  ro  3  0  O o cn H"  Cu  o  cn fD  cu  0  H O»  0O  cn HCu fD cn  OQ  fD cn  I O  J  OQ  h-  O O  1  cn (-"• (Cu D cn  f?  9  fD  pj i-i  M fD  Cu OQ H  O  JQ  fD Cu CU o  ON CO < O  0  fD cn  1 3  fD  1  H* O  l-ti  OQ  Co rt  cn  i-i C O rt C l-i Co  ro  rt  P<  P-  O  fD rt H-  0  HO  0  fD PJ rt  O  0  0  o  5  rt  C  I— Co rt  H  OJ  I  0  CD CO  H-  Cu  fD cn  o  cn H* CU ro cn  L-  1  PJ >-i rt Co cn 0  Co H  Boykinia  +  1  1  1  +  1  1  +  +  +  +  1  •1  +  +  +  +  1  1  1  +  +  +  1  1  +  +  +  1  +  +  +  +  +  + +  +  4-  1  elata  1  + + •+  1  major  •+ +  1  1  1  1  1  intermedia  ]  +  1  1  +  1  +  aconitifolia  (urceolate)  1  1  +  1  1  +  1  aconitifolia  (turbinate)  1  1  +  +  1  +  +  +  1  1  1  +  +  + +  +  1  1  1  1  1  +  1  +  1  l  richardsonii  +  1  1  1  +  +  1  1  1  1  1  jamesii  + +  1  1  +  +  1  + + +  1  h e u c h e r i f ormis  +  I  1  +  H  cn  lycoctonifolia rotundifolia  Peltoboykinia tellimoides I  1  1  1  1  1  +  1  1  1  1  1  tellimoides  1  1  1  1  1  I  +  I  1  ]  1  1  watanabei Suksdorfia  P  4  o  S3  H-  0  OQ  rt P fD 4  O  P  o  3 o Cu  cn rt l-i Hcr 0  rt  HO  0  +  1  1  1  +  1  +  +  +  1  +  1  ranunculifolia  o  +  1  1  1  +  1  1  +  1  1  +  1  yiolacea  cn  +  1  1  1  1  1  1  1  1  1  1  1  alchemilloides  8  Bolandra  +  +  1  1  1  1  +  1  +  +  +  oregana Sulliyantia  1  1  1  +  1  +  1  +  1  +  1  1  oregana  Mi  O  I  O  i-i rt  B rt  Ml  hCo  1  < O  0  o  H>  Cu  UT  172  mites i n the growth chamber were -JL. r i c h a r d s o n l i , B_. a c o n i t i f o l i a , and intermedia. Stability  These taxa l a c k the polymethylated  of  flavonoid  aglycones.  Expression P o p u l a t i o n a l sampling on a wide g e o g r a p h i c a l  w i t h s t u d i e s of the same m a t e r i a l from the u n i f o r m growth chamber, and  taken i n  t h a t , except i n the genus S u k s d o r f i a ,  p r o f i l e s of each taxon are q u a l i t a t i v e l y  constant.  v a r i a t i o n i n S u k s d o r f i a s p e c i e s i s minor and suggest t h a t i t may  scale,  be e n v i r o n m e n t a l l y  coupled  enviroment of  samples o f i n d i v i d u a l p o p u l a t i o n s  y e a r s , have r e v e a l e d  s u i t a b l e f o r taxonomic  the f l a v o n o i d  Inter-populational  there i s some e v i d e n c e to  controlled.  The  important  concluare  use,  Conclusions The  major c o n c l u s i o n i s t h a t many and v a r i o u s  l a r i t i e s e x i s t among the ships.  the  successive  s i o n s to be drawn.from these r e s u l t s o v e r a l l , i s t h a t f l a v o n o i d s  Taxonomic  B.  taxa, s u g g e s t i n g  They are o b v i o u s l y  flavonoid simi-  somewhat r e t i c u l a t e  a group a c l o s e l y r e l a t e d s p e c i e s .  relationMuch of  t h i s k i n d of v a r i a t i o n r e s u l t s from e v o l u t i o n a r y p a r a l l e l i s m , a phenomenon common i n the S a x i f r a g i n a e . allies  (Bohm and W i l k i n s  CBohm and  Ornduff 1978;  The  group i s d i s t i n c t  1978a, b ) , and Bohm and W i l k i n s  from Heuchera and i t s  from J e p s o n i a 1976)  combinations.  are shared by S a x i f r a g a and  and  3<-0-methyl-  a c e t y l a t i o n of  3 - 0 - M e t h y l a t i o n and  ( M i l l e r and Bohm 1980)  Peltiphyllum  i n possessing  a t i o n , 6-oxygenation, charged a c y l a t i n g f u n c t i o n s , and glycosides i n various  and  6-oxygenation  Chrysosplenjum CBohm  C o l l i n s 1979.) . D e a l i n g wjtth i n f r a s p e c i f i c taxonomy, the common s i t u a t i o n i n  the S a x i f r a g i n a e i s f o r species^ to show" q u a l i t a t i v e l y i n v a r i a n t p r o f i l e s Ce. g. Bohm and W i l k i n s 1978b) ,- T h i s p a t t e r n -'is maintained- i n B o y k i n i a :  o c c i d e n t a l i s where populations- sampled throughout i t s ^ range d i s p l a y e d  173  i d e n t i c a l flavonoids. segregate  species  a constant  Thus no  support  CRydBerg 1905).  i s g i v e n to the r e c o g n i t i o n of  S i m i l a r l y , B o y k i n i a a c o n i t i f o l i a shows  f l a v o n o i d p r o f i l e ; t h i s argues a g a i n s t r e c o g n i t i o n of v a r i a n t s  from AlaBama and  G e o r g i a , w i t h t u r b i n a t e r a t h e r than campanulate  at s p e c i f i c l e v e l as B,  capsules,  t u r B i h a t a CRydBerg 1905).  T h i r d , the r e c o g n i t i o n of B_, i n t e r m e d i a  at s p e c i f i c  Cones 1936)  r a t h e r than as a v a r i e t y of B_. major C ^ i p e r 1899;  e t a l . 1961)  i s supported  By  f l a v o n o i d data.  Thus B.  level Hitchcock  intermedia  lacks  3-0-methylation, 4'-0-methylation and 6-oxygenation, a l l of which t e r i z e B_. major. hyBrtd  charac-  B o y k i n i a i n t e r m e d i a has Been suggested to have had  o r i g i n , d e r i v e d from B. major and B. o c c i d e n t a l i s CPiper  However, i t s f l a v o n o i d p r o f i l e does not r e p r e s e n t i t s putative parents.  The  a  1899).  a summation of those  of  s y n t h e s i z e d h y B r i d B_. major x B_, o c c i d e n t a l i s  does show such a d d i t i v e i n h e r i t a n c e .  A h y b r i d o r i g i n f o r B_.  intermedia  Is therefore rejected. F o u r t h , i t i s c l e a r t h a t B o y k i n i a j a m e s i i has onoid  complement to t h a t o f B_. h e u c h e r i f o r m i s .  a similar  However, the small, amount  of m a t e r i a l a v a i l a b l e d i d not r e v e a l d e f i n i t e l y whether the f l a v o n o i d s were p r e s e n t i t would support  or not i n B_. j a m e s i i .  polymethylated  I f t h e i r absence i s genuine,  the r e c o g n i t i o n of both taxa at s p e c i f i c l e v e l  1897), r a t h e r than as two v a r i e t i e s CEngler With r e g a r d  flav-  (Rydberg  1928).  to s p e c i e s r e l a t i o n s h i p s and g e n e r i c l i m i t s ,  f l a v o n o i d d a t a are ambiguous,  the  Thus, v a r i a t i o n w i t h i n the v e r y n a t u r a l  B o y k i n i a s e c t i o n B o y k i n i a i s s u c h t h a t some of the other taxa s t u d i e d could Be  comfortaBly  accommodated w i t h i n i t .  I t does, however,  support  the placement of ff. j a m e s i l and;K,.heucheriformis i n B o y k i n i a , r a t h e r than i n S a x i f r a g a as suggested By Jones O-910) and H a r r i n g t o n  C1954).  Saxifraga  174  has  so f a r not been r e p o r t e d to c o n t a i n the  charged or a c e t y l a t e d f l a v o n -  o i d s which are found i n B o y k i n i a . E n g l e r .(1928) p l a c e d R.  j a m e s i i and B. h e u c h e r i f o r m i s  s e c t i o n R e n i f o l i u m along w i t h B_. r i c h a r d s o n i i . d a t a suggest that they r e p r e s e n t  the end  chemical  One  separate  lines  of  l i n e , whose c h i e f  t r e n d has been r e d u c t i o n , l e a d s to the f l a v o n e - r i c h B_. r i c h a r d -  s o n i i , endemic to the Yukon and A l a s k a . gone much chemical  considered  The  other  l i n e , which has  d i v e r s i f i c a t i o n , l e a d s to the p o l y m e t h y l a t e d  b e a r i n g B_. h e u c h e r i f o r m i s best  However, the f l a v o n o i d  p o i n t s of two  e v o l u t i o n , t r a c e a b l e back to s e c t i o n B o y k i n i a .  in  i n the Rocky Mountains.  as b e l o n g i n g  to two  separate  The  F l a v o n o i d d a t a a l s o suggest t h a t B^  flavonol-  s p e c i e s are  sections within  under-  thus  Boykinia.  lycoctonifolia is well-  p l a c e d i n s e c t i o n B o y k i n i a , r a t h e r than i n a montypic genus N e o b o y k i n i a (Hara 1937). F l a v o n o i d d a t a support a distinct  genus.  The  the r e c o g n i t i o n of P e l t o b o y k i n i a as  s i m p l i c i t y of i t s f l a v o n o i d p r o f i l e i s unique i n  t h i s group of genera, although i n d i c a t e that i t i s r e l a t e d .  the p r e s e n c e of charged f l a v o n o i d s F l a v o n o i d d a t a do not  r e l a t i o n s h i p w i t h P e l t i p h y l l u m (Bohm and W i l k i n s has  suggest any  certainly  close  1976), a genus which a l s o  p e l t a t e l e a v e s , and w i t h which an a s s o c i a t i o n has been p o s t u l a t e d i n  the p a s t  ( E n g l e r 1891). The  ilarities,  two  North American S u k s d o r f i a s p e c i e s show v e r y many sim-  i n c l u d i n g f o r example 4 ' - 0 r - g l y c o s y l a t i o n .  s p e c i e s haye sometimes- been put Engler  i n separate'genera,  A l t h o u g h the  Hamjeya and  two  Suksdorfia,  (1891) , on the §asl;s- of; t h e i r -vegetative s i m i l a r i t y , u n i t e d them  under S u k s d o r f i a ; f l a v o n o i d s unquestionablyThe  support  a congeneric  alignment.  thj-rd 'Suksd6rf 1 a s p e c i e s , SV a l c h e m i l l o i d e s - . l a c k s  f l a v o n e s t y p i c a l of the o t h e r two  species-, a l t h o u g h  the  i n v i e w of the v a r i a t i o n  175  found i n the r e l a t e d B o y k i n i a , ance.  t h i s absence i s not n e c e s a r t l y o f i m p o r t -  Indeed, the s i m i l a r f l a y o n o l t r i g l y c o s i d e  components i n b o t h S.  v i o l a c e a and S_. a l c h e m j l l o i d e s suggests a c l o s e r e l a t i o n s h i p , a t l e a s t i n this  respect. The  s e p a r a t i o n o f S u k s d o r f i a from B o y k i n i a ,  chiefly  on.the  B a s i s o f the former's b u l b i f e r o u s rhizome, i s supported by the absence o f b o t h 6-oxygenation and d i g l y c o s i d e s i n v o l v i n g arabihose  and/or x y l o s e  a l l S u k s d o r f i a s p e c i e s b u t t h e i r presence i n almost a l l B o y k i n i a S u l l i v a n t i a i s c l o s e l y r e l a t e d to Boykinia both and  chemically.  species.  morphologically  S i m i l a r i t i e s i n f l a v o n o i d s i n c l u d e 6-oxygenation and  a c e t y l a t e d monoglycosides.  Soltis  (1980) d i d n o t r e p o r t the l a t t e r i n h i s  study of the genus based on t h e use o f paper chromatography.  Differences  between the genera i n c l u d e the absence o f b o t h 3 , 7 - p o l y g l y c o s i d e s charged a c y l a t e d f l a v o n o i d s from S u l l i v a n t i a . Soltis  from  (1980) r e p o r t e d  a f l a v o n o l glucuronide  and  I t i s i n t e r e s t i n g that from S u l l i v a n t i a .  This i s  a charged compound b u t o b v i o u s l y d i f f e r s from those found i n t h i s  study.  Indeed, no charged compounds were found i n my m a t e r i a l of S_. oregana. Evolutionary Interpretation The operating  twin trends  o f r e d u c t i o n and d i v e r s i f i c a t i o n appears t o be  i n t h i s group of genera.  Thus, t h e r e i s a l o s s o f m y r i c e t i n ,  r e t a i n e d o n l y i n B o y k i n i a a c o n i t i f o l i a and B o l a n d r a oregana, and a tendency towards development o f f l a v o n e s  i n B_. o c c i d e n t a l i s , B. r o t u n d i f o l i a ,  B. r i c h a r d s o n l i , S u k s d o r f i a and S u l l i v a n t i a .  Superimposed on t h i s  reduc-  t i o n i s the d i v e r s i f i c a t i o n o f the f l a y o n o i d p r o f i l e by the development of 6^oxygenation and O r m e t h y l a t i o n •Cespecially a t p o s i t i o n 3)  f  Bearing i n  nijnd the d r e a d f u l p i t f a l l s - a w a i t i n g purveyors of f l a v o n o i d phylogeny ( G o r n a l l and. Bohm 1978) the most p r i m i t i v e p r o f i l e s are p r o b a b l y by B o y k i n i a  a c o n i t i f o l i a . B. i n t e r m e d i a ,  exhibited  P e l t o b o y k i n i a and to some extent  176  B o l a n d r a oregana.  Sugar e v o l u t i o n may  o r may  not be matched  w i t h t h a t i,n  a g l y c o n e s , and the i n t e r p r e t a t i o n of whether a predominance of g l u c o s e and rhamnose over o t h e r sugars is- p r i m i t i v e or advanced (see  i s l e f t to the r e a d e r  G o r n a l l and Bohm 1978),  Parallelism The genera s t u d i e d e x h i b i t f l a v o n o i d p r o f i l e s which show l a r g e amounts of p a r a l l e l v a r i a t i o n .  P r o b a b l y the most remarkable  t h i s p a r a l l e l i s m i s the d i s t r i b u t i o n of 3-0-methylation and i n the S a x i f r a g i n a e . Chyrsosplenium Sullivantia (the  One  (Bohm and  ( S o l t i s 1980)  present study).  example of 6-oxygenation  or both of these s t r u c t u r a l f e a t u r e s occurs i n C o l l i n s 1979) , S a x i f r a g a ( M i l l e r and Bohm and now  1980)  B o y k i n i a , B o l a n d r a , and S u k s d o r f i a  It is difficult  to say whether these f e a t u r e s e v o l v e d  i n d e p e n d e n t l y or whether some o f the genera d i d indeed e v o l v e from a common a n c e s t o r which a l s o possessed them.  The  absence  or r a r i t y of these  s t r u c t u r a l f e a t u r e s from p u t a t i v e l y p r i m i t i v e syndromes, v i z , i n B_. i n t e r media, B. a c o n i t i f o l i a and P e l t o b o y k i n i a , suggests an independent  origin.  P a r a l l e l i s m has i t s b a s i s i n an a n c e s t r a l gene p o o l ; the more c l o s e l y related  the taxa a r e , the g r e a t e r i s the resemblance  their variation.  i n the n a t u r e of  I t c l e a r l y supports the i n c l u s i o n of  whose p o s i t i o n has been i n doubt,  i n the s u b t r i b e  Chrysosplenium,  Saxifraginae.  The d i s t r i b u t i o n of 3-0-methylation and 6-oxygenation a c r o s s the t r a d i t i o n a l l y - r e c o g n i z e d d i v i s i o n of the s u b t r i b e based placentation;  Chrysosplenium,  cuts on  along w i t h Heuchera and a l l i e s , have p a r i -  e t a l p l a c e n t a t i o n , whereas the B o y k i n i a ^ ? a x l f r a g a group o f genera have axile placentation.  Howeyer, the f l a y o n o i d s i n q u e s t i o n have so f a r been  r e p o r t e d o n l y from t a x a w i t h ^ m o n o t e l i c i n f l o r e s c e n c e s .  ChrysospTe'nl-um,'  t h e r e f o r e , w i t h lts> p a r i e t a l p l a c e n t a t i o n coupled w i t h a m o n o t e l l e  inflor-  escence and p o l y m e t h y l a t e d f l a y o n o i d s , makes a good b r i d g e between the  177  'polytelic-parietal'  tteucheroid  and the ' m o n o t e l i c - a x l l e ' ylated  flavonoids.  genera, l a c k i n g p o l y m e t h y l a t e d  flavonoids,  S a x i f r a g o i d genera, some o f which have polymethr-  178  XIII,  ECOLOGY  I n this- c h a p t e r , h a b i t a t p r e f e r e n c e s , seed d i s p e r s a l mechanisms , p o l l i n a t i o n syndromes and r e l a t i o n s h i p s w i t h d i s e a s e s and p r e d a t o r s w i l l Be d i s c u s s e d . • The i n f o r m a t i o n i s based on my own f i e l d i n western North  America and on data on annotated  herbarium  observations specimens.  HABITATS' Boykinia I n North America, s p e c i e s o f s e c t i o n B o y k i n i a grow i n wet, woodland h a b i t a t s along the margins o f streams, r i v e r s , ponds and l a k e s i n the Western C o r d i l l e r a and the A p p a l a c h i a n Mountains.  The s p e c i e s do n o t  grow i n deep shade b u t f a v o u r semi-open o r green shade s i t u a t i o n s . are d i f f e r e n t i a t e d somewhat i n t h e i r a l t i t u d i n a l t o l e r a n c e s :  They  t h u s , B.  a c o n i t i f o l i a occurs between 300 - 1000m; _B_. i n t e r m e d i a i s u s u a l l y found below 700m; B_. o c c i d e n t a l i s ranges from s e a l e v e l t o  1400m,^exceptionally  1700m; B. r o t u n d i f o l i a occurs between 800 - 2000m; and B^ major u s u a l l y grows above 1000m b u t below 2200m, b e i n g '"commonest a t around 1700m. B o y k i n i a i n t e r m e d i a has a l a r g e moisture  requirement,  and i s r e s t r i c t e d to  c o a s t a l areas o f Washington and n o r t h e r n Oregon where the r a i n f a l l 250cm p e r annum•. water.  One p o p u l a t i o n ( G o r n a l l 24) was found  exceeds  growing i n running  Both B_. o c c i d e n t a l i s and B_. r o t u n d i f o l i a show some p r e f e r e n c e f o r  d i s t u r b e d , bare s o i l , e.g. t r a i l s i d e s and mud The  remaining  slides.  s p e c i e s o f s e c t i o n B o y k i n i a , B. l y c o c t o n i f o l i a ,  i n h a b i t s s e a s o n a l l y wet, snow-bed g r a s s l a n d i n the a l p i n e zone o f mountains i n n o r t h e r n Japan (Ohwi 1965; I s h i z u k a 1974). 2000 - 2800m, on v o l c a n i c s o i l s  I t i s u s u a l l y found between  (Kara 1959) .  B o y k j n i a r i c h a r d s o n l i ( s e c t i o n R e n i f o l i u m ) , an A l a s k a - Yukon endemic, grows i n stream-side  g u l l i e s and snow-bed g r a s s l a n d communities  i n p r o t e c t e d d e p r e s s i o n s where snow l i e s f r e q u e n t l y i n t o mid-June. I t s  179  h a b i t a t appears to be v e r y  s i m i l a r to t h a t o f B_. l y c o c t o n i f o l j a .  Boykinia  r i c h a r d s o n i i i s found i n open s i t u a t i o n s , a l t h o u g h i t o f t e n occurs w i t h a p a r t i a l l y - s h a d i n g w o o d y f l o r a comprised c h i e f l y o f S a l i x s p e c i e s . hexaploid  cytodeme (2n = 6x = 36) o f B_. r i c h a r d s o n i i i n h a b i t s the A l a s k a  Range and i s found between 1000 - 1700m i n b a s i c s o i l s 14-ploid  The  and f l u s h e s .  The  cytodeme (2n = 14x = 84), o c c u r r i n g on the a r c t i c s l o p e o f the  Brooks Range, grows c h i e f l y on e a s t and n o r t h - e a s t  f a c i n g slopes  that are  a l s o r i c h i n bryophytes and l i c h e n s .  I t grows from near s e a l e v e l up t o  400m.  Regarding edaphic p r e f e r e n c e s ,  P r o f . J . G. Packer (in. l i t t . ) r e -  ported  t h a t B^ r i c h a r d s o n i i i s a c a l c i c o l e wherever he has found i t . How-  ever Dr. D. F. Murray ( i n l i t t . )  stated that:  h a r d n o t to be around c a l c a r e o u s  substrates,  range i s a s e r i e s o f l i m e s t o n e s . approaching an a c i d s i t e , B o y k i n i a l e c t i o n that Boykinia but  N e v e r t h e l e s s when we do f i n d i s there  also.  something  I n f a c t , i t i s my  aspects o f the s u b s t r a t e :  B o y k i n i a would be r e s t r i c t e d to o n l y  dry.  recolsites,  t h a t these h i g h l y  l i m e s t o n e s are o f t e n f r a c t u r e d and r e t a i n l i t t l e  ure so t h a t the h a b i t a t s a r e e x c e e d i n g l y  Therefore,  s i n c e the backbone o f t h a t  i s l e s s common on the most s t r o n g l y c a l c a r e o u s  t h i s may be due t o other  calcareous  " I n the Brooks Range, i t i s  surface  I n such i n s t a n c e s  moist-  then,  the most*moist and s h e l t e r e d  sites.  I have concluded that i t i s m o i s t u r e r a t h e r than carbonates  t h a t determines the d i s t r i b u t i o n o f the s p e c i e s " . Species  o f s e c t i o n T e l e s o n i x , B_. j a m e s i i and B.  heucheriformis,  grow i n c r e v i c e s o f rock f a c e s and among stones o f t a l u s s l o p e s  i n high  a l p i n e s i t u a t i o n s , mainly i n the Rocky Mountains between A l b e r t a and C o l orada.  The h a b i t a t can be. an e x c e e d i n g l y  r e t a i n s very  d r y one, s i n c e the substratum  l i t t l e Jnoisture, and t h i s " p o s s i b l y accounts f o r the s p e c i e s  rather succulent  leaves.  open, although there  L i k e B. r i c h a r d s o n i i , b o t h s p e c i e s  can be a p a r t i a l  canopy o f c o n i f e r o u s  grow i n the  tree  species.  180  B o y k i n i a j a m e s i i and B_. h e u c h e r i f ormis are commonest between 2500. r- 3500m, although  the l a t t e r  can be  found as low  as 1500m i n the more n o r t h e r l y p a r t s  of i t s range ( A l b e r t a ) and i n p l a c e s w i t h s u i t a b l y c o l d Boykinia heucheriformis  microclimates.  i s a c a l c i p h i l e throughout i t s range, whereas B.  j a m e s i i has been found o n l y on g r a n i t e  outcrops.  Peltoboykinia P e l t o b o y k i n i a t e l l i m o i d e s grows among the herbaceous undergrowth of the b r o a d - l e a v e d ,  deciduous Fagus c r e n a t a f o r e s t s at low  t i o n s i n the mountains of Japan (Hara 1959; i n a l l i m i t i s 1500m (Hara 1959).  The  10°C  Ohwi 1965). isotherm  1958,  Shikoku i s bounded by  I t s upper  altitud-  correlates w e l l with  the d i s t r i b u t i o n of s s p . t e l l i m o i d e s on Honshu, w h i l e ssp. watanabei on Kyushu and  eleva-  the d i s t r i b u t i o n of  the 15°C  isotherm  (Hara  1959).  Suksdorfia S u k s d o r f i a s p e c i e s are found i n the r a i n shadow areas mountains of the Western and  i n e a r l y s p r i n g , from the r a i n s and two  to a v o i d those which f a c e n o r t h . tend  places  snow-melt, but becoming dry by  North American s p e c i e s grow on mossy, rocky  violacea sites  the  Andean C o r d i l l e r a s of North and South America.  The h a b i t a t s have the s p e c i a l c h a r a c t e r i s t i c of b e i n g seepy, wet  The  of  F i e l d observations  to dry out e a r l i e r than those  I t i s , however, p o s s i b l e to f i n d the two  outcrops  but  suggest t h a t  summer.  appear S.  of S_. r a n u n c u l i f o l i a .  s p e c i e s growing i n adjacent  and  even s l i g h t l y mixed p o p u l a t i o n s , so c l o s e are t h e i r e c o l o g i c a l p r e f e r e n c e s . Both s p e c i e s are u s u a l l y found i n the open, although;S. v i o l a c e a can be  found under a p a r t i a l  growing up  through'layers  canopy of Plnus of d u f f .  ponderosa or Pseudotsuga m e n z i e s l i ,  N o t h i n g has been r e p o r t e d o f the  t a i l e d e c o l o g i c a l b e h a v i o u r of the Andean S. a l c h e m j l l o i d e s , and arium l a b e l s are  also  disappointinglyuninformative.  de-  the h e r b -  181  The  two North American s p e c i e s can be found on b o t h a c i d i c and  b a s i c s o i l s or flushes. is  found v e r y  c l o s e to s a l t water i n the B e l l a Coola  B r i t i s h Columbia. the  Kermode (1918) r e p o r t e d t h a t  ranunculifolia c o a s t a l area of  T h i s i s another example o f a montane p l a n t growing on  coast i n the n o r t h e r n p a r t o f i t s range.  Altitudinally  American s p e c i e s range from near s e a l e v e l to 3000m.  the N o r t h  Suksdorfia alchemjll-  o i d e s has been c o l l e c t e d between 3000 and 3800m. Bolandra Bolandra  i s found on mossy rocks  i n shaded, humid p l a c e s i n  woodland h a b i t a t s , u s u a l l y c l o s e to a water course.  The m l c r o h a b i t a t i s  t y p i c a l l y w e l l - d r a i n e d and can be q u i t e d r y between r a i n f a l l s , e s p e c i a l l y marked i n summer months. on b a s a l t i c rock, and a c c o r d i n g occurs in  on g r a n i t e .  Bolandra  a condition  oregana i s f r e q u e n t l y found  to Jepson (1936), B_. c a l i f o r q i c a always  A l t i t u d i n a l l y , IL oregana grows from about s e a l e v e l  the ''Columbia R i v e r Gorge to about 1400m i n the B i t t e r r o o t Mountains o f  Idaho.  Bolandra  c a l i f o r n i c a i s found between 1600 - 2800m i n t h e S i e r r a  Nevada o f c e n t r a l C a l i f o r n i a . Sullivantia Rosendahl (1927) r e p o r t e d on d r i p p i n g wet c l i f f s calciphiles. it  o r canyon w a l l s .  From o b s e r v a t i o n s  A t l e a s t the e a s t e r n s p e c i e s a r e  o f S_. oregana i n the Columbia R i v e r Gorge,  seems t h a t deep shade and a constant h i g h h u m i d i t y ,  w a t e r f a l l s , i s a requirement. to  t h a t a l l S u l l i v a n t i a s p e c i e s grow  The a s s o c i a t e d f l o r a i s r e s t r i c t e d  v a r i o u s mosses and l i v e r w o r t s .  erate competition  e.g. t h e spray o f chiefly  I t may be t h a t S u l l i v a n t i a cannot  tol-  from other v a s c u l a r p l a n t s .  Discussion I n g e n e r a l terms the h a b i t a t s o f the s p e c i e s s t u d i e d can be grouped' i n t o s i x c a t e g o r i e s :  182  1) ;/ r i p a r i a n woodland, becoming snow-bed grassland'above line  . . . . . . . . . .  the t r e e  B o y k i n i a s e c t s . B o y k i n i a and  2)  a l p i n e rock f a c e s and  talus  . . . . .  3)  spring-wet,  4)  shaded, humid r o c k s , t e n d i n g t o be summer-dry  5)  forest floor  . . . . . . .  6)  d r i p p i n g wet  cliffs  summer-dry b l u f f s  Renfolium  Boykinia sect. Telesonix  . . . . . . .  Suksdorfia . . Bolandra  Peltoboykinia Sullivantia  These groups c o r r e l a t e q u i t e w e l l w i t h g e n e r i c l i m i t s , and  the  possibility  of e v o l u t i o n from a common a n c e s t r a l s t o c k by a d a p t i v e r a d i a t i o n i n t o f e r e n t h a b i t a t s appears  an a t t r a c t i v e p r o p o s i t i o n .  that e v o l u t i o n i n the f a m i l y was  Savile  (1961)  dif-  proposed  l i n k e d to a s e r i e s of r a d i a t i o n s f o l l o w i n g  delayed and l i m i t e d p e n e t r a t i o n of p h y s i o g r a p h i c or c l i m a t i c b a r r i e r s . r e c o g n i z e d , too, t h a t most genera have d i f f e r e n t i a t e d to p a r t i c u l a r h a b i t a t s , and he r e l a t e d  through  adaptation  t h i s t o seed d i s p e r s a l and  t i o n mechanisms, although each genus o f t e n has  He  pollina-  a characteristic vegetative  f a c i e s as w e l l . E c o l o g i c a l d i f f e r e n t i a t i o n at the g e n e r i c l e v e l i s a phenomenon which has not been commonly r e p o r t e d ( S t e b b i n s 1974: e c o l o g i c a l b a s i s of taxonomic d i v e r s i t y , S t e b b i n s groups h a v i n g primary  (1974) h y p o t h e s i z e d t h a t  l i k e deserts or  A c c o r d i n g to t h i s p r o p o s a l , the most p r i m i t i v e  c l o s e s t to the a n c e s t r a l s t o c k ) are l i k e l y  h a b i t a t s , and r a d i a t i o n s toward more mesic, s h o u l d be  ecotones  to more mesic c o n d i t i o n s , and  the o t h e r to t a x a s u i t e d to more extreme s i t u a t i o n s  (presumably  D i s c u s s i n g the  a d a p t a t i o n s to m a r g i n a l environments or  gave r i s e on the one hand to taxa adapted  alpine habitats.  181).  arctic-  genera  to occupy m a r g i n a l  or toward h a r s h e r , environments  correlated with morphological s p e c i a l i z a t i o n .  i n the S a x i f r a g i n a e t h e r e i s some tendency  on  I n the p r e s e n t case, .  f o r the most p r i m i t i v e  genera,  Heuchera and S a x i f r a g a , to occupy the more m a r g i n a l environments (steep  183  s l o p e s and  rocks w i t h  landslides, etc.).  t h i n s o i l cover and  Admittedly,  but i t i s n e v e r t h e l e s s  areas p r e v i o u s l y denuded by  S a x i f r a g a has  a wide e c o l o g i c a l  true t h a t m o r e s p e c i a l i z e d genera grow e i t h e r i n  more mesic woodland or i n more x e r i c , summer-dry h a b i t a t s . support  amplitude,  i s thus g i v e n to S t e b b i n ' s  Tentative  hypothesis.  SEED DISPERSAL Savile  (19 75), r e v i e w i n g  the seed d i s p e r s a l syndromes i n some  members of the herbaceous S a x i f r a g a c e a e ,  noted t h a t the two  N o r t h American  S u k s d o r f i a s p e c i e s and B o y k i n i a s e c t i o n T e l e s o n i x have a censer mechanism whereby the f l e x i b l e stems wave i n the wind and f a c t , a l l Boykinia, Pe1toboykinia,  throw out  Suksdorfia, Bolandra  t h e i r seeds.  and  s p e c i e s have t h i s mechanism, the commonest i n the f a m i l y .  In  Sullivantia In some v e r y  dwarf specimens of B o y k i n i a j a m e s i i and B_. h e u c h e r i f ormis the stems are r a t h e r more r i g i d  and  where the wind induces  d i s p e r s a l i s b e t t e r d e s c r i b e d as the v i b r a t o r type a resonant  v i b r a t i o n , bouncing the seeds out.  mechanism i s e s p e c i a l l y w e l l - d e v e l o p e d  This  i n some S a x i f r a g a s p e c i e s ( S a v i l e  1975). Other d i s p e r s a l mechanisms i n the f a m i l y i n c l u d e the cup  type,  found i n Chrysosplenium, T e l l i m a and M i t e l l a , and  board type found i n T i a r e l l a .  These a d a p t a t i o n s  evolved  under the canopies  exerted  a great s e l e c t i v e stimulus The  as an a d a p t a t i o n port.  He  are thought to have  of humid f o r e s t s where l a r g e drops of water ( S a v i l e and Hayhoe 1978).  Savile'(1975)  regarded  diffi-  the winged seeds i n S u l l i v a n t i a  to c l i f f h a b i t a t s and"presumably, t h e r e f o r e , a e r i a l  a l s o suggested t h a t the h o r i z o n t a l l y - f l a r e d  i n e n t l y t u b e r c u l a t e seeds of Tolmiea are a d a p t a t i o n s transport  the s p r i n g -  e c o l o g i c a l f u n c t i o n o f seed coat ornamentation i s  c u l t to i n t e r p r e t .  splash-  capsules  trans-  withCprom-  to b i r d or mammal  (the seeds presumably c a t c h i n g i n f e a t h e r s or f u r , r a t h e r than  184  being ingested). applicable.  However, t h i s e x p l a n a t i o n s c a r c e l y seems u n i v e r s a l l y  There seems to be l i t t l e  c o r r e l a t i o n between.?testa  ornament-  a t i o n , seed d i s p e r s a l mechanism and h a b i t a t ; i n d e e d , many t h i n g s , o t h e r than d i s p e r s a l mode, c o u l d be important  i n a f f e c t i n g the type of  s u r f a c e , i n c l u d i n g f o r example, m o i s t u r e  testa  r e t e n t i o n and s u r f a c e area/volume  ratio. I t was  observed  ( t u b e r c u l a t e or not) trichomes  found on  severely r e s t r i c t s PHENOLOGY AND  t h a t on b e i n g shed from the c a p s u l e s , the seeds  f r e q u e n t l y s t i c k to the dense a r r a y of g l a n d u l a r  the i n f l o r e s c n e c e s i n a l l s p e c i e s s t u d i e d . the d i s p e r s a l  This obviously  potential.  POLLINATION  D e t a i l s of f l o w e r i n g ' t i m e s ' o f a l l s p e c i e s are g i v e n i n T a b l e XXIII. with  S u k s d o r f i a and B o l a n d r a  the r a i n f a l l  and snow-melt.  oregana f l o w e r i n the s p r i n g , c o i n c i d i n g A l l the o t h e r s p e c i e s f l o w e r i n the  summer. V a r i o u s members of the herbaceous S a x i f r a g a c e a e have been the s u b j e c t of a n t h e c o l o g i c a l s t u d i e s , and t i o n s are shown i n T a b l e XXIV.  the main r e s u l t s of these  Flower v i s i t o r s  observa-  ( p o l l i n a t o r s ? ) to some of  the s p e c i e s i n the p r e s e n t study have been c o l l e c t e d and p a r t i a l l y i d e n t ified.  In most cases  c o l l e c t i o n s were made d u r i n g the e a r l y a f t e r n o o n i n  sunshine; p o p u l a t i o n s of B o y k i n i a r i c h a r d s o n i i and B o l a n d r a seen o n l y d u r i n g r a i n y weather and no T a b l e XXV  oregana were  f l o w e r v i s i t o r s were ever  shows t h a t the v i s i t o r s are a d i v e r s e group.  Yeo  observed.  (1971) r e p o r t e d  t h a t B e r g e n i a s p e c i e s show some p o l l i n a t o r s p e c i a l i z a t i o n , and  Savile  C1975) noted t h a t the green-yellow f l o w e r e d s p e c i e s of M j t e l l a are v i s i t e d mainly by mosquitoes and other lower D i p t e r a w h i l e the s p e c i e s are v i s i t e d by visitors  the h i g h e r D i p t e r a .  to B o y k i n i a and  a l l i e s may  white-flowered  Thus, c l o s e r . s t u d i e s of the  r e v e a l some i n t e r s p e c i f i c or  inter-  185  TABLE X X I I I .  F l o w e r i n g times i n B o y k i n i a , F e l t o b o y k l n i a ,  Suksdorfia,  B o l a n d r a and S u l l i v a n t i a .  Species  Flowering Period*  Boykinia a c o n i t i f o l i a B. i n t e r m e d i a B. l y c o c t o n i f o l i a B. major B. o c c i d e n t a l i s B. r o t u n d i f o l i a B. r i c h a r d s o n l i B. h e u c h e r i f o r m i s B. j a m e s i i  June June July June June June June. June July  Peltoboykinia tellimoides ssp. t e l l i m o i d e s s s p . watanabei  May - J u l y June - J u l y  Suksdorfia alchemilloides S.' r a n u n c u l i f o l i a * S. v i o l a c e a *  January - A p r i l May - June (- August) (March -) May - June  B o l a n d r a oregana B. c a l i f o r n i c a  May - June June - J u l y  S u l l i v a n t i a oregana  (May -) June - J u l y  -  August August August August (- September) August July August August August  A l t i t u d e can a f f e c t the f l o w e r i n g p e r i o d , p o p u l a t i o n s of a s p e c i e s at lower e l e v a t i o n s f l o w e r i n g e a r l i e r than t h o s e a t h i g h e r e l e v a t i o n s . I n contiguous o r mixed p o p u l a t i o n s of S u k s d o r f i a v i o l a c e a and S_. r a n u n c u l i f o l i a , the former b e g i n s f l o w e r i n g about two weeks e a r l i e r than the l a t t e r .  TABLE XXIV.  Summary o f f l o w e r v i s i t o r s  ( p o l l i n a t o r s ? ) o f the S a x i f r a g i n a e .  Reference l e a d s are g i v e n as s u p e r s c r i p t s where a p p r o p r i a t e . Flower V i s i t o r s P l a n t Genus  Order  Family  Genus ( i f known)  Bergenia*  Hymenoptera  Eumenidae  Odynerus"*  Andrenidae  Andrena"''"'"  Megachilidae  Osmia  Anthophoridae  Anthophora  Apidae  A p i s ~ \ Bombus"*  NymptLalidae  Aglais"''^  Pieridae  Colias^  Lepidoptera  186  TABLE XXIV, cont'd. Flower V i s i t o r s P l a n t Genus  Order  Family  Jepsonia  Diptera  Syrphidae  Hymenoptera  Halictidae  Dialictus^  Apidae  Apis"*"  Dermestidae  Anthrenus  Nitidulidae  Meligethes  Curculionidae  Miarus  Chironomidae  Lymnophyes^, 4  Saxifraga  Coleoptera  Call ref. 5 except where noted) Diptera  Empididae Dolichopodidae Syrphidae  Genus C i f known)  Smittia Empis 4 Ascia, Carposcalls , Cheilosia,  Eristalis,  H e l o p h i l u s , Melangy4 na  , Melanostoma,  Melithreptus,  Spheg-  ina, S y r i t t a * , SyrAnthomyiidae Muscidae Calliphoridae'  phus. Scatophaga* 9 A r i c i a , Spilogona 4 Borellus  Hymenoptera  , Calli-  Cephidae  phora"', L u c i l i a " *  Ichneumonidae  Cephus H e m i t e l e s , Orthocen-  Formicidae^  trus  Eumenidae  Odynerus  Sphecidae Halictidae  Halictus  Andrenldae  Andrena  Megachilidae  Megachile  Apidae  A p i s , Bombus '^'^  Lepidoptera Trichoptera  Phryganeidae  187  TABLE XXIV,  cont'd. 'Flower V i s i t o r s Family  Genus ( i f known)  P l a n t Genus  Order  Chrysosplenium•  Pulmonata (Gastropoda)  Succineidae  Succinea  Coleoptera  Staphylinidae  Lathrimaeum, Tachyporus  Diptera  Phalacridae  Olibrus  Lathridiidae  Corticaria  Co c c i n e l l i d ae  Coccinella  Curculionidae  Apion  Chironomidae  Chironomus  Simuliidae  Simulium  Mycetophilidae  Exechia  Sciaridae  Sciara  Cecidomyiidae  Cecidomyia  Lonchop t e r i d a e  Lonchoptera  Syrphidae  Melanostoma  Sepsidae  Sepsis  Sciomyzidae  Sciomyza  Chloropidae  Chlorops  Muscidae  Mus ca  Hemlptera Hymenoptera  Lepidoptera  Ichneumonidae Cynipidae  Eucoila  Formicidae  L a s i u s , Myrmica  Andrenidae  Andrena  Moths  Thysanoptera Lithophragma  10  Hymenoptera  Lepidoptera  Halictidae  Chloralictus  Andrenidae  Andrena  Megachilidae  Osmia  Apidae  Apis  Incurvaritdae  Lampronia  188  TABLE XXIV, cont'd. Flower V i s i t o r s P l a n t Genus  Order  Family  Heuchera .  Kymenoptera  Colletidae  Genus ( i f known) 2 Colletes  Halictidae  Halictus  Apidae  Apis* Tellima*"  Mitella  Kymenoptera  Apidae  Lepidoptera  Pieridae  Pieris  Culicidae  Diptera  Syrphidae  7,8  Apoidea^  Kymenoptera *  Apis  o b s e r v a t i o n s made on p l a n t s i n c u l t i v a t i o n away from n a t u r a l h a b i t a t ,  References:  1.  Ganders  4.  Kevan 1972; 5.  8.  Spongberg  TABLE XXV.  inlitt.;  Knuth  1972; 9.  2.  1908; 6.  Swales  Flower v i s i t o r s  G r a e n i c h e r 1907; 3.  Ornduff 1971; 7.  1979; 10.  Hadac 1961;  S a v i l e 1975;  T a y l o r 1965; 11.  Yeo 1966.  ( p o l l i n a t o r s ? ) o f some B o y k i n i a and S u k s d o r f i a  species. Flower  Visitors  Plant*  Order  Family  B. i n t e r m e d i a  Diptera  Empididae  B. major  Coleoptera Hymenoptera  Coccinellidae Formicidae Halictidae Apidae  B. o c c i d e n t a l i s  Coleptera Diptera  Hymenoptera  Staphylinidae Empididae Syrphidae Otitidae Muscidae Apidae  B. h e u c h e r i f o r m i s  Diptera  Empididae  S.' r a n u n c u l i f o l i a  Diptera Trichoptera  Empididae Phryganeidae  S. v i o l a c e a  Diptera  Syrphidae  Genus No. C o l l e c t e d ( i f known) me hour 1 1 2 6 1  Myrmica Bomb us a  Q  b 1 1 0 1 0  Bombus  11 1  189  TABLE XXV, *  cont'd.  P o p u l a t i o n s s t u d i e d : ,B_. i n t e r m e d i a - G o r n a l l 23; B. major - G o r n a l l .212; B^ o c c i d e n t a l i s - a: G o r n a l l 227; b: G o r n a l l 216; ;B. h e u c h e r i formis - G o r n a l l 344; S. r a n u n c u l i f o l i a - G o r n a l l 332; S_. - y i o l a c e a B m 1112.  generic p o l l i n a t o r  stratification.  F l y i n g v i s i t o r s u s u a l l y a l i g h t on the edge of a f l o w e r where the anthers  are l o c a t e d , and  d r i n k the n e c t a r .  proceed to crawl over the f l o w e r and ably e f f e c t e d .  A f t e r a s h o r t w h i l e many  i t i s then t h a t p o l l i n a t i o n i s presum-  Flowers w i t h a more t u b u l a r hypanthium, e.g.  o c c i d e n t a l i s , enforce  a more p r e c i s e p o s t u r e  Boykinia  on the l a r g e r i n s e c t s , i f the  n e c t a r i s to be reached than do the more bowl-shaped f l o w e r s , e.g. major and S u k s d o r f i a  ranunculifolia.  Flower c o l o u r and  scent  are obvious candidates  d i f f e r e n t i a t i n g a p o l l i n a t o r fauna. species r e f l e c t u l t r a - v i o l e t l i g h t  I n the p r e s e n t (UV).  The  study,  although  these  too r e f l e c t UV  for a role i n p e t a l s of a l l  r i n g of yellow anthers,  i n some s p e c i e s , the y e l l o w n e c t a r i f e r o u s d i s c , seem to be guides,  Boykinia  and  the only honey  light.  Regarding the scent of the western N o r t h American s p e c i e s , o n l y i n B o y k i n i a major, B_. i n t e r m e d i a  and  odour d e t e c t a b l e by human b e i n g s . by  Suksdorfia ranunculif o l i a i s f l o r a l  In t h i s connection  the g l a n d u l a r trichomes should be mentioned.  the odour produced  I t i s quite spicy, similar  to c o r i a n d e r , and e s p e c i a l l y pronounced j u s t a f t e r a shower of r a i n .  As  f a r as can be judged, i t i s s i m i l a r , but not i d e n t i c a l , i n a l l s p e c i e s of Boykinia. it  In Suksdorfia, Bolandra,  i s b a r e l y p e r c e p t i b l e , i f at a l l ,  odour from i t s trichomes although Boykinia.  The  aromatic  the trichomes and may  P e l t o b o y k i n i a and S u l l i v a n t i a oregana Jepsonia  a l s o produces a s t r o n g  i t is a little  different  from t h a t i n  p r i n c i p l e s are l o c a t e d i n the g l a n d u l a r heads of  s e r v e as an a d d i t i o n a l a t t r a c t a n t to i n s e c t v i s i t o r s ,  190  the glands b e i n g e s p e c i a l l y dense on the f l o w e r p e d i c e l s . trichomes  c e r t a i n l y f u n c t i o n as e f f e c t i v e i n s e c t t r a p s :  f l i e s have been seen glued to the p e d i c e l s and G l a n d u l a r trichomes plants. Rod  are.not  The  many s m a l l dead  stems o f the p l a n t s .  the o n l y i n s e c t t r a p s found on  the  At l e a s t i n B o y k i n i a major and S u k s d o r f i a r a n u n c u l i f o l i a . Golden  S p i d e r s (Mlsumena v a t l a C l e r c k ) p o s i t i o n themselves atop  or among the anthers and  capture s u i t a b l e f l o w e r v i s i t o r s .  ensuing s t r u g g l e s p o l l i n a t i o n appears to be v i s i t o r s were allowed DISEASE AND  to d r i n k the n e c t a r  the  stigmata  During  the  f a r more c e r t a i n than i f the  undisturbed.  PREDATORS The  use of p a r a s i t e s i n i d e n t i f y i n g h o s t r e l a t i o n s h i p s has  d i s c u s s e d by Davis  and Heywood (1963: 256)  (1961, 1975,  has  rusts  sticky  1976)  and by S a v i l e  (1979).  been  Savile  r e c o r d e d a l i n e a g e of s h o r t - c y c l e d , h e t e r o e c i o u s  ( P u c c i n i a spp., U r e d i n a l e s ) on 11 genera and  87 s p e c i e s of herbaceous  Saxifragaceae, i n c l u d i n g M i t e l l a , T i a r e l l a , Bergenia, Saxifraga, splenium, Heuchera, T o l m i e a , Elmera,  Chryso-  T e l l i m a , Lithophragma and C o n i m i t e l l a .  However, P u c c i n i a has not y e t been r e c o r d e d on B o y k i n i a and i t s a l l i e s . P e s t s of the herbaceous S a x i f r a g a c e a e i n c l u d e l e a f  miners.  F i v e s p e c i e s of Phytomyza (Agromyzidae, D i p t e r a ) are known t o a t t a c k the family  (Griffiths  1972).  Three s p e c i e s are c o n f i n e d to S a x i f r a g a and  o t h e r s occur on T i a r e l l a , T o l m i e a  and M i t e l l a .  recorded from the genera under study h e r e ,  two  Although none have been  they should be sought.  i s a l s o h o s t to l a r v a e of the I n c u r v a r i i d a e ( L e p i d o p t e r a ) and  Saxifraga  to some  C o l e o p t e r a (Huber 1963). L i t t l e i n f o r m a t i o n i s a v a i l a b l e on the g r a z i n g of herbaceous S a x i f r a g a c e a e by b i r d s o r mammals, and i t seems t h a t these p l a n t s are g e n e r a l l y unimportant  i n s u p p o r t i n g such w i l d l i f e .  t i o n s are of i n t e r e s t however.  The  f o l l o w i n g observa-  191  I n A l a s k a and the Yukon, B o y k i n i a r i c h a r d s o n i i i s r e p o r t e d t o be a f a v o u r i t e o f the G r i z z l y Bear (Ursus h o r . r i b i l i s O r d . ) , fragments h a v i n g been found i n i t s droppings  (Murie 1944; White 1974).  My own o b s e r -  v a t i o n s o f these animals i n A l a s k a showed t h a t they move s i n g l y , or i n groups o f up t o f o u r , s l o w l y a c r o s s the t u n d r a , g r a z i n g r a t h e r l i k e  cattle.  They graze mainly on newly-emergent green shoots and d i g f o r f l e s h y  roots,  e s p e c i a l l y those o f Hedysarum alpinum L.  Although I d i d n o t see B o y k i n i a  r i c h a r d s o n i i b e i n g e a t e n , i t would seem t o be an obvious c a n d i d a t e . I t s b r i g h t green l e a v e s are q u i t e s u c c u l e n t and are v e r y conspicuous a g a i n s t the ochres o f the tundra l i c h e n s . f o r o t h e r animals.  B o y k i n i a r i c h a r d s o n i i may p r o v i d e food  I o f t e n observed t h a t the seed c a p s u l e s had h o l e s  punched i n t h e i r b a s e s , and t h i s b i r d s e a t i n g the seeds.  c o u l d be a t t r i b u t a b l e to s m a l l p a s s e r i n e  A l t e r n a t i v e l y , i t might be caused by i n s e c t  larvae  e a t i n g t h e i r way out a f t e r h a t c h i n g i n the o v a r i e s o f the f l o w e r s . The o n l y o t h e r s p e c i e s i n the p r e s e n t study f o r which t h e r e i s any i n f o r m a t i o n are B o y k i n i a j a m e s i i and B^ h e u c h e r i f o r m i s .  According to  Craighead e t a l . (1963) t h e r e i s e v i d e n c e that these s p e c i e s a r e e a t e n by e l k (Cervus canadensis E r x l e b e n ) and deer p l a n t s are a c c e s s i b l e .  ( O d o c o i l e u s s p p . ) , when t h e  My own o b s e r v a t i o n s suggest that B i g Horn Sheep  (Ovis canadensis Shaw) o c c a s i o n a l l y may a l s o f e e d on these p l a n t s .  192  XIV.  GENERIC LIMITS AND TAXONOMY  Taxonomic Approach The  taxa i n the p r e s e n t  study have been a s s i g n e d  f e r e n t genera a t one time o r another.  These a r e :  one  o f i t s s p e c i e s was o r i g i n a l l y d e s c r i b e d i n B o y k i n i a .  so few t a x a s h o u l d have been a s s i g n e d  S i x o f these  The obvious q u e s t i o n i s why  to so many genera.  and p o s s i b l y the major, has been the b e l i e f  At l e a s t one  t h a t genera s h o u l d be  d e f i n e d on the b a s i s o f f l o r a l and seed c h a r a c t e r s , v e g e t a t i v e t a k i n g a secondary r o l e . has  persisted until  that  to be monotypic, depending on where the l i n e  between v a r i e t y and s p e c i e s has been drawn.  reason,  Hieronymusia  A f u r t h e r genus, S u l l i v a n t i a , i s a l s o c l o s e enough  genera have been c o n s i d e r e d  dif-  Saxifraga, Boykinia,  P e l t o b o y k i n i a . N e o b o y k i n i a , T e l e s o n i x . Hemieva, S u k s d o r f i a , and B o l a n d r a .  to n i n e  characters  T h i s dogma was h e l d by Rafinesque (1837) and  comparatively  recently.  Thus i n the p r e s e n t  case,  where v a r i a t i o n i n f l o r a l morphology between the s p e c i e s i s widespread, i t i s n o t s u r p r i s i n g t h a t so many genera were r e c o g n i z e d .  Particularly  v a r i a b l e are c h a r a c t e r s such as ovary p o s i t i o n , stamen number, f r e e hypanthium l e n g t h , hypanthium shape and p e t a l shape and c o l o u r . suggested t h a t t h i s v a r i a t i o n i n f l o r a l morphology i n v o l v e d  S a v i l e (1961) adaptations  i n seed d i s p e r s a l o r p o l l i n a t i o n mechanism, and t h a t t h e r e f o r e i t might h i d e r e l a t i o n s h i p s or suggest them i n convergent forms.  He suggested  t h a t v e g e t a t i v e c h a r a c t e r s might p r o v i d e  to r e l a t i o n s h i p s .  the b e s t guides  Indeed, the p r e s e n t work c e r t a i n l y emphasizes v e g e t a t i v e c h a r a c t e r s f a r more than has been customary i n the group, b u t i t does n o t i g n o r e f e a t u r e s i f they seem u s e f u l . day  systematics  and  attempts to  floral  The approach i s c o n v e n t i o n a l i n modern  i n t h a t i t draws on evidence  from a v a r i e t y of sources  i n t e g r a t e i t to produce a d e f e n s i b l e  An i n t e r e s t i n g v i e w p o i n t  classification.  on g e n e r i c d e l i m i t a t i o n which has  193  been found to be v e r y u s e f u l was  suggested by B u r t t .(1964).  He  discussed  s i t u a t i o n s where some groups are d e f i n e d on the b a s i s of a s i n g l e t e r , the composition  of the group v a r y i n g w i t h  Crowson (1953) c a l l e d t h i s  the c h a r a c t e r  the "non-congruence p r i n c i p l e " .  charac-  chosen. The  problem i s  c l e a r l y r e l e v a n t to s i t u a t i o n s where e v o l u t i o n a r y p a r a l l e l i s m i s widespread as i s the case i n the S a x i f r a g i n a e . might be guished  B u r t t (1964) suggested t h a t  there  a b i o l o g i c a l d e f i n i t i o n of a genus as "a group of s p e c i e s from r e l a t e d groups by  s p e c i f i c d i f f e r e n t i a t i on". a l characters  ters i n dealing with  a change i n emphasis i n the c h a r a c t e r s  Thus i f s p e c i e s were s e p a r a t e d  then i t might be  distin-  m o s t l y on  rewarding to emphasize v e g e t a t i v e  generic delimination.  The  of flor-  charac-  r e v e r s e would a l s o h o l d  true. Three p r i n c i p l e s were f o l l o w e d i n " t r e a t i n g the genera.  The  f i r s t p r i n c i p l e emphasizes that the focus of the modem genus concept i s naturalness,  a genus b e i n g d e f i n e d by  constituent species. with  diversity:  of every  taxon.  the mutual r e l a t i o n s h i p s of i t s  I t a l s o acknowledges, however, that taxonomy d e a l s  i t i s t h e r e f o r e unnecessary to maximize the homogeneity As B u r t t  (1964) has  explained,  p a t t e r n o f v a r i a b i l i t y i n a genus i s the key p r i n c i p l e , a convention between two  due  to u n d e r s t a n d i n g .  T h i s serves  as a guard  r e c o g n i t i o n of too many s m a l l or monotypic genera.  nomenclatural  second  size  against Thirdly,  the u n d e s i r a b i l i t y of unnecessary  changes.  Regarding i n f r a - g e n e r i c c l a s s i f i c a t i o n ,  tinuous  The  taxonomic groups should be i n v e r s e l y p r o p o r t i o n a l to the  r e g a r d i s g i v e n to t r a d i t i o n and  used:  the  of convenience, i s that the degree of d i f f e r e n c e  of the groups (Davis and Heywood 1963). the needles  an a p p r e c i a t i o n of  s e c t i o n , s p e c i e s and  subspecies.  Species  c o r r e l a t e d v a r i a t i o n i n at l e a s t two  three c a t e g o r i e s  are s e p a r a t e d  characters  by  are  discon-  (one of which must  194  be m o r p h o l o g i c a l ) 1958;  Davis  and o f t e n by s t a t i s t i c a l d i f f e r e n c e s i n o t h e r s  1978).  Subspecies  (Hedberg  a r e used f o r those p a r t s of a s p e c i e s which  are s e p a r a t e d by d i s c o n t i n u o u s v a r i a t i o n i n one c h a r a c t e r , or by  incomplete  d i s c o n t i n u i t i e s i n o t h e r s , and which have a c e r t a i n g e o g r a p h i c a l  coherence.  Although  the genus B o y k i n i a i s s m a l l , s e c t i o n s have been r e c o g n i z e d  because o f h i s t o r i c a l precedent  partly  and p a r t l y to emphasize groups of c l o s e l y  related species. The  r e l a t i v e u s e f u l n e s s o f the v a r i o u s taxonomic c h a r a c t e r s i s  commented on i n the a p p r o p r i a t e chapter.  The u t i l i t y  of a p a r t i c u l a r f e a -  t u r e may v a r y depending on c o n t e x t , and each case i s judged A summary o f the d i s t r i b u t i o n o f the most important  individually.  c h a r a c t e r s among the  genera r e c o g n i z e d i s g i v e n i n T a b l e XXVI. Synopsis  o f the Genera and S p e c i e s PELTOBOYKINIA  P e l t o b o y k i n i a ( E n g l e r ) Hara, B o t . Mag. Tokyo 51: 250-253.  1937.  B o y k i n i a s e c t . P e l t o b o y k i n i a E n g l e r i n E n g l e r & P r a n t l , Nat. Pflanzenfam, Type s p e c i e s :  ed. 2.  18a: 120.  1928.  S a x i f r a g a t e l l i m o i d e s Maximowicz, B u l l . Acad. Imp. S c i . S t . Petersb.  15: 215-216.  1871.  (= P e l t o b o y k i n i a t e l l i m o i d e s  (Maxim.) Hara. The ifraga  two t a x a i n t h i s genus were o r i g i n a l l y d e s c r i b e d i n Sax-  (Maximowicz 1871; Yatabe 1892) where they remained u n t i l  E n g l e r t r a n s f e r r e d them t o B o y k i n i a . P e l t o b o y k i n i a to•• accomodate them.  1919 when  I n 1928, E n g l e r c r e a t e d the s e c t i o n  L a t e r , Hara (1937) e l e v a t e d the s e c t i o n  to g e n e r i c rank, P e l t o b o y k i n i a , on t h e b a s i s of i t s g l o s s y , p e l t a t e l e a v e s and y e l l o w i s h , g l a n d u l a r , dentate p e t a l s . The  genus P e l t o b o y k i n i a can be r e c o g n i z e d i n i t s own r i g h t  because i t d i f f e r s  from S a x i f r a g a i n p o s s e s s i n g a f r e e hypanthium and has  TABLE XXVI.  Summary o f the d i s t r i b u t i o n o f some  Character Rhizome Trichomes glandular unicellular chaffy  Peltoboykinia  Boykinia  Thick  Thick  Uniseriate  Multiseriate + +  +  rtant taxonomic c h a r a c t e r s among the genera. Bolandra  Suksdorfia  Bulbiferous  Bulbiferous  Sullivantia Slender  Uni/Multiseriate Uni/Multiseriate Multiseriate +/(+)/-  Hypanthium shape  Tuberularcampanulate  Campanulate  Tubularcampanulate  Campanulate  Campanulate  Petal  Spatulate, toothed  Spatulate/ orbicularclawed  Subulate  Ovoid/spatulate /orbicularclawed  Spatulate  shape  Stamens  10  Stylar transmitting tissue  +  5/10  5  5  +  +  +  Pollen - . apertures tectum  Perforate  Seeds shape  Ovoid  Ovoid  Echinate  Tuberculate/ smooth  testa Chromosome base number Flavonoids 6-oxygenation diglycosides with arabinose or xylose  Colporoidate  11  -  Colporoidate/ colporate Reticulate/ occluded  Colporate  Colporate  Reticulate  Reticulate  C o l p o r a t e (2 orae/colpus) Reticulate  Narrowly ellipsoid Tuberculate  Ovoid  Ovoid-winged  Tuberculate  Smooth  6/7  7/9 spp. 8/9 spp.  + +  +  196  coloporoidate ated  p o l l e n with  from B o y k i n i a by  a p e r f o r a t e tectum.  I t can a l s o be w e l l  i t s p e l t a t e l e a v e s which are g l o s s y on t h e i r  surfaces, yellowish glandular p e t a l s , u n i s e r i a t e glandular of u n i c e l l u l a r e g l a n d u l a r tall  spines, a very  trichomes,  simple  lower  trichomes,  lack  t h e i r r e g u l a r rows of  f l a v o n o i d p r o f i l e based on monoglycosides of  q u e r c e t i n and k a e m p f e r o l , and cross-incompatible  i t s seeds w i t h  separ-  i t s chromosome base number of  with Boykinia  11.  I t i s also  o c c i d e n t a l i s , the o n l y h y b r i d i z a t i o n a t -  tempted. Maximowicz (1871) and E n g l e r  (1891: 61)  b o t h suggested a pos-  s i b l e a f f i n i t y of P e l t o b o y k i n i a w i t h P e l t i p h y l l u m . presumably on the of p e l t a t e l e a v e s  and  t e n stamens i n both t a x a .  Although the genera a l s o  share g l a n d u l a r , u n i s e r i a t e trichomes they are p r o b a b l y related. with  P e l t i p h y l l u m has  not v e r y c l o s e l y  a base chromosome number of x = 17,  x = 11 i n P e l t o b o y k i n i a .  p h y l l u m w i t h B e r g e n i a and Mukdenia which a l s o have x = 17.  conditions quite opposite  compared  T h i s would suggest the a l l i a n c e of  a l s o l a c k s a f r e e hypanthium, has  f r e e c a r p e l s and  a s i n g l e integument,  to those found i n P e l t o b o y k i n i a .  Peltoboykinia  r  charged f l a v o n o i d s ) and  number, trichome complement and p o s i t i o n between the S y n o p s i s and Key The 1.  chromosome number), and  genera t h e r e f o r e b e s t  to the S p e c i e s genus  some w i t h  of  Saxifraga an  l a . Leaf  (stamen  intermediate  Peltoboykinia  i s monotypic. Tokyo 51:  250-  1937.  Two Leaf  and  reflects i t s relationships.  P e l t o b o y k i n i a t e l l i m o i d e s (Maxim.) Hara, Bot. Mag. 253.  1.  two  Pelti-  Peltiphyllum  instead"shows "some :.af f i n l t i e s , ' w i t h ' B o y k i n i a - (stamen ,number, p o l l e n f l o w e r morphology and  basis  s ub specites-'-aref r e c o g n i z e d .  l o b e s o v a t e - d e l t o i d , b r o a d e r than l o n g . l a . s s p . t e l l i m o i d e s lobes n a r r o w l y cuneate-ovate,  longer  197  than b r o a d  l b . s s p . watanabei  la. Peltoboykinia Saxifraga  tellimoides  (Maxim.) Hara s s p . t e l l i m o i d e s  t e l l i m o i d e s Maximowicz,  P e t e r s b . 15: 215, 216. Boykinia  tellimoides  Pflanzenreich Type:  lb. Peltoboykinia  S c i . St.  1871.  (Maxim.) E n g l e r i n E n g l e r & Irmscher,  69: 675.  1919.  C o l l e c t o r unknown t o me, meridionalis"  B u l l . Acad. Imp.  " I n p r o v i n c i a Owari Nippon  (LE?).  tellimoides  (Maxim.) Hara s s p . watanabei (Yatabe)  G o r n a l l s t a t . nov. S a x i f r a g a watanabei Yatabe, B o t . Mag. t . 2.  1892.  Saxifraga Bot.  t e l l i m o i d e s Maxim, v a r . watanabei (Yatabe) Makino,  Mag.  Boykinia  Toyko 6: 7, 43, 44.  Tokyo 15: 12.  tellimoides  1901.  (Maxim.) E n g l . v a r . watanabei (Yatabe)  E n g l e r i n E n g l e r & Irmscher, P f l a n z e n r e i c h Boykinia 14.  1919.  watanabei (Yatabe) Makino, J o u r n . J a p . B o t . 3: 13, 1926.  Peltoboykinia  tellimoides  (Maxim.) H a r a v a r . watanabei (Yatabe)  Hara, D i s t r i b . Maps Flow. P l a n t s Book Co., Tokyo. Type:  69: 675.  Jap. 1. Map  17.  Inoue  1958.  Kano Watanabe s.n., 12th J u l y 1889, "At Nanokawa-mura i n the p r o v i n c e o f T o s a " , Japan ( ? ) .  Peltoboykinia as s e p a r a t e s p e c i e s  c o n t a i n s two t a x a which have been t r e a t e d  ( e . g . Ohwi 1965) or as v a r i e t i e s of a s i n g l e  (Makino 1901; Matsumura 1912; E n g l e r 1919; H a r a 1958). e d l y u s e f u l i n d i s t i n g u i s h i n g the two t a x a i n c l u d e l o b i n g , p e t a l colour  and shape.  either  species  Characters  report-  the e x t e n t o f l e a f  P o p u l a t i o n s from c e n t r a l and n o r t h e r n  198  Honshu ( s s p . t e l l i m o i d e s ) are r e p o r t e d to have l e a v e s w i t h wider than l o n g , and Populations  flowers with  the  lobes  creamy-white, b r o a d l y - o b l a n c e o l a t e p e t a l s .  from the i s l a n d s of Shikoku and Kyushu i n the south  watanabei) are r e p o r t e d to have l e a v e s w i t h  (ssp.  the lobes l o n g e r than wide  flowers with pale yellow, narrowly-oblanceolate  petals.  and  However, i t has  been shown t h a t t h e r e i s a c t u a l l y no d i f f e r e n c e i n p e t a l c o l o u r between the two  t a x a , and  (Ch. X I ) .  t h a t t h e r e i s complete i n t e r g r a d a t i o n i n p e t a l shape  I n v e s t i g a t i o n of the l e n g t h of the f r e e hypanthium, a c h a r a c t e r  h i t h e r t o overlooked,  r e v e a l e d t h a t i t tended to be s h o r t e r i n s s p . watan-  a b e i than i n s s p . t e l l i m o i d e s , but I t was  a l s o found  that the d i f f e r e n c e i n l e a f l o b i n g a p p l i e s o n l y to  last-formed: basal, leaves In view of the f a c t character only  t a x a show d i s c o n t i n u o u s v a r i a t i o n i n one  ( l e a f l o b i n g ) and have an a l l o p a t r i c d i s t r i b u t i o n p a t t e r n ,  s t r o n g l y supported  by  them as s u b s p e c i e s .  Their conspecific status i s  their identical flavonoid p r o f i l e s .  i n a e t y p i c a l l y show l i t t l e  The  Saxifrag-  or no i n f r a s p e c i f i c q u a l i t a t i v e v a r i a t i o n (Bohm  1978b). BOYKINIA  Taxonomic Treatment Boykinia N u t t a l l 114.  (nom.  c o n s . ) , J o u r n . Acad. Nat.  S c i . P h i l a d . 7:  113,  1834. Therofon  the  ( C h . . I I I ) . Both taxa have 11 p a i r s of chromosomes.  that the two  i t seems b e s t to t r e a t  and W i l k i n s  a g a i n t h e r e i s complete i n t e r g r a d a t i o n .  Rafinesque,  Neobot. 66.  ("1836"),  T e l e s o n i x Rafinesque. Neoboykinia  Neophyt. Bot.  1828  ( ? ) ; New  F l . N. Am.  IV.  1838.  F. T e l l . 2: 69.  Hara, Bot. Mag.  ("1836"),  Toyko 51: 253.  Rev.  Pursh.  1817;  1837.  1937.  Non  B o y k i n i a Rafinesque,  Neogenyton 2.  Non  B o y k i n i a N u t t a l l ex A r n o t t , Hook. J o u r n . Bot.  3: 276.  1825. 1840.  199  Type s p e c i e s :  B o y k i n i a a c o n i t i f o l i a N u t t a l l , J o u r n . Acad. Nat. S c i . P h i l a d . 7: 113,  The  genus was  114.  1834.  named a f t e r Dr.  Samuel B o y k i n (1786  - 1848), a  p l a n t e r , p h y s i c i a n and n a t u r a l i s t  from M i l l e d g e v i l l e , G e o r g i a ,  must have been a v e r y p o p u l a r  because h i s name has been g i v e n to gen-  e r a i n the L y t h r a c e a e thereby  causing  and  published  name was  1825).  The  s p e c i e s i s now  (1945) who  l a t e r misprinted  later  B o y k i n i a N u t t . was aconitifolia.  published  and was  (Rafinesque c.f. pub-  conserved a g a i n s t i t s  Lanjouw e t a l . 1952).  The  c i t e d as the m i s p r i n t e d Boykiana of R a f i n e s q u e  However, Rafinesque (1838) claimed  ton B o t a n i k o n or New  (Rafinesque  B o y k i n i a N u t t . was  e s t a b l i s h e d to accommodate a new  1828).  first  (syn. R o t a l a L.,  sphalm. ? ) .  (Green 1940;  t h i s s p e c i e s i n the new  (Rafinesque  as Boykiana h u m i l i s  ( N u t t a l l 1834)  e a r l i e r homonym, B o y k i n i a Raf. nomen r e j i c i e n d u m was  Saxifragaceae,  as B o y k i n i a h u m i l i s  p l a c e d i n Ammannia L.  g i v e s "Rotoea L.",  l i s h e d eleven years  He  confusion.  f o r a s p e c i e s of the L y t h r a c e a e ,  The  B.  as w e l l as the  to Rafinesque'(1840) the name B o y k i n i a was  1817).  Hulten  Cucurbitaceae  considerable  According  man  U.S.A.  The work he  genus T h e r o f o n ,  r e f e r r e d to was  saxifrage species, t h a t he had  previously  as T_. n a p e l l o i d e s  most l i k e l y h i s "Neophy-  P l a n t s of N o r t h America" (Kuntze 1891).  I have not been a b l e to t r a c e any  (1825).  Unfortunately  copies of t h i s book, and even F i t z p a t -  r i c k (1911) i n h i s d e t a i l e d b i b l i o g r a p h y of Rafinesque's w r i t i n g s , c o u l d g i v e no  i n d i c a t i o n of i t s whereabouts.  c a t i o n date of 1828  f o r T h e r o f o n Raf.  Rafinesque (1838), i n h i s "New  U n t i l a copy t u r n s up,  F l o r a of N o r t h America", p r o v i d e d  c i t i n g Boykinia a c o n i t i f o l i a Nutt. but  a c t u a l l y i s s u e d i n 1838  publi-  should be viewed w i t h s u s p i c i o n .  (?) d e t a i l e d d e s c r i p t i o n of T h e r o f o n and  1836  the  a second  i t s s p e c i e s T_. n a p e l l o i d e s ,  as a synonym.  (Barnhart  1907)  I t i s t h i s work  (dated  which i s u s u a l l y c i t e d  200  as the o r i g i n a l p l a c e of p u b l i c a t i o n o f T h e r o f o n when B o y k i n i a N u t t . was 1940).  ( M e r r i l 1949) , as i t was  conserved a g a i n s t that name (Dandy 1935;  I f the o r i g i n a l v a l i d p u b l i c a t i o n of T h e r o f o n was  c o n s e r v a t i o n o f B o y k i n i a N u t t . (1834) was mention  o f T h e r o f o n has been dropped  unnecessary,  Green  i n 1838  then  and i n d e e d a l l  from the "Nomina g e n e r i c a  conservanda  et r e j i c i e n d a " i n I n t e r n a t i o n a l Codes o f B o t a n i c a l Nomenclature from  1961  onwards. However, s h o u l d the p u b l i c a t i o n of T h e r o f o n t u r n out to be 1828,  B o y k i n i a N u t t . would p r o b a b l y not need to be conserved a g a i n s t i t  because  t h e r e i s e v i d e n c e t h a t the names may  Nuttall  (1834) c r e d i t e d  l a t e arduous was  be based  on the same type.  the d i s c o v e r y o f h i s B o y k i n i a a c o n i t i f o l i a to "the  and e c c e n t r i c P r u s s i a n c o l l e c t o r , Mr. K i n " , whose  s t a t e d to be i n Muhlenberg's herbarium, where i t was  as a Heuchera.  Rafinesque  apparently l a b e l l e d  (1838) a l s o s t a t e d t h a t the s p e c i e s was  e r e d by " K i n " and c i t e d a specimen herbarium.  specimen  discov-  l a b e l l e d Heuchera palmata i n C o l l i n ' s  I t i s known t h a t K i n d i s t r i b u t e d h i s c o l l e c t i o n s to C o l l i n s ,  as w e l l as to Muhlenberg (Stuckey 1971)  and i t i s t h e r e f o r e e n t i r e l y  s i b l e that N u t t a l l and Rafinesque based  t h e i r d e s c r i p t i o n s on  The  fact  isotypes.  that b o t h specimens are l a b e l l e d as Heuchera c o u l d support  a view.  A comparison  specimen  i s now  i f not l o s t , may  of the two  c o l l e c t i o n s would be u s e f u l .  of  h i s herbarium  such  Muhlenberg's  a t PH and t h e r e i s a s l i m chance t h a t C o l l i n ' s be at P where the remains  pos-  specimen,  are lodged  (Stuckey 19 71). A c c o r d i n g to A r t . 14, note 3 o f the Code, h a v i n g a l r e a d y been conserved once a g a i n s t B o y k i n i a Raf., B o y k i n i a N u t t . i s a u t o m a t i c a l l y a l s o c o n s e r v e d . a g a i n s t T h e r o f o n Raf. i f , as seems l i k e l y ,  they are based  on the same type. I f i t can be shown t h a t Rafinesque d e s c r i b e d T h e r o f o n i n  1828  201  r a t h e r than 1838, there i s an u n f o r t u n a t e type s p e c i e s .  consequence f o r the name o f the  Under A r t s . 63 and 67 o f the Code, the s p e c i f i c e p i t h e t  n a p e l l o i d e s should be used when T. n a p e l l o i d e s i s put i n the genus B o y k i n i a s i n c e i t i s the e a r l i e s t Nuttall's  l e g i t i m a t e e p i t h e t a v a i l a b l e a t t h a t rank.  (1834) e p i t h e t a c o n i t i f o l i a i s n o m e n c l a t u r a l l y  superfluous.  How-  e v e r , s i n c e the p r i o r i t y of T_. n a p e l l o i d e s i s i n doubt, and s i n c e the book where i t was p u b l i s h e d may no l o n g e r e x i s t , i t i s a d v i s a b l e to r e t a i n t h e f a m i l i a r and commonly-used name B o y k i n i a a c o n i t i f o l i a N u t t . species.  Should v a l i d p u b l i c a t i o n o f T_. n a p e l l o i d e s Raf.  f o r the type  p r i o r to 1834  be demonstrated, the combination B o y k i n i a n a p e l l o i d e s ought t o be made. U n t i l such time, i t i s p r o b a b l y  b e t t e r to l e t s l e e p i n g dogs l i e .  P o s s i b l y aware t h a t R a f i n e s q u e ' s T h e r o f o n o f 1828 made h i s B o y k i n i a o f 1834 a synonym, N u t t a l l for  a species of Cucurbitaceae.  (1840) d e s c r i b e d another B o y k i n i a  T h i s was B o y k i n i a t r i s p o r a N u t t . ex Am.  In f a c t Rafinesque (1838) was a l r e a d y aware o f t h i s name, although he a t t r i b u t e d i t to Wray and N u t t u a l l but as y e t u n p u b l i s h e d . N u t t . has been conserved, B o y k i n i a N u t t . ex Am. is  Since  Boykinia  i s a l a t e r homonym and  t h e r e f o r e to be r e j e c t e d ( A r t . 64 o f the Code).  The  Cucurbitaceous  s p e c i e s i n now'placed i n Cayaponia S. Manso. B o y k i n i a i s a genus d i f f i c u l t  to d e f i n e , s i n c e i t approaches  a number of o t h e r s i n the range o f i t s v a r i a t i o n .  I t can be c h a r a c t e r i z e d  by i t s t h i c k , s c a l y rhizome; presence o f b o t h m u l t i s e r i a t e g l a n d u l a r and u n i c e l l u l a r eglandular  trichomes;  campanulate or t u r b i n a t e f l o r a l  a f r e e hypanthium; s p a t u l a t e o r clawed e g l a n d u l a r p e t a l s ; a s t y l e transmitting t i s s u e ; p o l l e n with and  a r e t i c u l a t e exine  colporate or colporoidate apertures;  with  (sometimes occluded)  and a f l a v o n o i d p r o f i l e which  u s u a l l y includes d i g l y c o s i d e s i n v o l v i n g arabinose monoglycosides and 6-oxygenated  cup w i t h  aglycones.  or xylose,  charged  202  Many of the s p e c i e s were f i r s t d e s c r i b e d i n S a x i f r a g a , but because of t h e i r f i v e stamens, a f r e e hypanthium and ance they have s i n c e been removed. c e l l u l a r e g l a n d u l a r trichomes  The  (absent  B o y k i n i a s e c t i o n T e l e s o n i x has  appear-  presence on a l l taxa of the u n i -  from S a x i f r a g a ) , and  colporoidate p o l l e n (colpate i n Saxifraga) provides f o r t h e i r s e p a r a t i o n as a d i s t i n c t  a different  group.  Only w i t h  t h e r e been any  c o l p o r a t e or  further strong the two  s p e c i e s of  l i n g e r i n g doubt, owing t o  the presence of t e n stamens (as i n S a x i f r a g a ) , p u r p l e f l o w e r s , and p h i l i c habit.  1928)  chasmo-  O r i g i n a l l y put i n S a x i f r a g a ( T o r r e y 1827), o c c a s i o n a l f l o r a s  s t i l l p r e f e r to keep the s p e c i e s t h e r e 1954).  support  (Jones  1910;  Weber 1953;  Harrington  Most, however, e i t h e r f o l l o w e d Rosendahl (1905) or E n g l e r i n p l a c i n g them i n B o y k i n i a , or as seems more p o p u l a r  f o l l o w i n g Rafinesque  (1838) by a s s i g n i n g them to t h e i r own  (e.g. H i t c h c o c k e t a l . 1961;  Scoggan 1978).  (1919,  recently, i n genus, T e l e s o n i x  S i n c e the s p e c i e s have a l l  the c h a r a c t e r s of B o y k i n i a i n c l u d i n g a v e r y t y p i c a l f l a v o n o i d p r o f i l e , they must c e r t a i n l y be p l a c e d h e r e .  The  fact that synthetic hybrids  B. o c c i d e n t a l i s have been made adds p o w e r f u l The  two  support  to t h i s  with  conclusion.  s p e c i e s of s e c t i o n T e l e s o n i x , B_. j a m e s i i and B_. h e u c h e r i f o r m i s .  d i f f e r from the core members of the genus i n s e c t i o n B o y k i n i a by t e n r a t h e r than f i v e stamens, crimson-purple a s i m p l e r i n f l o r e s c e n c e , smooth seeds, p h i l i c habit. reminiscent  The  of the c o n d i t i o n s found  r a t h e r than w h i t e f l o w e r s ,  c o l p o r o i d a t e p o l l e n and  s i m p l e r i n f l o r e s c e n c e , and i n B.  r i c h a r d s o n i i , a s p e c i e s whose  ence and l e a f shape l e d b o t a n i s t s from the b e g i n n i n g  s o n i x (Hooker 1832).  r i c h a r d s o n i i and  Rosendahl  a chasmo-  c o l p o r o i d a t e p o l l e n are  p o s i t i o n i n B o y k i n i a has never s e r i o u s l y q u e s t i o n e d .  r e l a t i o n s h i p between B^  having  the two  In f a c t  infloresc-  to p o s t u l a t e a c l o s e  s p e c i e s of s e c t i o n T e l e -  (1905) even e r e c t e d a s p e c i a l s e c t i o n ,  s e c t . R e n i f o l i u m , to accommodate the t h r e e s p e c i e s , an arrangement which  203  Engler and  (1928) f o l l o w e d .  However, evidence  suggests t h a t J L  the s p e c i e s i n s e c t i o n T e l e s o n i x e v o l v e d  richardsonii  independently.  Boykinia  r i c h a r d s o n i i i s c h a r a c t e r i z e d by a f l a v o n o i d p r o f i l e emphasizing chromosome numbers of 36 seeds.  and  84,  f i v e stamens, and m i n u t e l y  Complex f l a v o n o l s , a chromosome number of 14,  flavones,  tuberculate  ten stamens  and  smooth seeds are found i n B_. .j a m e s i i and B_. h e u c h e r i f ormis.  I t seems  p r e f e r a b l e to r e g a r d  thus a s s i g n  them to s e p a r a t e Renifolium  the two  groups as s e p a r a t e  l i n e a g e s and  s e c t i o n s . B o y k i n i a r i c h a r d s o n i i should remain i n s e c t i o n  and B_. j a m e s i i and J L  h e u c h e r i f ormis are put  T e l e s o n i x , based on the genus d e s c r i b e d by Rafinesque  i n the new  (1838)  F o l l o w i n g i t s o r i g i n a l d e s c r i p t i o n as a S a x i f r a g a 1832), B o y k i n i a r i c h a r d s o n i i has been s e p a r a t e d previous  occasion  (Rafinesque  1837), and  e c c e n t r i c p u l v e r i s a t i o n of genera and Rydberg (1905).  from B o y k i n i a on o n l y  s p e c i e s p r a c t i s e d by Small i t to Hemieva (here  on grounds of f l o r a l morphology, but  treatment has not been adopted s i n c e . morpholoby, p o l l e n morphology and in  (Hooker one  t h i s i s s u r p r i s i n g i n view of  Rafinesque (1837) a s s i g n e d  synonymous w i t h S u k s d o r f i a )  section,  The  the  and considered this  trichome complement, rhizome  f l a v o n o i d p r o f i l e argue f o r i t s p o s i t i o n  Boykinia. The  o t h e r s p e c i e s of B o y k i n i a whose g e n e r i c p o s i t i o n has  been i n doubt i s B_. l y c o c t o n i f o l i a . (Maximowicz 1886), i t was  O r i g i n a l l y described i n Saxifraga  soon r e c o g n i z e d  as a B o y k i n i a  (Engler  However, because of the customary s e p a r a t i o n of S u l l i v a n t i a and  1891). Suksdorfia  from B o y k i n i a , H a r a (1937) e l e v a t e d J B ^ l y c o c t o n i f o l i a to g e n e r i c rank, N e o b o y k i n i a . on grounds of c o n s i s t e n c y .  H i s new  on i t s s m a l l , g r e e n i s h , p e r s i s t e n t p e t a l s , and thium.  As has  l a b i l e and  monotypic genus was  based  i t s v e r y s h o r t f r e e hypan-  been shown (Ch. V I ) , p e t a l marcescence i s  environmentally  i s e x h i b i t e d by B o y k i n i a s p e c i e s i n growth chamber  cultivation.  204  Furthermore, B o y k i n i a  i s q u i t e v a r i a b l e i n i;ts?:fLoral morphology,  enough to accommodate B. a t y p i c a l Boykinia  lycoctonifolia.  flavonoid p r o f i l e .  Boykinia  In f a c t  certainly  l y c o c t o n i f o l i a also  has  the s p e c i e s b e l o n g s to  the  core of the genus i n s e c t i o n B o y k i n i a based on i t s i n f l o r e s c e n c e s t r u c t u r e , p o l l e n , anther and and  i t s general  not  even h i m s e l f  seed morphology, i t s trichome complement, l e a f  appearance.  No  one  has  f o l l o w e d Hara's (1937) treatment,  (e.g. Hara 1959).  Regarding the d i s t i n c t i o n of B o y k i n i a  from S u l l i v a n t i a , i t i s  s u r p r i s i n g t h a t so few exchanges o f taxa between such s i m i l a r been proposed. described  The  only  as a B o y k i n i a  Sullivantia differed sepals.  case was  i s environmentally  (Brandegee 1899).  form B o y k i n i a  clearly  Rydberg  (Ch. VI)  Furthermore, B o y k i n i a  used i n any  These p e t a l and  generic  distinction,  two  genera are m o r p h o l o g i c a l l y  Rosendahl (1927) noted t h a t they were e a s i l y  winged seeds, campanulate c a p s u l e s ,  l y c o c t o n i f o l i a has sepal  im-  characters  and n e i t h e r do  flavon-  v e r y . s i m i l a r indeed,  confused.  He  l a c k of brown, c h a f f y trichomes  less  seeds, u r c e o l a t e seed c a p s u l e s , brown c h a f f y h a i r s and trichomes i n B o y k i n i a .  cymes.  this l i s t  B o y k i n i a has  can be  and  trichomes i n S u l l i v a n t i a v s . wing-  He  dense  array  a l s o noted a tendency f o r  lower p a r a c l a d i a of the i n f l o r e s c e n c e i n S u l l i v a n t i a chasial helicoid  to  characteristics:  s p a r s e o c c u r r e n c e of u n i c e l l u l a r  of u n i c e l l u l a r  and  declined  very  To  imbricate  t h a t p e t a l marcescence  them, however, c i t i n g the f o l l o w i n g d i s t i n g u i s h i n g  oped.  that  chromosome numbers support s e p a r a t e r e c o g n i t i o n . The  unite  originally  (1905) s t a t e d  i n i t s marcescent p e t a l s and  (Maximowicz 1886).  cannot be  o i d s nor  labile.  genera have  when S u l l i v a n t i a p u r p u s i i was  However, i t has been demonstrated  b r i c a t e sepals  shape  the  to develop as t r i -  t h i s tendency much l e s s w e l l  devel-  added an important p a l y n o l o g i c a l f e a t u r e .  Thus w h i l e the p o l l e n i n B o y k i n i a  i s colporate  or c o l p o r o i d a t e ,  in  205  S u l l i v a n t i a each colpus  i s a s s o c i a t e d w i t h , not one,  orae (Hideux and Ferguson 1976).  but  two  or  three  Furthermore, t h e r e i s a d i f f e r e n c e i n  f l o r a l anatomy i n t h a t S u l l i v a n t i a l a c k s s t y l a r t r a n s m i t t i n g t i s s u e , c o n d i t i o n unique among the genera i n v e s t i g a t e d h e r e . of S u l l i v a n t i a are i n t e r c r o s s a b l e ( S o l t i s i n l i t t . ) with  those  of B o y k i n i a , although  inter-generic pollination. with  Leaf  the ovules  In a d d i t i o n , species but  cross-incompatible  sometimes do s w e l l f o l l o w i n g  t e x t u r e i n the  two  the lower s u r f a c e s of B o y k i n i a l e a v e s h a v i n g  S u l l i v a n t i a l e a v e s are smooth.  a  genera i s d i f f e r e n t ,  prominent v e i n r i d g e s ;  Based upon h i s t o r i c a l precedent and  the  above c o l l e c t i o n of d i f f e r e n c e s , i t seems j u s t i f i a b l e , t h e r e f o r e , to keep the two  genera  separate.  Probably  the g r e a t e s t d i f f i c u l t y i n the d e l i m i t a t i o n of  l i e s i n i t s r e l a t i o n s h i p with S u k s d o r f i a . p o l l e n i n Suksdorfia i s 3-colporate, those  of B o y k i n i a , and  i n two  Boykinia  L i k e most B o y k i n i a s p e c i e s ,  the seeds are r e l a t i v e l y s i m i l a r  of i t s s p e c i e s  (S^ v i o l a c e a and  l o i d e s ) the i n f l o r e s c e n c e bears a dense a r r a y of u n i c e l l u l a r  the to  S_. a l c h e m j l trichomes.  The  l a t t e r f e a t u r e e s p e c i a l l y , emphasizes the c l o s e a f f i n i t y between the  two  genera.  provide  Chromosome numbers (x = 7 i n S u k s d o r f i a ; x = 6,  l i t t l e help i n determining  awns on the anthers on the anthers  generic l i m i t s .  7 in  Boykinia)  Furthermore, the  apical  of s p e c i e s i n B o y k i n i a s e c t i o n B o y k i n i a are a l s o found  o f _S_. a l c h e m j l l o i d e s .  between B o y k i n i a major and and Jepson (1925, 1936)  The  s i m i l a r i t y i n f l o r a l morphology  Suksdorfia r a n u n c u l i f o l i a i s quite  i n c l u d e d the two  striking  s p e c i e s i n the same subgenus,  Hemleva. of B o y k i n i a . In view of these r a n u n c u l i f o l i a was Gray 1840; B.  considered  Gray 1842).  occidentalis.  s i m i l a r i t i e s i t i s not  s u r p r i s i n g that  e a r l y as a p o s s i b l e B o y k i n i a  Gray (1876) l i s t e d i t as a p r o b a b l e  Greene (1891) l a t e r made the formal  (Torrey  S. and  synonym of  combination B o y k i n i a  206  r a n u n c u l i f o l i a , but few authors 1925, 1936, Jones 1938).  have f o l l o w e d him ( E n g l e r 1919; Jepson  F l o r a l morphology a l s o T e d R a f i n e s q u e (1838) t o  i n c l u d e B o y k i n i a r i c h a r d s o n i i w i t h S u k s d o r f i a r a n u n c u l i f o l i a i n h i s genus Hemieva  (here regarded Despite  as synonymous w i t h  these m o r p h o l o g i c a l  Suksdorfia).  s i m i l a r i t i e s , none of the three  S u k s d o r f i a s p e c i e s would f i t comfortably  i n Boykinia.  Thus  Suksdorfia  r a n u n c u l i f o l i a would.have to be put i n s e c t i o n B o y k i n i a on grounds of i n f l o r e s c e n c e s t r u c t u r e and p o l l e n morphology. trichomes and awned anthers  However, i t l a c k s the c h a f f y  t y p i c a l of t h a t s e c t i o n .  Indeed, i t l a c k s the  u n i c e l l u l a r trichomes found i n a l l B o y k i n i a s p e c i e s .  Suksdorfia alchemil-  l o i d e s and S. v i o l a c e a could be put i n s e c t i o n R e n i f o l i u m or s e c t i o n T e l e s o n i x on the b a s i s of i n f l o r e s c e n c e s t r u c t u r e although the same p o l l e n and seed morphologies as those  sections.  e q u a l l y out of p l a c e i n s e c t i o n B o y k i n i a because they and _S_. v i o l a c e a l a c k s awned anthers  as w e l l .  they do not have They would be  lack chaffy  Thus i n order  trichomes,  to accommodate  the t h r e e S u k s d o r f i a s p e c i e s i n B o y k i n i a , the proposed i n f r a g e n e r i c d i v i s i o n o f B o y k i n i a would have to be abandoned and r e p l a c e d e i t h e r by no d i v i s i o n at a l l ,  i . e . an amorphous a r r a y of s p e c i e s , or by a s u b g e n e r i c  i s i o n w i t h subgenus B o y k i n i a .on the one hand w i t h i t s p r e s e n t and  subgenus S u k s d o r f i a on the o t h e r .  E i t h e r way,  div-  sections  the v a r i a t i o n p a t t e r n  i n B o y k i n i a would be d i s t o r t e d , e s p e c i a l l y by the i n c l u s i o n of a s p e c i e s l a c k i n g u n i c e l l u l a r trichomes. a l s o p r o v i d e no support  The i n t e r g e n e r i c b r e e d i n g r e l a t i o n s h i p s  f o r the i n c l u s i o n of S u k s d o r f i a i n B o y k i n i a .  Boykinia, i n t r a - s e c t i o n a l hybrids somewhat l e s s s o .  In  are e a s i l y made and i n t e r s e c t i o n a l ones  At l e a s t i n the N o r t h American S u k s d o r f i a s p e c i e s ,  i n t r a - g e n e r i c crosses f a i l ,  as do c r o s s e s w i t h B o y k i n i a s e c t i o n B o y k i n i a .  The i n c l u s i o n of S u k s d o r f i a i n B o y k i n i a i s i n c o n s i s t e n t w i t h pattern of i n t r a g e n e r i c breeding r e l a t i o n s h i p s .  the l a t t e r ' s  207  Furthermore, B o y k i n i a and S u k s d o r f i a have a d i s t i n c t  appear-  ance such t h a t b o t h a r e c l e a r l y r e c o g n i z a b l e i n the f i e l d as s e p a r a t e groups.  The f e a t u r e s c o n t r i b u t i n g most to the appearance o f S u k s d o r f i a  are i t s g e n e r a l s l e n d e r n e s s , b a s a l l e a v e s w i t h venation  relatively  uncomplicated  and c a u l i n e l e a v e s which a r e o f t e n adnate to prominent f o l i a c e o u s  stipules.  B o y k i n i a i s much more r o b u s t , and i t s l e a v e s have a more com-  p l e x v e n a t i o n p a t t e r n ; a l l b u t two s p e c i e s l a c k f o l i a c e o u s s t i p u l e s . F l a v o n o i d d a t a a l s o suggest a d i v i s i o n between genera:  a l l but two Boy-  k i n i a s p e c i e s have 6-oxygenated f l a v o n o i d s , and a l l b u t one have d i g l y c o sides i n v o l v i n g arabinose  or x y l o s e ; these  substances a r e absent  Suksdorfia.  E c o l o g i c a l l y , the spring-wet,  is different  from the r i p a r i a n woodland o r c l i f f  i s t i c of Boykinia.  summer-dry S u k s d o r f i a h a b i t a t face"habitats character-  I t i s t o the more x e r i c c o n d i t i o n s t h a t  presumably owes i t s b u l b i f e r o u s rhizome.  from  Suksdorfia  T h i s c h a r a c t e r i s t h e most im-  p o r t a n t one i n s e p a r a t i n g S u k s d o r f i a from B o y k i n i a , which has a t h i c k , s c a l y rhizome. How much weight s h o u l d be g i v e n t o t h i s c h a r a c t e r ? B o y k i n i a and S u k s d o r f i a , s p e c i e s are s e p a r a t e d grounds o f f l o r a l morphology.  Burtt  from each o t h e r  I n both l a r g e l y on  (1964) suggested t h a t i n t h i s k i n d  of s i t u a t i o n , e v o l u t i o n a r y , or " b i o l o g i c a l " , genera might be r e c o g n i z e d on the b a s i s o f c h a r a c t e r s d i f f e r e n t  from those which a r e used i n s e p a r -  a t i n g the s p e c i e s o f the r e s p e c t i v e genera. f o r e , the rhizome d i f f e r e n c e may r e f l e c t between B o y k i n i a and S u k s d o r f i a .  I n the p r e s e n t  case,  a genuine e v o l u t i o n a r y  there-  schism  Thus, many o f the s i m i l a r i t i e s i n f l o r a l  morphology and i n f l o r e s c e n c e s t r u c t u r e can be a t t r i b u t e d t o p a r a l l e l i s m . Both genera show the same ranges o f v a r i a t i o n i n f l o r a l morphology, some species having  l a r g e , showy f l o w e r s w i t h prominent n e c t a r i e s (e.g. Boy-  k i n i a ma.jor and S u k s d o r f i a r a n u n c u l i f o l i a ) , and some h a v i n g  inconspicuous  208  f l o w e r s w i t h i n c l u d e d p e t a l s and no n e c t a r i e s (e.g. B_. r o t u n d i f o l i a and S. a l c h e m j l l o i d e s ) .  Variation i n inflorescence structure i s also  S. r a n u n c u l i f o l i a has the same k i n d o f b r a n c h i n g  similar:  p a t t e r n as i n s e c t i o n  B o y k i n i a , and the s i m p l e r s t r u c t u r e s i n S. a l c h e m j l l o i d e s and S. v i o l a c e a are m i r r o r e d by s i m i l a r p a t t e r n s i n B o y k i n i a s e c t i o n s R e n i f o l i u m and T e l e sonix, r e s p e c t i v e l y . Bolandra elongate  can be d i s t i n g u i s h e d from B o y k i n i a by i t s minute,  seeds, tubular-campanulate f l o w e r s w i t h s u b u l a t e p e t a l s , a b u l b -  i f e r o u s rhizome, absence o f u n i c e l l u l a r trichomes and i t s ibility  at least with section Boykinia.  cross-incompat-  The s i z e of the gap between  Bolan-  dra and B o y k i n i a i s s i m i l a r t o t h a t s e p a r a t i n g S u l l i v a n t i a and B o y k i n i a . Synopsis  1.  and Key t o the S e c t i o n s and S p e c i e s  of Boykinia  I n f l o r e s c e n c e composed o f p a r a c l a d i a i n the form o f many-flowered, u s u a l l y h e l i c o i d , d i c h a s i a l cymes; brown, c h a f f y trichomes  present  at l e a s t on the l o w e r stem nodes; seeds f i n e l y t u b e r c u l a t e , with  a p i c a l awns  Section Boykinia  anthers 3.  l a . I n f l o r e s c e n c e composed o f p a r a c l a d i a i n the form o f simple d i c h a s i a , or reduced to one o r two f l o w e r s ; brown c h a f f y trichomes u s u a l l y absent; seeds + smooth; anthers without . a p i c a l awns 2.  Flowers preddminently white ( r o s e v e i n s may be p r e s e n t ) ; stamens 5 . . .  2a. Flowers p u r p l e - c r i m s o n ; 3.  2.  S e c t i o n R e n i f o l i u m : 1L r i c h a r d s o n i i stamens 10 . . . . S e c t i o n T e l e s o n i x  Upper s t i p u l e s £ f o l i a c e o u s , o f t e n adnate t o the c a u l i n e l e a v e s o r b r a c t s ; n e c t a r y prominent, y e l l o w  4.  3a. Upper s t i p u l e s reduced to s m a l l f l a p - l i k e e x t e n s i o n s f r i n g e d w i t h brown b r i s t l e s ; n e c t a r y 4.  8.  o f the p e t i o l e ,  obscure, g r e e n i s h  . . .  P e t a l s ovate t o o r b i c u l a r , narrowed a b r u p t l y t o s h o r t claws;  5. inflor-  209  escence somewhat corymbiform; n o t s t o l o n i f e r o u s ; 2n = 2 6  . . . .  B_. major 4a. P e t a l s obovate t o s p a t u l a t e , a t t e n u a t e  t o the c l a w s ; . i n f l o r e s c e n c e  more p y r a m i d a l than corymbiform; s t o l o n i f e r o u s ; 2n = 14 B.  intermedia  5. ' P e t a l s w h i t e , much l o n g e r  than s e p a l s  6.  5a. P e t a l s w h i t e o r g r e e n i s h ,  about e q u a l l i n g o r s h o r t e r than s e p a l s . 7.  6.  Leaves r e n i f o r m ,  cleft  t o deeply  d i v i d e d ; 2n = 12  .  B. a c o n i t i f o l i a  6a. Leaves o r b i c u l a r , crenate 7.  Petals greenish;  to c l e f t ; 2n = 14 . .  leaves reniform,  cleft  B^ o c c i d e n t a l i s  to deeply  divided; free  hypanthium s h o r t e r than f i l a m e n t l e n g t h ; s t o l o n i f e r o u s B^ l y c o c t o n i f o l i a 7a.  Petals white; leaves o r b i c u l a r , crenate-dentate;  f r e e hypanthium  l o n g e r than f i l a m e n t l e n g t h ; n o t s t o l o n i f e r o u s . B_. r o t u n d i f o l i a 8.  P e t a l s crimson-purple, styles  o n l y j u s t exceeding s e p a l s  connate about % t h e i r l e n g t h  8a. P e t a l s crimson, about twice least  I.  B_. h e u c h e r i f o r m i s  as l o n g as s e p a l s ; s t y l e s connate a t  3/4 t h e i r l e n g t h  B_. j a m e s i i  S e c t i o n B o y k i n i a Rosendahl ex E n g l e r i n E n g l e r zenfam.  ed. 2. 18a: 120.  Euboykinia  ( i f at a l l ) ;  1928.  & P r a n t l , Nat. P f l a n -  ("Euboykinia").  Rosendahl, B o t . J a h r b . 37, B e i b l . 83: 61.  1905.  Nomen  nudum. 1.  B o y k i n i a a c o n i t i f o l i a N u t t a l l , Journ. 7:  113, 114.  1834.  Acad. N a t . S c i . P h i l a .  210  Therofon n a p e l l o i d e s R a f i n e s q u e , Neophyt. Bot. 1828?; New F l . Am.  IV. Neobot. 66.  ("1836"),  Saxifraga a c o n i t i f o l i a p l . 57.  1838.  (Nutt.) F i e l d i n g & Gardner,  Sert. P l .  1844.  Therofon a c o n i t i f o l i a U n i v . A g r i c . Exp.  (Nutt.) M i l l s p a u g h , B u l l . West V i r g . Sta.  2: 361.  1892.  Type: K i n s.n. " I n the v a l l e y s of the A l l e g h a n y (Holotype PH). as "Unaka Mts.  Rafinesque  mountains"  (1838) g i v e s the type  or I r o n Mts. of N o r t h  Therophon t u r b i n a t u m Rydberg, N. Am.  locality  Carolina".  F l . 22:  124.  1905.  B o y k i n i a t u r b i n a t a (Rydb.) Fedde, J u s t . Bot. J a h r e s b . 33(1): 607.  ("1905"),  1906.  Type: S. B. B u c k l e y s.n., September 1840, mountains,  Banks o f a stream among  N. Alabama, U n i t e d S t a t e s (Holotype  NY!).  The n o m e n c l a t u r a l problems a s s o c i a t e d w i t h B o y k i n i a a c o n i t i f o l i a have been d i s c u s s e d e a r l i e r .  The  s p e c i e s i s v a r i a b l e i n l e a f shape  and d e n t i t i o n i n the same manner as many of the o t h e r s p e c i e s i n s e c t i o n B o y k i n i a (Ch. I l l ) ; may  i t a l s o v a r i e s i n the shape of i t s seed c a p s u l e , which  range from t u r b i n a t e to u r c e o l a t e .  Rydberg (1905) d e s c r i b e d a  s p e c i e s from n o r t h e r n Alabama, Therophon t u r b i n a t u m . based ter"  and on the shape of the l e a f t e e t h .  c r i b e d as h a v i n g rounded-ovate 8mm  long.  Therophon t u r b i n a t u m was  A group  always  ( F i g . 44).  c a p s u l e up t o  of f r u i t i n g specimens was  e r a l l e a f and hypanthium dimensions diagram  des-  d i s t i n g u i s h e d by i t s l a n c e o l a t e ,  or t r i a n g u l a r l e a f t e e t h and a s p h e r i c a l or rounded 1905).  charac-  l e a f t e e t h and a t u r b i n a t e c a p s u l e up to  B o y k i n i a a c o n i t f o l i a was  l o n g (Rydberg  on t h i s  new  ovate  4mm  measured f o r sev-  and the r e s u l t s p l o t t e d on a s c a t t e r  The r e s u l t s show t h a t l e a f and  c a p s u l e shape are not  c o r e l l a t e d and t h a t both c h a r a c t e r s show a continuous range of  211  FIGURE 44. Variation i n characters used to distinguish Boykinia  turbinata  from B. a c o n i t i f o l i a . 24  o 20  U 3  o  o  o  1-2  3-4  5-6  7-8  Hypanthium length  6-5 CU  o  •-  16  4-1  O  z 3  U  o  12 O  o  O  • o-  o  O  o  L>  cfl  o-  AAQ-  O  v.  o-  90  100  110  — i —  120  130  140  150  Capsule base angle Plant origins •  Alabama  •  Georgia  •  South Carolina  O Tennessee • North. Carolina V Kentucky  capsule  leaf tooth  A  base  curvature  angle  = a/b x 100%  Virginia  160  212  variation.  Moreover although t h e r e i s some tendency f o r p l a n t s  t u r b i n a t e capsules  t o be found i n Alabama and G e o r g i a  North; C a r o l i n a and V i r g i n i a . can be found i n Alabama. i n chapter  S i m i l a r l y , plants with  having  they occur urceolate  The v a r i a t i o n i n l e a f shape has been  also i n capsules  discussed  I I I , and has been shown t o have some developmental component.  I n view o f these r e s u l t s , no r e c o g n i t i o n of a segregate s p e c i e s , o r even an i n f r a s p e c i f i c taxon, from  2.  Boykinia intermedia 168.  a c o n i t i f o l i a i s warranted.  ( P i p e r ) G. N. Jones, U n i v . Wash. Pub. B i o l . 5:  1936.  B o y k i n i a major i n t e r m e d i a  Piper, Erythea  7: 172, 173. 1899.  T h e r o f o n intermedium ( P i p e r ) H e l l e r , Muhlenbergia 1: 53. T h e r o f o n majus intermedium 11:  1904.  ( P i p e r ) P i p e r , C o n t r i b . U.S. Nat.: Herb.  311. 1906.  Type: Frank H. Lamb 1176, 10th June 1897,.New London, C h e h a l i s Co., Washington ( L e c t o t y p e Piper  US!, I s o t y p e F ! ) ,  (1899) c i t e d F. H. Lamb 1267 as the type.  mens e x i s t , one a t US and one a t F.  Two s p e c i -  The one a t F, however, i s c l e a r l y  marked " F . H. Lamb 1167", n o t 1267; on the specimen a t US the c o l l e c t i o n number i s ambiguous and c o u l d read bears the a n n o t a t i o n  1167 o r 1267.  The US specimen a l s o  " B o y k i n i a major Gray v a r . i n t e r m e d i a  specimen." and i s t h e r e f o r e chosen as the l e c t o t y p e .  Piper.  Type  Its collection  number should p r o p e r l y be i n t e r p r e t e d as 1167 i n view o f the c l e a r annot a t i o n on the i s o t y p e a t F. Although o r i g i n a l l y d e s c r i b e d  as an i n f r a s p e c i f i c taxon under  B o y k i n i a major ( P i p e r (1899) used a t r i n o m i a l i n t h e p r o t o l o g u e ) ,  there  are s u f f i c i e n t  (1915)  reasons t o f o l l o w H e l l e r (1904), P i p e r and B e a t t i e  and Jones (1936) i n r e c o g n i z i n g B. i n t e r m e d i a  as a d i s t i n c t  species.  F i r s t , the chromosome number o f B. i n t e r m e d i a  i s 2n = 14 whereas t h a t o f  213  B. major i s 2n = 26. u l a t e than  Second, B.  s t o l o n s , p e t a l s more s p a t -  clawed, much s m a l l e r f b l i a c e o u s s t i p u l e s and  escence shape from B_. major. t y p i c a l of B.  a different  T h i r d i t l a c k s the p o l y m e t h y l a t e d  inflor-  flavonols  major.  Piper product  i n t e r m e d i a has  (1899) suggested  t h a t B. i n t e r m e d i a was  the p o s s i b l e  of h y b r i d i z a t i o n between B. major and B_. o c c i d e n t a l i s .  Heller  (1904) q u e r i e d t h i s view, and i n the p r e s e n t  study, e v i d e n c e  and  of the s y n t h e t i c B_. major x  the comparative morphology and  chemistry  from c y t o l o g y  B. o c c i d e n t a l i s h y b r i d combine e m p h a t i c a l l y to r e j e c t a h y b r i d  3.  Boykinia l y c o c t o n i f o l i a Pflanzenfam.  (Maxim.) E n g l e r , i n E n g l e r & P r a n t l .  I I I . 2a: 52.  31: 41,  42.  Therophon l y c o c t o n i f o l i a 114.  Nat.  1891.  S a x i f r a g a l y c o c t o n i f o l i a Maximowicz, B u l l . Acad. Imp. Petersb.  hypothesis.  S c i . St.  1886. (Maxim.) Takeda, J o u r . B o t .  (Lond.) 49:  1911.  Neoboykiiiia l y c o c t o n i f o l i a  (Maxim.) Hara, Bot. Mag.  Toyko 51:  253.  1937. Type;  Yatabe s.n.,  2nd  August 1882,  "Senano prov.  a l p e Komaga-take,  Nippon" ( T I ? ) . In f l o r a l structure, Boykinia l y c o c t o n i f o l i a i s rather  differ-  ent  from i t s v i c a r i o u s r e l a t i v e s . B. i n t e r m e d i a and B. a c o n i t i f o l i a .  has  s m a l l g r e e n i s h p e t a l s , a v e r y reduced  f u l l y i n f e r i o r ovary,  f r e e hypanthium w i t h an almost  and i m b r i c a t e s e p a l s .  I t was  these d i f f e r e n c e s  which l e d Hara (1937) to e l e v a t e the s p e c i e s to g e n e r i c rank. been d i s c u s s e d , however, B_. l y c o c t o n i f o l i a possesses necessary  It  f o r membership i n s e c t i o n B o y k i n i a .  a l l the  As  has  characters  214  4.  B o y k i n i a o c c i d e n t a l i s T o r r e y & Gray, F l . Ni' Am. 1: 577, 698. Therofon o c c i d e n t a l i s  1840.  (T. & G.) K u n t z e , Rev. Gen. 1: 227. 1891.  B o y k i n i a e l a t a v a r . o c c i d e n t a l i s Rosendahl, B o t . J a h r b . 37, B e i b l . 83: 36.  1905.  Nomen nudum.  B o y k i n i a e l a t a (Nutt.) Greene v a r . o c c i d e n t a l i s  (T. & G.) Rosendahl  ex E n g l e r , i n E n g l e r & P r a n t l Nat. P f l a n z e n f a m . ed. 2. 18a: 120. 1928. Type: Douglas s.n. C a l i f o r n i a o r Oregon  (Holotype GH!) T o r r e y & Gray  (1840) g i v e the date of Douglas' c o l l e c t i o n as 1835, and the p l a c e as Oregon.  Douglas d i e d i n 1834, however.  S a x i f r a g a e l a t a N u t t a l l i n T o r r e y & Gray, F l . N. Am. 1: 575.  1840.  B o y k i n i a o c c i d e n t a l i s T. & G. v a r . e l a t a (Nutt.) A. Gray, P r o c .  Am.  Acad. A r t s S c i . 8: 383. . 1872. B o y k i n i a e l a t a ( N u t t . ) Greene, F l . F r a n . 190. 1891. T h e r o f o n elatum (Nutt.) Greene, Man. B o t . Reg. San F r a n . Bay 121.  1894.  B o y k i n i a n u t t a l l i i J . M. Macoun, Can. Rec. S c i . 6: 408.  1896.  Type: N u t t a l l s.n., Columbia R i v e r by Chinhook i n v e r y wet p l a c e (Rolotype BM!).  The date o f the c o l l e c t i o n i s almost c e r t a i n l y  4th J u l y 1835 ( G r a u s t e i n 1967:  312).  Therophon c i n c i n n a t u m Rosendahl & Rydberg i n Rydberg, N. Am. F l . 22: 124.  18/12/1905.  B o y k i n i a o c c i d e n t a l i s T. & G. v a r . c i n c i n n a t a Rosendahl, B o t . J a h r b . 37, B e i b l . 83: 61. cincinnata  (Rosend. & Rydb.) 22/12/1905. ("var.  (Rvdb. & Rosend.) n. comb.").  Boykinia cincinnata 33(1): 607.  (Rosend. & Rydb.) Fedde, J u s t B o t . J a h r e s b .  ("1905"), 1906.  215  B o y k i n i a e l a t a (Nutt.) Greene v a r . c i n c i n n a t a (Rosend. & Rydb.) E n g l e r , i n E n g l e r & P r a n t l Nat. P f l a n z e n f a m . ed. 2. 18a:.120. 1928.  ("var.  c i n c i n n a t a (Rosend.) n. comb.").  Type: C. G. P r i n g l e , J u l y 1882, rocky banks o f streams,  Santa  Cruz,  C a l i f o r n i a (CAN'.). Therophon vancouverense  Rydberg, N. Am. F l . 22: 125. 1905.  B o y k i n i a vancouverensis  (Rydb.) Fedde, J u s t . B o t . J a h r e s b .  607.  33(1):  ("1905"), 1906.  Type: H a r r y Edwards w.n., August 1874, Vancouver I s l a n d  (Holotype N Y l ) .  Rydberg (1905) gave the c o l l e c t i o n date as "Aug. 1 s t , 1874". The  taxonomic h i s t o r y o f B o y k i n i a o c c i d e n t a l i s i s complex.  Most o f the d i f f i c u l t i e s a r e the r e s u l t both o f i t s g r e a t v a r i a b i l i t y and the f a c t  that i t was d e s c r i b e d s i m u l t a n e o u s l y as two d i f f e r e n t s p e c i e s  i n s e p a r a t e genera:  S a x i f r a g a e l a t a N u t t a l l i n T o r r e y and Gray (1840)  and B o y k i n i a o c c i d e n t a l i s T o r r e y and Gray (1840).  A l l subsequent  have t r e a t e d the p l a n t s as b e l o n g i n g t o B o y k i n i a (or T h e r o f o n ) .  works As more  m a t e r i a l was c o l l e c t e d , the range o f v a r i a t i o n i n c r e a s e d such t h a t Rydberg (1905) f e l t  compelled  t o r e c o g n i z e f o u r s p e c i e s , and Rosendahl  one w i t h two v a r i e t i e s .  A comparison  d i s t r i b u t i o n o f supposedly  o f these c l a s s i f i c a t i o n s  (1905) two,, and the  d i a g n o s t i c c h a r a c t e r s i s shown i n T a b l e XXVII.  U n t i l now, e x p e r i m e n t a l e v i d e n c e on the s p e c i e s has been lacking. the degree ility,  The o b s e r v a t i o n s i n t h e p r e s e n t work on the amount o f pubescence, of p e d i c e l curvature, inflorescence structure,  cross-compatib-  p o l l e n and seed morphology, c y t o l o g y and f l a v o n o i d p r o f i l e i n pop-  u l a t i o n s from Vancouver I s l a n d t o C a l i f o r n i a are reviewed The  here.  f l a v o n o i d p r o f i l e s o f a l l p o p u l a t i o n s were q u a l i t a t i v e l y  s i m i l a r , a c o n d i t i o n common i n many o t h e r w i d e l y - d i s t r i b u t e d s p e c i e s o f Saxifraginae  (Bohm and W i l k i n s 1978b).  Conversely, d i f f e r e n t species  216  TABLE XXVII.  C h a r a c t e r s supposedly  Boykinia occidentalis Torr.  diagnostic  & Gray, i n c l u d i n g  of taxa s e g r e g a t e d a comparison of the  f i c a t i o n s of Rydberg (1905) and Rosendahl (1905). Rydberg  on  (1905).  I n f l o r e s c e n c e dichotomouscymose, p e d i c e l s curved . . .  Rosendahl  2.  Hypanthium d e n s e l y g l a n d u l a r , p u r p l i s h ; l e a f blades firm; petioles hirsute  cincinnatum  2a. Hypanthium s p a r i n g l y g l a n d occidentalis u l a r , green; l e a f b l a d e s t h i n ; p e t i o l e s glabrous or w i t h a few s c a t t e r e d h a i r s  occidentalis var. cincinnatum occidentalis  l a . I n f l o r e s c n e c e i r r e g u l a r l y cymose, pedicels straight 3. 3.  classi-  The key i s based  Rydberg  2.  from  S e p a l s i n f r u i t r e f l e x e d or spreading; inflorescence p u b e r u l e n t , green  3a. Sepals i n f r u i t e r e c t or ascending; i n f l o r e s c e n c e densely g l a n d u l a r , p u r p l i s h  vancouverense  elata  elatum  elata  217  o f t e n have d i f f e r e n t p r o f i l e s .  No  tween p o p u l a t i o n s i n terms o f seed  significant  d i f f e r e n c e s were found  and p o l l e n morphology.  h i g h l y v a r i a b l e , p a r t l y a r e s u l t of heteroblasty. are o f t e n l e s s d i v i d e d than those  formed l a t e r .  than those  growing i n b r i g h t e r l i g h t , and  typically plastic  There i s a complete  f o r t h i s c h a r a c t e r tp be pheno-  (e.g. Evans and Hughes 1961).  A l l c r o s s e s between d i f the progeny were  D e n s i t y of pubescence has been shown to be  i n i t s v a r i a t i o n and v a r i a b l e w i t h variation. continuous  age.  fully  Inflorescence structure i s also environmentally v a r i a t i o n makes i t i m p o s s i b l e  continuous  labile,  and  to r e c o g n i z e d i s c r e e t groups.  (1906) comment t h a t the v a r i a n t c i n c i n n a t u m  i s only " f e e b l y d i s The  col-  of the i n f l o r e s c e n c e branches and hypanthium i s v a r i a b l e o f t e n  between p l a n t s of the same p o p u l a t i o n , and d a t i o n from red to green. The  14  S e p a l r e f l e x i o n shows s i m i l a r  t i n g u i s h a b l e " and not worthy of r e c o g n i t i o n i s endorsed h e r e . ouring  fully  A l l p o p u l a t i o n s s t u d i e d were u n i f o r m l y d i p l o i d w i t h 2n =  chromosomes.  Piper's  In terms of  t o have t h i n n e r l e a v e s  f e r e n t p o p u l a t i o n s were h i g h l y s u c c e s s f u l , and fertile.  L e a f shape i s  Lowermost b a s a l l e a v e s  i n t e r g r a d a t i o n i n shape and o u t l i n e between p o p u l a t i o n s . l e a f t h i c k n e s s , i t i s common f o r shaded.plants  be-  t h e r e i s a complete  Sometimes o n l y one  s i d e of the stem i s c o l o u r e d .  c o l o u r a t i o n i s a t t r i b u t a b l e to anthocyanins  whose c o n c e n t r a t i o n i n a  p l a n t i s g r e a t l y a f f e c t e d by b o t h environment and The  data presented  intergra-  age.  here i n d i c a t e t h a t a s i n g l e , v a r i a b l e spe-  cies^ s h o u l d be r e c o g n i z e d , a path f o l l o w e d by a l l r e c e n t r e g i o n a l f l o r a s ( H i t c h c o c k e t a l . 1961;  Peck 1961;  Regarding the  Munz 1959).  c o r r e c t name f o r the s p e c i e s , the e a r l i e s t names  were p u b l i s h e d s i m u l t a n e o u s l y  ( T o r r e y and Gray 1840):  N u t t a l l and B o y k i n i a o c c i d e n t a l i s T o r r e y and Gray. the c h o i c e of e p i t h e t i s the p e r o g a t i v e of the f i r s t  Saxifraga elata  In t h i s  circumstance  author who  combines  218  the names ( A r t . 57 o f the Code).  I n the supplement t o t h e i r f l o r a ,  Torrey  and Gray (1840) commented t h a t N u t t a l l ' s S. e l a t a "proves t o be a t r u e B o y k i n i a " and s t r o n g l y i m p l i e d t h a t i t was synonymous w i t h t h e i r B. occidentalis.  Indeed, Gray (1842) s t a t e d t h a t _§_. e l a t a was r e f e r r e d t o  B. o c c i d e n t a l i s i n t h e supplement.  However, i t was n o t u n t i l  1872 t h a t  a f o r m a l combination was made, whereby S. e l a t a N u t t . was g i v e n v a r i e t a l s t a t u s under _B_. o c c i d e n t a l i s (Elihu Hall  (Gray 1872) .  U n h a p p i l y , the specimen  158) a c t u a l l y b e l o n g s t o B o y k i n i a major, a f a c t l a t e r  n i z e d by Gray (1876).  cited recog-  H i s d i f f i c u l t y stemmed from u n c e r t a i n t y over whet-  h e r B^ major graded i n t o J B .  o c c i d e n t a l i s i n Oregon, and i f s o , he thought  t h a t p l a n t s there might be r e f e r r a b l e to N u t t a l l ' s taxon (Gray 1876). However, he continued t o r e g a r d S. e l a t a as " w h o l l y o r p a r t l y w i t h B_. o c c i d e n t a l i s  (Gray 1876).  synonymous"  Macoun (1883) u n e q u i v o c a l l y reduced  S. e l a t a to complete synonymy under JB_. o c c i d e n t a l i s . a l t h o u g h he l a t e r changed h i s mind  (Macoun  Boykinia N u t t a l l i i .  1896) when he renamed N u t t a l l ' s S_. e l a t a as  The new name was deemed n e c e s s a r y because  (1891) had made the combination B o y k i n i a e l a t a w i t h  Greene  o c c i d e n t a l i s as a  synonym. Although almost a l l r e c e n t treatments have used B o y k i n i a e l a t a - ( N u t t . ) Greene  ( H i t c h c o c k e t a l . 1961; Peck 1961; Munz 1959), the  c o r r e c t name i s B o y k i n i a o c c i d e n t a l i s T o r r . & Gray, which i s the name used i n the i n v e n t o r y o f the B r i t i s h Columbia f l o r a  ( T a y l o r and MacBryde  1977).  5.  B o y k i n i a r o t u n d i f o l i a P a r r y i n Gray, P r o c . Am. Acad. A r t s S c i . 13: 371.  1878.  T h e r o f o n r o t u n d i f o l i a Wheelock, B u l l . T o r r . B o t . Club 23: 70. 1896. Type:  C. C. P a r r y and J . G. Lemmon 113, 1876, San B e r n a r d i n o Co.,  219  C a l i f o r n i a (Holotype GH!).  The t y p e ' l o c a l i t y  i s apparently i n  Waterman Canon on the s o u t h e r n s l o p e s of the San B e r n a r d i n o Mountains 6.  ( P a r i s h 7147,  8 July  B o y k i n i a major A. Gray, Bot. C a l i f .  W.  H. Brewer 2166?, 1863,  ft  ( L e c t o t y p e GHl); Syntypes:  Gen.  (GH!), i b i d ,  Grove,  California  California  1: 227.  CAN).  F l a t , N.  1891. C a l i f o r n i a , 4000  Thomas B r i d g e s 126,  California  Meadows, B i g - t r e e Grove Yo-  (GH!)'-, i b i d , M a r i p o s a  (Fl), Elihu Hall  and  1876.  Robinson's  ( G H l ) , H. N. Bolander 4982, 1866, semite  specimens a t OSC  1: 196.  T h e r o f o n major (Gray) Kuntze, Rev. Type:  1909,  158,  1871,  Sequoia  I n the rocky bed  of S i l v e r Creek or margins, Oregon (GHl F ! ) . In h i s p r o t o l o g u e , Gray comparison  (1876) c i t e d f o u r specimens.  Careful  o f the d e s c r i p t i o n w i t h these specimens shows t h a t they were  a l l used i n i t s c o n s t r u c t i o n , , a l t h o u g h o n l y B o l a n d e r 4982'("Yosemite") at GH i s annotated " B o y k i n i a major Gray n. sp.". isosyntypes of Bolander's c o l l e c t i o n . are  There appear  to be  U n c e r t a i n t y e x i s t s because  two  both  i d e n t i f i e d as B o y k i n i a o c c i d e n t a l i s T. & G. and y e t bear the number  4982 as i n the specimen at GH has ink  i d e n t i f i e d as IS_. maj o r .  the c o l l e c t i o n number e n t e r e d i n p e n c i l r a t h e r than the b l a c k  of the r e s t o f the l a b e l .  different.  The p u t a t i v e i s o s y n t y p e  The  The h a n d w r i t i n g of the number a l s o seems  s u s p e c t e d i s o s y n t y p e a t F bears an a u t h e n t i c c o l l e c t i o n  number, 4982, b u t the l o c a t i o n i s g i v e n as "Mariposa Sequoia r a t h e r than " B i g - t r e e Grove"  as i n the GH syntype.  Grove"  The g e n e r a l s i m i l a r -  i t y of the specimens suggests t h a t they do r e p r e s e n t the same c o l l e c t i o n . The syntypes i n b e s t c o n d i t i o n are those o f Brewer and H a l l . o f a l e c t o t y p e s h o u l d p r e f e r a b l y be from these two, of  the d i f f e r e n c e s i n the a n n o t a t i o n  Hall  c o l l e c t i o n was  Selection  especially  of B o l a n d e r ' s c o l l e c t i o n .  i n view The  E.  c i t e d f o l l o w i n g the p u b l i s h e d combination o f B o y k i n i a  220  o c c i d e n t a l i s T. & G. v a r . e l a t a p o s s i b l e source  (Nutt.) Gray (1872), and i s t h e r e f o r e a  o f c o n f u s i o n should  i t be chosen as the l e c t o t y p e o f B.  major. The b e s t c o l l e c t i o n t o serve as t h e l e c t o t y p e i s t h e r e f o r e W.H. Brewer's from Robinson's F l a t , C a l i f o r n i a . to be 2166 but the h a n d w r i t i n g  The c o l l e c t i o n number appears  i s unclear.  B o y k i n i a major i s the l a r g e s t s p e c i e s i n t h e genus. I n f l o r e s c ence s t r u c t u r e can v a r y somewhat, r a t h e r as i n B. o c c i d e n t a l i s , w i t h some populations having so.  r e g u l a r dichotomous b r a n c h i n g w h i l e i n o t h e r s i t i s l e s s  No i n f r a s p e c i f i c  warranted.  taxa have been d e s c r i b e d on t h i s b a s i s , n o r a r e they  I n a few p o p u l a t i o n s  toothed p e t a l s .  t h e r e i s a polymorphism f o r e n t i r e v s .  Some p o p u l a t i o n s a l s o have f l o w e r s w i t h p u r p l e b l o t c h e s a t  the base o f the p e t a l s . Taxonomic r e c o g n i t i o n a t the l e v e l o f forma c o u l d be  g i v e n , although  one might merely be l a b e l l i n g s i n g l e genes.  I therefore  p r e f e r simply t o r e c o r d the e x i s t e n c e o f these v a r i a n t s r a t h e r than  give  them f o r m a l s t a t u s .  II.  S e c t i o n R e n i f o l i u m Rosendahl ex E n g l e r i n E n g l e r & P r a n t l , Nat. P f l a n zenfam. ed. 2. 18a: 120. 1928. R e n i f o l i u m Rosendahl, B o t . J a h r b . 37, B e i b l . 83: 62. 1905. Nomen nudum.  7.  Boykinia r i c h a r d s o n i i  (Hook.) Rothrock, Ann. Rep. Board Reg. Smiths.  I n s t . 1867: 447. 1868. S a x i f r a g a R i c h a r d s o n i i Hooker, F l . Bor. Am. 1: 247. 1832. Hemieva r i c h a r d s o n i i  (Hook.) R a f i n e s q u e ,  F l . T e l l . 2: 70.  ("1836"), 1837. Therofon  R i c h a r d s o n i i (Hook.) Kuntze, Rev. Gen. 1: 227. 1891.  Saxifraga Nelsoniana  auct non D. Don:  Beechey's Voy. 124. t.29.  1832.  Hooker & A r n o t t , B o t .  221  Type:  Lay and K'.).  C o l l i e s.n.,  Dr. Richardson  Imarook, Kotzebue's Sound (Isosyntypes s.n.,  A r c t i c Sea  shore  Hooker (1832) r e p o r t e d t h a t Richardson's between the Mackenzie and K bears  Coppermine Rivers..  (Syntype K l ) .  specimen came from  Lay and  the name "Beechey" r a t h e r than t h e i r own,  GH,  C o l l i e ' s specimen a t  but must have been  coll-  e c t e d by them as they were the e x p e d i t i o n ' s n a t u r a l i s t s . T h e i r c o l l e c t i o n s were made between J u l y and  September, 1826  (Hooker and A r n o t t 1 8 3 2 ) .  c u r r e n t works d e a l i n g with, the A r c t i c f l o r a c r e d i t Gray (1876) w i t h t r a n s f e r of the s p e c i e s i n t o B o y k i n i a . although f i r s t made t h i s  III.  i t was  All the  Rothrock (1868)  who  combination.  Section Telesonix  (Raf.) G o r n a l l , s t a t .  T e l e s o n i x Rafinesque,  nov.  F l . T e l l . 2: 69.  Reasons f o r t r e a t i n g the two  ("1836"),  1837.  taxa as members of a s e p a r a t e  sect-  i o n i n B o y k i n i a , r a t h e r than i n s e c t i o n R e n i f o l i u m or i n S a x i f r a g a or T e l e s o n i x have a l r e a d y been d i s c u s s e d .  The problem remaining  should be a t t r i b u t e d to each o f them. each o t h e r and berg  (1897) was  O r i g i n a l l y they were confused  t r e a t e d as a s i n g l e , v a r i a b l e s p e c i e s the f i r s t  (Hooker 1832).  to d e s c r i b e t h e i r d i f f e r e n c e s and  v a r i a n t s at s p e c i f i c l e v e l w i t h the e p i t h e t s j a m e s i i and ("heucheraeforme").  The  g r e a t e r c a r p e l f u s i o n i n j a m e s i i than i n heuch-  under j a m e s i i .  a wide range of herbarium m a t e r i a l has  Examination.of  reduced  heucheriformis  to E n g l e r  who  to v a r i e t a l  e n t i t i e s show d i s c o n t i n u o u s v a r i a t i o n i n p e t a l s i z e  show s t a t i s t i c a l d i f f e r e n c e s i n c a r p e l f u s i o n and (Table X X V I I I ) .  both  heucheriformis  (1891) or to B a c i g a l u p i . ( 1 9 4 7 )  ratio  Ryd-  recognised  These d i s t i n c t i o n s were a p p a r e n t l y not so i m p r e s s i v e  e a l e d t h a t the two  with  d i s t i n c t i o n i s based on the much l a r g e r p e t a l s i z e ,  d i f f e r e n t p e t a l c o l o u r and eriformis .  i s what rank  anther/filament  status revand  length  Furthermore, p o p u l a t i o n s w i t h s m a l l p e t a l s have p o l l -  222  TABLE XXVIII.  M o r p h o l o g i c a l v a r i a t i o n between B o y k i n i a j a m e s i i and B.  heucheriformis.  Data a r e g i v e n as the mean + one s t a n d a r d  d e v i a t i o n , and  the range ( i n b r a c k e t s ) . B. j a m e s i i (N = 10)  B. h e u c h e r i f o r m i s  Fetal/Sepal length r a t i o  2.15 + 0.48 (1.78 - 3.33)  1.04 + 0.18 (0.67 - 1.31)  Petal/Sepal width r a t i o  2.01 + 0.56 (1.18 - 2.90)  0.86 + 0.21 (0.53 - 1.29)  Degree o f c a r p e l freedom*  0.13 + 0.06 (0.06 - 0.24)  0.40 + 0.10 (0.20 - 0.49)  122 (110  148 (130  P e t a l claw  angle ** 0  + 10 - 140)  + 12 - 170)  Free hypanthium l e n g t h i n mm  1.8 + 0.3 (1.4 - 2.5)  2.3 + 0 . 4 (1.9 - 3.4)  Anther/Filament length r a t i o  0.96 + 0.21 (0.60 - 1.22)  1.40 + 3.4 (0.90 - 1.83)  *  C a r p e l freedom = f r e e c a r p e l l e n g t h / t o t a l c a r p e l l e n g t h .  **  Claw angle = a  en w i t h a d e n s e l y with occluded  r e t i c u l a t e tectum.  (N  Those w i t h l a r g e p e t a l s have a tectum  luminae such t h a t i t appears t o be continuous  (Ch. V I I ) .  The  e c o l o g i c a l and g e o g r a p h i c a l ranges o f the two v a r i a n t s do n o t o v e r l a p , B. jamesii being r e s t r i c t e d  to the g r a n i t e peaks o f Colorado  and New Mexico,  and B_. h e u c h e r i f ormis b e i n g more widespread on c a l c a r e o u s s u b s t r a t e s from A l b e r t a t o Wyoming, South Dakota and Utah, and d i s j u n c t i n southern T h e r e f o r e , I propose t o t r e a t the v a r i a n t s as s e p a r a t e  Nevada.  s p e c i e s based on  c o r r e l a t e d d i s c o n t i n u o u s v a r i a t i o n i n two c h a r a c t e r s and q u a n t i t a t i v e d i f f erences 8.  i n others.  B o y k i n i a j a m e s i i ( T o r r . ) E n g l e r i n E n g l e r & P r a n t l , Nat. Pflanzenfam. III.  2a: 51.  1891.  S a x i f r a g a J a m e s i i T o r r e y , Ann. L y c . Nat. H i s t . New York 2: 204. ' 1827. Telesonix j a m e s i i (Torr,) Rafinesque,  F l . Tell.  2: 69.  ("1836"),  223  1837.  (in part).  Therofon J a m e s i i ( T o r r . ) Wheelock, B u l l . T o r r . B o t . Club 23: 70. Type:  1896.  Dr. James.8, Rocky Mountains ( L e c t o t y p e NY'. I s o t y p e s K'. COLO'.). The NY specimen comes from T o r r e y ' s herbarium  "n. sp.".  I t i s t h e r e f o r e d e s i g n a t e d as the l e c t o t y p e .  was  made on 14th J u l y 1820 a t P i k e ' s Peak, Colorado  9.  Boykinia heucheriformis 62.  (Rydb.) Rosendahl,  and i s annotated James' c o l l e c t i o n  (Osterhout  1920).  B o t . Jahrb. 37, B e i b l . 83:  1905. ("heucheriforme"). Therofon heucheraeforme Rydberg, B u l l . T o r r . B o t . Club 24: 247, 248.  1897.  Telesonix j a m e s i i (Torr.) Rafinesque, 1837.  F l . Tell.  2: 69.  ("1836"),  (in part).  T e l e s o n i x h e u c h e r i f o r m i s (Rydb.) Rydberg, N. Am. F l . 22: 126. 1905. B o y k i n i a J a m e s i i v a r . h e u c h e r i f o r m i s Rosendahl. B e i b l . 83: 34.  B o t . J a h r b . 37,  1905. Nomen nudum.  S a x i f r a g a heucheriforme  M.E. Jones, B u l l . U n i v . Mont. 61: 32.  1910. Boykinia Jamesii (Torr.) Engl. var. heucheriformis E n g l e r i n E n g l e r & P r a n t l , N a t . Pflanzenfam. 120.  (Rydb.)  ed. 2. 18a:  1928.  T e l e s o n i x J a m e s i i ( T o r r . ) R a f . v a r . heucheriforme  (Rydb.)  B a c i g a l u p i , L e a f l . West. B o t . 5: 71. 1947. S a x i f r a g a J a m e s i i a u c t . non T o r r e y : 247. Syntypes:  Hooker, F l . Bor. Am. 1:  t.84. 1832.  Flodman 514, 28th J u l y 1896, B r i d g e r Mountains, Montana (NY);  224  Rydberg 2677, 23rd J u l y 1895, F. Tweedy 255, The "Drummond s.n.,  1887,  Bozeman Canon, Montana  East Boulder,  Montana  (NY);  (NY).  specimen on which Hooker's (1832) d e s c r i p t i o n was Rocky Mountains" (K'.).  the Canadian R o c k i e s . ( G r a u s t e i n  1967:  I t was 218,  based i s  almost,certainly collected i n  273).  SUKSDORFIA Taxonomic Treatment S u k s d o r f i a A. Gray (nom.  c o n s . ) , P r o c . Am.. Acad. A r t s S c i . 15:  Hemieva R a f i n e s q u e .  F l . Tell.  Hieronvmusia E n g l e r , Not. 7: 265-267. Type s p e c i e s :  2:  Kon.  70.  ("1836"),  Bot. G a r t . Mus.  42.  1879.  1837. Berlin-Dahlem  Acad. A r t s S c i . 15:  1879.  S u k s d o r f i a , as c i r c u m s c r i b e d here, monotypic segregate  42.  1918.  S u k s d o r f i a v i o l a c e a A. Gray, P r o c . Am. 41,  41,  i s composed of t h r e e  genera: Hemieva, S u k s d o r f i a and Hieronymusia.  former Their  h i s t o r y began i n 1832.when Hooker d e s c r i b e d some p l a n t s from the K e t t l e F a l l s o f the Columbia R i v e r as S a x i f r a g a r a n u n c u l i f o l i a .  He noted t h a t the  d i f f e r e d g r e a t l y from a l l o t h e r S a x i f r a g a s p e c i e s but he was speculate  which might be  i t s nearest  ally.  genus Hemieva to accommodate the s p e c i e s . a r d s o n i i Hook..in h i s new f l o r a l d i s c were important ima  r e l u c t a n t to  R a f i n e s q u e (1837) e r e c t e d  the  also included Saxifraga  rich-  He  genus, c o n s i d e r i n g t h a t the f i v e stamens and c h a r a c t e r s which suggested a f f i n i t i e s  and Heuchera r a t h e r than w i t h  Saxifraga.  T o r r e y and  with  to B o y k i n i a , and  wrote to Hooker t h a t both _S_. r i c h a r d s o n i i and  S. r a n u n c u l i f o l i a "would  r e s p e c t i v e l y made the n e c e s s a r y  Rothrock (1868) and  combinations.  the Tell-  Gray (1840) however,  suggested t h a t S_j_ r a n u n c u l i f o l i a c o u l d belong  more c o r r e c t l y t r a n s f e r r e d . to Boykinia";.  species  Gray  (1842) be  Greene (1891)  225  In 1878,  Suksdorf  c o l l e c t e d from the Columbia Gorge i n Washing-  ton a s a x i f r a g e which he a s s i g n e d d o u b t f u l l y t o B o y k i n i a  (Suksdorf,  w i t h h o l o t y p e ) , but which Gray (1879) r e c o g n i s e d as b e l o n g i n g naming  i t Suksdorfia violacea.  i n note  to a new  genus,  Gray, however, d i d n o t e a p o s s i b l e r e l a t i -  onship w i t h B o y k i n i a , i n which he presumably i n c l u d e d ' S. r a n u n c u l i f o l i a . In the same y e a r , G r i s e b a c h illoides  (1879) d e s c r i b e d S a x i f r a g a alchem-  from the Andes of n o r t h e r n A r g e n t i n a .  Later, Engler  (1891) con-  s i d e r e d S a x i f r a g a r a n u n c u l i f o l i a , S. a l c h e m i l l o i d e s and S u k s d o r f i a v i o l a c e a to have s u f f i c i e n t v e g e t a t i v e s i m i l a r i t y t o warrant t h e i r u n i f i c a t i o n under Suksdorfia. Hemieva,  Wheelock  (1896) t r a n s f e r r e d the N o r t h American s p e c i e s  on grounds of p r i o r i t y .  Hemieva ( B r i q u e t 1906).  Engler  S u k s d o r f i a was  l a t e r conserved  into  against  (1918) then e l e v a t e d _S_. a l c h e m i l l o i d e s to  g e n e r i c rank as Hieronymusia, on grounds o f f l o r a l  morphology.  Because o f the absence of any c o o r d i n a t e d study o f t h e i r c t i v e b i o l o g i e s , t w e n t i e t h century works d e a l i n g w i t h _S_. v i o l a c e a and r a n u n c u l i f o l i a have t r e a t e d them i n d i f f e r e n t ways. galupi  (1944, 1952), Peck  Rydberg  S.  (1905), B a c i -  (1961) and Moss (1959) r e t a i n e d them i n s e p a r a t e  monotypic genera ( S u k s d o r f i a and Hemieva), whereas o t h e r s regarded congeneric,  respe-  them as  e i t h e r i n Hemieva ( H e l l e r 1900; Howell 1903; P i p e r 1906;  1915 and Peck 1941) or i n S u k s d o r f i a (Rosendahl 1905; Dandy 1927;  Henry  Hitchcock  et a l . 1961; Scoggan 1978).  Jepson (1925, 1936) r e t a i n e d S. r a n u n c u l i f o l i a  i n B o y k i n i a i n h i s treatment  o f the C a l i f o r n i a f l o r a ; JS^ v i o l a c e a was not  c o n s i d e r e d because i t does ,not occur i n C a l i f o r n i a . l l o i d e s has c o n t i n u e d ception  Hieronymusia a l c h e m i -  to be r e c o g n i s e d as a monotypic genus s i n c e i t s i n -  (Dandy 1927; E n g l e r  1928).  Is t h e r e any j u s t i f i c a t i o n f o r the r e c o g n i t i o n o f the t h r e e monotypic genera?  H i s t o r i c a l l y . t h e y have been s e p a r a t e d  f l o r a l c h a r a c t e r s alone  (Table XXIX).  on the b a s i s o f  These c h a r a c t e r s a r e the same ones  226  TABLE XXIX.  F l o r a l differences separating Suksdorfia r a n u n c u l i f o l i a ,  S. a l c h e m j l l o i d e s and  S. v i o l a c e a .  Character  Species S.  White, sometimes p u r p l i s h at base, f a d i n g to y e l l o w white  Rose, sometimes white  P i n k , s ome times white  P e t a l shape  Spatulate,  Oval,  Spatulate,  Petal  Longer than c a l y x  Less than, or equalling caylx  Longer than  Nectary d i s c  Present  Absent  Absent  Anthers  Awnless, on filaments  Awned, on filamnets  Awnless, almost sessile  Stigmata  Capitate  Capitate  Truncate  Ovary  Ca.  Inferior  Ca. 2/3 or more inferior  Petal  colour  length  inferior  which are used to separate trends  clawed  alchemilloides  S. v i o l a c e a  S. r a n u n c u l i f o l i a  clawless  d i f f e r e n t s p e c i e s of B o y k i n i a ,  i n f l o r a l v a r i a t i o n are s i m i l a r i n the two  filament lengths, nectary changes i n b r e e d i n g  and  p e t a l shape and  system and p o l l i n a t o r  genera.  Beginning with  protologue  Engler  the  r e l a t e d to  fauna. characters  are exam-  the rhizome, i t i s i n t e r e s t i n g t h a t a l t h o u g h i n the  of S a x i f r a g a a l c h e m i l l o i d e s , G r i s e b a c h  "tuberiforme",  indeed  caylx  Differences i n  c o l o u r c o u l d be  A d i f f e r e n t p i c t u r e emerges i f v e g e t a t i v e ined.  and  clawed  (1891, 1918)  believed  (1879) d e s c r i b e d  i t as  t h a t b u l b i l s were l a c k i n g .  Presumably t h i s i n f l u e n c e d h i s d e c i s i o n to c r e a t e the monotypic Hieronymusia.  Examination  study has  revealed  o f a v a i l a b l e m a t e r i a l of S_. a l c h e m i l l o i d e s i n the  t h a t the s p e c i e s d e f i n i t e l y p o s s e s s e s a b u l b i f e r o u s  zome, thereby c l e a r l y s u p p o r t i n g Further All  present  a l i n k with  i t s N o r t h American  rhi-  counterparts.  evidence of a c l o s e a f f i n i t y comes from the trichome complement.  t h r e e s p e c i e s possess m u l t i s e r i a t e g l a n d u l a r  trichomes.  In a d d i t i o n ,  227  b o t h jS_. v i o l a c e a and common i n B o y k i n i a . v i o l a c e a and  a l c h e m i l l o i d e s have the u n i c e l l u l a r trichomes so They p r o v i d e  good support  S_. a l c h e m i l l o i d e s .  f o r the c o n g e n e r i c  Perhaps the most obvious  between the t h r e e s p e c i e s i s t h e i r u n i f o r m i t y of h a b i t . with  crenate  to d e e p l y - d i v i d e d  v a r i a t i o n occurs  i n Boykinia)  to t h e c a u l i n e l e a v e s . morphologies and  The  , cordate and  status  of  similarity  They are  slender,  to r e n i f o r m b a s a l l e a v e s  (similar  f o l i a c e o u s , s t i p u l e s which are o f t e n adnate  s p e c i e s a l s o share v e r y s i m i l a r p o l l e n and  e c o l o g i c a l preferences.  seed  C y t o l o g i c a l l y o n l y the North Ame-  r i c a n s p e c i e s are known, b o t h w i t h 2n = 14 chromosomes.  In the i n f l o r e s c e n c e  t h e r e i s a p r o g r e s s i o n from the complex s t r u c t u r e w i t h p a r a c l a d i a of d i c h otomous h e l i c o i d cymes i n chemi1loides,  d own  r a n u n c u l i f o l i a , to a s i m p l e r v e r s i o n i n S. a l -  to the v e r y simple  c o n d i t i o n i n S. v i o l a c e a .  ence v a r i a t i o n thus p a r a l l e l s t h a t i n B o y k i n i a . file,  r a n u n c u l i f o l i a and  f l a v o n e s and  polymethylated  what d i s t i n c t glycosides.  flavonols.  i n l a c k i n g f l a v o n e s , 3-0I t does however, share  and  4'-0-  methylation  7-0-methylation and  violacea.  Again,  and  4'-0-  an a r r a y of  similar total  trigly-  variation  i n Boykinia.*  the t h r e e s p e c i e s .  Admittedly  of v a r i a t i o n i n morphology and in Boykinia. Boykinia. and  possessing  S u k s d o r f i a a l c h e m i l l o i d e s i s some-  A l t o g e t h e r , t h e r e f o r e , the evidence een  In terms o f f l a v o n o i d p r o -  S_. v i o l a c e a are v e r y s i m i l a r indeed,  c o s i d e s s i m i l a r to those, i n ^ occurs  Infloresc-  The  i n d i c a t e s an a f f i n i t y betw-  they are v e r y d i s t i n c t i v e but  chemistry  V a r i a t i o n i n . p o l l e n and  i s s i m i l a r t o , and p a r a l l e l s  v i o l a c e a are t h e r e f o r e b e s t t r e a t e d by ( E n g l e r 1891)•.  that,  seed morphology i s l e s s than t h a t i n  r e l a t i o n s h i p s between S^ a l c h e m i l l o i d e s ; S.  genus, S u k s d o r f i a  the.range  ranunculifolia  the r e c o g n i t i o n of a s i n g l e  In. view of the v a r i a t i o n w i t h i n  t h r e e monotypic s e c t i o n s c o u l d be r e c o g n i s e d , ment of the same v a r i a t i o n i n B o y k i n i a .  commensurate w i t h  Suksdorfia.  the  treat-  However, r a t h e r than c l u t t e r  the  228  nomenclature, i t seems s u f f i c i e n t  to r e a l i s e t h a t the t h r e e s p e c i e s  are  distinct.  discussed.  The  s e p a r a t i o n of S u k s d o r f i a from B o y k i n i a has  The  d i s t i n c t i o n s between S u k s d o r f i a and  S a x i f r a g a are the same  as those which d i s t i n g u i s h B o y k i n i a from S a x i f r a g a .  S u k s d o r f i a i s separated  from S u l l i v a n t i a on the bases o f rhizome morphology, seed o l o g y , s t y l e anatomy, e c o l o g y between S u k s d o r f i a and  and  Bolandra  g e n e r a l appearance.  cussed under Synopsis 1.  The  i n many ways  and Key  to the Species of S u k s d o r f i a  l a . B a s a l l e a v e s 1-2  times  a nectary  ranunculifolia present;  and  stigmata  2  V  with  awned, on f i l a m e n t s  .  2. S^ a l c h e m i l l o i d e s  2a. P e t a l s about twice as l o n g as s e p a l s ; anthers  and  stigmata l a c k i n g  u n i s e r i a t e e g l a n d u l a r h a i r s ; anthers unawned, + s e s s i l e 1.  flowers  . . . . . . . . .  P e t a l s e q u a l l i n g or s h o r t e r than s e p a l s ; anthers u n i s e r i a t e e g l a n d u l a r h a i r s ; anthers  l a c k i n g ; flowers  1.  c r e n a t e ; u n i c e l l u l a r trichomes  u s u a l l y p i n k or r o s e , w i t h o u t  .  Bolandra  distinction is dis-  B a s a l l e a v e s t e r n a t e l y d i v i d e d ; u n i c e l l u l a r trichomes  :  distinction  Bolandra.  w h i t e w i t h prominent n e c t a r y d i s c  2.  and p o l l e n morph-  The  i s l e s s c l e a r - c u t j and  i s i n t e r m e d i a t e between S u k s d o r f i a and B o y k i n i a .  a l r e a d y been  Suksdorfia r a n u n c u l i f o l i a n z e n f a m i l i e n I I I . 2a: 52.  . S^ v i o l a c e a  (Hook.) E n g l e r i n E n g l e r & P r a n t l , Nat.  Pfla-  1891.  S a x i f r a g a r a n u n c u l i f o l i a Hooker, F l . B o r . Am.  1: 246,  247.  t.83.  1832. Hemieva r a n u n c u l i f o l i a ("1836"),  (Hook.) R a f i n e s q u e ;  F l . Tell.  2:  70.  1837.  B o y k i n i a r a n u n c u l i f o l i a . ( H o o k . ) Greene, F l . F r a n . 190,  191.  1891.  229  Type: Douglas s.n.,  1826,  on the Columbia  rocks on the mountains near K e t t l e  ( L e c t o t y p e BM!  Only on the BM the p r o t o l o g u e , and  Wet  I s o t y p e K'.).  specimen does the a n n o t a t i o n match t h a t g i v e n i n  t h i s i s therefore designated  sheet i s a mixed c o l l e c t i o n .  The  the l e c t o t y p e .  two)  remaining  p l a n t s on the sheet  (the bottom  Columbia R i v -  types.  Suksdorfia a l c h e m i l l o i d e s (Griseb.) Engler i n Engler & P r a n t l , Pflanzenfam.  I I I . 2a: 52.  G o t t . 24:  142,  Nat.  1891.  S a x i f r a g a a l c h e m i l l o i d e s G r i s e b a c h , Abhandl. Kon. 143.  Ges.  Wiss.  1879.  Hieronymusia a l c h e m i l l o i d e s ( G r i s e b . ) E n g l e r , Not. Kon. G a r t . Mus.  Berlin-Dahlem  Type: G. Hieronymus & P.G.  7: 265-267.  L o r e n t z s.n.,  der S i e r r a de Tucuman, A r g e n t i n a The by him;  to  These p l a n t s a r e ,  are from.a s e c o n d ' c o l l e c t i o n by Douglas: "Douglas 1830,  e r " , and are not 2.  The  K  1. 575" which r e f e r s  the same Douglas c o l l e c t i o n c i t e d by Hooker (1832). isotypes.  The  top f i v e p l a n t s a r e . i d e n t i f i e d as "Sax-  i f r a g a r a n u n c u l i f o l i a Hook., T o r r . Gr. F l . N. Am.  therefore, probably  Falls  specimen a t K was  i t i s therefore designated  d e s t r o y e d i n 1943,  Bot.  1918.  January  1874,  L a Cienega  in  ( L e c t o t y p e K'. I s o t y p e US'.).  sent by G r i s e b a c h h i m s e l f and the l e c t o t y p e .  i s annotated  The h o l o t y p e at B  was  but an i s o t y p e from the B e r l i n Herbarium e x i s t s a t  the c o l l e c t i o n d e t a i l s conforming  US,  i n a l l r e s p e c t s w i t h those i n the p r o t o -  logue . 3.  S u k s d o r f i a v i o l a c e a A. Gray, P r o c . Am.  Acad. A r t s S c i . 15: 41, 42.  Hemieva v i o l a c e a (Gray) Wheelock, B u l l . T o r r . Bot. Club 71. Type: Suksdorf  23:  1896. 19, A p r i l 1878,  Washington. T e r r i t o r y  (Holotype GH'.) .  1879.  230  Howeli  s.n., June 1879, Wasco-Co., Oregon, near the Columbia  (Paratype GH'. F'.) . Gray (1879) gave the p r e c i s e type l o c a l i t y as " wet r o c k s on the Columbia  R i v e r , i n Washington T e r r i t o r y , near the j u n c t i o n o f t h e  White Salmon R i v e r " .  BOLANDRA Taxonomic  Treatment  B o l a n d r a A.. Gray, P r o c . Am. Acad. A r t s S c i . 7: 341, 342. 1867. Type s p e c i e s : B o l a n d r a c a l i f o r n i c a A. Gray, P r o c . Am. Acad. A r t s . S c i . 7: 341,  342. 1867.  B o l a n d r a c a l i f o r n i c a was d e s c r i b e d (Gray 1867) t o accommodate a new s a x i f r a g e c o l l e c t e d by B o l a n d e r .  Watson (1879a) d e s c r i b e d a second  s p e c i e s , B^ oregana, b u t . d i d not.comment on g e n e r i c r e l a t i o n s h i p s .  Conven-  t i o n a l l y , the genus has been a l l i e d w i t h those which share a x i l e p l a c e n t a t i o n , e s p e c i a l l y w i t h B o y k i n i a , S u k s d o r f i a and S u l l i v a n t i a Rosendahl  1905). The  cussed.  ( E n g l e r 1891;  The p r e s e n t study has confirmed such an a l l i a n c e . s e p a r a t i o n o f B o l a n d r a from B o y k i n i a has a l r e a d y been d i s -  The problem  o f d i s t i n g u i s h i n g B o l a n d r a and S u k s d o r f i a remains.  F i r m c o n c l u s i o n s cannot be reached y e t because the v a r i a t i o n w i t h i n B o l a n d r a has n o t been a s s e s s e d c o m p l e t e l y : more i n f o r m a t i o n i s needed on B^ c a l i f o r n i c a which was i n c l u d e d o n l y o c c a s i o n a l l y i n the p r e s e n t study. genera have b u l b i f e r o u s rhizomes,  and they a l s o share a s i m i l a r  morphology, seed t e s t a ornamentation  and f o l i a c e o u s s t i p u l e s .  pollen A l s o they  both have m u l t i s e r i a t e trichomes w i t h p a p i l l o s e g l a n d u l a r heads. of u n i c e l l u l a r trichomes i n B. oregana ranunculifolia.  Both  The l a c k  i s matched by t h e i r absence i n S.  L e a f shape and o u t l i n e i n B. c a l i f o r n i c a . i s s i m i l a r i n  many r e s p e c t s t o t h a t i n S u k s d o r f i a , although, l e a f v e n a t i o n ( a t l e a s t i n B. oregana) i s more complex.  231  The most.striking,differences--.between  the. genera..are i n f l o r a l  morphology.. Bolandra-has a tubular-campanulate .hypanthium. and s u b u l a t e p e t a l s , whereas, i n S u k s d o r f i a , the hypanthium - i s campanulate ..and  the p e t a l s  o v o i d , s p a t u l a t e o r o r b i c u l a r and - c l a w e d B o l a n d r a a l s o has .narrowly p s o i d seeds r a t h e r than, the o v o i d ones i n S u k s d o r f i a .  Chemically,  oregana p o s s e s s e s 6-oxygenated f l a v o n o i d s , compounds absent from The  two genera a l s o have d i s t i n c t i v e g e n e r a l  Bolandra  Suksdorfia.  The d i f f e r e n c e s  l a r g e l y ovary p o s i t i o n which i s s u p e r i o r i n  and i n f e r i o r . t o . v a r y i n g degrees i n S u k s d o r f i a .  t h i s c h a r a c t e r t o be important genera, o f the S a x i f r a g i n a e .  Bolandra  appearances.  Do t h e d i f f e r e n c e s outweigh t h e s i m i l a r i t i e s ? i n hypanthium shape r e f l e c t  elli-  I do n o t c o n s i d e r  because i t v a r i e s g r e a t l y w i t h i n  other  However, g e n e r a l appearance, supported  by the  d i f f e r e n c e s i n . s e e d and p e t a l shape, and c h e m i s t r y may be used t o d i s t i n g u i s h the.genera.  Ornduff  i f r a g i n a e a r e separated c r i t e r i a ' , although 1  (1969b) observed t h a t many genera o f the Sax-  by "somewhat weak and a r b i t r a r y  morphological  the r e c o g n i t i o n t h a t the s u b t r i b e i s i n . the p r o c e s s o f  a c t i v e e v o l u t i o n can be used as j u s t i f i c a t i o n f o r t h i s Therefore,  ( S a v i l e 1961).  u n t i l the v a r i a t i o n w i t h i n Bolandra  b e t t e r , I marginally p r e f e r to continue  i s described  t o r e c o g n i s e Bolandra. as a s e p a r a t e  genus. Synopsis 1.  and Key t o the S p e c i e s  o f Bolandra  ( a f t e r B a c i g a l u p i 1944) .  Lobes and t e e t h o f l e a v e s rounded, mucronulate; s e p a l s 3-4mm l o n g ; c a r p e l s connate 1/3 t h e i r l e n g t h  . . 1. B_. c a l i f o r n i c a .  l a . Lobes and t e e t h o f l e a v e s t r i a n g u l a r , acute; c a r p e l s connate a t most 1/4 t h e i r l e n g t h 1.  Bolandra 1867.  s e p a l s 6-10mm l o n g ;  . . . . 2. B_. oregana.  c a l i f o r n i c a A. Gray, P r o c . Am. Acad.' A r t s S c i . 7: 341, 342.  232  Type: H.N.  Bolander 4898, 1 s t J u l y 1866, On r o c k s , M a r i p o s a  Yosemite V a l l e y , C a l i f o r n i a  (Holotype GH'.) ;  Toxrey s.n.,  F o o t h i l l s o f the S i e r r a Nevada (Paratype, whereabouts 2.  B o l a n d r a oregana S. Watson, P r o c . Am. Hemieva oregana  unknown).  Acad. A r t s S c i . 14: 292.  1879.  (Wats.) N e l s o n and Macbride, p r i n t e d a n n o t a t i o n  on specimen c o l l e c t e d by Henderson Type: J . Howell s.n., June 1877, Wet type GH'.).  trail,  s.n., 29th J u l y  r o c k s , Milwaukee,  Oregon  1896  (RM) .  (Holo-  Watson (1879a) gave the p r e c i s e l o c a l i t y as ''Willam-  e t t e R i v e r , near Oregon C i t y ,  Oregon".  B o l a n d r a imnahaensis Peck, Rhodora  36: 266.  1934.  B o l a n d r a oregana Wats. v a r . imnahaensis (Peck) Peck, Man. P l a n t s Oregon  371.  High.  1941.  Type: Peck 18495, 4th J u l y 1933, Wet  cliff  i n s m a l l canyon a l o n g Imnaha  R i v e r , 3 m i l e s above Imnaha, Wallowa Co., Oregon  (Isotypes  GH'.  WS'.) . I h e s i t a t e to r e c o g n i s e the two t a x a here as d i s t i n c t Measurements o f a l i m i t e d number of specimens c h a r a c t e r s g i v e n by Rydberg the  two taxa (Table XXX).  show t h a t the d i a g n o s t i c  (1905) and B a c i g a l u p i It i s likely,  s p e c i f i c d i s t i n c t i o n i s warranted.  species.  (1944) i n t e r g r a d e between  t h e r e f o r e , t h a t o n l y an i n f r a -  \  However, a f i n a l d e c i s i o n s h o u l d await  f u r t h e r measurements and o t h e r taxonomic d a t a on B_. c a l i f o r n i c a . A t h i r d s p e c i e s , B o l a n d r a imnahaensis. was (1934) »  He s t a t e d i t was  d e s c r i b e d by Peck  c l o s e l y r e l a t e d t o J . oregana and c l a i m e d t h a t i t  d i f f e r e d , i r i i t s . " m o r e numerous.flowers, a narrower c a l y x tube not u r c e o l a t e , and i n the n e a r l y s e p a r a t e c a r p e l s " .  becoming  I t i s c l e a r from T a b l e XXX  t h a t v a r i a t i o n i n these and o t h e r c h a r a c t e r s i s continuous w i t h B_. oregana, and t h a t even v a r i e t a l s t a t u s of  (Peck 1941,  1961)  B_. oregana from Idaho and e a s t e r n Oregon  i s unwarranted.  Populations  are more v a r i a b l e than from e l s e -  233  TABLE XXX.  Morphological v a r i a t i o n  C a l i f o r n i a have been r e c o g n i z e d  i n Bolandra.  Populations  as B_. c a l i f o r n i c a A. Gray, those  Columbia R i v e r Gorge area as B^ oregana Wats, and those Salmon R i v e r drainages Values  from from t h e  from the Snake and  as both JB^ oregana Wats, and B. imnhaensis Peck.  are g i v e n as the mean + one s t a n d a r d  d e v i a t i o n , with  the range i n  brackets.  California* Basal leaf division/#  52 (50  + 6'  Leaf t o o t h c u r v a t u r e ##  22 (17  + 4  Number o f flowers Sepal mm  5 (3  -  61)  -  29)  Snake - Salmon* 56 (38  + 2 - 8)  + 11  Columbia Gorge*  + 3  68)  59 (56  12 (5  + 6 - 29)  5 (0  ±  17 (5  + 10  12 (8  + 4 - 20)  -  -  37)  -  -  63) 3 8)  5.4 (3.9  + 0.9 - 6.5)  7.0 (5.5  + •1.2 - 9.2)  + 0.76  -  1:35  + 0.14  (1.10  1.54)  1.33 (1.21  + 0.13  3.2)  -  + 0.09 - 0.60)  0.29 (0.29  + 0.50  0.47)  -  3.6)  5.3 (4.3  + 0.6 - 5.8)  -  0.71 (0.63  + 0.06  0.72)  length  3.8 (2.4  Petal/Sepal length r a t i o  1.86 (1.40  Anther length  Filament ratio  0.33 (0.27  + 0.08  Free hypanthium l e n g t h mm  3.3 (2.8  + 0.3  Degree o f c a r p e l 0.64 freedom** (0.56  + .0.07  0.9  - 4.7)  0.38 (0.29 4.4 (3.5 0.68 (0.61  + 0.7 - 5.4) + 0.04 - 0.75)  -  1.57)  -  0.50)  -  0.78)  *  Number o f c o l l e c t i o n s measured: 5 from C a l i f o r n i a , 12 from the Snake - Salmon a r e a and 6 from the Columbia Gorge a r e a .  **  Free c a r p e l l e n g t h / t o t a l c a r p e l l e n g t h . ## L e a f t o o t h c u r v a t u r e i s d e f i n e d as i n F i g . 44, p.211.  Basal leaf d i v i s i o n = b/a x:100% where b i s the shortest i n t e r s i n u s l e n g t h on ^ a leaf.  where, and B_. imnahaensis should be c o n s i d e r e d simply as p a r t o f t h i s variation;  i t i s not a d i s t i n c t  taxon.  234  XV. In t h i s  PHYTOGEOGRAPHY AND EVOLUTION  chapter  t h e d i s t r i b u t i o n s and e v o l u t i o n a r y  s h i p s o f the d i f f e r e n t genera and s p e c i e s a r e d i s c u s s e d .  relation-  Much o f what i s  s a i d has to be s p e c u l a t i v e i n view o f the poor f o s s i l r e c o r d . to  propose hypotheses  The aim i s  a g a i n s t which t h e taxonomic c o n c l u s i o n s  can be  viewed. Methods o f Approach In the determination  of hypothetical evolutionary  s h i p s , two approaches have been used here. proach whereby t h e b r a n c h i n g to for  a common a n c e s t o r .  i s a c l a d i s t i c ap-  p a t t e r n s o f p h y l e t i c l i n e s a r e t r a c e d back  Two g e n e r a l methods o f a n a l y s i s have been proposed  the c o n s t r u c t i o n o f such e v o l u t i o n a r y t r e e s .  taxa a r e grouped t o g e t h e r it  The f i r s t  relation-  on t h e b a s i s o f shared  I n the f i r s t  method,  advanced c h a r a c t e r s t a t e s ;  assumes that e v o l u t i o n proceeds by the s i m p l e s t r o u t e , i . e . t h a t p a r a l -  l e l i s m s and r e v e r s a l s have been minimal (the parsimony p r i n c i p l e ) . The s e c ond method i s based on c h a r a c t e r c o m p a t i b i l i t y a n a l y s i s and assumes t h a t the most probable acters. nature  e v o l u t i o n a r y t r e e i s the one supported  by t h e most  char-  Both methods make assumptions about the p r i m i t i v e v s . advanced of character s t a t e s .  Some o f the concepts b e h i n d  approach have been d i s c u s s e d by E s t a b r o o k (1978). procedures have been reviewed by Funk and Steussy The  the c l a d i s t i c  The v a r i o u s  analytical  (1978).  c l a d i s t i c approach has generated much c o n t r o v e r s y ,  a l l y i n the pages o f " S y s t e m a t i c "Nature" ( H a l s t e a d  1978; G a r d i n e r  especi-  Z o o l o g y " and, most e n t e r t a i n i n g l y , i n e t a l . 1979; F i n k and Wiley  1979; H a l -  s t e a d e t a l . 1979; and Panchen 1979). The axiomatic  second approach has a p h y t o g e o g r a p h i c emphasis.  o f Neo-Darwinist theory  It is  t h a t s p e c i a t i o n i s most l i k e l y t o occur  d u r i n g p e r i o d s o f e n v i r o n m e n t a l i n s t a b i l i t y , e.g. d u r i n g c l i m a t i c and/or  235  orogenic events.  The phytogeographic  approach  uses the p a l a e o n t o l o g i c a l  r e c o r d to e x p l a i n p r e s e n t d i s t r i b u t i o n p a t t e r n s and hence t o suggest  rela-  t i o n s h i p s between s p e c i e s and genera. The  C l a d i s t i c Approach F i f t e e n c h a r a c t e r s were i d e n t i f i e d which showed v a r i a t i o n be-  tween t h e genera. XXXI).  Sequences o f c h a r a c t e r s t a t e s were i d e n t i f i e d  (Table  I t was n o t p o s s i b l e to a s s i g n p r i m i t i v e or advanced s t a t e s t o f i v e  of the c h a r a c t e r sequences because o f i n s u f f i c i e n t e v i d e n c e .  Only f o r  p o l l e n a p e r t u r e type was f o s s i l e v i d e n c e a v a i l a b l e t o support the e v o l u t i o n ary i n t e r p r e t a t i o n o f the sequence.  F o r the remaining t e n c h a r a c t e r s two  k i n d s o f e v i d e n c e were used t o a s s i g n p h y l o g e n e t i c grades states:  1) developmental  t o the c h a r a c t e r  e v i d e n c e , whereby t h e e a r l y stages o f a c h a r a c -  t e r ' s development a r e i n t e r p r e t e d t o r e p r e s e n t a p r i m i t i v e c o n d i t i o n , and the l a t e r stages an advanced c o n d i t i o n .  The e v i d e n c e f o r t h i s  assumption  has been reviewed by Stebbins (1974); 2) the common ground p l a n , by which the p r i m i t i v e c o n d i t i o n i s p o s s e s s e d by most taxa (Estabrook 1977). must be sure t o conduct  a s u f f i c i e n t l y wide taxonomic s u r v e y , o u t s i d e the  group at hand, f o r the approach The  One  t o be v a l i d  ( S t e b b i n s 1974).  a n a l y s i s used here i s the v e r y s i m p l e , manual method  c r i b e d by Johnson and B r i g g s (1975).  des-  Although i t c o u l d be c l a s s e d as one  of the "minimum e v o l u t i o n " methods, i t has t h e advantage o f n o t n e c e s s a r i l y a c c e p t i n g the most parsimonious  t r e e i f t h e r e a r e grounds f o r b e l i e v i n g  t h a t an a l t e r n a t i v e c o u l d be b e t t e r j u s t i f i e d . flexibility. XXXI.  I t thus has a c e r t a i n  The r e s u l t s o f t h e c l a d i s t i c approach  a r e shown i n T a b l e  They w i l l now be d i s c u s s e d w i t h r e f e r e n c e to p h y t o g e o g r a p h i c a l and  p a l a e o n t o l o g i c a l d a t a to d i s c o v e r whether any i n c o n s i s t e n c i e s Summary o f Some Important The  arise.  T e r t i a r y Events i n t h e N o r t h e r n Hemisphere  Rocky Mountains u p l i f t e d i n the Eocene - e a r l y  Oligocene  236  TABLE XXXI.  P u t a t i v e l y e v o l u t i o n a r y i n t e r p r e t a t i o n o f some c h a r a c t e r  state  sequences; t h e i r d i s t r i b u t i o n among the genera are shown, t o g e t h e r w i t h a hypothetical evolutionary tree constructed simonious s h a r i n g o f advanced c h a r a c t e r  Character  Character  on the b a s i s  o f t h e most  states.  S t a t e Sequence*  Comment o r Evidence**  1.  Rhizome  Bulbiferous  2.  Stipules  F o l i a c e o u s (a)  B r i s t l e s o r expanded p e t i o l e bases  3.  Trichomes  Multiseriate  Uniseriate  4.  Ovary  I n f e r i o r (a)  Superior  D, CGP 4 CGP  5.  Free hypanthium  P r e s e n t (a)  Absent  CGP  4  6.  Stamens  4  7.  Seeds  Tuberculate  8.  P o l l e n apertures  C o l p o r a t e (a)  Colporoidate or c o l p a t e  CGP 2 D .6  9.  Flavonoids: myricetin  Absent (a)  Present  D, CGP  10.  Chromosomes  P o l y p l o i d (a)  Diploid  D, CGP  11.  Inflorescence (cyme)  Complex, coid  Simple  Reversible  (a)  (a)  Not b u l b i f e r o u s  10  5 (a) (a)  heli-  par-  Smooth  D, CGP 4 CGP  4  J  L  12.  Nectary  Present  Absent  No  13.  Petals  Large  Small  Reversible  14.  P o l l e n tectum  Reticulate  Perforate  15.  Flavonoids: e x t r a oxygenat i o n , methylation  Present  Absent  data  L c  Reversible" Reversible  (a) = advanced s t a t e , the a l t e r n a t i v e i s p r i m i t i v e . D = developmental e v i d e n c e ; CGP = common ground p l a n ; F = f o s s i l evidence. I f the same p u t a t i v e t r e n d occurs i n the angiosperms, o r e s p e c i a l l y i n the Saxifragaceae, a reference i s provided: 1. G o r n a l l and Bohm (1978); 2. Hara (1957); 3. Stebbins (1971); 4. Stebbins (1974); 5. Walker and Doyle (1975); 6. Wolfe, Doyle and Page (1975).  237  TABLE XXXI, cont'd. Genus  Advanced C h a r a c t e r S t a t e s - P r e s e n t Throughout  Saxifraga Peltoboykinia Boykinia Sullivantia Bolandra Suksdorfia  5, 3, 3, 1, 1,  7, 4, 5, 2, 2,  9, 5 6, 3, 3,  cd •H cd •rl  00 cd 5-i  MH •H  crj  >>  o o 4-1  cd •rl  cd u  •rl  ti cd  ti  rH CU  o  rH  o  pq  10 8, 9 5, 6, 7, 8 4, 5, 6, 7, 8, 9  cd •rl m rl  o  CO  3  cd •rl 4J  3  cd  >  •rl rH rH  CO  3  ANCESTOR H y p o t h e t i c a l E v o l u t i o n a r y Tree  ( c a . 38 m.y. BP) ( E i s b a c h e r 1977).  A t about the same time t h e r e was a  s i g n i f i c a n t drop i n t h e mean annual temperature 1978;  ( M a c G i n i t i e 1974; Wolfe  Rouse & Mathews 1980) which r e s u l t e d i n t h e e x t i n c t i o n o f many  elements o f the f o r m e r l y widespread p a r a t r o p i c a l / s u b t r o p i c a l  rainforest.  A mixed mesophytic f o r e s t had r e p l a c e d i t by the O l i g o c e n e (Wolfe and Leopold 1967).  T h i s temperate b r o a d - l e a v e d f o r e s t was continuous between  e a s t e r n A s i a and e a s t e r n North America 1980)  u n t i l the l a t e Miocene  (Wolfe 1969; Rouse and Mathews  (10 - 14 m.y. B P ) .  I n the l a t e  Miocene  there was a sharp f a l l i n summer temperatures n e a r l y everywhere  (Rouse  238  and Mathews 1980), a b o r e a l f o r e s t formed I n A l a s k a 1967;  Wolfe 1969)  1968) .  and  (Wolfe  and  the B e r i n g S t r a i t s opened f o r the f i r s t  Leopold time  (Hulten  Thus the mixed mesophytic f o r e s t became d i s j u n c t between e a s t e r n  A s i a and N o r t h America. I n P l i o c e n e times and  BP)  t h e r e was  a general cooling  a marked summer-dry c l i m a t e e v o l v e d i n the P a c i f i c Northwest  1969) .  The  generated  summer dryness was  probably  by the u p l i f t of the Coast  (Coney 1972).  and  by r a i n shadow e f f e c t s  S i e r r a Nevada - Cascade Ranges  a tundra v e g e t a t i o n formed (Johnson and Packer  p r o g r e s s i v e c o o l i n g culminated  10,000 y.  accentuated  (Wolfe  The B o r e a l f o r e s t degenerated i n the n o r t h i n response to  the c o o l i n g , and The  (2 - 10 m.y.  1967).  i n the P l e i s t o c e n e g l a c i a t i o n s  (2 m.y.  -  BP).  E v o l u t i o n of B o y k i n i a and  Allies  S a v i l e (1975) has  suggested  a s c e n a r i o f o r the e v o l u t i o n and  biogeography of the S a x i f r a g i n a e u s i n g i n f o r m a t i o n from the r u s t p a r a s i t e s . He proposed an o r i g i n i n e a s t e r n A s i a d u r i n g the O l i g o c e n e then e a r l y m i g r a t i o n s l a t t e r m i g r a t i o n was  i n t o the Himalyas and western N o r t h America. suggested  and The  t o have g i v e n r i s e to the v a r i o u s North  American genera by e x t e n s i v e r a d i a t i o n . begun l i f e  or l a t e r ,  S a x i f r a g a was  considered  to have  i n N o r t h America as a " p r o t o - S a x i f r a g a " which r e - c r o s s e d  B e r i n g S t r a i t s back i n t o Japan b e f o r e becoming d i f f e r e n t i a t e d .  the  Radiations  o c c u r r e d from h e r e p a r t l y i n t o the Himalayas (again) and on i n t o Europe, and p a r t l y back i n t o N o r t h America a c r o s s B e r i n g i a . It lity ing  i s beyond the scope of t h i s t h e s i s to d i s c u s s the  plausibi-  of t h i s s c e n a r i o , but i t s c h i e f problem would seem to be i n r e c o n c i l the t i m i n g of a l l t h i s s a x i f r a g a c e o u s  g e n i c and  climatic  events.  t r a f f i c w i t h the t i m i n g of  oro-  23?  B o y k i n i a , t o g e t h e r w i t h genera l i k e T i a r e l l a , M i t e l l a and s e c t i o n s of S a x i f r a g a show d i s j u n c t d i s t r i b u t i o n s between the the Western C o r d i l l e r a , A l a s k a and Japan. (Wood 1970,  1972)  probably  T h i s and  some  Appalachians,  similar disjunctions  r e s u l t e d from the d i s r u p t i o n of the widespread  mixed mesophytic f o r e s t by  c l i m a t i c change, v u l c a n i s m  and  brogenic  events  (Graham 1972).  genera of the S a x i f r a g i n a e had  probably  evolved  at l e a s t by  Thus these  l a t e Miocene times when the f o r e s t became d i s j u n c t between  e a s t e r n A s i a and N o r t h America. u n t i l e a r l y Miocene times and 10 and 24 m i l l i o n y e a r s  Herbs do not  appear i n the f o s s i l  so B o y k i n i a p r o b a b l y  had  record  i t s o r i g i n between  ago.  H a r a (1959) suggested t h a t B o y k i n i a and P e l t o b o y k i n i a share common Miocene a n c e s t o r . f o l l o w i n g the Oligocene  By Miocene times Japan had become a p e n i n s u l a f o r m a t i o n of the Japan Sea  middle Miocene t h e r e was which c o i n c i d e d w i t h  a phase of o r o g e n i c  (Maekawa 1974).  In  the  a c i t i v i t y i n eastern Asia  the i n v a s i o n by the sea of the Japan p e n i n s u l a ,  t i n g the l a n d mass i n t o a s e r i e s of s m a l l i s l a n d s (Maekawa 1974). probably  a  split-  It  was  at t h i s time t h a t P e l t o b o y k i n i a and B o y k i n i a became d i f f e r e n t i a t e d  from t h e i r common a n c e s t o r , p r o b a b l y  a taxori which had  diverged  little  from a p r i m i t i v e S a x i f r a g a ( T a b l e XXXI). Subsequent e v o l u t i o n was gene p o o l i n N o r t h America. ( F i g . 47)  The  based on r a d i a t i o n s from the  present  d i s t r i b u t i o n of  as a s e r i e s of h i g h l y d i s j u n c t p o p u l a t i o n s  suggests a f o r m e r l y  continuous  the new  across  the U.S.A.  An e a r l y d i f f e r e n t i a t i o n of  S u k s d o r f i a and B o l a n d r a  i n the P l i o c e n e i n response to the new Northwest and  Sullivantia  d i s t r i b u t i o n , q u i t e p o s s i b l y as a component  of the Miocene mixed mesophytic f o r e s t . v a n t i a i s thus e n v i s a g e d .  Sulli-  most p r o b a b l y  evolved  summer-dry c l i m a t e i n the  Pacific  h a b i t a t s c r e a t e d by the u p l i f t of the Coast  S i e r r a - Cascade Ranges.  Boykinia  The b u l b i f e r o u s rhizomes may  be  and  seen as an  adapt-  240  a t i o n t o the new  environments.  onoid p r o f i l e and  I n view of i t s woodland h a b i t a t , i t s f l a v -  i t s o v e r a l l appearance, B o l a n d r a  l i k e f e a t u r e s than does S u k s d o r f i a .  r e t a i n s more B o y k i n i a -  However, the l a t t e r has  u n i c e l l u l a r trichomes of B o y k i n i a and  the  a s i m i l a r f l o r a l morphology.  t i o n s h i p s between the t h r e e genera are somewhat r e t i c u l a t e and that both Bolandra  and  Suksdorfia evolved  I suggest  (Table XXXI).  Generally, therefore, phytogeographical  considerations  complement the h y p o t h e t i c a l e v o l u t i o n a r y t r e e produced by the  approach.  Rela-  at about the same time i n the  P l i o c e n e from a common B o y k i n i a - l i k e a n c e s t o r  and  typical  support cladistic  I n the f o l l o w i n g d i s c u s s i o n , each genus w i l l be examined i n  more d e t a i l . Boykinia D i s t r i b u t i o n s of B o y k i n i a s p e c i e s are shown i n F i g s . 45 -  47.  S i n c e B o y k i n i a s e c t i o n B o y k i n i a i s d i s j u n c t between Japan and N o r t h Ameri c a , i t presumably had  e v o l v e d by l a t e Miocene times, when t h i s d i s j u n c t i o n  o c c u r r e d i n the mixed mesophytic f o r e s t .  I suggest t h a t the a n c e s t o r  s e c t i o n B o y k i n i a e x i s t s today, i n a v e r y much Telesonix.  modified  of  form, as s e c t i o n  T h i s s e c t i o n r e t a i n s the most p r i m i t i v e f e a t u r e s ( c o l p o r o i d a t e  p o l l e n , t e n stamens, smooth seeds) i n the The  two  genus.  s p e c i e s of s e c t i o n T e l e s o n i x , B_. j a r o e s i i and B.  heu-  c h e r i f o r m i s . have become d i f f e r e n t i a t e d e d a p h i c a l l y as w e l l as morphologically.  At p r e s e n t , B_. h e u c h e r i f o r m i s  grows on  calcareous  the Rocky Mountains from A l b e r t a south to Wyoming and Black H i l l s ada;  of South Dakota and  A formerly  continuous  substrates i n  d i s j u n c t l y i n the  the C h a r l e s t o n Mountains of southern  range i s t h e r e f o r e i n d i c a t e d .  Nev-  Although mostly  e l i m i n a t e d from the Canadian Rockies by the P l e i s t o c e n e g l a c i a t i o n s t h e r e i s good evidence  t h a t at l e a s t one  p o p u l a t i o n managed to s u r v i v e , t h e r e ,  i n the Mountain Park a r e a , near J a s p e r , A l b e r t a (Packer  and V i t t  1974).  241  242  FIGURE 46.  The  distributions  of B o y k i n i a major, B_. i n t e r m e d i a and  Bolandra.  243  FIGURE 47 . The tellimoides  distributions  of Boykinia l y c o c t o n i f o l i a , P e l t o b o y k i n i a  (from H a r a (1958)), and S u l l i v a n t i a  (from Wood  (1971)).  244  I t i s l i k e l y that present  populations  i n t h i s p a r t of A l b e r t a d e r i v e from  t h i s r e f u g i a l gene p o o l .  A s i m i l a r refugium  may  have e x i s t e d f o r B_.  c h e r i f o r m i s i n the Waterton Lakes a r e a of southern  A l b e r t a (Packer  A f t e r the P l e i s t o c e n e ended, a warm H y p s i t h e r m a l  1971).  p e r i o d ensued  d u r i n g which B_. h e u c h e r i f ormis r e t r e a t e d northwards, l e a v i n g behind disjunct populations. mountains of the Utah  R e l i c t populations  may  the  s t i l l be d i s c o v e r e d I n  rado F r o n t Range, v i z . P i k e s Peak and  the mountains of "the  Mexico.  disjunctly i n  As w i t h B_j, h e u c h e r i f ormis,  d i s j u n c t i o n suggests a f o r m e r l y continuous  the warm H y p s i t h e r m a l  i n a t e d most of these southernmost  the  d i s t r i b u t i o n southwards, pos-  s i b l y as a r e s u l t of m i g r a t i o n i n response to the P l e i s t o c e n e As b e f o r e ,  Colo-  neighbouring  and Rocky Mountain N a t i o n a l Park, and  the Santa Fe mountains of New  (Whitehead 1972).  the  Plateaux.  B o y k i n i a j a m e s i i grows on the h i g h g r a n i t e peaks of the  P l a t t e R i v e r drainage  heu-  climates  p e r i o d probably  elim-  populations.  B o y k i n i a r i c h a r d s o n i i of s e c t i o n R e n i f o l i u m i s p r e s e n t l y endemic to A l a s k a and  the Yukon, where i t i s p a r t of the  alpine-tundra  flora.  I t i s not p o s s i b l e to be v e r y s p e c i f i c about i t s o r i g i n s  one may  speculate  t h a t i t e v o l v e d e i t h e r as a component of the b o r e a l  f o r e s t i n l a t e Miocene times (Tolmachev 1959) tundra f l o r a f o l l o w i n g the degeneration.?of cene.  two  or as a component of  the b o r e a l f o r e s t i n the  the Plio-  B o y k i n i a r i c h a r d s o n i i almost c e r t a i n l y s u r v i v e d the P l e i s t o c e n e  g l a c i a t i o n s i s o l a t e d i n the u n g l a c i a t e d areas (Huit en  although  1968).  I t may  cytodemes, one  of A l a s k a and  the Yukon  have been at t h i s time t h a t i t d i f f e r e n t i a t e d i n t o  i n the A l a s k a Range (2ri = 6x = 36)  Range (2n = 14x = 84). u l a t i o n s from the Yukon.  and  Chromosome numbers have y e t to be The h i g h p o l y p l o i d y may  one  i n the Brooks  counted i n pop-  be i n t e r p r e t e d as  an  a d a p t a t i o n toward the r e c o l o n i z a t i o n of h a b i t a t s a f t e r the i c e r e t r e a t e d ;  245  i n d e e d , the 14x cytodeme has invaded f a r more g l a c i a t e d t e r r i t o r y than the 6x cytodeme ( F i g . 45).  I t i s not p o s s i b l e to say whether one  gave r i s e to the o t h e r , and i t i s s i m p l e r to suggest indepently.  has  cytodeme  t h a t they e v o l v e d  At p r e s e n t , B_. r i c h a r d s o n i i has a U-shaped d i s t r i b u t i o n i n  A l a s k a and the Yukon T e r r i t o r y , i n h a b i t i n g the A l a s k a Range, the R i v e r drainage and the Brooks Range.  Porcupine  Rydberg (1905) and E n g l e r (1928) r e -  p o r t e d B_. r i c h a r d s o n i i from e a s t e r n S i b e r i a ; H u l t e n (1945, 1968)  d i d not  r e c o r d i t from A s i a , although he admitted t h a t the o c c u r r e n c e o f such popu l a t i o n s seemed " v e r y p r o b a b l e " .  However the s p e c i e s i s not l i s t e d i n the  F l o r a S.S.S.R. ( L o z i n a - L o z i n s k a y a 1939)  nor i n the more r e c e n t S i b e r i a n  F l o r a (Tolmachev 1966). D e a l i n g w i t h s e c t i o n B o y k i n i a i n more d e t a i l , B. l y c o c t o n i f o l i a i n Japan has p r o b a b l y been i s o l a t e d genus s i n c e l a t e Miocene times.  Its closest  as noted  earlier,  from the r e s t o f the  r e l a t i v e s on m o r p h o l o g i c a l  grounds ( e s p e c i a l l y i n l e a f morphology and s t o l o n p r o d u c t i o n ) are B. media i n the Olympic The  inter-  P e n i s u l a and B_. a c o n i t i f o l i a i n the A p p a l a c h i a n s .  t h r e e s p e c i e s p r o b a b l y c o n s t i t u t e the remnants of the f o r m e r l y wide-  ranging a n c e s t r a l s t o c k o f s e c t i o n B o y k i n i a , and vicariads. cene times  There have been few  can be  c o n s i d e r e d as  changes i n the Japanese f l o r a s i n c e P l i o -  (Wolfe and L e o p o l d 1967); the P l i o c e n e c o o l i n g and P l e i s t o c e n e  g l a c i a t i o n s d i d not a f f e c t Japan as much as they d i d N o r t h America Europe.  or  Even,during maximum g l a c i a t i o n , g l a c i e r s were r e s t r i c t e d to a  few h i g h peaks i n c e n t r a l Honshu and Hokkaido (Hara 1959).  Boykinia lycoc-  t o n i f o l i a became adapted  i t s present  t o v o l c a n i c s u b s t r a t e s and adopted  d i s t r i b u t i o n i n the mountains o f c e n t r a l and n o r t h e r n Honshu and Hokkaido (Fig.  47). B o y k i n i a a c o n i t i f o l i a ranges  a t p r e s e n t through the  chians from the V i r g i n i a s south to G e o r g i a and Alabama.  Appala-  I t presumably  246  became i s o l a t e d from the r e s t of the genus by the d r y i n g of the P l a i n s and by P l e i s t o c e n e e x t i n c t i o n s but has e r n American f o r e s t s . times but was  I t probably  Great  s u r v i v e d i n the mesic e a s t -  ranged f u r t h e r n o r t h i n p r e - P l e i s t o c e n e  e l i m i n a t e d as the g l a c i e r s  advanced.  I n western N o r t h America, the three d i p l o i d B. i n t e r m e d i a . B. o c c i d e n t a l i s and B^  (2n = 14)  r o t u n d i f o l i a p r o b a b l y began t h e i r  d i f f e r e n t i a t i o n i n response to the P l i o c e n e mountain b u i l d i n g . i n t e r m e d i a has two  the most p r i m i t i v e morphology and  s p e c i e s have p r o b a b l y  evolved  species,  chemistry  from i t s gene p o o l .  to the e v o l u t i o n of B^ o c c i d e n t a l i s and B.  and  Boykinia the  other  A f u r t h e r stimulus  r o t u n d i f o l i a was  probably  pro-  v i d e d by the southward v e g e t a t i o n a l s h i f t i n the P l e i s t o c e n e , when many new  n i c h e s were opened as s p e c i e s from v a r i o u s o r i g i n s came i n t o  contact  (whitehead 1972). B o y k i n i a i n t e r m e d i a i s p r e s e n t l y endemic to the western s i d e of the Olympic P e n i n s u l a i n Washington and Oregon.  I t p r o b a b l y possessed  the Coast  Range of n o r t h e r n  a w i d e r , more n o r t h e r l y range p r e c e d i n g  P l e i s t o c e n e g l a c i a t i o n s , although why  i t d i d not s u r v i v e south of the i c e  sheets i n g r e a t e r numbers, as d i d r e l a t e d s p e c i e s , i s a problem, i t s present  requirement The  of a h i g h r a i n f a l l may  r e p o r t o f the o c c u r r e n c e  Idaho ( B a c i g a l u p i 1952)  be  although  relevant i n this  probably  prompted by the mis-  q u o t a t i o n of the type l o c a l i t y by Rydberg (1905) as b e i n g i n Idaho. t o B.  respect.  of B_j i n t e r m e d i a i n n o r t h e r n  i s erroneous and was  c o l l e c t i o n s from Idaho belong  All  major.  B o y k i n i a o c c i d e n t a l i s i s a wide-ranging  but e s s e n t i a l l y c o a s t a l  element, p r e s e n t l y o c c u r r i n g on the western s l o p e s of the Coast from n o r t h e r n Vancouver I s l a n d to southern  California.  Ranges  Some p o p u l a t i o n s  have managed to p e r s i s t i n the S i e r r a - Cascade a x i s c h i e f l y i n c e n t r a l Oregon, the Klamath Ranges and  the  the Yuba and American R i v e r  drainages.  247  The  d i s j u n c t i o n s between the Coast  r e c e n t o r i g i n , and  result  and S i e r r a - Cascade Ranges may  from the H y p s i t h e r m a l  c a u s i n g i n c r e a s e d d r y i n g between the two  be  warming (ca.. 6000 y.  mountain ranges.  of BP)  R a i n shadow  e f f e c t s may  have c o n t r i b u t e d a l s o to the e l i m i n a t i o n of mesic elements.  Populations  of B_^ o c c i d e n t a l i s i n the N o r t h Washington Cascades, the  e r n B r i t i s h Columbia Coast  Range and Vancouver I s l a n d a l s o may  south-  be of r e -  cent o r i g i n , i n v a d i n g a f t e r the P l e i s t o c e n e i c e r e t r e a t e d , although  coastal  r e f u g i a on Vancouver I s l a n d d u r i n g the P l e i s t o c e n e are a p o s s i b i l i t y . B o y k i n i a o c c i d e n t a l i s has not been found although  they form a v e r y l i k e l y refugium  on the Queen C h a r l o t t e I s l a n d s , ( C a l d e r and T a y l o r  1968).  B o y k i n i a r o t u n d i f o l i a i s a narrow endemic, and w i t h ;one t i o n i s p r e s e n t l y r e s t r i c t e d to the s o u t h e r n Mountains i n the T r a n s v e r s e  excep-  s l o p e s of the San G a b r i e l  Ranges of southern  California.  The  exception  i s a s i n g l e p l a n t d i s c o v e r e d i n San Diego County, about 130km f u r t h e r south.  T h i s may  might suggest  the r e s u l t o f a chance l o n g - d i s t a n c e d i s p e r s a l or i t  a f o r m e r l y wider d i s t r i b u t i o n southwards i n the  Ranges of B a j a The Boykinia.but  be  Peninsula  California. species i s morphologically very d i s t i n c t i v e i n s e c t i o n  i t s c l o s e s t r e l a t i v e i s B. o c c i d e n t a l i s , ' a s p e c i e s w i t h s i m i -  l a r l y o r b i c u l a r l e a v e s and w i t h which i t shows g r e a t e s t c r o s s - c o m p a t i b i l ity.  I t may  t h e r e f o r e be h y p o t h e s i z e d  d e n t a l i s share present  two  t h a t B_. r o t u n d i f o l i a and B_.  a common a n c e s t o r which became d i f f e r e n t i a t e d i n t o  s p e c i e s d u r i n g the P l i o c e n e mountain b u i l d i n g and  P l e i s t o c e n e southward m i g r a t i o n s . a r e l i c t i n the T r a n s v e r s e  B o y k i n i a r o t u n d i f o l i a now  occithe  during  the  s u r v i v e s as  Ranges.  B o y k i n i a major grows on the western s l o p e s of the S i e r r a Cascade a x i s from c e n t r a l Oregon to c e n t r a l C a l i f o r n i a , and i n the B i t t e r r o o t Mountains of Idaho and Montana.  disjunctively  I n p l a c e s i t has  man-  248  aged to p e r s i s t i n t h e Coast California.  Ranges, e.g. the Salmon Mountains o f n o r t h e r n  L i k e many o t h e r s p e c i e s , B o y k i n i a major p r o b a b l y s u f f e r e d  e l i m i n a t i o n from the n o r t h e r l y p a r t s o f i t s range d u r i n g t h e P l e i s t o c e n e , the p r e s e n t day d i s j u n c t i o n b e i n g one r e s u l t o f t h i s . B o y k i n i a major i s t e t r a p l o i d  (2n = 26) and one o f the genome  c o n t r i b u t o r s was p r o b a b l y a p l a n t l i k e B_. i n t e r m e d i a (Ch. X ) . The second c o n t r i b u t o r may have been t h e common a n c e s t o r o f I K o c c i d e n t a l i s and B. r o t u n d i f o l i a , i . e . a p l a n t s i m i l a r t o B^ i n t e r m e d i a (Ch. X ) . major would thus q u a l i f y as a segmental a l l o t e t r a p l o i d The  Boykinia  ( S t e b b i n s 1950).  time o f o r i g i n o f B_. major presumably must have been a f t e r the d i f f e r -  e n t i a t i o n o f the a n c e s t o r o f B_. i n t e r m e d i a i n t o two t a x a , i . e . p r o b a b l y l a t e Pliocene or l a t e r . Reports handle  (Cooley  of B_. major from the Juneau a r e a o f the A l a s k a pan-  1892) a r e erroneous.  A l l o f Cooley's  specimens b e l o n g t o  Heuchera g l a b r a W i l l d . Peltoboykinia P e l t o b o y k i n i a i s endemic t o Japan ( F i g . 4 7 ) . I t s p o s s i b l e Miocene o r i g i n has a l r e a d y been d i s c u s s e d .  Hara (1959) b e l i e v e s t h a t i t  was d u r i n g the c l i m a t i c f l u c t u a t i o n s and t o p o g r a p h i c tocene  changes o f t h e P l e i s -  t h a t many Japanese p l a n t groups d i f f e r e n t i a t e d i n t o new t a x a .  c o u l d have been a t t h i s time i t s two s u b s p e c i e s .  that P e l t o b o y k i n i a t e l l i m o i d e s s p l i t  Subspecies  It  into  watanabei d i f f e r e n t i a t e d i n the Shikoku  and Kyushu mountains and became i s o l a t e d from ssp. t e l l i m o i d e s by t h e f o r m a t i o n o f the Seto  (Inland) Sea.  Ssp. t e l l i m o i d e s p r e s e n t l y grows i n  c e n t r a l and n o r t h e r n Honshu. Bolandra The  d i s t r i b u t i o n of Bolandra  i s shown i n F i g . 46.  I t i s sim-  i l a r i n many r e s p e c t s t o t h a t o f B o y k i n i a major and p r o b a b l y has a s i m i l a r  249  explanation.  One  may  envisage  a f o r m e r l y continuous  distribution  along  the S i e r r a - Cascade a x i s c r o s s i n g i n t o the mountains of Idaho i n the more n o r t h e r l y p a r t of i t s range.  P l e i s t o c e n e c l i m a t i c and  l i k e l y e l i m i n a t e d the northernmost between the r e s t .  The  genus now  p o p u l a t i o n s and  occupies  R i v e r drainages  events  caused d i s j u n c t i o n s  three areas.  i s d i s j u n c t between the Columbia R i v e r Gorge and  glacial  Bolandra  the Snake and  oregana  Salmon  of the Wallowa and B i t t e r r o o t Mountains i n e a s t e r n Oregon,  and n o r t h e r n Idaho; B o l a n d r a  c a l i f o r n i c a i s r e s t r i c t e d to h i g h e l e v a t i o n s  on the western s l o p e s of the c e n t r a l S i e r r a Nevada i n C a l i f o r n i a . c a l i f o r n i c a i s not v e r y d i s t i n c t  from B_. oregana and p r o b a b l y  Bolandra  owes i t s  taxonomic r e c o g n i t i o n to the P l e i s t o c e n e e x t i n c t i o n of i n t e r m e d i a t e popu l a t i o n s i n the S i e r r a - Cascade a x i s . Suksdorfia The  d i s t r i b u t i o n s of S u k s d o r f i a s p e c i e s are shown i n F i g . 48.  The  genus shows a remarkable d i s j u n c t i o n between the C o r d i l l e r a s of N o r t h  and  South America.  The  two N o r t h American s p e c i e s are d i s c u s s e d  first.  The major problem i n t r a c i n g the h i s t o r y o f S u k s d o r f i a r a n u n c u l i f o l i a S. v i o l a c e a i s i n e x p l a i n i n g how  they became d i f f e r e n t i a t e d .  At  and  present  they show a c o n s i d e r a b l e o v e r l a p i n t h e i r d i s t r i b u t i o n s , and u n l e s s s p e c i a t i o n was  sympatric  (an uncommon phenomenon ( S t e b b i n s 1950:  assume t h a t at some time they have o c c u p i e d  238))  largely allopatric  one must  ranges.  Some i n d i c a t i o n of t h e i r former a l l o p a t r i c d i s t r i b u t i o n s i s g i v e n by the f a c t t h a t S_. r a n u n c u l i f o l i a r e t a i n s p o p u l a t i o n s i n the  coast-  a l b e l t , from the Skeena R i v e r i n n o r t h e r n B r i t i s h Columbia to the Klamath Ranges i n n o r t h e r n  California.  These occur west of the area i n h a b i t e d ' -  by _S. v i o l a c e a which predominates i n the Monashee, S e l k i r k and Ranges of B r i t i s h Columbia and Idaho  and Montana.  The  t h e i r extensions  present  Purcell  i n a d j a c e n t Washington,  l a r g e degree of o v e r l a p of the two  spe-  250  251  cies probably  r e s u l t s from southward m i g r a t i o n s  during  from subsequent northward i n v a s i o n s of f o r m e r l y Pleistocene  to the n o r t h of the i c e sheets disjunct populations  R i v e r v a l l e y , and  one  d e r i v e d from a now  glaciated t e r r i t o r y i n post-  1968).  have s u r v i v e d the g l a c i a t i o n s  as w e l l as to the south.  I t presently  on the B r i t i s h Columbian c o a s t :  i n the B e l l a Coola R i v e r v a l l e y .  one  i n the  These may  has Skeena  have been  e x t i n c t p o p u l a t i o n on the Queen C h a r l o t t e I s l a n d s ,  p l a c e where s e v e r a l s p e c i e s p r o b a b l y Taylor  and  times. S u k s d o r f i a r a n u n c u l i f o l i a may  two  the P l e i s t o c e n e  s u r v i v e d the P l e i s t o c e n e  a  (Calder  and  I t i s i n t e r e s t i n g t h a t a somewhat s i m i l a r d i s t r i b u t i o n  p a t t e r n i s shown by Heuchera c h l o r a n t h a P i p e r  ( C a l d e r and  S a v i l e 1959).  S u k s d o r f i a v i o l a c e a can have s u r v i v e d the i c e ages o n l y of the g l a c i a l boundary i n Idaho and Washington.  south  Although s t a t i o n s i n the  n o r t h Washington Cascades were at the l i m i t s of the i c e sheets  and  there-  f o r e presumably v e r y i n h o s p i t a b l e , t h e r e i s some e v i d e n c e t h a t r e f u g i a sepa r a t e d by v a l l e y g l a c i e r s may  have e x i s t e d h e r e , e.g.  i n the Wenatchee  Mountains ( C a l d e r and S a v i l e 1959). The  highly disjunct Suksdorfia  alchemilloides i s presently  found on the h i g h e a s t e r n s l o p e s of the Andean C o r d i l l e r a i n Argentina  and  southern B o l i v i a .  According  i c a l d i s j u n c t i o n s between N o r t h and distance d i s p e r s a l .  Although he was  northern  to Raven (1963) many amphitrop-  South America can be  a t t r i b u t e d to  concerned p r i m a r i l y w i t h  desert plant species, h i s generalizations nevertheless  coastal  I n summary h i s arguments were t h a t 1) N o r t h and  American p o p u l a t i o n s  are c l o s e l y r e l a t e d .  are almost w i t h o u t e x c e p t i o n  u n i c e l l u l a r trichomes).  self-compatible  South  c l o s e s t r e l a t i v e of _S_j_ a l -  c h e m i l l o i d e s i s _S_. v i o l a c e a (both have s i m i l a r l e a f shapes and cnece s t r u c t u r e s , p i n k p e t a l s and  and  seem to be a p p l i c -  able to S u k s d o r f i a .  The  long-  and  inf lores-  2) The  o f t e n autogamous.  plants Judging  252  by  f l o r a l morphology, the  S.  a l c h e m i l l o i d e s suggest t h a t i t i s s e l f - c o m p a t i b l e  ous.  c l o s e j u x t a p o s i t i o n o f anthers and  S u k s d o r f i a v i o l a c e a i s both (Ch. X I ) .  distance  stigmata  -  d i s p e r s a l would thus be  and  p o s s i b l y autogam-  Establishment a f t e r long-  facilitated  (Baker 1955).  3) The  c o n s t i t u t e an unbalanced assemblage e n t i r e l y u n r e p r e s e n t a t i v e e x t r a t r o p i c a l areas. t r i b e Saxif raginae  are w i t h the N o r t h e r n Hemisphere.  almost e x c l u s i v e l y i n open communities, e.g.  sea  snow-melt, summer-dry environment; and i s above the t r e e l i n e and  i n the open.  cases among t e r r e s t r i a l v e r t e b r a t e s f l o r a s of the two  middle Cretaceous and  and  sub-  p l a n t s grow  be  wet  Suksdorfia  vio-  S_. alchemi 1 l o i d e s subject  to a s p r i n g  growing above 3000m means t h a t i t 5) There are no very  few  corresponding  among .the i n s e c t s .  areas have been d i s t i n c t  are s t i l l very  two  coast or s e a s o n a l l y  Since  o c c u p i e s h i g h e a s t e r n mountain s l o p e s , i t too may  the  of the  4) The  or woodland a s s o c i a t i o n s .  l a c e a i s found i n open, spring-wet h a b i t a t s .  a l l y , 6)  species  I n t h i s case"the p r i m a r y a s s o c i a t i o n s of the  h a b i t a t s , r a t h e r than i n scrub  in  Fin-  at l e a s t s i n c e  d i f f e r e n t eat p r e s e n t .  The  only  the  explan-  a t i o n t h a t accounts f o r a l l these f a c t s seems to be  t h a t most of the  reached t h e i r d i s j u n c t areas r e l a t i v e l y r e c e n t l y by  long d i s t a n c e d i s p e r s a l .  For temperate s p e c i e s tocene would be  l i k e S.  a l c h e m i l l o i d e s . the l a t e P l i o c e n e  the most l i k e l y time, j u s t a f t e r the P l i o c e n e  or  plants  Pleis-  uplift  of  the Andes (Raven 1972). Such i n f o r m a t i o n  as i s a v a i l a b l e , t h e r e f o r e , suggests t h a t  the  p r o p e r t i e s of S. v i o l a c e a and_S. a l c h e m i l l o i d e s conform w i t h Raven's (1963) generalizations favouring pothesize  t h a t the  long-distance•dispersal.  common a n c e s t o r  of the two  One  may  s p e c i e s was  where i t has  and became e s t a b l i s h e d i n the B o l i v i a n and evolved  i n i s o l a t i o n f o r the  last 2 - 4  hy-  transported  b i r d s from the Rocky Mountains sometime i n the l a t e P l i o c e n e Pleistocene  therefore  by  or e a r l y  Argentinian  m i l l i o n years,  Andes, and  253  i s presently recognized  as S_. a l c h e m i l l o i d e s .  I n view o f the s e a s o n a l l y wet h a b i t a t , t h e r e i s a good chance t h a t b i r d s o f the o r d e r C h a r a d i i f o r m e s ,  which comprises about 70% o f the  s p e c i e s m i g r a t i n g between the c o n t i n e n t s the d i s p e r s a l v e c t o r s  (Raven 1963).  (Cruden 1966), c o u l d have been  Although evidence  S u k s d o r f i a by b i r d s i s l a c k i n g , c i r c u m s t a n t i a l evidence r e l a t e d B o y k i n i a . r i c h a r d s o n i i (Ch. X I I I ) . is  a b e t t e r candidate  f o r p r e d a t i o n on does e x i s t  I t might be n o t e d t h a t  f o r l o n g - d i s t a n c e d i s p e r s a l than some o t h e r  because i t can be t r a n s p o r t e d b o t h as seeds and/or as b u l b i l s .  f o r the Suksdorfia genera  As t e s t i - .  mony t o i t s m o b i l i t y , i t i s a l s o the o n l y genus which has r e c o l o n i z e d g l a c i a t e d t e r r i t o r y t o any g r e a t  extent.  254  XVI.  ETHNOBOTANY  Horticulture Many s p e c i e s of the S a x i f r a g i n a e , e s p e c i a l l y of S a x i f r a g a Bergenia, favoured  are p o p u l a r  as garden ornamentals.  as house p l a n t s .  j a m e s i i has been w i d e l y ful  are  Of the s p e c i e s i n the p r e s e n t study, o n l y B. -  grown.  p i n k - p u r p l e f l o w e r s and  Some, l i k e T o l m i e a ,  and  I t has been j u s t l y admired f o r i t s b e a u t i -  attractive foliage.  There has  been some de-  bate i n the p a s t over i t s r e a d i n e s s to f l o w e r i n the garden (Heath 1964), and s p e c i a l c o n d i t i o n s , i n c l u d i n g r e s t r i c t i n g (Preece in  1937).  the r o o t s are r e q u i r e d  H i t c h c o c k e t a l . (1961) r e p o r t e d t h a t B_. j a m e s i i does w e l l  Great B r i t a i n but i s u n t h r i f t y i n gardens i n the P a c i f i c Northwest.  T h i s d i f f e r e n c e may  be s i m p l y a c o n f u s i o n o f IK j a m e s i i w i t h JK_ h e u c h e r i -  f o r m i s by gardeners.  Boykinia heucheriformis  B r i t a i n at a l l (B. Mathew, Kew western N o r t h America. heucheriformis, r e g u l a r l y but  r e s u l t s would be had  grown i n  Gardens, i n l i t t . ) whereas i t i s grown i n  P r o f . J . G. Packer ( i n l i t t . )  t r a n s p l a n t e d from M i e t t e Hot  does not  i s a p p a r e n t l y not  reported that  S p r i n g s , near J a s p e r ,  f l o u r i s h i n h i s Edmonton garden.  B. flowers  Perhaps b e t t e r  i n the northwest i f IK j a m e s i i were to be more w i d e l y  grown. Although  d i s c o v e r e d i n 1820  h o r t i c u l t u r a l p o t e n t i a l of IK when i t was in  ( T o r r e y 1827), i n t e r e s t i n the  j a m e s i i was  shown f o r the f i r s t  not roused  u n t i l the  time by Dr. G u i s e p p i i n 1935  1930's,  (B. Mathew  litt.). Other s p e c i e s have been suggested  including Boykinia a c o n i t i f o l i a  (Gray  to h o r t i c u l t u r a l promise,  1842), J K  major, B.  occidentalis,  S u k s d o r f i a r a n u n c u l i f o l i a and_S. v i o l a c e a ( H i t c h c o c k e t a l . 1961). t h i s l i s t I would add P e l t o b o y k i n i a t e l l i m o i d e s (both s u b s p e c i e s ) Boykinia richardsonii.  P e l t o b o y k i n i a has  To and  a t t r a c t i v e p e l t a t e leaves  and  255  showy, d e l i c a t e l y - c o l o u r e d , y e l l o w f l o w e r s .  Boykinia richardsonii i s a  t r u l y handsome p l a n t w i t h i t s deep green l e a v e s and l a r g e white often with red veins i n the p e t a l s .  flowers,  However, i t would p r o b a b l y be r e l u c -  t a n t to f l o w e r i n gardens a t a more s o u t h e r l y l a t i t u d e than i t s n a t i v e arctic. Medicine The  l e a v e s o f B o y k i n i a o c c i d e n t a l i s a r e eaten by t h e Q u i l e u t e  I n d i a n s o f the Olympic P e n i n s u l a , Washington, as a treatment losis  (Gunther 1973).  f o r tubercu-  None o f the o t h e r s p e c i e s i n the p r e s e n t  been r e p o r t e d t o have m e d i c i n a l p r o p e r t i e s .  study has  256  REFERENCES Agababyan, V.C. 1961. M a t e r i a l f o r the p a l y n o s y s t e m a t i c study o f the f a m i l y S a x i f r a g a c e a e s . l . ( i n R u s s i a n ) . Acad. S c i . Rep. Arm., S c i B i o l . , E r i v a n 14(2): 45-61. B a c i g a l u p i , R. 1944. S a x i f r a g a c e a e . I n : Abram's i l l u s t r a t e d f l o r a o f the P a c i f i c s t a t e s 2: 349-384. S t a n f o r d U n i v . P r e s s , C a l i f o r n i a . B a c i g a l u p i , R. 1947. A new combination Bot. 5: 71. B a c i g a l u p i , R. 1952. S a x i f r a g a c e a e . Brown Co., Dubuque, Iowa.  i n the genus T e l e s o n i x .  L e a f l . West.  I n : R.J. D a v i s , F l o r a o f Idaho.  W.C.  Baker, H.G. 1955. S e l f - c o m p a t i b i l i t y and e s t a b l i s h m e n t a f t e r " l o n g d i s t a n c e " dispersal. E v o l u t i o n 9: 347-348. Barnhart, J.H. 1907. The d a t e s . o f Rafinesque's ana. T o r r e y a 7: 177-182.  New F l o r a and F l o r a  Telluri-  Bate-Smith, E.C. 1962. vThe p h e n o l i c c o n s t i t u e n t s o f p l a n t s and t h e i r taxonomic s i g n i f i c a n c e . 1. D i c o t y l e d o n s . J . L i n n . Soc. (Bot.) 58: 95-173. B e n s e l , C R . and B.F. P a l s e r . 1975. F l o r a l anatomy i n the S a x i f r a g a c e a e sensu l a t o . I I . S a x i f r a g o i d e a e and I t e o i d e a e . Amer. J . B o t . 62: 661-675. Bohm, B.A. 1979. F l a v o n o i d s . o f Tolmiea m e n z i e s i i .  Phytochem. 18: 1079-1080.  Bohm, B.A. and F.W. C o l l i n s . 1979. F l a v o n o i d s o f some s p e c i e s o f Chrysosplenium. Biochem..Syst. E c o l . 7: 195-201. Bohm, B.A. and R. O r n d u f f . 1978. Chemotaxonomic s t u d i e s i n the S a x i f r a g a c e a e s . l . 9. F l a v o n o i d s o f J e p s o n i a . Madrono 25: 39-43. Bohm, B.A. and C.K. W i l k i n s . 1976. F l a v o n o i d s and g a l l i c a c i d d e r i v a t i v e s from P e l t i p h y l l u m p e l t a t u m . Phytochem. 15: 2012-2013. Bohm, B.A. and C.K. W i l k i n s . 1978a. Chemotaxonomic s t u d i e s i n the S a x i f r a g aceae s . l . 10. The f l a v o n o i d s o f Heuchera c y l i n d r i c a . Can. J . B o t . 56: 1174-1176. Bohm, B.A. and C.K. W i l k i n s . 1978b. Chemosystematic s t u d i e s i n the S a x i f r a g aceae sensu l a t o . 8. The f l a v o n o i d s o f Elmera racemosa (Watson) Rydberg. B r i t t o n i a 30: 327-333. Brandegee, T.S. 1899. New s p e c i e s o f western p l a n t s .  B o t . Gaz. 27: 447-448.  B r i q u e t , J . 1906. Regies i n t e r n a t i o n a l e s de l a nomenclature botanique adoptees p a r l e congres i n t e r n a t i o n a l de botanique de Vienne 1905. V e r l a g von G. F i s c h e r , Jena. B u r t t , B.L. 1964. Angiosperm taxonomy i n p r a c t i c e . 16.  S y s t . Assoc.  P u b l . 6: 5-  257 C a l d e r , J.A. and D.B.O. S a v i l e . 1959. S t u d i e s Heuchera c y l i n d r i c a complex i n and a d j a c e n t B r i t t o n i a 11: 49-67.  i n S a x i f r a g a c e a e . 1. The t o B r i t i s h Columbia.  C a l d e r , J.A. and R.L. T a y l o r . 1968. F l o r a o f the Queen C h a r l o t t e I s l a n d s . V o l . 1. Systematics o f the v a s c u l a r p l a n t s . Queen's P r i n t e r , Ottawa. C a r l q u i s t , S. 1961. Comparative p l a n t anatomy. New York. Chambers, K.L. 1964.  Holt, Rinehart  Saxifraga e s c h s c h o l t z i i Sternb.  & Winston,  Madrono 17: 203-204.  C o l l i n s , F.W. and B.A. Bohm. 1974. Chemotaxonomic s t u d i e s i n the Saxifrage: .•aceae s . l . 1. The f l a v o n o i d s o f T e l l i m a g r a n d i f l o r a . Can. J . B o t . 52: 307-312. C o l l i n s , F.W., B.A. Bohm and C K . W i l k i n s . 1975; F l a v o n o l g l y c o s i d e g a l l a t e s from T e l l i m a g r a n d i f l o r a . Phytochem. 14: 1099-1102. Coney, P . J . 1972. C o r d i l l e r a n t e c t o n i c s , and North American p l a t e motion. ' Amer; J . S c i . 22: 603-628. C o n o l l y , A.P.. 1972. Scanning e l e c t r o n microscope photographs o f s a x i f r a g e seeds. Watsonia 9: 58. C o n o l l y , A.P. 1976. Use o f t h e scanning e l e c t r o n microscope f o r t h e i d e n t i f i c a t i o n o f seeds w i t h s p e c i a l r e f e r e n c e t o S a x i f r a g a and Papaver. F o l i a Quat. 47: 29-38. Cooley, G.E. 1892.  Impressions o f A l a s k a .  B u l l . T o r r . B o t . Club 19: 178-189.  C o r r e n s , C. 1928. Neue Uhtersuchungen an s e l b s t s t e r i l e n P f l a n z e n . menziesii. B i o l . Z b l . 48: 759-768. C o u l t e r , J.M. 1892.  S u l l i v a n t i a Hapemani.  C o u l t e r , J.M. and E.M. F i s h e r . 1892. Bot. Gaz. 17: 348-352.  1. Tolmiea  B o t . Gaz. 17: 421.  Some new North American p l a n t s 1.  C r a i g h e a d , J . J . , F.C. Craighead and R.J. D a v i s . 1963. A f i e l d Mountain w i l d f l o w e r s . Houghton M i f f l i n Co., Boston.  guide to Rocky  Crowson, R.A. 1953. A p o s s i b l e new p r i n c i p l e i n taxonomy, and i t s e v o l u t i o n ary i m p l i c a t i o n s . Nature 171: 883. Cruden, R.W. 1966. B i r d s as agents o f l o n g d i s t a n c e d i s p e r s a l f o r d i s j u n c t p l a n t groups o f t h e temperate western hemisphere. E v o l u t i o n 20: 517-532. Dahlgren, K.V.O. 1930. Zur embryologie der S a x i f r a g o i d e e n . T i d s k r . 24: 429-448.  Svensk. B o t .  Dandy, J . E . 1927. The genera o f S a x i f r a g a c e a e . Pp. 107-118 i n J . H u t c h i n s o n , C o n t r i b u t i o n s towards a p h y l o g e n e t i c c l a s s i f i c a t i o n o f f l o w e r i n g p l a n t s : VI. B u l l . M i s c . I n f . Kew 1927: 100-118.  258  Dandy, J . E . 1935. B o y k i n i a . P. 87 i n J.S.L. Gilmour e t a l . V. A d d i t i o n s and ammendments t o the i n t e r n a t i o n a l r u l e s o f b o t a n i c a l nomenclature. B u l l . M i s c . I n f . Kew 1935: 65-92.D a r l i n g t o n , C.D. 1939. E v o l u t i o n o f g e n e t i c systems. Edinburgh.  O l i v e r & Boyd,  D a r l i n g t o n , C.D. and L.F. L a Cour. 1976, ed. A l l e n & Unwin, London.  o f chromosomes. 6th  D a v i s , G.L. 1966.  Systematic  The h a n d l i n g  embryology o f t h e angiosperms.  Wiley,  New York.  D a v i s , P.H. 1978. The moving s t a i r c a s e : a d i s c u s s i o n o f taxonomic rank and affinity. Notes R. B o t . Gard. E d i n b . 36: 325-340. • D a v i s , P.H. and V.H. Heywood. 1963. P r i n c i p l e s o f angiosperm taxonomy. O l i v e r & Boyd, Edinburgh. Eaton, R.D. 1973. The e v o l u t i o n o f seed i n c o m p a t i b i l i t y i n P r i m u l a . P h y t o l . 72: 855-860.  New  E i s b a c h e r , G.H. 1977. M e s o z o i c - T e r t i a r y b a s i n models f o r t h e Canadian Cordi l l e r a and t h e i r g e o l o g i c a l c o n s t r a i n t s . Can. J . E a r t h S c i . 14: 2414-2421. E n g l e r , A. 1891. S a x i f r a g a c e a e . I n : A. E n g l e r and K. P r a n t l , D i e n a t u r l i c h e n P f l a n z e n f a m i l i e n I I I . 2a: 41-93. E n g l e r , . A . 1918. Hieronymusia E n g l . , e i n e neue Gattung der S a x i f r a g a c e e n . N o t i z . K o n i g l . B o t . G a r t . Mus. B e r l i n - D a h l e m 7: 265-267. E n g l e r , A. 1928. S a x i f r a g a c e a e . I n : A. E n g l e r and K. P r a n t l , D i e n a t u r l i c h e n P f l a n z e n f a m i l i e n ( e d . 2) 18a: 74-226. E n g l e r , A. and E . Irmscher. 1916-1919. S a x i f r a g a c e a e P f l a n z e n r e i c h 67 (1916) and 69 (1918-1919).  - Saxifraga.  Erdtman, G. 1960. The a c e t o l y s i s method. A r e v i s e d d e s c r i p t i o n . Bot. T i d s k r . 54: 561-564. Estabrook, G.F. 1977. Does -common e q u a l p r i m i t i v e ?  Das  Svensk.  S y s t . B o t . 2: 36-42.  E s t a b r o o k , G.F. 1978. Some concepts f o r t h e e s t i m a t i o n o f 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 n s y s t e m a t i c botany. S y s t . B o t . 3: 146-158. Evans, G.C. and A.P. Hughes. 1961. P l a n t growth and the a e r i a l , environment. 1. E f f e c t o f a r t i f i c i a l shading on Impatiens p a r v i f l o r a . New P h y t o l . 60: 150 Fedde, F. 1906. Novorum generum, specierum, v a r i e t a t u m , formarumque, Siphonogamarum Index. J u s t ' s B o t . J a h r e s b e r . 33 ( 1 ) : 361-643. 1905. Ferguson, I.K. and D.A. Webb. 1970. P o l l e n morphology i n the genus S a x i f r a g a a n d . i t s taxonomic s i g n i f i c a n c e . B o t . J . L i n n . Soc. 63: 295-311. ;  F e r r i s , R.S. 1960. Orobanchaceae. I n : Abram's I l l u s t r a t e d f l o r a o f t h e P a c i f i c s t a t e s 4: 3-10. S t a n f o r d U n i v . P r e s s , C a l i f o r n i a .  259  F i e l d i n g , J . and G. Gardner. 1844. F i n k , W.L.  and E.O. W i l e y . 1979.  Saxifraga a c o n i t i f o l i a .  Cladism  defended.  F i t z p a t r i c k , T . J . 1911. R a f i n e s q u e : a s k e t c h H i s t o r i c a l Dept. o f Iowa, Des Moines.  Sert. P l . t57.  Nature 280: 542.  of h i s l i f e with  bibliography.  Funk, V.A. and T.F. S t u e s s y . 1978. C l a d i s t i c s f o r the p r a c t i c i n g p l a n t taxonomist. S y s t . B o t . 3: .159-178. G a r d i n e r , B.G. e t d . 1979. Nature 277: 175-176.  The salmon, the l u n g f i s h and the cow: a r e p l y .  G o r e n f l o t , R. 1971. I n t e r e t taxinomique e t phylogenique des c a r a c t e r e s stomatiques ( a p p l i c a t i o n a l a t r i b u des S a x i f r a g e e s ) . B o i s s i e r a 19: 181192. G o r e n f l o t , R. and F. Moreau. 1970. Types stomatiques e t p h y l o g e n i e des Saxifraginees (Saxifragacees). C.R. Acad. S c i . , P a r i s 270: 2802-2805. G o r n a l l , R.J. and B.A. Bohm. 1978. Angiosperm f l a v o n o i d e v o l u t i o n : a reappraisal. S y s t . B o t . 3: 353-368. G r a e n i c h e r , S. 1907. W i s c o n s i n f l o w e r s and t h e i r p o l l i n a t i o n . II.- S a x i f r a g aceae and G r o s s u l a r i a c e a e . B u l l . Wis. N a t . H i s t . Soc. I I . 5: 84-95. Graham, A. 1972. O u t l i n e o f the o r i g i n and h i s t o r i c a l r e c o g n i t i o n o f f l o r i s t i c a f f i n i t i e s between A s i a and e a s t e r n North America. I n : A. Graham (ed.) F l o r i s t i c s and p a l e o f l o r i s t i c s o f A s i a and e a s t e r n North America. E l s e v i e r , New York. Grant, V. 1971.  P l a n t s p e c i a t i o n . Columbia U n i v . P r e s s , New York.  G r a u s t e i n , J . E . 1967. Thomas N u t t a l l , n a t u r a l i s t ; e x p l o r a t i o n s i n America 1808-1841. Harvard U n i v . P r e s s , Cambridge. Gray, A. 1842. Notes o f a b o t a n i c a l e x c u r s i o n . t o the mountains o f North C a r o l i n a e t c . ; w i t h some remarks on the botany o f the h i g h e r A l l e g h a n y Mountains. Amer. J . S c i . 42: 1-49. Gray, A. 1867. C h a r a c t e r s o f new p l a n t s o f C a l i f o r n i a and elsewhere p r i n c i p a l l y o f those c o l l e c t e d by H.N. Bolander i n the S t a t e G e o l o g i c a l Survey. P r o c . Amer. Acad. A r t s S c i . 7: 327-401. Gray, A. 1872. B o t a n i c a l c o n t r i b u t i o n s I I . D e t e r m i n a t i o n o f a c o l l e c t i o n o f p l a n t s made i n Oregon by E l i h u H a l l d u r i n g t h e summer o f 1871, w i t h chara c t e r s o f some new s p e c i e s and v a r i o u s n o t e s . P r o c . Amer. Acad. A r t s S c i .  8: 3 6 5 - 4 1 2 .  Gray, A. 1876. S a x i f r a g a c e a e . I n : Botany o f C a l i f o r n i a Co., U n i v . P r e s s , Cambridge, Mass.  1.  Welch, Bigelow &  Gray, A. 1878. C o n t r i b u t i o n s t o t h e botany o f North America. Acad. A r t s S c i . 13: 361-374.  Proc.  Amer.  260 Gray, A. 1879. 25-52.  Botanical contributions.  P r o c . Amer. Acad. A r t s S c i . 15:  Green, M.L. 1940. X I I . A d d i t i o n a l nomina g e n e r i c a conservanda and phanerogamae). Kew B u l l . 1940: 81-132. Greene, E.L. 1891.  Flora franciscana.  (Pteridophyta  San F r a n c i s c o .  Greene, E.L. 1894. Manual o f the botany o f the r e g i o n o f San F r a n c i s c o Bay. Cubery & Co., San F r a n c i s c o . G r i f f i t h s , G.C.D. 1972. S t u d i e s on b o r e a l Agromyzidae ( D i p t e r a ) 1. Phytomyza miners on S a x i f r a g a c e a e . Quaest. Entomol. 8: 67-80. G r i s e b a c h , A.H.R. 1879. Symbolae ad f l o r a m a r g e n t i n i a m . Zweite b e a r b e i t u n g a r g e n t i n i s c h e r p f l a n z e n . Abhandl. Kon. G e s e l l . Wiss. G o t t i n g e n 24.: 3-345. Gunther, E. 1973. Ethnobotany o f western Washington. The knowledge and use of i n d i g e n o u s p l a n t s by n a t i v e Americans. R e v i s e d ed., U n i v . Washington Press, Seattle. Hadac, E . 1961. Ants as p o l l i n a t o r s o f p l a n t s . T a t r a N a t i o n a l Park, B r a t i s l a v a 4: 255-256.  T r e a t i s e s c o n c e r n i n g the  H a l s t e a d , L.B. 1978. The c l a d i s t i c r e v o l u t i o n - can i t make the grade? Nature 276: 759-760. H a l s t e a d , L.B., E . I . White and G.T. M a c l n t y r e . 1979. Reply t o "The salmon, the l u n g f i s h and the cow: a r e p l y " . Nature 277: 176. Hamel, J . L . 1948. Notes p r e l i m i n a i r e s a une etude .caryologique des S a x i f v;.ragaeees. 1. Les chromosomes de P e l t i p h y l l u m p e l t a t u m ( T o r r . ) E n g l , e t de B o y k i n i a t e l l i m o i d e s (Maxim.) E n g l . B u l l . Mus. H i s t . N a t . P a r i s I I . 20: 198-200. :  Hamel, J . L . 1953. C o n t r i b u t i o n a l ' e t u d e c y t o - t a x i n o m i q u e des S a x i f r a g a c e e s . Rev. C y t o l . e t B i o l . Veg. 14: 113-313. Hamel, J . L . 1958. M a t e r i a u x pour I* "etude caryo-taxinomique des S a x i f r a g a c e e s . 5. Le noyau e t l e s chromosomes somatiques du B o l a n d r a oregana S. Wats. B u l l . Mus. H i s t . Nat; P a r i s I I . 30: 522-524. Hara, H. 1937. Two new genera o f S a x i f r a g a c e a e i n Japan. 51: 250-253. Hara, H. 1957. Synopsis o f the genus Chrysosplenium L . Tokyo I I I . 7: 1-90. Hara, H. 1958. Co., Tokyo.  B o t . Mag., Tokyo  J , Fac. S c i . Univ.  D i s t r i b u t i o n maps o f f l o w e r i n g p l a n t s i n Japan 1.  Hara, H. 1959. An o u t l i n e o f the phytogeography  o f Japan.  D i s t r i b u t i o n maps o f f l o w e r i n g p l a n t s i n Japan.2.  Collectors' alpines.  I n : H. Hara,  Inoue Book Co., Tokyo.  H a r r i n g t o n , H.D. 1954. Manual o f t h e p l a n t s o f C o l o r a d o . Heath, R.A. 1964.  Inoue Book  Sage Books,  C o l l i n g r i d g e , London.  Denver.  261 Hedberg, 0. 1958. The taxonomic treatment A r s s k r . 6: 186-195.  of v i c a r i o u s t a x a .  Uppsala  Univ.  H e l l e r , A.A. 1900.. Catalogue of North American p l a n t s n o r t h o f Mexico, e x c l u s i v e of the lower cryptogams, ed. 2. H e l l e r , A.A.  1904.  Western s p e c i e s , new  and  old - I I .  Muhlenbergia  1: 47-62,  Henry, J.K. 1915. F l o r a o f southern B r i t i s h Columbia and Vancouver I s l a n d . W.J. Gage & Co., T o r o n t o . H e s l o p - H a r r i s o n , J . 1975. I n c o m p a t i b i l i t y and Ann. Rev. P l a n t P h y s i o l . 26: 403-425.  the p o l l e n - s t i g m a  H e s l o p - H a r r i s o n , Y. and K.R. Shivanna. 1977.. The angiosperm stigma. Ann. Bot. 41: 1233-1258.  interaction.  r e c e p t i v e s u r f a c e of the  H i c k e y , L.F. .1973. C l a s s i f i c a t i o n of the a r c h i t e c t u r e o f l e a v e s . Amer. J . Bot. 60: 17-33.  dicotyledonous  Hideux, M.J. and I.K. Ferguson.'1976. The s t e r e o s t f u c t u r e of the e x i n e and i t s e v o l u t i o n a r y s i g n i f i c a n c e i n S a x i f r a g a c e a e sensu l a t o . I n : I.K. Ferguson and J . M u l l e r . (eds.) The e v o l u t i o n a r y s i g n i f i c a n c e of the e x i n e . L i n n . Soc. Symp. S e r . 1. Academic P r e s s , London. H i t c h c