{"http:\/\/dx.doi.org\/10.14288\/1.0093501":{"http:\/\/vivoweb.org\/ontology\/core#departmentOrSchool":[{"value":"Science, Faculty of","type":"literal","lang":"en"},{"value":"Botany, Department of","type":"literal","lang":"en"}],"http:\/\/www.europeana.eu\/schemas\/edm\/dataProvider":[{"value":"DSpace","type":"literal","lang":"en"}],"https:\/\/open.library.ubc.ca\/terms#degreeCampus":[{"value":"UBCV","type":"literal","lang":"en"}],"http:\/\/purl.org\/dc\/terms\/creator":[{"value":"Pojar, Jim","type":"literal","lang":"en"}],"http:\/\/purl.org\/dc\/terms\/issued":[{"value":"2010-02-01T22:27:09Z","type":"literal","lang":"en"},{"value":"1974","type":"literal","lang":"en"}],"http:\/\/vivoweb.org\/ontology\/core#relatedDegree":[{"value":"Doctor of Philosophy - PhD","type":"literal","lang":"en"}],"https:\/\/open.library.ubc.ca\/terms#degreeGrantor":[{"value":"University of British Columbia","type":"literal","lang":"en"}],"http:\/\/purl.org\/dc\/terms\/description":[{"value":"Four plant communities of southwestern British Columbia were studied in an attempt to answer the following related questions: (1) do communities of harsh physical environments exhibit any characteristic phytosociological features? (2) are there any correlations between environmental harshness and certain synecological properties of such communities ? (3) are species of such communities selected for reproductive specializations that tend to reduce their genetic variability? The four communities (representing three types of herbaceous or semi-shrubby vegetation) were a salt marsh, two coastal sphagnum bogs, and a subalpine meadow. In answer to the first two questions, the findings of this investigation indicate that: (a) species population structure becomes more aggregated as environmental heterogeneity and physical stress increase, and less aggregated as succession proceeds and interspecific\r\ncompetition increases. (b) interspecific association and correlation, both negative and positive, increase as environmental heterogeneity and competition increase. (c) levels of polyploidy within communities appear to be correlated with environmental rigor (broadly defined).\r\n\r\n(d) the most abundant species within a community are the most variable and presumably have the largest niches ; niche size and population variability decrease as interspecific\r\ncompetition increases. (e) within a community, ecological distinctiveness reduces interspecific competition; communities under the least stress (especially seasonal stress) have the most ecologically dissimilar1 species. (f) dominance decreases as species diversity increases, and species diversity is roughly correlated with overall environmental severity. In answer to the last question: (a) all four communities are dominated by predominantly outcrossing species; there is no major shift to self-pollination or apomixis in any of the communities. (b) an index of potential recombination was devised, embodying a number of aspects of reproductive biology, according to which there is no significant difference in potential recombination, on the average, between species of the four different communities. Plant communities and their constituent species both respond to evolutionary forces, but more or less independently, at different rates, and often in different or even opposite ways. Environmental stress has a powerful effect on the structure and function of plant communities, but in an evolutionary sense there is little difference between normal (mesic, zonal) environments and extreme or azonal environments\r\n\r\nto an individual species. Different selection pressures have been operating in salt marshes, sphagnum bogs, and subalpine meadows, but the present study indicates that, on the average, the resultant evolutionary strategies of the species of these communities are equivalent.","type":"literal","lang":"en"}],"http:\/\/www.europeana.eu\/schemas\/edm\/aggregatedCHO":[{"value":"https:\/\/circle.library.ubc.ca\/rest\/handle\/2429\/19524?expand=metadata","type":"literal","lang":"en"}],"http:\/\/www.w3.org\/2009\/08\/skos-reference\/skos.html#note":[{"value":"C- I THE RELATION OF THE REPRODUCTIVE BIOLOGY OF PLANTS TO THE STRUCTURE AND FUNCTION OF FOUR PLANT COMMUNITIES by JIM POJAR B . S c , U n i v e r s i t y of Minnesota, 1969 M.Sc. , U n i v e r s i t y of Minnesota, 1970 A THESIS SUBMITTED IN.PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY i n the Department of Botany We accept t h i s t h e s i s as conforming to the r e q u i r e d standard THE UNIVERSITY OF BRITISH COLUMBIA In presenting th is thes is in p a r t i a l fu l f i lment of the requirements fn an advanced degree at the Un ivers i ty of B r i t i s h Columbia, I a g r e e that the L ibrary sha l l make it f ree ly ava i lab le for reference and s t udy I fur ther agree that permission for extensive copying of th is t h e s i s for scho lar ly purposes may be granted by the Head of my Department or by his representat ives . It is understood that copying or p u b l i c a t i o n of th is thes is fo r f i n a n c i a l gain sha l l not be allowed without my writ ten pe rm i ss i on . Department of \"\\j> The Univers i ty of B r i t i s h Columbia Vancouver 8, Canada Date \u00b0\\LaAX \\ , V H *r Abstract F o u r p l a n t c o m m u n i t i e s o f s o u t h w e s t e r n B r i t i s h C o l u m b i a were s t u d i e d i n an a t t e m p t t o a n s w e r t h e f o l l o w i n g r e l a t e d q u e s t i o n s : (1) do c o m m u n i t i e s o f h a r s h p h y s i c a l e n v i r o n m e n t s e x h i b i t any c h a r a c t e r i s t i c p h y t o s o c i o l o g i c a l f e a t u r e s ? (2) a r e t h e r e any c o r r e l a t i o n s b e t w e e n e n v i r o n m e n t a l h a r s h n e s s and c e r t a i n s y n e c o l o g i c a l p r o p e r t i e s o f s u c h c o m m u n i t i e s ? (3) a r e s p e c i e s o f s u c h c o m m u n i t i e s s e l e c t e d f o r r e p r o d u c t i v e s p e c i a l i z a t i o n s t h a t t e n d t o r e d u c e t h e i r g e n e t i c v a r i a b i l i t y ? The f o u r c o m m u n i t i e s ( r e p r e s e n t i n g t h r e e t y p e s o f h e r b a c e o u s o r s e m i - s h r u b b y v e g e t a t i o n ) w e r e a s a l t m a r s h , two c o a s t a l sphagnum b o g s , and a s u b a l p i n e meadow. I n a n s w e r t o t h e f i r s t two q u e s t i o n s , t h e f i n d i n g s o f t h i s i n v e s t i g a t i o n i n d i c a t e t h a t : ( a ) s p e c i e s p o p u l a t i o n s t r u c t u r e becomes more a g g r e g a t e d as e n v i r o n m e n t a l h e t e r o g e n e i t y and p h y s i c a l s t r e s s i n c r e a s e , and l e s s a g g r e g a t e d as s u c c e s s i o n p r o c e e d s and i n t e r -s p e c i f i c c o m p e t i t i o n i n c r e a s e s . (b) i n t e r s p e c i f i c a s s o c i a t i o n and c o r r e l a t i o n , b o t h n e g a t i v e and p o s i t i v e , i n c r e a s e as e n v i r o n m e n t a l h e t e r o g e n e i t y a n d c o m p e t i t i o n i n c r e a s e . ( c ) l e v e l s o f p o l y p l o i d y w i t h i n c o m m u n i t i e s a p p e a r t o be c o r r e l a t e d w i t h e n v i r o n m e n t a l r i g o r ( b r o a d l y d e f i n e d ) . (d) the most abundant s p e c i e s w i t h i n a community are the most v a r i a b l e and presumably have the l a r g e s t niches ; niche s i z e and p o p u l a t i o n v a r i a b i l i t y decrease as i n t e r -s p e c i f i c competition i n c r e a s e s . (e) w i t h i n a community, e c o l o g i c a l d i s t i n c t i v e n e s s reduces i n t e r s p e c i f i c c o m p e t i t i o n ; communities under the l e a s t s t r e s s ( e s p e c i a l l y seasonal s t r e s s ) have the most e c o l o g i c a l l y d i s s i m i l a r 1 s p e c i e s . ( f ) dominance decreases as s p e c i e s d i v e r s i t y i n c r e a s e s , and species d i v e r s i t y i s roughly c o r r e l a t e d w i t h o v e r a l l environmental s e v e r i t y . In answer to the l a s t q u e s t i o n : (a) a l l f o u r communities are dominated by predominantly o u t c r o s s i n g s p e c i e s ; there i s no major s h i f t t o s e l f -p o l l i n a t i o n or apomixis i n any of the communities. (b) an index of p o t e n t i a l recombination was d e v i s e d , embodying a number of aspects of r e p r o d u c t i v e b i o l o g y , a c c o r d i n g to which 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 p o t e n t i a l recombination, on the average, between s p e c i e s of the f o u r d i f f e r e n t communities. Pl a n t communities and t h e i r c o n s t i t u e n t s p e c i e s both respond to e v o l u t i o n a r y f o r c e s , but more or l e s s independently, at d i f f e r e n t r a t e s , and o f t e n i n d i f f e r e n t or even op p o s i t e ways. Environmental s t r e s s has a powerful e f f e c t on the s t r u c t u r e and f u n c t i o n of p l a n t communities, but i n an e v o l u t i o n a r y sense there i s l i t t l e d i f f e r e n c e between normal (mesic, zonal) environments and extreme or azo n a l environments to an i n d i v i d u a l s p e c i e s . D i f f e r e n t s e l e c t i o n p r e s s u r e s have been o p e r a t i n g i n s a l t marshes, sphagnum bogs, and subalpine meadows, but the present study i n d i c a t e s t h a t , on the average, the r e s u l t a n t e v o l u t i o n a r y s t r a t e g i e s of the s p e c i e s of these communities are e q u i v a l e n t . TABLE OF CONTENTS Section Page I Introduction 1 II Description 5 A. Sampling methods 5 B. Sampling r e s u l t s 7 C. Vegetation of the four communities 10 S a l t marsh 10 Sphagnum bogs 14 Subalpine meadow 19 III Theory and Results 23 A. Pop u l a t i o n s t r u c t u r e of i n d i v i d u a l species (D\/d index of aggregation) 23 B. I n t e r s p e c i f i c r e l a t i o n s h i p s 30 A s s o c i a t i o n 30 X 2 and Cole's index of a s s o c i a t i o n 30 Species c o n s t e l l a t i o n s 33 Corre latdon x: 40 C. Chromosome numbers and p o l y p l o i d y 51 D. Flowering phenology 6 9 E. P o l l i n a t i o n ecology 7 3 F. D i s p e r s a l ecology 115 G. Vapiability^'and-v'niche--w-id-th. 12 7 H. Niche d i f f e r e n t i a t i o n 136 I. Dominance, d i v e r s i t y , and s t a b i l i t y 147 J. Index of p o t e n t i a l recombination 159 v i . Section Page IV Summary, D i s c u s s i o n , and Conclusions 190 V L i t e r a t u r e Cited 202 Appendix 1. Species l i s t s 227 Appendix 2. Summary of some s t a t i s t i c s 234 Appendix 3. Summary of reproductive biology of each species 288 v i i . , LIST OF TABLES TABLE Page 1 Summary of vegetation sampling 6 2 Mean number of species per quadrat f o r the various quadrat s i z e s 8 3 Frequency, mean cover, and importance values of the vascular p l a n t species of the S a l t Marsh 10 4 Frequency, mean cover, and importance values of the vasc u l a r o p l a n t species of Wade's Bog 15 5 Frequency, mean cover, and importance values of the vascu l a r p l a n t species of Ogg's Bog 16 6 Frequency, mean cover, and importance values of the vascu l a r p l a n t species of B l a c k w a l l Meadow 21 7 D\/d index of aggregation 2 5 8 Weighted average community D\/d values 29 9 Summary of species i n t e r a c t i o n s 4 7 10 Chromosome numbers, p o l y p l o i d y , and importance values of species of the four study communities.... 60 11 Summary of l e v e l s of p o l y p l o i d y 6 6 12 Community mode of p o l l i n a t i o n as percentage of the f l o r a and vegetation 7 3 13 Outline of diaspore types 116 14 Percentages of diaspore types 117 15 Percentages of d i s p e r s a l methods; f l o r a \/ v e g e t a t i o n . 121 16 Morphological v a r i a t i o n i n nine grass species 131 17 E c o l o g i c a l u n i t characters 138 v i i i , TABLE Page 18 Number of s p e c i e s , two i n d i c e s of dominance, and an index of d i v e r s i t y f o r the f o u r study communities.. 149 19 Index o f P o t e n t i a l Recombination 175 20 Average community I.P.R.'s 183 21 Combinations of opposing r e g u l a t o r y f a c t o r s 185 22 R e s u l t s and i n d i c a t i o n s 191 LIST OF FIGURES Figure Page 1 M o d i f i e d s p e c i e s - a r e a curves \u2022\u2022 9 2 S a l t Marsh 12 3 S a l t Marsh 12 4 Deschampsia c e s p i t o s a 13 5 Wade's Bog. i 13 6 B l a c k w a l l Meadow 2 0 7 B l a c k w a l l Meadow 20 8 S a l t Marsh - s p e c i e s c o n s t e l l a t i o n 33 9 Wade's Bog - sp e c i e s c o n s t e l l a t i o n 35 10 Ogg's Bog - sp e c i e s c o n s t e l l a t i o n 36 11 B l a c k w a l l Meadow - s p e c i e s c o n s t e l l a t i o n 38 12 Frequency d i s t r i b u t i o n s of c o r r e l a t i o n c o e f f i c i e n t s 49 13 .Number:, of .var i a b l e p a i r s vs. Ar 49 14 Deschampsia c e s p i t o s a , n - 13 53 15 F e s t u c a r u b r a , n = 21 53 16 T r i g l o c h i n maritimum, n - 48 53 17 S a l i c o r n i a v i r g i n i c a , n = 18 53 18 P l a n t a g o m a r i t i m a , n - 6 53 Figure Page 19 J u n a u s b a l t i c u s 3 n = 40 53 20 C a v e x l y n g b y e i 3 n = 36 53 21 G l a u x m a v i t i m a 3 n = 15 53 22 P o t e n t i l l a p a c i f i c a 3 n - 14 53 23 A g v o s t i s e x a v a t a 3 n - - 14. 53 24 S t e l l a v i a h u m i f u s a 3 n = 13 53 25 T v i f o l i u m w o v m s k j o l d i i 3 n - 16 53 26 S c i v p u s c e v n u u s 3 n = 30 5 3 27 P u c c i n e l l i a p u m i l a 3 n = 21 53 28 S p e r g u l a r i a c a n a d e n s i s 3 n = 18 53 29 H o r d e u m b r a c h y a n t h e v u m 3 n - 14 53 30 L i l a e o p s i s o c c i d e n t a H s 3 n = 22 53 31 Myrica. g a l e 3 n - 48 54 32 A p a v g i d i u m b o v e a l e 3 n - 9 54 33 C a v e x o b n u p t a 3 n = 37 54 34 C a v e x p l u v i f l o v a 3 n - 26 54 35 A g v o s t i s a e q u i v a l v i s 3 n = 7 54 36 S a n g u i s o v b a o f f i c i n a l i s 3 n - 14 54 37 Ledum g v o e n l a n d i c u m 3 n = 13 54 38 V a c c i n i u m o x y c o c c u s 3 n = 24..\u2022 54 39 D v o s e v a v o t u n d i f ' o l i a 3 n - 10 \u2022 54 40 K a l m i a p o l i f o l i a 3 n = 12 54 41 Empetvum n i g v u m 3 n = 13 54 42 T v i e n t a l i s a v c t i c a 3 n = c a . ~42-44 54 43 C a v e x c a n e sc ens 3 n = 28 54 44 T o f i e l d i a g l u t i n o s a 3 n = 15 54 45 L i n n a e a b o v e a l i s 3 n = 16 54 Figure 46 R h y n c h o s p o r a a l b a 3 n - 13 47 V l a n t a g o m a o r o o a r p a 3 n - 12 48 G a u l t h e r i a s h a l l o n 3 n = 44 49 G e n t i a n a s o e p t r u m , nv - 13 50 C a l a m a g r o s t i s n u t k a e n s i s 3 n = 14 51 M a i a n t h e m u m d i l a t a t u m 3 n = 18 52 S a i r p u s c e s p i t o s u s 3 n = c a . 52 53 C o p t i s a s p l e n i f o l i a 3 n - 9 . . . . 54 C o p t i s t r i f o l i a 3 n - 9 55 J u n o u s s u p i n i f ' o r m i s 3 n = c a . 56 56 G e n t i a n a d o u g l a s i a n a 3 n = 13 57 V a c c i n i u m o v a t u m 3 n - 1 2 . . . . 58 V a o o i n i u m v i t i s - i d a e a 3 n = 12 59 V a o o i n i u m u l i g i n o s u m 3 n = 24 60 N e p h r o p h y l l i d i u m o r i s t a - g a l l i 3 n 61 E r i o p h o r u m p o l y s t a c h i o n 3 n = 30 62 C a r e x p a u o i f l o r a 3 n - c a . 37 63 V a l e r i a n a s i t o h e n s i s 3 n = c a . 48 64 L u p i n u s l a t i f o l i u s 3 n = c a . 48 65 F e s t u c a v i r i d u l a 3 n - 14 66 E r i g e r o n p e r e g r i n u s 3 n - 9 . . . 67 Anemone o c o i d e n t a l i s 3 2n - 16 68 E r y t h r o n i u m g r a n d i f l o r u m , 2n - 24 69 V o t e n t i l l a f l a b e l l i f o l i a 3 n - 14 70 V a o o i n i u m s o o p a r i u m 3 n = 12 71 C l a y t o n i a l a n o e o l a t a 3 2n = 16 72 A r e n a r i a o a p i l l a r i s 3 n - 11 F i g u r e 73 A n t e n n a v i a l a n a t a , n - 14.. 74 V e r o n i c a c u s i c k i i , n = 36.. 75 A g o s e r i s a u r a n t i a c a , n - 18 76 P h l e u m a l p i n u m 3 n = 1 4 . . . . 77 A r n i c a l a t i f o l i a 3 n = 19.. 78 L u z u l a h i t c h c o e k i i , n - 12 79' T h a l i c t r u m o c c i d e n t a l e 3 n - 28 80 A c h i l l e a m i l l e f o l i u m 3 n - 27 81 T r i s e t u m s p i c a t u m 3 n = 14... 82 E l y m u s g l a u c u s 3 n = 14 83 S i l e n e p a r r y i 3 n = 24 84 A r n i c a m o l l i s , n = c a . 38... 85 P e n s t e m o n p r o c e r u s 3 n = 8... 86 P o a c u s i c k i i 3 n - 14 87 S e n e c i o i n t e g e r r i m u s 3 n = 20 88 E i e r a c i u m g r a c i l e 3 n - 9.. 89 L u z u l a s p i c a t a 3 n = 1 2 . . . . 90 Sedum l a n c e o l a t u m 3 n = 8.. 91 C a s t i l l e j a m i n i a t a 3 n = 12 92 P e d i c u l a r i s b r a c t e o s a 3 n - 8 93 P h l o x d i f f u s a , n - 7 94 P o t e n t i l l a d i v e r s i f o l i a , n - c a . 95 C a r e x s p e c t a b i l i s ; n = c a . 42 96 E p i l o b i u m a l p i n u m , n = 1 8 . . . . 97 D e l p h i n i u m n u t t a l l i a n u m , n = 16 98 C a s t i l l e j a p a r v i f l o r a , n = 12 99 R a n u n c u l u s e s c h s c h o l t z i i , n - 16 x i i . Figure Page 100 S i b b a l d i a p r o c u m b e n s 3 n - 7 58 101 J u n o u s d r u m m o n d i i , n - c a . 60 58 102 E y d r o p h y l l u m f e n d l e r i 3 n = 18 58 103 S e n e c i o t r i a n g u l a r i s 3 n - 20 58 104 V a o o i n i u m d e l i c i o s u m 3 n - 24 58 105 M i t e l l a p e n t a n d r a 3 n - 7 58 106 L u e t k e a p e c t i n a t a , n - 9 58 107 P e d i c u l a r i s r a c e m o s a , n - 8 58 108 P h y l l o d o c e e m p e t r i f o r m i s 3 n = 24 58 109 V e r a t r u m v i r i d e 3 n - 16 58 110 V e r o n i c a w o r m s k j o l d i i 3 n - 9 58 111 F lower ing phenology 7 0 112 S p e r g u l a r i a c a n a d e n s i s , s e l f - p o l l i n a t i n g f l o w e r . . . . 75 113 D r o s e r a r o t u n d i f o l i a 3 s e l f - p o l l i n a t i n g f lower 75 114 P o t e n t i l l a f l a b e l l i f o l i a , bowl-shaped blossom 77 115 P l a n t a g o m a r i t i m a , s t r o n g l y protogynous, anemophil -ous f lowers 77 116a S a l i c o r n i a v i r g i n i c a 3 f lowers i n female s t a g e . . . . . . 82 116b S a l i c o r n i a v i r g i n i c a , f lowers i n male stage 82 117 J u n c u s b a l t i c u s , s t r o n g l y protogynous, anemophilous f lowers 8 3 118 S c i r p u s c e s p i t o s u s , s t r o n g l y protogynous, anemo-p h i l o u s f lowers 8 3 119a T h a l i c t r u m o c c i d e n t a l e , male f lowers 86 119b T h a l i c t r u m o c c i d e n t a l e , female f lowers 86 120 S a n g u i s o r b a o f f i c i n a l i s , f l y - p o l l i n a t e d f l o w e r s . . . . 87 121 T r i e n t a l i s a r c t i c a , f l y - p o l l i n a t e d f lowers 87 x i i i . F i g u r e Page 122 Coptis tvifolia, f l y - p o l l i n a t e d f lower 89 123 Veratrum viride , f l y - p o l l i n a t e d f lowers 89 124 Nephrophyllidium crista-galli, c a r r i o n f l y - p o l l i -nated f lower 91 125 Lysiehitum americanum, m e p h i t i c , c a r r i o n f l y - p o l l i -nated f lowers 91 126 Veronica c u s i o k i i , f l owers p o l l i n a t e d by s y r p h i d f l i e s 92 127 Valeriana sitchensis , \"cornucopian\" f lowers 92 128 Apargidium boreale , \"cornucopian\" f lowers 94 129 Erigeron p e r e g r i n u s , \"cornucopian\" f lowers 94 130 Glaux m a r i t i m a , a n t - p o l l i n a t e d f lowers 96 131 Delphinium n u t t a l i i a n u m , bombophilous f lowers 96 132a Gentiana sceptrum, view i n t o i n t e r i o r o f a bombo-p h i l o u s f lower 98 132b Gentiana s c e p t r u m , c l o sed f lowers i n r a i n y weather. 98 133 Lupinus latifolius, bombophilous f lowers w i t h p i s t o n mechanism of p o l l e n p r e s e n t a t i o n 100 134 Pedicularis b r a c t e o s a , bombophilous f lowers w i t h short-beaked galeas 10 0 135 Pedicularis r a c e m o s a , bombophilous f lowers w i t h t w i s t e d , long-beaked galeas 10 2 136 Pedicularis g r o e n l a n d i c a , bombophilous f lowers w i t h extremely long-beaked galeas 10 2 137 Gentiana douglasiana, whi te f lowers w i t h b lue nec tar guides 104 138 Vaccinium ovatum, u r c e o l a t e , l i g h t - p i n k f l o w e r s . . . . 104 Figure Page 139 V a c o i n i u m o x y c o c c u s }convergent f l o r a l e v o l u t i o n . . . . 106 140 D'ode oath eon j e f f r ^ y i 141a S i l e n e p a r r y i (day) 108 141b S i l e n e p a r r y i (n igh t ) 108 142 A g o s e r i s a u r a n t i a c a , b u t t e r f l y - p o l l i n a t e d f l o w e r s . . HO 143 Phlox d i f f u s a , b u t t e r f l y - p o l l i n a t e d f lowers 110 144 Castilleja mini at a, hummingb i rd -po l l ina ted f l o w e r s . 112 145 Anemone o o c i d e n t a l i s , w ind-d i spe r sed f r u i t s 112 146 Empetrum n i g r u m , i n b l a c k - b e r r i e d f r u i t 120 147 P l a n t a g o macrooarpa , f r u i t i n g sp ike 120 148 Carex lyngbyei , f r u i t i n g sp ike of t h i c k - w a l l e d p e r i g y n i a 123 149 Coptis a s p l e n i f o l i a , . f o l l i c l e s adapted f o r s p l a s h -cup d i s p e r s a l 123 150 I . V . v s . C V . f o r n ine spec ies of grasses 133 151 S t rength of i n t e r s p e c i f i c compe t i t i on v s . average e c o l o g i c a l d i s t i n c t i v e n e s s 146 152 Dominance - d i v e r s i t y curves 148 153 D . I . , N v s . H ' , X . 150 XV. Acknowiedgements I thank Dr. K.I. Beamish for her supervision, of this project-, and for reading, the thesis throughout, i t s production,, keeping, a c r i t i c a l eye and lo g i c a l r e i n on i t s developments Discussions with Drs. R, Cruden, R. Foreman, J. Maze, G. Person, P, Raven, W, Schofield, G. Scudder, and R, Taylor were helpful. Thanks are extended to Dr. G. Eaton and Mrs. D,. lauriente for writing computer programs,, and to H.E. Mill i r o n , Canada Dept. of Agriculture, for identifying some bumble bees. I much appreciated the use of equipment, belonging, to Drs. R. Foreman, V. Krajina, C. Marchant, and G. Scudder, and the housing and l o c a l guidance provided i n Tbfino by Mr. A. Guppy. My brother,, Jerry, did good work as f i e l d assistant in 1971, and took most of the photographs i n the thesis;. The f i e l d work was done with the cooperation of the National and Historic Parks Branch, Department of Indian Affairs and Northern Development-, Canada, and the Brit i s h Columbia Department of Recreation and Conservation, Parks Branch. The research was supported by the Committ.ee on Research, University of Brit i s h Columbia, Grant 219554 to Dr. K.I.. Beamish, by a Grant.-in-Aid from the Society of the Sigma Xi, by a Univ. of B r i t i s h Columbia Graduate fellowship, and by the Department, of. Botany, Univ. of Brit i s h Columbia.. I. INTRODUCTION 1 In t ima t ions of i n t e r r e l a t i o n s and c o r r e l a t i o n s between the r ep roduc t ive b i o l o g y of a s i n g l e spec ies and the type o f community i t grows i n have been s u r f a c i n g i n the recent gnomic l i t e r a t u r e of e v o l u t i o n , eco logy , and sy s t ema t i c s . As the f i e l d s o f taxonomy, eco logy , p o p u l a t i o n b i o l o g y , biogeography, and e v o l u t i o n a r y b i o l o g y , fo rmer ly separate mainstreams of whole organism b i o l o g y , impinge more and more upon one another , c e r t a i n m u l t i d i s c i p l i n a r y ques t ions are be ing posed. For example, i n v e s t i g a t i o n s by S a l i s b u r y (1942) , Grant (1958) , C a r l q u i s t (1966) , Mosquin (1966) , R o l l i n s (1967) , Ornduff (1969) , Wel l s (1969) , Whitehead (1969) , Kevan (1970) , Baker , Cruden, and Baker (1971) , Baker (1972) , and S a v i l e (1972) have d e a l t , e i t h e r d i r e c t l y or i n c i d e n t a l l y , w i t h the r e l a t i o n s h i p between c e r t a i n fea tures of the r ep roduc t ive b i o l o g y of angiosperms and the nature o f the p l a n t community. Other s t ud i e s have sought to i n t e g r a t e r ep roduc t i ve and p o p u l a t i o n b i o l o g y ( e g . , L e v i n and Anderson 19 70; McNaughton and Wolf 19 70; Bradshaw 1971, 19 72; Mosquin 1971; B e a t t i e , Breed love , and E h r l i c h 1973; and a number of papers by D . A . L e v i n and by^Levin d aM wH^Wv^Kerster, E c o l o g i c a l fea tures e i t h e r found or to be expected i n c e r t a i n types o f vege t a t i on have been e l u c i d a t e d by Gre ig -Smi th (19 64) , Kershaw (1964) , Smith and Cottam (1967) , Odum (1969) , Byer (1970), Mcin tosh (1970) , and McNaughton and Wolf (1970). Furthermore, Baker (1966a) , Harper (1967) , Langford and B u e l l (1969) , Mcintosh (19 70) , and Whi t t ake r and Woodwell (1972) have commented (wi th somewhat d i f f e r e n t i n t e r p r e t a t i o n s ) on p r o p e r t i e s 2 o f c o o r d i n a t i o n , i n t e r d e p e n d e n c e , and h o m e o s t a s i s i n p l a n t c o m m u n i t i e s , p r o p e r t i e s t h a t h a v e i m p o r t a n t e c o l o g i c a l and e v o l u t i o n a r y i m p l i c a t i o n s f o r b o t h t h e c o m m u n i t y and i t s s p e c i e s . M o s t o f t h e a b o v e p a p e r s c o n t r i b u t e d i n d i v i d u a l l y t o t h e i n i t i a l s t i m u l u s f o r t h i s s t u d y , and c o l l e c t i v e l y t o t h e r e a l i z a t i o n t h a t w h a t was n e e d e d was a s y n t h e t i c a p p r o a c h . T h e r e f o r e , t h e i n v e s t i g a t i o n h a s b e e n a d d r e s s e d t o t h r e e r e l a t e d q u e s t i o n s : (1) Do p l a n t c o m m u n i t i e s o f h a r s h p h y s i c a l e n v i r o n m e n t s e x h i b i t a n y c h a r a c t e r i s t i c \" p h y t o s o c i o l o g i e a l f e a t u r e s ? ( 2 ) A r e t h e r e a n y c o r r e l a t i o n s b e t w e e n e n v i r o n m e n t a l h a r s h n e s s and c e r t a i n s y n e c o l o g i c a l p r o p e r t i e s o f s u c h c o m m u n i t i e s ? ( 3 ) A r e s p e c i e s o f s u c h c o m m u n i t i e s s e l e c t e d f o r r e p r o -d u c t i v e s p e c i a l i z a t i o n s t h a t ( a c c o r d i n g t o e s t a b l i s h e d e v o l u t i o n -a r y t h e o r y ) t e n d t o r e d u c e g e n e t i c v a r i a b i l i t y ? F o u r c o m m u n i t i e s r e p r e s e n t i n g t h r e e t y p e s o f h e r b a c e o u s o r s e m i - s h r u b b y v e g e t a t i o n ( s a l t m a r s h , sphagnum b o g , s u b a l p i n e meadow) u s u a l l y t h o u g h t o f a s o c c u r r i n g i n e x t r e m e p h y s i c a l e n v i r o n m e n t s w ere c h o s e n as s t u d y a r e a s , a n d t h e i r v e g e t a t i o n d e s c r i b e d by some o f t h e s t a n d a r d m e t h o d s o f d e s c r i p t i v e p l a n t e c o l o g y . The f o l l o w i n g c o n c e p t s o r d i s c i p l i n e s w e r e t h e n b r o u g h t t o b e a r upon t h e f i r s t t w o q u e s t i o n s : ( 1 ) s p e c i e s p o p u l a t i o n s t r u c t u r e ( 2 ) i n t e r s p e c i f i c r e l a t i o n s h i p s w i t h i n c o m m u n i t i e s a. ) a s s o c i a t i o n b. ) c o r r e l a t i o n ( 3 ) c o m m u n i t y l e v e l s o f p o l y p l o i d y ( 4 ) c o mmunity p a t t e r n s o f f l o w e r i n g p h e n o l o g y , p o l l i n a t i o n 3 eco logy , and d i s p e r s a l ecology (5) the theory of the n iche and c o r o l l a r i e s d e a l i n g w i t h n iche s i z e , p o p u l a t i o n v a r i a b i l i t y , and n iche d i f f e r e n t i a t i o n (6) dominance, d i v e r s i t y , and s t a b i l i t y . Regarding the t h i r d q u e s t i o n , Mosquin (19 66) has recommended t ha t \" I f we wish to d i s c o v e r whether o r not d i f f e r e n t f l o r i s t i c or e c o l o g i c a l zones have s i g n i f i c a n t l y d i f f e r e n t l e v e l s of gene t i c v a r i a b i l i t y . . . then the r ep roduc t i ve f a c t o r s i n each spec ies of a r e g i o n or zone must f i r s t be i d e n t i f i e d and e v a l u a t e d . . . \" The same l o g i c a p p l i e s to vege t a t i on types or i n d i v i d u a l communities. There fo re , i n a t tempt ing to answer the t h i r d q u e s t i o n , d e t a i l s of the s p e c i e s ' b reeding systems have been worked out and spec ies and community l e v e l s of p o t e n t i a l recombina t ion assessed. When I f i r s t s t a r t e d the i n v e s t i g a t i o n , I b e l i e v e d ( p a r t l y fo r the sake of argument) t h a t , i n order o f i n c r e a s i n g ha r sh -ness or r i g o r of the p h y s i c a l environment, the study communities should have been ranked: suba lp ine meadow, sphagnum bogs, s a l t marsh. Wi th t h i s r ank ing i n mind I expected c e r t a i n r e s u l t s . Some of the expec ta t ions were my own, others were e i t h e r s t a ted or i m p l i e d i n the l i t e r a t u r e . The i n i t i a l expec ta t ions are o u t l i n e d below ( d e t a i l s w i l l be g iven i n the appropr i a t e s ec t ions of the t h e s i s ) : (1) Species popu la t ions should be more aggregated as envi ronmenta l he te rogene i ty and p h y s i c a l s t r e s s i n c r e a s e , l e s s so as success ion proceeds and i n t e r s p e c i f i c compe t i t i on i n c r e a s e s . (2) The s t rengths of i n t e r s p e c i f i c a s s o c i a t i o n and c o r r e l a t i o n should inc rease as p h y s i c a l s t r e s s , environmental 4 h e t e r o g e n e i t y , and i n t e r s p e c i f i c c o m p e t i t i o n i n c r e a s e . ( 3 ) L e v e l s o f p o l y p l o i d y s h o u l d be c o r r e l a t e d more w i t h t h e h i s t o r i c a l t h a n t h e p h y s i c a l a s p e c t s o f e n v i r o n m e n t a l r i g o r . ( 4 ) The c o m m u n i t i e s u n d e r t h e most s t r e s s s h o u l d h a v e t h e l e a s t i n t e r s p e c i f i c c o m p e t i t i o n f o r f l o w e r i n g t i m e , p o l l i n a t i o n v e c t o r s , a n d d i s p e r s a l a g e n t s . ( 5 ) a.) The most a b u n d a n t s p e c i e s w i t h i n a c o m m u n i t y s h o u l d be t h e most v a r i a b l e ( h a v e t h e l a r g e s t n i c h e ) . b. ) A v e r a g e w i t h i n - c o m m u n i t y n i c h e s i z e and p o p u l a t i o n v a r i a b i l i t y s h o u l d d e c r e a s e w i t h i n c r e a s i n g c o m p e t i t i o n and l o w e r s t r e s s . c. ) W i t h i n a c o m m u n i t y , e c o l o g i c a l d i s t i n c t i v e n e s s s h o u l d r e d u c e i n t e r s p e c i f i c c o m p e t i t i o n ; t h e c o m m u n i t i e s u n d e r t h e l e a s t s t r e s s s h o u l d h a v e t h e most e c o l o g i c a l l y d i s s i m i l a r s p e c i e s . ( 6 ) S p e c i e s d i v e r s i t y w i l l be g r e a t e s t , and d o m i n a n c e o f t h e c o m m u n i t y by r e l a t i v e l y few s p e c i e s w e a k e s t , i n t h e c o m m u n i t i e s o f t h e most b e n i g n p h y s i c a l e n v i r o n m e n t s . ( 7 ) The most d i v e r s e c o m m u n i t i e s w i l l be t h e m o s t s t a b l e . ( 8 ) C o m m u n i t i e s o f r i g o r o u s p h y s i c a l e n v i r o n m e n t s w i l l be composed o f s p e c i e s w i t h r e p r o d u c t i v e s p e c i a l i z a t i o n s t e n d i n g t o w a r d g e n e t i c u n i f o r m i t y . F i n a l l y , I s h o u l d a d d t h a t , i n t h e f o l l o w i n g e x p o s i t i o n , c o m p a r i s o n s h a v e u s u a l l y b e e n c o n f i n e d t o t h e f o u r c o m m u n i t i e s u n d e r s t u d y , i n a s m u c h as t h e r e i s l i t t l e o r no p r e c e d e n t f o r much o f t h e i n v e s t i g a t i o n . II. DESCRIPTION 5 A. M e t h o d s o f v e g e t a t i o n s a m p l i n g . F o u r s i t e s w e r e s a m p l e d d u r i n g t h e s p r i n g a n d summer o f 19 71. A s a l t m a r s h and two sphagnum b o g s ( l a b e l l e d Wade's Bog and Ogg's Bog) w e r e c h o s e n i n t h e v i c i n i t y o f T o f i n o , on t h e w e s t c o a s t o f V a n c o u v e r I s l a n d , B r i t i s h C o l u m b i a . B o t h bogs a r e l o c a t e d i n P a c i f i c R i m N a t i o n a l P a r k . A s u b a l p i n e meadow s i t e ( B l a c k w a l l Meadow) was c h o s e n i n M a n n i n g P r o v i n c i a l P a r k , p a r t o f t h e e x t e n s i v e meadow v e g e t a t i o n t h a t s t r e t c h e s n o r t h o f B l a c k w a l l P e a k f o r a b o u t 15 m i l e s t o N i c o m e n L a k e . The v e g e t a t i o n o f a p a r t i c u l a r s i t e i s h e r e a f t e r r e f e r r e d t o as a p l a n t c o m m u n i t y . No i m p l i c a t i o n as t o a c o m m u n i t y ' s o b j e c t i v e e x i s t e n c e as a d i s c r e t e v e g e t a t i o n a l u n i t i s i n t e n d e d , i n v i e w o f t h e w i d e s p r e a d c o n t r o v e r s y o v e r t h e c o m m u n i t y c o n c e p t . S i m i l a r l y , g r o u p i n g s o f s p e c i e s w i t h i n a c o mmunity may be t e r m e d a s s o c i a t i o n s , b u t w i t h t h e same r e s e r v a t i o n s . A l l c o m m u n i t i e s w e r e s a m p l e d w i t h q u a d r a t s l o c a t e d by 2 r andom number p a i r s . C e n t r a l l y n e s t e d w i t h i n e a c h 1 m q u a d r a t , 2 2 a 50K-50 cm ( 0 . 2 5 m ) and a 10x10 cm ( 0 . 0 1 m ) q u a d r a t w e r e a l s o u s e d . P r e s e n c e a n d c o v e r e s t i m a t e s w e r e r e c o r d e d f o r e a c h v a s c u l a r p l a n t s p e c i e s i n e a c h q u a d r a t . An i m p o r t a n c e v a l u e ( I . V . ) f o r e a c h s p e c i e s was c a l c u l a t e d a s t h e sum o f r e l a t i v e f r e q u e n c y and r e l a t i v e c o v e r ( K u r a m o t o and B l i s s 1 9 7 0 ) . I n a d d i t i o n , e a c h c o m m u n i t y was s a m p l e d by a c r i s s - c r o s s o f b e l t t r a n s e c t s 30.5 cm ( 1 f t ) w i d e . S p e c i e s o c c u r r e n c e i n c o n t i g u o u s 3 0.5 cm s q u a r e q u a d r a t s was r e c o r d e d a l o n g t h e s e t r a n s e c t s . The 6 sampling scheme i s summarized i n Table 1. TABLE 1. Summary of v e g e t a t i o n sampling. Dimensions of Number Length and Sum of sampling area of number of t r a n s e c t S i t e (m) a quadrats t r a n s e c t s (m) le n g t h (m) S a l t Marsh 244x61 70 183 (2) 457.5 30.5 (3) Wade's Bog 122x61 50 122 (1) 244 61 (2) Ogg's Bog 76x23 77 79.3 (1) 217 53x23 b 46 (1) 23 (4) B l a c k w a l l 122x61 50 122 (1) 244 Meadow 61 (1) aThe corresponding areas are a l l 8000-12000 m (0.8-1.2 ha). DTwo r e c t a n g u l a r areas at an angle to one another, with a common corner. A complete s p e c i e s l i s t f o r each s i t e can be found i n Appendix 1. Voucher specimens of each s p e c i e s are on f i l e i n the Herbarium o f the U n i v e r s i t y o f B r i t i s h Columbia. A l l s p e c i e s names and a u t h o r i t i e s are as i n Hi t c h c o c k et a l . (1955-1969), except where noted. B. Sampling r e s u l t s . 7 The number o f . s p e c i e s per quadrat f o r a l l quadrat s i z e s i s l i s t e d i n Table 2. An i n t e r e s t i n g fea ture o f these data i s tha t 2 at the sma l l e s t quadrat s i z e (0.01 m ) , the mean number of spec ies i s about the same f o r a l l four communities. This suggests tha t the lower l i m i t of the number o f species tha t can be packed i n t o a givensspace_isaapproximate.ly-.the--. same:: f o r the s a l t marsh, both sphagnum bogs, and the subalp ine meadow. The g rea te r o v e r a l l \" c a r r y i n g capac i t y \" of the suba lp ine meadow and bogs i s a f u n c t i o n of a g rea te r degree of .change i n species compos-i t i o n as quadrat s i z e i n c r e a s e s . Th i s change r e f l e c t s i n par t the g rea te r h a b i t a t and micro topographic he te rogene i ty o f the meadow and bogs. The s i t u a t i o n i s analogous (on a much sma l l e r s c a l e ) to the inc rease i n between-habi ta t or be ta species d i v e r s i t y tha t r e s u l t s from an inc reased change i n spec ies compos i t ion a long environmental g rad ien t s (Whi t taker 1969, 1970a, 1972). The mod i f i ed spec i e s - a r ea curves o f F igure 1 i n d i c a t e tha t 2 the 1 m quadrats a c c u r a t e l y es t imate the t o t a l spec ies compos i t ion o f the communit ies , s ince the curves l e v e l o f f 2 between the 0.45 and 1 m quadrat s i z e s . As quadrat s i z e i n c r e a s e s , the number of d i f f e r e n t species added w i t h s i z e becomes p r o g r e s s i v e l y lower . Spec ies -a rea curves u s u a l l y p l o t 8 TABLE 2. Mean number of species per quadrat f o r the various 2 quadrat s i z e s . The 1.00, 0.25, and 0.01 m quadrats 2 were randomly l o c a t e d ; the 0.45 and 0.09 m quadrats are t r a n s e c t segments. l x l 1.5x0.3 0.5x0.5 0.3x0.3 0.1x0.1 1.00 0.45 0.25 0.09 0.01 S a l t Marsh x = 7\\ 5 6.5 6.1 4.7 4.1 Wade's Bog 11. 5 11.2 9.5 7.6 4.7 Ogg's Bog 14. 4 13. 5 11.1 7.7 4.5 B l a c k w a l l 15.6 15.2 12 .1 8.8 4.6 Meadow Quadrat dimensions (m) Quadrat area Cm ) the t o t a l \"number of species v s . area (Cain 1938 ; Kershaw 1 9 6 4 ) ; I have chosen to p l o t average number of species vs. area as a more accurate i n d i c a t i o n of the comparative community taxonomic d i v e r s i t y . F i g . 1. M o d i f i e d s p e c i e s - a r e a c u r v e s . C. D e s c r i p t i o n o f communities. 10 S a l t marsh The f l o r a of the marsh i s sma l l (18 spec ies ) but v a s c u l a r p l an t cover averages about 8 8%. The vege t a t i on i s dominated ( i n the present d i s c u s s i o n , importance value i s taken to be a measure o f dominance) by grasses and g r a s s - l i k e herbs ( F i g s . 2 & 3 ) . The dominant species i n the s a l t marsh i s Desoh amp si a oespitosa ( F i g . 4) , which has the h ighes t average cover of any s p e c i e s , 28.1% (Table 3 ) . Trigloohin maritimum i s the commonest spec ies (100% frequency) but w i t h an average cover of 6.8% i s not as concent ra ted as Desohampsia. Sub-dominant species are F e s t u o a r u b r a v a r . l i t t o r a l i s , S a l i o o r n i a v i r g i n i o a 3 P l a n t a g o m a r i t i m a , J u n c u s b a l t i c u s , C a r e x l y n g b y e i , and G l a u x m a r i t i m a (Table 3 ) . TABLE 3. Frequency, mean cove r , and importance va lues o f the v a s c u l a r p l a n t spec ies of the Sa l t2Marsh . The values were c a l c u l a t e d from da\u00a3a from 1 m quadra ts ; values for the 0.25 and 0.01 m quadrats were a l s o c a l c u l a t e d but are not g iven here s i n c e they agree w e l l w i t h the f i g u r e s below. Species Frequency Mean cover Importance value (acronym) (%) (%) (I.V.) D e s o h a m p s i a o e s p i t o s a 85.7 28.1 38.6 (DCS) F e s t u o a r u b r a (FRU) 70.0 19.2 24.5 T r i g l o o h i n m a r i t i m u m 100.0 6.8 21.0 (TMA) TABLE 3. (Continued) 11 Species (acronym) Frequency Mean cover (%) (%) Importance value ( I . V . ) S a l i c o r n i a v i r g i n i c a 67. 1 11. 7 17 . 8 (SPA) P l a n t a g o m a r i t i m a (PMA) 61. 4 12. 6 16. 9 J u n c u s b a l t i c u s (JBA) 50. 0 17 . 9 16. 8 Carex lyngby.ei..(CLY) 74. 3 5. 3 14. 3 Glaux m a r i t i m a (GMA) 62. 8 7. 8 13. 9 Pot e n t i l l a p a c i f i c a (PPA) 27 . 1 11. 9 7. 2 A g v o s t i s e x a r a t a (AEX) 24. 3 14. 3 7. 1 S t e l l a r i a h u m i f u s a (SHU) 38. 6 2. 0 6 . 0 T r i f o l i u m w o r m s k j o l d i i 15. 7 8. 3 3 . 5 (TWO) S c i r p u s o e r n u u s (SCE) 24 . 3 0. 5 3. 3 P u e o i n e l l i a p u m i l a (PPU) 18 . 6 1. 2 2. 7 S p e r g u l a r i a c a n a d e n s i s 14. 3 0. 7 2. 0 (SCA) D i s t i e h l i s s p i o a t a (DSP) 10. 0 4. 6 1. 8 H o r d e u m b r a c h y a n t h e r u m 7. 1 10. 7 1. 8 (HBR) L i l a e o p s i s o c c i d e n t a l i s 2. 8 0. 7 0. 4 (LOC) T o t a l p l a n t cove r : 88.3% I know of no such d e s c r i p t i o n o f s a l t marsh vege t a t i on of the no r th P a c i f i c Nor th American coas t . Chapman (19 60) and MacDonald (1969) g ive cu r so ry reviews o f western Nor th American s a l t marshes. Hanson (19 61) has desc r ibed s a l t marsh v e g e t a t i o n a long the coas t o f western A l a s k a ; Ca lde r and T a y l o r (19 68) g i v e spec ies l i s t s and v e g e t a t i o n notes f o r the s a l t marshes o f the Queen C h a r l o t t e I s l a n d s . Johannessen (1961, 1964) and F r a n k l i n and Dyrness (1969) l i s t key species and g ive b r i e f e c o l o g i c a l resume's of s a l t marshes i n Washington and Oregon. 12 cu F i g . 2. S a l t Marsh, mid-May, 1971. F i g . 3. T i p of S a l t Marsh, e a r l y May, 1971, from 20 m up a S i t k a spruce. L i g h t - c o l o r e d t u f t s to the l e f t of the bleached l o g are o f Desohampsia oespitosa; dark green masses are o f Junaus b a l t i c u s . A long the margins of the mud f l a t s are c i r c u l a r clumps of Puooinellia p u m i l a 3 S a l i o o r n i a v i r g i n i o a 3 and T r i g l o o h i n m a r i t i m u m . 13a\/ F i g . 4. Desohampsia oespitosa, dominant s p e c i e s of the S a l t Marsh. F i g . 5. View from i n t e r i o r of Wade's Bog, e a r l y May, 1971. Stunted t r e e s are mostly Pinus oontovta; l i g h t - c o l o r e d sedge leaves belong mainly to Cavex obnupta. 14 C a l i f o r n i a ' s s a l t marshes have been more i n t e n s i v e l y s t u d i e d , most no tab ly by Purer (1942) , Hinde (1954) , Vogl (1966) , and Barbour and Davis (1970). By i n t e r p o l a t i o n , the Tof ino marsh seems s i m i l a r t o o ther h i g h s a l i n i t y s a l t marshes a long the nor th P a c i f i c coas t of Nor th Amer ica . U n l i k e s a l t marshes of eas te rn North America and the southern C a l i f o r n i a c o a s t , there i s no Spartina here to form the lowermost zone of v e g e t a t i o n . In the Tof ino a r e a , Cavex lyngbyei i s the commonest species at the lowermost l e v e l s , e s p e c i a l l y a long the banks o f dra inage channels . At the mud f l a t margins of the marsh, Salicornia virginica, Triglochin maritimum, and Puccinellia pumila form l a r g e , more or l e s s c i r c u l a r clumps 0.3 to 3.0 m i n diameter (F igure 3D.'' Johannessen (19 64) main ta ins tha t such clumps i n d i c a t e tha t the marsh i s expanding r a p i d l y . Sphagnum bogs Both bogs a r e , f o r the most p a r t , dominated by the same s p e c i e s , but to v a r y i n g degrees ( c f . Tables 4 and 5 ) . In g e n e r a l , the v e g e t a t i o n o f Ogg's Bog has a h igher p r o p o r t i o n o f woody spec ies than Wade's Bog (45% v s . 28%). Most n o t i c e a b l e are the inc reased dominances of Myrica gale ( I . V . of 36.1 v s . 4 . 9 ) , T h u j a p l i c a t a (16.2 v s . 3 . 4 ) , L i n n a e a b o r e a l i s (4.0 v s . 1 .2 ) , and Gaultheria shallon (3.3 v s . 0.2) i n Ogg's Bog. Vascu l a r p l a n t cover i s about 5 3% i n Ogg's Bog and 6 3% i n Wade's Bog. Bryophyte ground cove r , due p r i m a r i l y to Sphagnum s p p . , i s 60-70% and f a i r l y cont inuous i n both bogs, so tha t t o t a l p l an t 15 TABLE 4. Frequency, mean cove r , and importance va lues of the v a s c u l a r p l an t species of Wade's Bog. Species Frequency Mean cover Importance va lue (acronym) (%) (%) ( I . V . ) C a r e x p l u r i f l o r a (CPL) 96. 0 14. 3 30. 1 A p a r g i d i u m b o r e a l e (ABO) 92. 0 13. 2 27 . 2 A g r o s t i s a e q u i v a l v i s 98. 0 4. 8 15 . 9 (AAE) C a r e x o b n u p t a (GOB) 80 . 0 7. 0 15 . 9 S a n g u i s o r b a o f f i c i n a l i s 84. 0 5. 9 15 . 1 (SMI) K a l m i a p o l i f o l i a (KPO) 98. 0 3 . 6 14. 0 V a c c i n i u m o x y c o c c u s (VOX) 98. 0 3 . 1 13. 3 C a r e x c a n e s c e n s (CCA) 48. 0 10. 1 11. . 9 D r o s e r a r o t u n d i f o l i a 88. 0 2. 5 11. , 2 (DRO) Ledum g r o e n l a n d i c u m (LGR) 70. 0 2. 8 9. ,1 T r i e n t a l i s a r c t i c a (TAR) 96. 0 0. 4 8. , 9 E m p e t r u m n i g r u m (END 26. , 0 7. 1 5. , 2 M y r i c a g a l e (MGA) 28 . , 0 5. , 7 4. , 9 T o f i e l d i a g l u t i n o s a (TGL) 48 . , 0 0. , 5 4, . 6 T h u j a p l i c a t a (TPL) 22. , 0 4. , 3 3 . .4 P i n u s c o n t o r t a (PCO) 20. . 0 4. . 7 3 . . 2 G e n t i a n a d o u g l a s i a n a 14. . 0 0. .1 1, .2 (GDO) L i n n a e a b o r e a l i s (LBO) 12. . 0 1. ,1 1. . 2 M a i a n t h e m u m d i l a t a t u m 10, .0 0, .5 0, . 9 (MDI) R h y n c h o s p o r a a l b a (RAL) 6, . 0 2, . 5 0, . 7 V a c c i n i u m v i t i s - i d a e a 4, . 0 0, .5 0 . 4 (VVI) S c i r p u s c e s p i t o s u s (SCS) 2, . 0 5. . 0 0 . 3 J u n c u s s u p i n i f o r m i s (JOR) 2 . 0 1, . 0 0 . 2 B l e c h n u m s p i c a n t (BSP) 2. . 0 0 .5 0 . 2 V a c c i n i u m u l i g i n o s u m 2 . 0 0 . 5 0 . 2 (VUL) G a u l t h e r i a s h a l l o n (GSH) 2 . 0 0 .3 0 . 2 G e n t i a n a s c e p t r u m (GSC) 2 . 0 0 . 3 0 . 2 C o p t i s a s p l e n i f o l i a (CAS) 2 . 0 0 .1 0 . 2 T o t a l p l an t cover : 63.1% 16 TABLE 5. Frequency, mean cover , and importance va lues of the v a s c u l a r p l a n t species of Ogg's Bog. Species Frequency Mean cover Importance va lue (acronym) (%) (%) ( I . V . ) Myvioa g a l e (MGA) 88 . 3 18. 1 36. 1 A p a v g i d i u m b o v e a l e (ABO) 77 . 9 8. 9 18. 4 T h u j a p l i c a t a (TPL) 70. 1 8. 6 16 . 2 C a v e x o b n u p t a (COB) 76. 6 5. 8 13. 7 C a v e x p l u v i f l o v a (CPL) 85. 7 3. 5 11. 6 A g v o s t i s a e q u i v a l v i s 79. 2 3. 5 10 . 7 (AAE) S a n g u i s o v b a o f f i c i n a l i s 93. 5 1. 6 9. 3 (SMI) Ledum g v o e n l a n d i o u m (LGR) 76. 6 1. 9 8. 1 V a o o i n i u m o x y o o o o u s (VOX) 81. 8 1. 2 7 . 5 D r o s e v a v o t u n d i f o l i a 76. 6 0. 6 6 . 1 (DRO) K a l m i a p o l i f o l i a (KPO) 57 . 1 1. 8 5 . 8 Empetvum n i g v u m (END 37 . 7 4. 1 5 . 5 T v i e n t a l i s a v c t i o a (TAR) 74. 0 0. 2 5 . 4 C a v e x o a n e s o e n s (CCA) 41. 5 2. ,7 5 . 0 T o f i e l d i a g l u t i n o s a (TGL) 61. 0 0. ,4 4 . , 7 L i n n a e a b o v e a l i s (LBO) 45 . ,4 1. , 0 4. , 0 R h y n o h o s p o v a a l b a (RAL) 23. .4 5. . 2 3 . , 9 P l a n t a g o m a c v o o a v p a (PMC) 37. ,7 1. , 7 3 . , 8 G a u l t h e v i a s h a l l o n (GSH) 37 . ,7 0. . 9 3 . . 3 B l e o h n u m s p i c a n t (BSP) 23. .4 1. . 0 2. . 6 G e n t i a n a s o e p t v u m (GSC) 33. . 8 0, .4 2 , . 5 C a l a m a g v o s t i s n u t k a e n s i s 18. . 2 2. . 3 2. . 0 (CNU) . 0 M a i a n t h e m u m d i l a t a t u m 27 , . 3 0, .2 2, (MDI) S c i v p u s o e s p i t o s u s (SCS) 15. .6 1 . 9 1 . 6 C o p t i s a s p l e n i f o l i a (CAS) 14 . 3 1 . 5 1 . 4 C o p t i s t v i f o l i a (CTR) 16 .9 0 .7 1 . 4 J u n o u s s u p i n i f o v m i s (JOR) 7 . 8 3 . 6 1 . 1 D e s o h a m p s i a o e s p i t o s a 9 .1 1 . 8 0 . 9 (DCS) 0 P i n u s o o n t o v t a (PCO) 7 . 8 2 .1 . 8 L y c o p o d i u m o l a v a t u m ( L C D 6 .5 1 . 3 0 . 6 G e n t i a n a d o u g l a s i a n a 7 . 8 0 . 3 0 . 6 (GDO) 0 V a o o i n i u m o v a t u m (VOV) 6 . 5 1 . 0 . 6 V a o o i n i u m v i t i s - i d a e a 5 . 2 0 . 6 0 .4 (VVI) 0 'Cornus- u n a l a s o h k e . n s i s 3 . 9 1 . 5 . 4 (CUN) 17 TABLE 5. CContinued) Species Frequency (acronym) (%) Mean cover (%) Importance value ( I . V . ) Vaccinium uliginosum 3.9 0.9 0 . 3 (VUL) 0 . 3 Juniperus communis (JCO) 2 . 6 3.0 Nephrophyllidium crista- 2 . 6 2.0 0 . 3 galli (FCG) 0 . 2 Lycopodium inundatum 2.6 1.1 (LIN) 0.1 Eriophorum polystachion 1.3 2.0 (EPO) Cavex pauciflora (CPA) 1.3 2.0 0.1 Lysichitum americanum 1.3 0.3 0.1 (LAM) T o t a l p l an t cover : 5 3.4% cover exceedstlO0%. Evergreen shrubs and h a l f - s h r u b s , cyperaceous spec ies and the sphagnum moss g ive both bogs t h e i r c h a r a c t e r i s t i c physiognomy ( F i g . 5 ) . Wade (1965) has p r e v i o u s l y c l a s s i f i e d the vege t a t i on of Wade's Bog. The sampling area i n t h i s study would f a l l most ly w i t h i n the Oxycoccus - Sphagnum papillosum and Carex pluriflora a s s o c i a t i o n s as mapped by Wade (1965, F i g 6 ) . Ogg's Bog con ta ins more of the Myrica gale and Vaccinium vitis-idaea v a r i a n t s and both sampling areas have very l i t t l e of the Scirpus - Sphagnum mendocinum a s s o c i a t i o n . Rigg (1925, 1940) , Osvald (1933) , Hansen (1947) , and Heusser (1960) have desc r ibed the vege t a t i on and sketched the h i s t o r y of a number o f sphagnum bogs o f no r th P a c i f i c Nor th Amer ica . A p p l i c a t i o n o f genera l p r i n c i p l e s o f bog development 18 o u t l i n e d by Gorham (1957) and Heinselman (1963) to the d e s c r i p -t i o n s of the above authors and to my own and Wade's observa t ions suggests tha t the two study bogs, and bogs i n genera l i n the T o f i n o - U c l u e l e t a r ea , are s u c c e s s i o n a l l y j u v e n i l e . The very shal low peat depths (averaging no more than 1 m) support t h i s sugges t ion , as does the y o u t h f u l age o f Wade's Bog, e s t a b l i s h e d as approximately 400190 years by rad ioca rbon d a t i n g (Wade 1965). The bogs probably have been formed by p a l u d i f i c a t i o n (swamping) o f p o o r l y d ra ined c o a s t a l f o r e s t , r a t h e r than by g r a d u a l , c e n t r i p e t a l f i l l i n g - i n o f water bod ie s . Wade (19 65) cons idered the recent format ion o f a h ighe r l i p o f land now covered by S i t k a spruce (Piaea sitchensis) and the format ion of an u n d e r l y i n g hardpan to be the l i k e l i e s t b a r r i e r s to the drainage of the c o a s t a l t e r r a c e upon which the bogs are perched. Success ion i n the bogs appears to be toward development o f a h i g h moor or r a i s e d bog o f sphagnum moss, sedges , and e r icaceous shrubs w i t h f u r t h e r bogaexpansion at the expense o f the surrounding wet f o r e s t . Of the two bogs, Wade's Bog seems to be more advanced i n t h i s succes s ion . The average peat depth i n Wade's Bog i s 1 m (Wade 19 65) ; i n Ogg's Bog, 0.5 m. Average t r e e cover i s a l s o l e s s i n Wade's Bog, as are the I . V . ' s o f species such as L i n n a e a b o v e a l i s C o r n u s u n a l a s c h k e n s i s 3 G a u l -t h e r i a s h a l l o n 3 and L y a o p o d i u m a l a v a t u m , which are more t y p i c a l o f f o r e s t undergrowth than sphagnum bogs. Furthermore, the dominat ion of Ogg's Bog by Myriaa gale (Table 5) i s very s i g n i f i c a n t , s ince Myriaa grows p o o r l y i n h i g h l y a c i d i c h i g h moor c o n d i t i o n s ( K r a j i n a , pe r sona l communication). Subalpine meadow 19 The f l o r a of B l a c k w a l l Meadow i s the r i c h e s t of the four communit ies , w i t h 45 spec ies present i n the sample. The v e g e t a t i o n i s co-dominated by three s p e c i e s : Valeriana sitch-e n s i s ( I . V . of 2 4 . 9 ) , F e s t u o a v i r i d u l a ( 2 4 . 3 ) , and L u p i n u s l a t i f o l i u s ( 2 1 . 8 ) . E r i g e r o n p e r e g r i n u s . Anemone o c c i d e n t a l i s m and Potentilla flabellifolia are secondary co-dominants (Table 6) . At the he igh t of the shor t growing season, the community c o n s i s t s :of -.a,, lushc mixture \u201eof _ showy-flowered forbscand more inconspicuous g ra s se s , sedges, and wood rushes ( F i g s . 6 S 7 ) . Vege ta t i on cover approximates 93%; the bare s o i l areas are due main ly to the a c t i v i t i e s of a sma l l pocket gopher tha t tunne l s below the snow dur ing the w i n t e r , j u s t beneath the s o i l su r face . In con t r a s t to many o ther mountainous a reas , the subalpine vege t a t i on o f nor th P a c i f i c Nor th America has been l i t t l e s tud ied by p l an t e c o l o g i s t s u n t i l recen t yea r s . Fonda and B l i s s (1969) and Kuramoto and B l i s s (1970) have desc r ibed communities i n the suba lp ine of the Olympic Mountains of Washington. B r i n k (1959, 1964), Arche r (1963), and Brooke, al. (1970) have s t ud i ed the suba lp ine v e g e t a t i o n of the southwestern Coast Range of B r i t i s h Columbia. Eady (1971) d i scussed the a l p i n e - s u b a l p i n e zone o f B ig White Mounta in , i n the Okanagan Highland of southern B r i t i s h Columbia. Douglas (1972) has desc r ibed i n d e t a i l the suba lp ine p lan t communities of the western Nor th Cascades, Washington. Comparison o f these s tud ie s leads me to conclude tha t the B l a c k w a l l Meadow vege ta t i on most c l o s e l y resembles the 20 ou F i g . 6. B l a c k w a l l Meadow, e a r l y August , 19 72. F i g . 7. B l a c k w a l l Meadow, e a r l y Augus t , 19 72. In both p i c t u r e s , the whi te c o l o r i s p r i m a r i l y due to i n f l o r e s c e n c e s o f V a l e r i a n a sitchensis and ^mop-tops\" or seed heads of Anemone Occidentalis ; the b l u e , to L u p i n u s l a t i f o l i u s ; the y e l l o w , to P o t e n t i l l a f l a b e l l i f o l i a 3 A r n i c a l a t i f o l i a 3 and A r n i c a m o l l i s . 21 d r i e r V a l e r i a n a s i t c h e n s i s - C a s t i l l e j a e l m e r i a s s o c i a t i o n of Eady (1971) , w i t h some a f f i n i t i e s to the mois te r V. sitchensis - Veratrum viride a s s o c i a t i o n of Douglas (1972). Th i s i n t e r -mediate c o n d i t i o n . i s to be expected , s ince B l a c k w a l l Meadow i s s i t u a t e d on the gen t l e west s lope of a nor th - sou th r i d g e a long the phys iog raph ic break between the east and west f l a n k s of the Cascade Range i n the Manning Park a rea . TABLE 6. Frequency, mean cover , and importance va lues o f the v a s c u l a r p l an t spec ies of B l a c k w a l l Meadow. Species Frequency Mean cover Importance va lue (acronym) (%) <%) ( I . V . ) V a l e r i a n a s i t c h e n s i s 76.0 24.6 24.9 (VSI) F e s t u c a v i r i d u l a (FVI) 100.0 16.7 24.3 L u p i n u s l a t i f o l i u s (LLA) 96.0 14.6 21.8 E r i g e r o n p e r e g r i n u s (EPE) 86.0 7.5 12.4 Anemone o c c i d e n t a l i s 80.0 8.0 11.9 (AOC) P o t e n t i l l a f l a b e l l i f o l i a 90.0 4.8 10.4 (PFL) E r y t h r o n i u m g r a n d i f l o r u m 7 0.0 2.0 7.0 (EGR) V a c c i n i u m s c o p a r i u m (VSC) 42.0 9.3 6.9 C l a y t o n i a l a n c e o l a t a 82.0 1.5 6.5 (CLA) A r e n a r i a c a p i l l a r i s (ACA) 72.0 2.2 6.3 A n t e n n a r i a l a n a t a (ALA) 54.0 4.8 6.2 V e r o n i c a c u s i c k i i (VCU) 54.0 3.1 5.3 A g o s e r i s a u r a n t i a c a (AAU) 60.0 1.5 4.8 P h l e u m a l p i n u m (PAL) 64.0 1.0 4.8 A r n i c a l a t i f o l i a (ALF) 50.0 2.7 4.7 L u z u l a h i t c h c o c k i i (LHI) 56.0 1.8 4.6 T h a l i c t r u m o c c i d e n t a l e 14.0 18.6 3.7 (TOO A c h i l l e a m i l l e f o l i u m 42.0 1.7 3.5 (AMI) T r i s e t u m s p i c a t u m (TSP) 44.0 0.8 3.2 22 TABLE 6. (Continued) Species Frequency Mean cover Importance va lue (acronym) (%) (%> ( I . V . ) C a r e x r o s s i i (CRO) E l y m u s g l a u o u s (EGL) S i l e n e p a r r y i (SPA) A r n i c a m o l l i s (AMO) P e n s t e m o n p r o c e r u s (PPR) P o a c u s i c k i i (PEP) S e n e c i o i n t e g e r r i m u s (SIN) E i e r a c i u m g r a c i l e (HGR) L u z u l a s p i c a t a (LSP) Sedum l a n c e o l a t u m (SLA) C a s t i l l e j a m i n i a t a (CMI) P e d i c u l a r i s b r a c t e o s a (PBR) P h l o x d i f f u s a (PDI) P o t e n t i l l a d i v e r si f o l i a (PDV) C a r e x s p e c t a b i l i s (CSP) E p i l o b i u m a l p i n u m (EAL) D e l p h i n i u m n u t t a l l i a n u m (DNU) C a s t i l l e j a p a r v i f l o r a (CAL) R a n u n c u l u s e s c h s c h o l t z i i (RES) S i b b a l d i a p r o c u m b e n s (SPI M i c r o s t e r i s g r a c i l i s (MGR) J u n c u s d r u m m o n d i i (JDR) H y d r o p h y l l u m f e n d l e r i (HFE) S e n e c i o t r i a n g u l a r i s (STR) V a c c i n i u m d e l i c i o s u m (VDE) S e l a g i n e l l a w a l l a c e i (SWA) T o t a l p l a n t cover : 9 3.3% 44. 0 0 . 5 2.8 10 . 0 17 . 6 2.5 28.0 1.5 2.2 22.0 2.8 2.1 24. 0 2.3 2.1 26.0 1.0 1.9 24. 0 1.4 1.9 20.0 1.9 1.7 20.0 0.4 1.4 18. 0 0 . 5 1.2 16 . 0 1.1 1.2 12. 0 1.0 0.9 .-8:0 3.8 0.8 \" 8 o 2.4 0.7 1 4v0 8.5 0.6 8.0 0 . 3 0.5 6.0 2.0 0.5 6 . 0 0.8 0.4 6.0 0.6 0.4 ) 4.0 1.3 0. 3 4.0 0.7 0.3 2.0 3.0 0.2 2.0 3 . 0 0.2 2.0 2.0 0.2 2.0 2.0 0.2 2.0 0.5 0.1 I I I . THEORY AND RESULTS 23 A . P o p u l a t i o n s t r u c t u r e of i n d i v i d u a l s p e c i e s : ,D\/d index o f aggrega t ion . The d i s t r i b u t i o n o f i n d i v i d u a l s of the same species i n a p a r t i c u l a r p l a n t community g e n e r a l l y i s non-random or aggregated (Gre ig -Smi th 1964; Kershaw 1964) , even i n apparen t ly homo-geneous v e g e t a t i o n . An aggregated p o p u l a t i o n s t r u c t u r e can r e s u l t from environmental he te rogene i ty o r mosa ic i sm, i n t e r -s p e c i f i c i n t e r a c t i o n s , and spec ies morphology (vege t a t i ve apomixis or v i c i n i s m i n seed d i s p e r s a l can l ead to clumped d i s t r i b u t i o n s ) . The non-randomness can take the form of a clumped (\"contagious\") d i s t r i b u t i o n of i n d i v i d u a l s , or an even ( \" r egu la r \" ) d i s t r i b u t i o n (Kershaw 1964). One measure of aggregat ion i s the D\/d index of McGinnies ( i n C u r t i s and Mcin tosh 1950). I have c a l c u l a t e d t h i s index f o r each species i n the present study from presence-absence da ta f o r 'l-i.^-m- 'and\" 0; 9-m segments o f the 0.3-m wide b e l t t r a n s e c t s , a f t e r the method o f Smith and Cottam (1967). The a c t u a l d e n s i t y (D) was c a l c u l a t e d by c o n s i d e r i n g each 0.3-m square w i t h i n each 1.5- or 0.9-m segment to be a p o t e n t i a l i n d i v i d u a l ; t h u s , the maximum d e n s i t y p o s s i b l e per 1.5-m segment was 5, and per 0.9-m segment 3. Frequency va lues were computed as the percentage o f a l l p o s s i b l e 1.5- or 0.9-m segments occupied by a p a r t i c u l a r s p e c i e s . The expected d e n s i t y (d) va lues cou ld then be determined from a b i n o m i a l curve (see Gre ig -Smi th 1964, Table 6 ) . The d va lue thus obta ined was the d e n s i t y (number o f 0.3-m 24 quadrats occupied per segment) t ha t would be expected i f the p o p u l a t i o n were a random one. U s u a l l y the expected dens i t y was l e s s than the a c t u a l d e n s i t y , g i v i n g a D\/d value g rea te r than 1 and i n d i c a t i n g aggregat ion (Table 7 ) . The s i z e o f the D\/d value g ives some i n d i c a t i o n of the type and degree of aggreg-a t i o n . A D\/d grea te r than 1 i n d i c a t e s a contagious d i s t r i b u t i o n ; a D\/d l e s s than 1 i n d i c a t e s a r e g u l a r d i s t r i b u t i o n . The on ly species to show markedly r e g u l a r d i s t r i b u t i o n s (both v i a 0.9-m segments) were T v i g l o c h i n m a v i t i m u m and F e s t u o a v i r i d u l a , the commonest spec ies i n the s a l t marsh and suba lp ine meadow, r e s p e c t i v e l y . U n f o r t u n a t e l y , there i s no way to determine the s i g n i f i c a n c e o f the D\/d index ' d e v i a t i o n from u n i t y (Beals 1968). I t has been suggested (Gre ig -Smi th 1964) tha t success ion i n vege t a t i on i s accompanied by an inc rease i n the s i z e of spec ies aggregat ions concomitant w i t h a decrease i n the i n t e n s i t y o f aggrega t ion . One would expec t , t h e n , e a r l y s u c c e s s i o n a l vege t a t i on to be composed of spec ies w i t h , on the average, h ighe r D\/d values than spec ies o f more advanced s u c c e s s i o n a l or c l imax communities. Regular d i s t r i b u t i o n s r e f l e c t e d by lower D \/ d ' s are sugges t ive o f a n e g a t i v e , probably compe t i t i ve i n t e r -a c t i o n between s p e c i e s , and would be expected i n more s t a b l e , c l imax communities (Beals 19 68) . To compare the average degree of non-randomness i n the study communities I have used the r a t i o o f the sum of observed d e n s i t i e s to the sum of expected d e n s i t i e s ( i . e . , Ed , \/ _ d ) , as recommended by C u r t i s and Mcin tosh (1950). Both observed and expected d e n s i t i e s are the same as those used i n the c a l c u l a t i o n of the D\/d index . Note tha t t h i s r a t i o i s weighted i n favor of 25 the more abundant s p e c i e s , which i s an a t t r a c t i v e f ea tu re . R a t i o s fo r both segment s i z e s f o r a l l four communities are presented i n Table 8. TABLE 7. D\/d index of aggrega t ion . Species D\/d (0.9-m segments) D\/d (1.5-m segments) S a l t Marsh D e s o h a m p s i a o e s p i t o s a F e s t u o a r u b r a T r i g l o o h i n m a r i t i m u m S a l i o o r n i a v i r g i n i o a P l a n t a g o m a r i t i m a J u n o u s b a l t i o u s G l a u x m a r i t i m a P o t e n t i l l a p a o i f i o a A g r o s t i s e x a r a t a S t e l l a r i a h u m i f u s a T r i f o l i u m w o r m s k j o l d i i S c i r p u s c e r n u u s P u o o i n e l l i a p u m i l a S p e r g u l a r i a c a n a d e n s i s D i s t i o h l i s s p i c a t a H o r d e u m b r a o h y a n t h e r u m L i l a e o p s i s o c o i d e n t a l i s 1.80 1.98 0. 77 1. 87 1. 2. 1. 1. 1, 1. 2. 1. 60 22 93 67 78 86 11 62 1. 55 72 53 08 1.39 mean 65 77 11 52 , 03 ,46 ,53 , 07 ,42 ,46 ,83 1.90 1.93 2.26 3.97 3.10 1.88 C a r e x p l u r i f l o r a A p a r g i d i u m b o r e a l e A g r o s t i s a e q u i v a l v i s C a r e x o b n u p t a S a n g u i s o r b a o f f i c i n a l i s K a l m i a p o l i f o l i a V a o o i n i u m o x y o o o o u s C a r e x c a n e s o e n s D r o s e r a r o t u n d i f o l i a Ledum g r o e n l a n d i o u m - 1. 76 mean = 2 .77 - 2.53 range: 1.11 \u2022 g 1.00 1. 31 1.29 1.61 0.92 1. 21 1.23 1.67 1.13 1.53 1.13 1.38 0.98 1. 24 1.47 1.62 1.03 1.38 1.29 1.74 TABLE 7. (Continued) 26 Species D\/d (0.9-m segments) D\/d (1.5-m segments) T r i e n t a l i s a r c t i o a 1 .16 1 . 53 E m p e t r u m n i g r u m 1 .63 1 .93 M y r i o a g a l e 1 .67 2 .19 T o f i e l d i a g l u t i n o s a 1 .15 1 .29 T h u j a p l i c a t a 1 . 63 1 . 81 P i n u s e o n t o r t a 1 . 39 1 . 82 G e n t i a n a d o u g l a s i a n a 1 .28 1 .29 L i n n a e a b o r e a l i s 1 .96 1 . 97 M a i a n t h e m u m d i l a t a t u m 1 .50 1 .60 R h y n o h o s p o r a a l b a 1 . 57 2 . 71 J u n o u s s u p i n i f o r m i s 2 . 33 2 .67 C o p t i s t r i f o l i a 1 .0 0 1 .11 mean = 1.35 mean = 1 range: 0. 92 - 2.33 range: 1. 11 Ogg's Bog M y r i o a g a l e 0. 96 1. 18 A p a r g i d i u m b o r e a l e 1. 20 1. 71 T h u j a p l i o a t a 1. 35 1. 81 C a r e x o b n u p t a 1. 50 2. 02 Carex p l u r i f l o r a 1. 10 1. 47 A g r o s t i s a e q u i v a l v i s 1. ,30 1. 62 S a n g u i s o r b a o f f i c i n a l i s 1. 13 1. ,47 Ledum g r o e n l a n d i c u m 1. ,26 1. 65 V a o o i n i u m o x y o o c o u s 1. ,43 1. , 83 D r o s e r a r o t u n d i f o l i a 1. ,38 1. ,79 K a l m i a p o l i f o l i a 1. . 27 1. ,60 E m p e t r u m n i g r u m 1. ,75 2. ,16 T r i e n t a l i s a r c t i o a 1. ,42 2. , 07 C a r e x c a n e s c e n s 1. ,41 1. ,69 T o f i e l d i a g l u t i n o s a 1. ,25 1. ,53 L i n n a e a b o r e a l i s 1. , 53 2. ,02 R h y n o h o s p o r a a l b a 2. .02 2. .85 P l a n t a g o m a c r o c a r p a 1. .75 2. .14 G a u l t h e r i a s h a l l o n 1. , 3 6 1. ,70 B l e c h n u m s p i o a n t 1. .37 1. .60 G e n t i a n a s o e p t r u m 1, .39 1. .40 C a l a m a g r o s t i s n u t k a e n s i s 1, .71 2, . 08 M a i a n t h e m u m d i l a t a t u m 1. .45 1, .77 S o i r p u s c e s p i t o s u s 1, . 59 1, .92 C o p t i s a s p l e n i f o l i a 1, . 98 2, .73 C o p t i s t r i f o l i a 1, .86 2, .07 J u n o u s s u p i n i f o r m i s 2, . 07 2, .90 D e s o h a m p s i a o e s p i t o s a 1, .33 1, . 94 TABLE 7. (Continued) 27 Species D\/d (0.9-m segments) D\/d (1.5-m segments) P i n u s c o n t o r t a L y c o p o d i u m c l a v a t u m G e n t i a n a d o u g l a s i a n a V a o o i n i u m o v a t u m V a o o i n i u m v i t i s - i d a e a V a o o i n i u m u l i g i n o s u m J u n i p e r u s c o m m u n i s 1.43 1.79 1.64 1.28 2.04 1.90 1.28 63 76 13 mean 1.50 range: 0.9 6 - 2.0 7 B l a c k w a l l Meadow 1. 31 2. 58 1. 94 1.68 mean = 1.88 range: 1.18 - 2.9 0 V a l e r i a n a s i t c h e n s i s 1. 80 2. 05 F e s t u c a v i r i d u l a 0. 63 1. 15 L u p i n u s l a t i f o l i u s 1. 12 1. 36 E r i g e r o n p e r e g r i n u s 1. 03 1. 46 Anemone o o c i d e n t a l i s 1. 30 1. 68 P o t e n t i l l a f l a b e l l i f o l i a 0. 97 1. 37 E r y t h r o n i u m g r a n d i f l o r u m 1. 19 1. 53 V a o o i n i u m s c o p a r i u m 1. 71 2. 29 C l a y t o n i a l a n c e o l a t a 1. 28 1. 82 A r e n a r i a o a p i l l a r i s 1. 55 2. 28 A n t e n n a r i a l a n a t a 1. 52 2. 04 V e r o n i c a c u s i c k i i 1. 37 1. 86 A g o s e r i s a u r a n t i a c a 1. 06 1. ,16 P h l e u m a l p i n u m 1. .01 1. 08 A r n i c a l a t i f o l i a 1. ,30 1. , 37 L u z u l a h i t c h c o c k i i 1. ,32 1. ,67 T h a l i c t r u m o c c i d e n t a l e 1. ,88 2. , 74 A c h i l l e a m i l l e f o l i u m 1. , 35 1. , 68 T r i s e t u m s p i c a t u m 1. .15 1. , 29 C a r e x r o s s i i 1. .29 1. ,42 E l y m u s g l a u c u s 1, .93 3. . 09 S i l e n e p a r r y i 1, .23 1, .25 A r n i c a m o l l i s 1. .60 2, . 53 P o a c u s i c k i i 1. . 01 1, .11 S e n e c i o i n t e g e r r i m u s 1 .30 1, . 53 H i e r a c i u m g r a c i l e 1. .70 1. .96 L u z u l a s p i c a t a 1 .16 1 . 24 Sedum l a n c e o l a t u m 1 .25 1 .48 C a s t i l l e j a m i n i a t a 1 .25 1 .37 P e n s t e m o n p r o c e r u s 1. . 55 2 .07 P e d i c u l a r i s b r a c t e o s a 1 . 58 1 . 52 P o t e n t i l l a d i v e r s i f o l i a 1 .40 1 . 53 TABLE 7. (Continued) 28 Species D\/d (0.9-m segments) D\/d (1.5-m segments) E p i l o b i u m a l p i n u m 1.69 1.65 R a n u n c u l u s e s c h s c h o l t s i i 1.31 1.62 S i b b a l d i a p r o c u m b e n s 1.62 2.08 M i o r o s t e r i s g r a c i l i s 2.00 2.08 S e n e c i o t r i a n g u l a r i s 1.65 2.28 V a c c i n i u m d e l i c i o s u m 1.90 3.07 S e l a g i n e l l a w a l l a c e i 2.06 1.79 V e r a t r u m v i r i d e 1.69 2.13 mean = 1.4 2 mean = 1.7 7 range: 0.6 3 - 2.0 6 range: 1.08 - 3.0 9 Wade's Bog has the l e a s t aggregated v e g e t a t i o n (Table 8 ) . Th i s i s expected , s ince t h i s bog appears to be nearest the c l imax s t a t e f o r the c o a s t a l sphagnum bog community t ype . At the o ther extreme, the S a l t Marsh has by f a r the h ighes t average D\/d v a l u e . Perhaps t h i s i s because the r i g o r o u s p h y s i c a l environment of the s a l t marsh imposes f a i r l y d i s c r e t e h a b i t a t zona t ion (promoting aggregat ion) and p laces a premium on mere s u r v i v a l , d i m i n i s h i n g i n t e r s p e c i f i c compe t i t i ve e f f e c t s , e s p e c i a l l y i n the p ioneer mud f l a t v e g e t a t i o n ( c f . C l a rke and Hannon 1970, 1971). I t cou ld a l s o be due i n par t t o the morphology o f the marsh s p e c i e s , many o f which have ex tens ive vege t a t i ve spread. Ogg's Bog has r e l a t i v e l y s t r o n g l y aggregated v e g e t a t i o n , w h i l e tha t of B l a c k w a l l Meadow i s on ly weakly aggregated. Both communities have d i v e r s e , m o s a i c - l i k e m i c r o -topograph ies . The fac t tha t Ogg's Bog has a h igher average D\/d 29 than B l a c k w a l l Meadow r e f l e c t s , I t h i n k , a much h igher degree of i n t e r s p e c i f i c compe t i t i on i n the suba lp ine meadow (see Sec t . I I I - B ) and probable advanced s u c c e s s i o n a l ' s t a t u s . TABLE 8. Weighted average community D\/d v a l u e s . a 0.9-m segments, 1.5-m segments, S i t e r a t i o = r a t i o = S a l t Marsh 1.39 1.97 Wade's Bog 1.12 1.44 Ogg's Bog 1.25 1.69 B l a c k w a l l Meadow 1.14 1.53 A s i m p l e , unweighted average was a l s o computed f o r each community, and the r e s u l t s were e q u i v a l e n t . However, i n a l l cases the weighted average was l e s s than the unweighted, i n d i c a t i n g tha t the r a r e r spec ies had h igher D \/ d ' s (were more aggregated) , which seems reasonable . C lose r examinat ion o f Table 8 r e v e a l s tha t a l l four communities e x h i b i t more in tense aggrega t ion at 1.5-m i n t e r v a l s , as would be expected merely from the inc rease i n i n t e r v a l l eng th from 0.9 m. However, the change i n segment s i z e does not a f f e c t a l l s i t e s e q u a l l y . In p a r t i c u l a r , the s a l t marsh species show a compara t ive ly g rea te r inc rease i n aggrega t ion at the 1.5-m l e v e l . T h i s may be i n d i c a t i v e of a secondary sca l e of p a t t e r n i n the S a l t Marsh. That i s , i n d i v i d u a l s o f the same spec ies tend to occur toge ther w i t h i n a r a d i u s of at l e a s t 0.9 m i n a l l communit ies; i n Ogg's Bog and the subalp ine meadow, and e s p e c i a l l y i n the s a l t marsh, there i s a tendency fo r the 30 i n d i v i d u a l s to form l a r g e r clumps. Th i s i s i n agreement w i t h B r e r e t o n ' s (19 71) f i n d i n g s t h a t , at l e a s t i n s a l t marsh s u c c e s s i o n , spec ies are i n i t i a l l y randomly d i s t r i b u t e d but l a t e r become more aggregated and e v e n t u a l l y the aggregat ions inc rease i n s i z e . B. I n t e r s p e c i f i c r e l a t i o n s h i p s . A s s o c i a t i o n I n t e r s p e c i f i c a s s o c i a t i o n can be detec ted by c o n s t r u c t i o n 2 of a 2x2 cont ingency t a b l e and c a l c u l a t i o n of x va lues based on j o i n t occurrence of p a r t i c u l a r spec ies p a i r s i n sample quadrats (Kershaw 1964). T h i s has been done f o r a l l p o s s i b l e species p a i r s i n each community, p rov ided each spec ies occur red 2 i n at l e a s t f i v e sampling u n i t s (see Appendix 2 ) . The x va lue shows whether the d i f f e r e n c e between the observed and expected number of quadrats o f co-occurrence i s s i g n i f i c a n t o r no t . The 2 s i g n of the x va lue i n d i c a t e s whether the a s s o c i a t i o n i s p o s i t i v e or n e g a t i v e . 2 Since c a l c u l a t i o n o f x \u00b1 s based on q u a l i t a t i v e presence\/ absence da t a , the type and l e v e l of a s s o c i a t i o n i s dependent on 2 quadrat s i z e (Gre ig -Smi th 1964; Kershaw 19 64) . Therefore x values have been c a l c u l a t e d f o r f i v e d i f f e r e n t quadrat s i z e s the three random quadrat s i z e s and the 1.5- and 0.9-m t r ansec t segments. I t i s hoped tha t t h i s approach w i l l more v a l i d l y detect t rends o f a s s o c i a t i o n and i n d i c a t e s ca l e s o f p a t t e r n i n the communit ies. 31 2 In most cases , the r e s u l t s o f the x t e s t s are c o n s i s t e n t throughout the d i f f e r e n t quadrat s i z e s . Occas iona l changes i n s i g n from one quadrat s i z e to another may be viewed as e f f ec t s o f quadrat s i z e and\/or number. C o l e ' s index of i n t e r s p e c i f i c a s s o c i a t i o n (Cole 1949; H u r l b e r t 19 69) was a l s o determined fo r a l l spec ies p a i r s . C o l e ' s 2 index g ives e s s e n t i a l l y the same i n f o r m a t i o n as a x value but reduces the values to a range from +1 to - 1 . These r e s u l t s are a l so presented i n Appendix 2. A s s o c i a t i o n as a s t a t i s t i c can suggest p o s s i b l e causes of s p e c i e s ' d i s t r i b u t i o n a l p a t t e r n s . H i g h l y nega t ive a s s o c i a t i o n s may i n d i c a t e d i s s i m i l a r h a b i t a t requirements o r compe t i t i ve e x c l u s i o n or a l l e l o p a t h y . H i g h l y p o s i t i v e a s s o c i a t i o n s may i n d i c a t e s i m i l a r h a b i t a t requi rements , absence of s t rong c o m p e t i t i o n , or environmental m o d i f i c a t i o n by one species to the advantage of another (Kershaw 1964; Smith and Cottam 1967; Gooda l l 1970). Byer (1970) main ta ins tha t s t rong p o s i t i v e and negat ive a s s o c i a t i o n i s c h a r a c t e r i s t i c o f the vege t a t i on o f heterogene'ous h a b i t a t s , and of extreme h a b i t a t s i n which s m a l l environmental d i f f e r e n c e s may be c r i t i c a l . Fur thermore, i t has been suggested (Gre ig -Smi th 1964; Kershaw 1964) tha t success ion i s accompanied by a decrease i n i n t e r a c t i o n between s p e c i e s . 2 t h i s should be r e f l e c t e d i n l e s s he te rogene i ty of x va lues i n more s t a b l e or s u c c e s s i o n a l l y advanced vege t a t i on the converse should a l s o h o l d . A s s o c i a t i o n - S a l t Marsh 32 I n t e r s p e c i f i c a s s o c i a t i o n i n the s a l t marsh i s marked, as can be seen i n Appendix 2 and F i g . 8. F i g . 8 i l l u s t r a t e s , the two main spec ies groupings . The aggregat ion dominated by D e s c h a m p s i a c e s p i t o s a , F e s t u c a r u b r a , J u n c u s b a l t i c u s , and Glaux maritima occurs on the h i g h e r , densely vegetated l e v e l s o f the marsh, whereas the group dominated by Salicornia v i r g i n i c a 3 C a r e x l y n g b y e i , and P l a n t a g o m a r i t i m a i s more c h a r a c t e r i s t i c of the lower l e v e l mud f l a t s . Desohampsia, C a r e x , P l a n t a g o , S a l i c o r n i a , S t e l l a r i a h u m i f u s a , and S c i r p u s cernuus e x h i b i t moderate p o s i t i v e a s s o c i a t i o n w i t h some species of both groupings . Triglochin maritimum i s a more or l e s s ub iqu i tous spec ies i n t h i s marsh, o c c u r r i n g so f r equen t ly as to f a l l i n n e i t h e r g rouping . There are o c c a s i o n a l i n c o n s i s t e n c i e s i n the r e s u l t s o f the 2 X t e s t s . For example, i n the species p a i r Carex lyngbyei* Plantago m a r i t i m a , s t rong p o s i t i v e a s s o c i a t i o n i s i n d i c a t e d by quadrat s i z e s l x l , 0 . 5 x 0 . 5 , and 1.5x0.3 m (Appendix 2 ) . At the l e v e l of the 0.3x0.3 m quadra ts , however, s t rong nega t ive a s s o c i a t i o n i s i n d i c a t e d . Perhaps t h i s i s i n d i c a t i v e o f compe t i t i on tha t was masked i n the l a r g e r quadrats but showed up i n the sma l l e r quadrat . In c o n t r a s t , the p a i r Juncus balticus Scirpus cernuus e x h i b i t s s i g n i f i c a n t nega t ive a s s o c i a t i o n v i a l x l m and 0 .5x0.5 m quadrats but s t rong p o s i t i v e a s s o c i a t i o n v i a 0.3x0.3 m quadrats (Appendix 2 ) . In t h i s case , the l a rge number of co-occurrences i n the 0.3x0.3 m quadrats may have been r e q u i r e d t o manifest a p o s i t i v e a s s o c i a t i o n on ly m i l d l y 33 s t a c h i o h 3 n - 30 F i g - 62 . C a r e x p a u c i f l o r a 3 n - ca . 37. 5Sb 60 61 6 2 5 6ox F i g . 63 . V a l e r i a n a s i t c h e n s i s 3 n - c a . 48. F i g . 64. L u p i n u s l a t i f o l i u s 3 n - c a . 48. F i g . 65. F e s t u c a v i r i d u l a , n - 14. F i g . 66. E r i g e r o n p e r e g r i n u s 3 n - 9. F i g . 67. Anemone o c c i d e n t a l i s 3 2n = 16. F i g . 68. E r y t h r o n i u m g r a n d i f l o r u m 3 2n = 24. F i g . 69 . P o t e n t i l l a f l a b e l l i f o l i a 3 n = 14. F i g . 70. V a c c i n i u m s c o p a r i u m 3 n - 12 . F i g . 71. C l a y t o n i a l a n c e o l a t a , 2n = 16. F i g . 12. A r e n a r i a c a p i l l a r i s 3 n - 11. F i g . 73. A n t e n n a r i a l a n a t a , n - 14 \u2022 F i g . 74. V e r o n i c a c u s i c k i i 3 n - 36 \u2022 F i g . 75. A g o s e r i s a u r a n t i a c a 3 n - 18. F i g . 76. F h l e u m a l p i n u m , n - 14. F i g . 77. A r n i c a l a t i f o l i a 3 n - 19. Stb 75 77 F i g . 78. L u z u l a h i t c h c o c k i i , n - 12. F i g . 79. T h a l i c t r u m o c c i d e n t a l e , n = F i g . 80 . A c h i l l e a m i l l e f o l i u m , n = 27 F i g - 81. T r i s e t u m s p i c a t u m , n = 14. F i g . 82 . E l y m u s g l a u c u s , n = 14. F i g . 83. S i l e n e p a r r y i , n = 24. F i g . 84. A r n i c a m o l l i s , n = ca . 38 . F i g . 85. P e n s t e m o n p r o c e r u s , n = 8. F i g . 86. P o a c u s i c k i i , n - 14. F i g . 87. S e n e c i o i n t e g e r r i m u s , n = 20 F i g . 88. E i e r a c i u m g r a c i l e , n - 9. F i g . 89. L u z u l a s p i c a t a , n - 12. F i g . 90. Sedum l a n c e o l a t u m , n - 8. F i g . 91. C a s t i l l e j a m i n i a t a , n - 12. F i g . 92. P e d i c u l a r i s b r a c t e o s a , n = 8 F i g - . 93. P h l o x d i f f u s a , n = 7. 94. P o t e n t i l l a d i v e r s i f o l i a 3 n - c a . 95. C a r e x s p e c t a b i l i s 3 n - c a . 42. 96. E p i l o b i u m a l p i n u m 3 n - 18. 97. D e l p h i n i u m n u t t a l l i a n u m 3 n - 16. 98. C a s t i l l e j a p a r v i f l o r a 3 n - 12. 99. R a n u n c u l u s e s c h s c h o l t z i i 3 n - 16. 100. S i b b a l d i a p r o c u m b e n s 3 n = 7. 101. J u n c u s d r u m m o n d i i 3 n c a . 60. 102. H y d r o p h y l l u m f e n d l e r i , n = 18. 103. S e n e c i o t r i a n g u l a r i s 3 n - 20. 104. V a c c i n i u m d e l i c i o s u m 3 n - 24. 105. M i t e l l a p e n t a n d r a 3 n = 7. 106. L u e t k e a p e c t i n a t a 3 n = 9. 107. P e d i c u l a r i s r a c e m o s a 3 n = 8. 108. 109. 110. P h y l l o d o c e e m p e t r i f o r m i s 3 V e r a t r u m v i r i d e 3 n - 16. 7eroniea w o r m s k j o l d i i 3 n n - 24 = 9. 59 f o r which no number has been repor ted p r e v i o u s l y . Each species has been ass igned a b a s i c chromosome number obta ined o r i n f e r r e d from the l i t e r a t u r e , e s p e c i a l l y D a r l i n g t o n and Wyl ie (1955) , subsequent IOBP Repor t s , and Tay lo r and M u l l i g a n (1968). B a s i c numbers which are themselves probably o f p o l y p l o i d o r i g i n ( e g . , x = 12 f o r many E r i c a c e a e ; x = 15 fo r T o f i e l d i a ; x = 10 f o r A r n i c a ) have been r e t a i n e d here as d i p l o i d . For each s i t e , percentages of p o l y p l o i d y i n both the f l o r a and v e g e t a t i o n have been c a l c u l a t e d and are l i s t e d at the bases o f the cor responding t ab l e s and i n Table 11. In view of the e x c e p t i o n a l cy to logy of the Cyperaceae and Juncaceae (Nordensk io ld 19 51; Davies 19 56; Stebbins 1971b), a b a s i c number of 10 has been a r b i t r a r i l y chosen for Carex, Soiryus, and Junous. This i s in tended as a compromise. Both H e i l b o r n (1939) and Wahl (1940) cons idered x = 7 to be the o r i g i n a l b a s i c number of the genus C a r e x , a l though Wahl a l s o de tec ted s e r i e s based on x = 5, 6, and 8. Love et al. (1957) proposed x = 5 as the pr imary b a s i c number of both Carex and J u n o u s , w h i l e Snogerup (1963) mainta ined t ha t i n Junous the pr imary b a s i c number can be as h igh as x = 25. The aneup lo id numbers i n Carex, Scirpus, and Junous have probably a r i s e n through a combinat ion o f t rue p o l y p l o i d y and p a r t i a l agmato-p l o i d y , or endonuclear p o l y p l o i d y . At any r a t e , the chromosome numbers of the above genera i n t h i s study are h i g h enough to be cons idered p o l y p l o i d even i f the -basic number i s as h igh as x = 20. The r e s u l t s i n d i c a t e tha t l e v e l s of p o l y p l o i d y i n these four communities are f a i r l y h i g h , and tha t the l e v e l w i t h i n a 60 TABLE 10. Chromosome numbers, p o l y p l o i d y , and importance va lues of -species of the four study communit ies. Species Chromosome Bas i c No. , n N o . , x P o l y -p l o i d Importance va lue ( I . V . ) S a l t Marsh D e s o h a m p s i a 13 7 + 38.6 c e s p i t o sa 24. 5 F e s t u c a r u b r a 21 7 + T r i g l o c h i n 48 6 . + 21. 0 m a r i t i m u m 17 . 8 S a l i c o r n i a 18 9 + v i r g i n i c a 16.9 P l a n t a g o m a r i t i m a 6 6 \u2014 J u n c u s b a l t i c u s 40 10 + 16 . 8 C a r e x lyngbyei 36 10 + 14. 3 G l a u x m a r i t i m a 15 15 - 13. 9 P o t e n t i l l a 14 7 + 7. 2 p a c i f i c a 7 . 1 A g r o s t i s e x a r a t a 14 7 + S t e l l a r i a h u m i f u s a 13 13 \u2014 6 . 0 T r i f o l i u m wormsk- 16 8 + 3 . 5 g o l d i i S c i r p u s c e r n u u s 30 10 + 3.3 P u c c i n e l l i a p u m i l a 21 7 + 2. 7 S p e r g u l a r i a 18 9 + 2 . 0 c a n a d e n s i s 1.8 D i s t i c h l i s s p i c a t a 2 0 a 5 H o r d e u m b r a c h y - 14 7 + 1. 8 a n t h e r u m L i l a e o p s i s 22 11 + 0.4 o c c i d e n t a l i s b 0.1 O e n a n t h e s a r m e n t o s a 22 D 11 + P l a n t a g o m a c r o c a r p a 12 6 + 0.1 17\/20 or 85% of f l o r a Sum of p o l y p l o i d i s p o l y p l o i d I . V . ' s \/ 2 = 81.5% of v e g e t a t i o n i s p o l y p l o i d Wade's Bog C a r e x p l u r i f l o r a 26 10 + 30.1 A p a r g i d i u m b o r e a l e 9 9 - 27.2 A g r o s t i s a e q u i - 7 7 - 15.9 v a l v i s C a r e x o b n u p t a 37, c a . 38 10 + 15.9 TABLE 10. (Continued) 61 Chromosome Bas i c P o l y - Import ance Species No. , n No. , x p l o i d va lue ( I . V . ) S a n g u i s o r b a 14 7 + 15. 1 o f f i c i n a l i s 14. 0 K a t m i a p o l i f o l i a 12 12 -V a o o i n i u m o x y o o o o u s 24 12 + 13. 3 C a r e x c a n e s o e n s 28 10 + 11. 9 D r o s e r a r o t u n d i - 10 10 - 11. 2 fo lia 1 Ledum g r o e n l a n d i o u m 13 13 - 9. T r i e n t a l i s a r c t i o a c a . 42-44 11(?) + 8 . 9 E m p e t r u m n i g r u m 13 13 \u2014 5 . 2 M y r i o a g a l e 48 8 + 4. 9 T o f i e I d i a 15 15 \u2014 4 . 6 g l u t i n o sa 13 G e n t i a n a 13 \u2014 1. 2 d o u g l a s i a n a L i n n a e a b o r e a l i s 16 8 + 1. 2 M a i a n t h e m u m 18 9 + 1. 0 d i l a t a t u m R h y n o h o s p o r a a l b a 13 13 - 0. 7 V a o o i n i u m v i t i s - 12 12 \u2014 0 . 4 i d a e a S c i r p u s o e s p i t o s u s c a . 5 2 10 + 0. 3 J u n o u s s u p i n i f o r m i s c a . 5 6 10 + 0 . 2 V a o o i n i u m 24 12 + 0. 2 u l i g i n o s u m 0. G a u l t h e r i a s h a t t o n 44 11 + 2 G e n t i a n a s o e p t r u m 13* 13 - 0 . 2 C o p t i s a s p l e n i - 9 9 - 0 . 2 f o l i a 0. ,1 C o p t i s t r i f o l i a 9 9 -V a o o i n i u m o v a t u m 12 12 0 . , 1 13\/27 or 48 .1% of Sum of p o l y p l o i d f l o r a i s I . V . ' s \/ 2 = = 51.6% p o l y p l o i d of v e g e t a t i o n i s p o l y p l o i d Ogg's Bo g M y r i o a gate 48 8 + 36 , .1 A p a r g i d i u m b o r e a t e 9 9 - 18 , . 4 C a r e x o b n u p t a 37 10 + 13. . 7 C a r e x p l u r i f l o r a 26 10 + 11 . 6 A g r o s t i s a e q u i - 7 7 \u2014 10 . 7 v a l v i s TABLE 10. (Continued) 62 Species Chromosome No. , n Bas i c No. , x P o l y -p l o i d Importance value ( I . V . ) S a n g u i s o r b a 14 7 + 9 . 3 o f f i c i n a l i s Ledum g r o e n l a n d i o u m 13 13 - 8.1 V a o o i n i u m o x y o o o o u s 24 12 + 7.5 D r o s e r a r o t u n d i - 10 10 - 6.1 f o l i a K a l m i a p o l i f o l i a 12 12 - 5.8 E m p e t r u m n i g r u m 13 13 - 5.5 T r i e n t a l i s a r c t i o a c a . 42-44 11(?) + 5.4 C a r e x c a n e s c e n s 28 10 + 5.0 To f i e I d i a 15 15 - 4.7 g l u t i n o s a 4.0 L i n n a e a b o r e a l i s 16 8 R h y n o h o s p o r a a l b a 13 13 - 3.9 P l a n t a g o m a c r o c a r p a 12 6 + 3.8 G a u l t h e r i a s h a l l o n 44 11 + 3.3 G e n t i a n a s c e p t r u m 13* 13 - 2 . 5 C a l a m a g r o s t i s 14 7 + 2.0 n u t k a e n s i s M a i a n t h e m u m 18 9 + 2 . 0 d i l a t a t u m S c i r p u s c e s p i t o s u s ca . 52 10 + 1.6 C o p t i s a s p l e n i - 9 9 - 1.4 fo lia C o p t i s t r i f o l i a 9 9 - 1.4 J u n c u s s u p i n i f o r m i s c a . 56 10 + 1.1 D e s o h a m p s i a 13 7 + 0.9 o e s p i t o s a 0.6 G e n t i a n a 13 13 -d o u g l a s i a n a V a o o i n i u m o v a t u m 12 12 - 0 . 6 V a o o i n i u m v i t i s - 12 12 - 0.4 i d a e a C o r n u s u n a l a s o h - 22 11 + 0.4 k e n s i s V a o o i n i u m 24 12 + 0.3 u l i g i n o s u m N e p h r o p h y l l i d i u m ca . 51 17 + 0.3 o r i s t a - g a l l i E r i o p h o r u m 30 10 + 0.2 poly s t a c h i o n C a r e x p a u o i f l o r a c a . 3 7 d 10 + 0.2 Ly s i o h i t u m 14 d 14(?) - 0.1 a m e r i c a n u m TABLE 10 (Continued) 63 Species Chromosome Bas ic N o . , n N o . , x P o l y -p l o i d Importance value ( I . V . ) Habenaria saccata Goody era oblongi-fo l i a Junous effusus Juncus ensif olius 21 15 c 40< 20 ( d 21 15 10 10 22\/39 or 56.4% of f l o r a i s p o l y p l o i d + + 0.1 0.1 0.1 0.1 Sum of p o l y p l o i d I .V. ' s \/ 2 = 54.5% o f vege t a t i on i s p o l y p l o i d B l a c k w a l l Meadow Valeriana c a . 48 8 + 24. 9 sitchensis Festuca viridula 14 7 + 24. 3 Lupinus latifolius c a . 48 12 + 21. 8 Erigeron 9 9 ' - 12. 4 peregrinus 11. Anemone 2n = 16 '8 - 9 occidentalis ( root t i p ) Potentilla 14 7 + 10. 4 f l a b e l l i f o l i a Erythronium 2n = 24 12 - 7. 0 grandiflorum (p re^meio t ic m i t o s i s ) Vaccinium \u202212* 12 - 6. 9 scoparium Claytonia 2n = 16 8 - 6 . 5 lanceolata ( p r e - m e i o t i c m i t o s i s ) Arenaria 11 11 - 6. 3 capillaris Antennaria lanata 14* 7 + 6. 2 Veronica cusickii 3 6* 9 + 5. 3 Agoseris 18 9 + 4. 8 aurantiaea Phleum alpinum 14 7 + 4. 8 Arnica l a t i f o l i a 19 10 + 4. 7 Luzula hitch- 12* 6 + 4. 6 cockii^ Thalictrum 2 8* 7 + 3. 7 occidentale AchiIlea 27 9 + 3. 5 mi Ilefolium Trisetum spicatum 14 7 + 3. 2 Carex rossii Not Not counted counted TABLE 10. (Continued) 64 Chromosome B a s i c P o l y - Importance Species N o . , n N o . , x p l o i d va lue ( I . V . ) E l y m u s g l a u c u s 14 7 + 2.5 S i l e n e p a r r y i 24 12 + 2.2 A r n i c a m o l l i s c a . 3 8 10 + 2.0 Pens.temon p r o c e r u s 8 8 - 2.0 P o a c u s i c k i i 14 7 + 1.9 S e n e c i o 20 10 + 1.8 i n t e g e r r i m u s E i e r a c i u m g r a c i l e 9 9 - 1.7 L u z u l a s p i c a t a 12 6 + 1.4 Sedum l a n c e o l a t u m 8 8 - 1.2 C a s t i l l e j a m i n i a t a 12 12 - 1.2 P e d i c u l a r i s 8 8 - 0.9 b r a c t e o s a P h l o x d i f f u s a 7 7 - 0 . 8 P o t e n t i l l a c a . 70 7 + 0.7 d i v e r s i f o l i a C a r e x s p e c t a b i l i s ca . 42 10 + 0.6 E p i l o b i u m a l p i n u m 18 9 + 0.5 D e I p h i n i u m 16* 8 + 0.5 n u t t a l l i a n u m C a s t i l l e j a 12 12 - 0.4 p a r v i f l o r a R a n u n c u l u s 16 8 + 0.4 e s c h s c h o l t z i i S i b b a l d i a 7 7 \u2014 0.3 p r o o u m b e n s J u n o u s d r u m m o n d i i c a . 6 0 10 + 0.2 H y d r o p h y I l u m 18 9 + 0.2 f e n d l e r i S e n e c i o 20 10 + 0.2 t r i a n g u l a r i s V a o o i n i u m 24* 12 + 0.2 d e l i c i o s u m M i t e l l a p e n t a n d r a 7 7 - 0.1 L u e t k e a p e o t i n a t a 9 9 - 0.1 P e d i c u l a r i s 8 8 - 0.1 r a o e m o s a P h y l l o d o c e 24 12 + 0.1 empetrif'or mis f S a x i f r a g a 1 9 r 10 + 0.1 o c c i d e n t a l i s 8 g T r o l l i u s laxus 8 - . 0.1 V e r a t r u m v i r i d e 16 8 + 0.1 TABLE 10. (Concluded) 65 Species Chromosome No. , n Bas ic No. , x P o l y - Importance p l o i d va lue ( I . V . ) V e r o n i c a wormsk- 9 9 0.1 j o l d i i 31\/50 or 62 . 0% Sum of p o l y p l o i d of f l o r a i s I . V . ' s \/ 2 = 70.3% p o l y p l o i d of v e g e t a t i o n i s p o l y p l o i d h S t e b b i n s and Love (1941). B e l l and Constance (1957); T a y l o r and M u l l i g a n (1968). ^ A f t e r Ca lder and Tay lo r (1968). As r epor ted i n Tay lo r and M u l l i g a n (1968). ^ A f t e r Hamet-Aht i (1971). Krause and Beamish ( in p r e s s ) . g P a c k e r (1964). p a r t i c u l a r area i s not d r a s t i c a l l y d i f f e r e n t whether computed as a percentage of the f l o r a or v e g e t a t i o n . Of the th ree vege t a t i on t y p e s c s t u d i e d , the s a l t marsh has the h ighes t l e v e l of p o l y p l o i d y (about 80%). The sphagnum bog v e g e t a t i o n has the lowest l e v e l (about 50%) and the suba lp ine meadow has an in te rmedia te l e v e l of about 65% (Tables 10 & 11) . Note tha t i f Carex and Seirpus are l e f t out of the c a l c u l a t i o n s , the p o l y p l o i d y l e v e l s fo r the bogs are d i s p r o p o r t i o n a t e l y lowered , i n c r e a s i n g the d i f f e r e n c e between l e v e l s i n the bogs and the other three s i t e s . Since a l l s i t e s are at e s s e n t i a l l y the same l a t i t u d e and were a l l covered by the P l e i s t o c e n e i c e sheet (Heusser 1960), d i f f e r e n c e s i n l e v e l s of p o l y p l o i d y cannot be exp la ined s a t i s f a c t o r i l y by d i f f e r e n t i a l e f f e c t s o f g l a c i a t i o n or 66 TABLE 11. Summary of l e v e l s of p o l y p l o i d y . No. of % . .polyploidy % p o l y p l o i d y S i t e spec ies ( f l o r a ) (vege ta t ion) S a l t Marsh 20 85.0 81.5 Wade's Bog 27 48.1 51. 6 Ogg's Bog 39 56.4 54. 5 B l a c k w a l l 50 62 . 0 70.3 Meadow p o s t g l a c i a l r e c o l o n i z a t i o n . R e c o l o n i z a t i o n of a l l s i t e s probably proceeded p r i m a r i l y from the area south o f the g l a c i a l boundary i n Washington, a l though the Tof ino area could conce ivab ly have been reached by c o a s t a l immigrants from r e f u g i a i n the Queen C h a r l o t t e I s l ands and P a c i f i c c o a s t a l A l a s k a (Heusser 1960; S c h o f i e l d 1969; Randhawa and Beamish 1972). Exp lana t ions based on v a r y i n g degrees o f average e n v i r o n -mental \"harshness\" , whether i n a c l i m a t i c or edaphic sense, are a l s o u n s a t i s f a c t o r y . Most impor tan t , comparisons o f measure-ments or es t imates of p h y s i c a l parameters among the four s i t e s would have l i t t l e meaning because of the d i s p a r i t y of t h e i r p h y s i c a l environments , assuming tha t such measurements would , i f comparable, have any f i n a l meaning (see Sect . I I I - J ) . However, c o r r e l a t i o n s w i t h environmenta l r i g o r can be made i f \" r i g o r \" i s def ined i n genera l terms and p a r t i t i o n e d i n t o s e v e r a l a spec t s . In l i n e w i t h the reasoning of Stebbins (1971b) and Johnson et al. (1965), l e v e l s o f p o l y p l o i d y can be r e l a t e d to the type and degree o f environmental d i s t u rbance . 67 In a d d i t i o n to average c o n d i t i o n s of the environmenta l complex such as c l i m a t i c means and n u t r i e n t l e v e l s , e s t i m a t i o n of environmental r i g o r must i n c l u d e the r e l a t i v e ampli tudes o f environmental f l u c t u a t i o n s and.-:ther , irr.eguiarity: or u n p r e d i c t a b i l i t y of these f l u c t u a t i o n s (S lobodk in and Sanders 19 69; Whi t t aker 19 72) . Regular environmenta l f l u c t u a t i o n s such as t i d a l f l o o d i n g and d i u r n a l o r seasonal temperature v a r i a t i o n s should not d i f f e r e n t i a l l y a f f e c t l e v e l s of p o l y p l o i d y , s ince p e r e n n i a l d i p l o i d s should be able to accommodate them as w e l l as comparable p o l y p l o i d s . However, l onge r - t e rm, more u n p r e d i c t -able d i s tu rbances ( shor t of long- te rm c l i m a t i c changes such as warming or c o o l i n g t rends) cou ld s e l e c t i v e l y favor p o l y p l o i d s . I would c l a s s the r a p i d and unp red i c t ab l e changes i n mobi le substratum and topography, c h a r a c t e r i s t i c of s a l t marshes, as f l u c t u a t i o n s of the l a t t e r type . Changes i n drainage and sedimenta t ion pa t te rns f r equen t ly occur i n s a l t marshes and are accompanied by simultaneous e r o s i o n and a c c r e t i o n of mud f l a t s and marsh (Johannessen 19 64; R e d f i e l d 1965, 1972). In a d d i t i o n , s a l t marsh vege t a t i on has been subject to e u s t a t i c r i s e and f a l l i n sea l e v e l r e l a t e d to P l e i s t o c e n e g l a c i a t i o n and de-g l a c i a t i o n (Cooper 1958, 1967; Heusser 1960). Such phys iograph ic changes would be accompanied by both r a p i d es tab l i shment and e l i m i n a t i o n of popu la t ions o f marsh species i n newly a v a i l a b l e or swamped s i t e s , r e s p e c t i v e l y . Subalpine meadows have exper ienced the i n t e r g l a c i a l advance and r e t r e a t of a l p i n e g l a c i e r s , and frequent but unp red i c t ab l e f i r e s (Douglas and B a l l a r d 1971) , l a t e snow packs , summer f r e e z e s , and s o i l d i s turbances by marmots and pocket gophers 68 a l l events tha t may be accompanied by c y c l e s o f e r o s i o n and es tabl i shment o f meadow v e g e t a t i o n i n new, d i s t u r b e d h a b i t a t s . These d i s tu rbances are s i m i l a r , i n type i f not degree, to those of the s a l t marsh. In c o n t r a s t , sphagnum bogs i n t h i s area have no s i m i l a r d i s t u r b a n c e s . Even the r e g u l a r hummock-ho l low c y c l e s of bogs (Gorham 1957; Lawrence 1958) are apparen t ly s e l f -gene ra t ed by the bog s p e c i e s , do not r e s u l t i n any d r a s t i c e r o s i o n , and are more or l e s s p r e d i c t a b l e and s e l f - c o n t a i n e d . I t appears , t hen , tha t the d i f f e r e n t l e v e l s of p o l y p l o i d y i n these three vege t a t i on types are best exp la ined by d i f f e r e n t degrees of environmental i n s t a b i l i t y . H i s t o r i c a l , m i g r a t i o n a l f a c t o r s have s u r e l y been a major cause o f the r e l a t i v e l y h i g h l e v e l of p o l y p l o i d y i n the o v e r a l l geographic a r ea , but have had minimal e f f e c t on d i f f e r e n c e s w i t h i n the a rea . Exp lana t i ons based on i n t r i n s i c p h y s i o l o g i c a l p r o p e r t i e s of p o l y p l o i d s tha t favor them i n extreme environments are t h e o r e t i c a l l y and p r a c t i c a l l y i n a p p r o p r i a t e . Environmenta l i n s t a b i l i t y or u n p r e d i c t a b i l i t y c o u l d , however, r e s u l t i n h igher l e v e l s o f p o l y p l o i d y both because such u n p r e d i c t a b i l i t y i m p l i e s c o n d i t i o n s (newly a v a i l a b l e d i s t u r b e d h a b i t a t s , r a p i d f l u c t u a t i o n s i n p o p u l a t i o n s i z e ) f avorab le f o r h y b r i d i z a t i o n , h y b r i d e s t ab l i shmen t , and es tabl i shment o f a l l o p o l y p l o i d d e r i v a t i v e s , and because the i n c r e a s e d , s t a b i l i z e d gene t i c v a r i a b i l i t y of the a l l o p o l y p l o i d -spec ies would p l ace them at a s e l e c t i v e advantage i n an unpred ic t ab l e environment. D. Flowering phenology. 69 One o f t e n - n e g l e c t e d aspect of p l a n t synecology i s r e p r o d u c t i v e phenology. A l l s p e c i e s t h a t are o u t c r o s s e d to any extent w i l l compete f o r p o l l i n a t i n g agents, whether the agents be b i o t i c or a b i o t i c . The s p e c i e s a l s o w i l l be competing f o r water, l i g h t , and n u t r i e n t s d u r i n g the growing season as they marshal resources f o r the r e p r o d u c t i v e e f f o r t . Phenology should r e v e a l a spectrum of peak f l o w e r i n g times f o r the species o f a p a r t i c u l a r community, a spectrum r e s u l t i n g from competition and an e v o l u t i o n toward niche d i f f e r e n t i a t i o n ( c f . Mosquin 1971). Fig u r e HlA-DJ d e p i c t s the f l o w e r i n g phenologies of a l l the angiospermous s p e c i e s of the s a l t marsh, bogs, and subalpine meadow. Each community was canvassed dur i n g the f l o w e r i n g season at l e a s t t w i c e , and o f t e n t h r e e to f o u r times per week. Each area was t r a v e r s e d and f l o w e r i n g behavior noted along a network of t r a n s e c t s . During the onset and c e s s a t i o n of flower-i n g , rough estimates were made of the p r o p o r t i o n of a p a r t i c u l a r s p e c i e s p o p u l a t i o n t h a t was i n flower. The h o r i z o n t a l bars i n F i g u r e 111A-D are tapered_at. both-, e n d s , p r o p o r t i o n a l to the r a p i d i t y with which the m a j o r i t y o f a g i v e n s p e c i e s p o p u l a t i o n commences and terminates f l o w e r i n g , but the p o p u l a t i o n dynamics probably would be more a c c u r a t e l y represented by more or l e s s normal, b e l l - s h a p e d curves. I t should be emphasized t h a t the blooming p e r i o d i s o n l y one phenophase of many, such as emergence or germination, stem or culm p r o d u c t i o n , formation of u n r i p e seeds and f r u i t s , d i s p e r s a l of r i p e seeds and f r u i t s , y e l l o w i n g of l e a v e s , and death or p a r t i a l dieback, t h a t could Fig. 111. Flowering phenology. 70 b Triglochin maritimum Carex lyngbyei Glaux maritima Stellaria humifusa Deschampsia cespitosa Juncus balticus Plantago maritima Puccinellia pumila Potentilla pacifica Trifolium wormskjoldii Scirpus cernuus Festuca rubra Spergularia canadensis Agrostis exarata Hordeum brachyantherum Lilaeopsis occidentalis Distichlis spicata Salicornia virginica Myrica gale Empetrum nigrum Coptis asplenifolia Carex pluriflora Scirpus cespitosus Coptis trifolia Carex obnupta Kalmia polifolia Gentiana douglasiana Apargidum boreale Trientalis arctica Vaccinium oxycoccus Carex canescens Vaccinium uliginosum Vaccinium vitis-idaea Ledum groenlandicum Agrostis aeguivalvis Tofieldia glutinosa Juncus supiniformis Rhynchospora alba Sanguisorba officinalis Linnaea borealis Drosera rotundifolia Gentiana sceptrum A S a l t M a r s h x = 4 6 d a y s B W a d e ' s B o g x = 3 2 d a y s \u2014i 1 r 10 20 30 APRIL 10 20 MAY __~r 31 \u2014i 1 1-10 20 30 JUNE T\" 10 20 JULY 31 10 20 AUGUST \u2014i\u2014 10 31 20 SEPTEMBER Myrica gale Empetrum nigrum Plantago macrocarpa Coptis asplenifolia Eriophorum polystachion Scirpus cespitosus Carex pluriflora Coptis t r i f o l i a Carex obnupta Kalmia polifolia Vaccinium ova turn Apargidium boreale Trientalis arctica Vaccinium oxycoccus Carex canescens Vaccinium uliginosum Vaccinium vitis-idaea Nephrophyllidium crista-galli Carex pauciflora Maianthemum dilatatum Ledum groenlandicum Gentiana douglasiana Agrostis aequivalvis Tofieldia glutinosa Gaultheria shallon Juncus supiniformis Cornus unalas$ch'ensis Rhynohospora alba Desohampsia cespitosa Sanguisorba officinalis Linnaea borealis Drosera rotundifolia Calamagrostis nutkaensis Gentiana sceptrum 's Bog 27 days -J O Q \u2014I ' 1 1 T 10 20 30 10 20 APRIL MAY 31 ^ 1 r 10 20 30 JUNE i 1\u2014 10 20 JULY 31 i r~ 10 20 AUGUST ~I ' r~ 31 10 20 SEPTEMBER 70 Anemone occldentalis Erythronium grandiflorum Claytonia lanceolata Luzula hitchcockii Vaccinium scoparium Phlox diffusa Antennaria lanata Potentilla f l a b e l l i f o l i a Sibbaldia procumbens Vaccinium deliciosum Microsteris gracilis Poa cusickii Ranunculus eschscholtzii Luzula spicata Thalictrum occidentale Carex rossii Senecio integerrimus Hydrophyllum fendleri Lupinus latifolius Epilobium alpinum Potentilla diversifolia Festuca viridula Valeriana sitchensis Arenaria capillaris Castilleja miniata Penstemon procerus Pedicularis bracteosa Delphinium nuttallianum Erigeron peregrinus Veronica cusickii Castilleja parviflora Phleum alpinum Veratrum viride Carex spectabilis Agoseris aurantiaca Juncus drummondii Trisetum spicatum Elymus glaucus Arnica l a t i f o l i a Hieracium gracile Sedum lanceolatum Arnica mollis Achillea millefolium Senecio triangularis Silene parryi Blackwall Meadow 23 days -| 1 1 r-10 20 30 10 ~l 1 \u00ab 1 ' ' 1 ' 1 1 1 r 31 10 20 30 10 20 31 10 20 31 10 20 20 APRIL MAY JUNE JULY AUGUST SEPTEMBER 71 be represented i n a complete phenodynamic s t r i p ( L i e t h 1970). F igu re 111A-D confirms the ca sua l impress ion t h a t , w i t h i n a g iven p l an t community, species f l o w e r i n g t imes form a sequence of ove r l app ing i n t e r v a l s o r , more a c c u r a t e l y , ove r l app ing cu rves . Analogous p h e n o l o g i c a l g rad ien t s have been repor ted by Mowbray and Oost ing (1968) and Mosquin (1971). The average blooming p e r i o d l eng th f o r species o f the S a l t Marsh, Wade's Bog, Ogg's Bog, and B l a c k w a l l Meadow i s 4-6, 32, 27, and 23 days , r e s p e c t i v e l y . These lengths r e f l e c t both the number of species per community and the l eng th o f the growing season. The number of angiospermous species i n each of these s i t e s i s , i n o r d e r , 18, 24, 30, and 45 showing an inve r se r e l a t i o n s h i p between the number of spec ies per community and the average l e n g t h of a s p e c i e s ' blooming p e r i o d . The f i r s t th ree s i t e s a l l have about the same long growing season; approximate ly a 5 1\/2 month p e r i o d from the beginning o f A p r i l to the middle of September. B l a c k w a l l Meadow, on the other hand, has a growing season compressed i n t o 9-10 weeks from l a t e June to e a r l y September, a t l e a s t i n l a t e snow- l i e y e a r s , as were 1971 and 1972. The short growing season, h i g h number of s p e c i e s , d e n s i t y of f l o w e r i n g i n d i v i d u a l s , and: percentage. ;.of . b i o t i c a l l y -p o l l i n a t e d s p e c i e s , and the consequent s i g n i f i c a n t i n t e r s p e c i f i c over lap of f l o w e r i n g per iods i n the subalp ine meadow suggest tha t meadow spec ies have evolved i n an environment i n t e n s e l y compe t i t i ve f o r animal ( e s p e c i a l l y i n s e c t ) p o l l i n a t o r s , much more so than a s a l t marsh or bog environment. S e l e c t i o n i n such h i g h l y compe t i t i ve c o n d i t i o n s would be f o r f l o r a l s p e c i a l i z a t i o n s 72 tha t would inc rease a f l o w e r ' s a t t r a c t i v e n e s s to animal p o l l i n -a to rs and the p o l l i n a t o r ' s constancy to the f l o w e r . There cou ld a l s o be s e l e c t i o n f o r autogamy ( c f . L e v i n 1972c')-, bu.i;at the expense of long term v a r i a b i l i t y . Thus, the b rea th t ak ing d i s p l a y o f showy f lowers of many forms, c o l o r s , and odors tha t i s c h a r a c t e r i s t i c of l u s h subalp ine meadows i s l o g i c a l i n an e v o l u t i o n a r y con tex t . One might i n t e r p r e t the longer average f l o w e r i n g per iods of the s a l t marsh spec ies as a r e s u l t i n par t of the \"pre-ponderance of w i n d - p o l l i n a t e d species i n the marsh. T h e o r e t i c a l l y , a longer blooming season would be of advantage to an anemo-p h i l o u s spec ies s ince wind i s a much l e s s r e l i a b l e p o l l i n a t i n g agent than most animals and a longer f l o w e r i n g p e r i o d would inc rease the chances of succes s fu l p o l l i n a t i o n . However, c a l c u l a t i o n s r e v e a l no s i g n i f i c a n t d i f f e r e n c e , e i t h e r w i t h i n one community or over a l l communit ies , between l eng th o f blooming t imes fo r anemophilous v s . entomophilous spec i e s . There i s , though, a d e f i n i t e tendency f o r autogamous spec ies such as Drosera r o t u n d i f o l i a , Spergularia canadensis, Elymus g.laucus 3 and Hordeum brachy antherum to have shor te r than average an thes i s t i m e s , which i s what one would expec t , s ince autogamy ensures succes s fu l p o l l i n a t i o n . E . P o l l i n a t i o n eco logy . 73 The f o l l o w i n g summary of the p o l l i n a t i o n ecology of the study communities i s based on two summers of f i e l d observa t ions i n 1971 and 1972, and on observa t ions and in fe rences c u l l e d from the l a rge l i t e r a t u r e on the sub jec t . During the growing season I v i s i t e d each of the study areas u s u a l l y two or three times a week to study f l o r a l b i o l o g y and observe and c o l l e c t f lower v i s i t o r s . Many a d d i t i o n a l observa t ions were made w h i l e I was doing the quadrat and t r an sec t p h y t o s o c i o l o g i c a l sampling on o ther occas ions . Table 12 summarizes the o v e r a l l p o l l i n a t i o n scheme i n each of the four sample s i t e s . Each species has been c l a s sed as being p o l l i n a t e d predominant ly by e i t h e r wind (anemophi ly) , i n s e c t s ( en tomophi ly ) , or b i r d s ( o r n i t h o p h i l y ) none are w a t e r - p o l l i n a t e d . The p o l l i n a t i o n mechanisms of- the i n d i v i d u a l species are d i scussed i n d e t a i l i n Appendix 3. TABLE 12. Community mode of p o l l i n a t i o n as percentage o f the f l o r a and v e g e t a t i o n . P o l l i n a t i o n B l a c k w a l l mechanism S a l t Marsh Wade's Bog Ogg's Bog Meadow Anemophily 67%\/83.4% a 36\/42.6 40\/48.2 25\/25 Entomophily 33\/16.6 64\/57.4 60\/51.8 72 .7 \/74 .4 O r n i t h o p h i l y 2 .3 \/0 .6 a% f l o r a \/ % vege t a t i on (determined from importance v a l u e s ) . 74 C l e a r l y , anemophily i s the major mode of p o l l i n a t i o n i n the s a l t marsh. Entomophily predominates i n the suba lp ine meadow, w h i l e both bogs have more of a balance between i n s e c t and wind p o l l i n a t i o n . The s i t u a t i o n i n the s a l t marsh i s not as c l e a r - c u t as i t seems, however. During both summers of f i e l d work I have observed bumble bee (Bombus terricola occi-dentalis Grne. ) p o l l e n fo rag ing on s i x t y p i c a l l y anemophilous s a l t marsh s p e c i e s : D e s o h a m p s i a o e s p i t o s a , F e s t u o a r u b r a v a r . l i t t o r a l i s , A g r o s t i s e x a r a t a , P l a n t a g o m a r i t i m a , J u n o u s b a l t i o u s , and Salioornia virginica (Pojar 1973b). In g e n e r a l , bumble bees are the most important animal p o l l i n a t i n g vec to r at a l l four s i t e s . D i p t e r a are extremely impor tan t , p a r t i c u l a r l y Syrph idae , Muscidae, and Bombylidae. L e p i d o p t e r a , e s p e c i a l l y b u t t e r f l i e s and sk ippe r s (Rhopalocera) , are a l s o of major s i g n i f i c a n c e i n the suba lp ine meadow, but i t i s s t range tha t there are very few b u t t e r f l i e s i n the Tof ino area . I t i s a l s o noteworthy tha t none of the nine spec ies (.Puccinellia p u m i l a , S p e r g u l a r i a c a n a d e n s i s ( F i g . 112) , H o r d e u m b r a o h y a n t h e r u m , D r o s e r a r o t u n d i f o l i a ( F i g . 113) , E l y m u s g l a u c u s , E p i l o b i u m a l p i n u m , M i o r o s t e r i s g r a c i l i s , and S i b b a l d i a prooumbens) t ha t are f u l l y or predominant ly autogamous ( s e l f - p o l l i n a t i n g ) are dominant elements i n t h e i r communities. From q u a l i t a t i v e es t imates of p o l l i n a t o r d e n s i t y and a c t i v i t y I f e e l t ha t i n a l l of these communities the i n s e c t s are the l i m i t i n g f a c t o r i n the r e p r o d u c t i o n of many entomo-p h i l o u s s p e c i e s . That i s , I b e l i e v e tha t there i s compe t i t i on among f lowers f o r p o l l i n a t o r s r a t h e r than among i n s e c t s f o r 75a, F i g . 112. Spergularia canadensis; chasmogamous, s e l f -p o l l i n a t i n g f l o w e r . I n d i v i d u a l p l an t s o f ten have both chasmogamous and cle is togamous f l o w e r s . F i g . 113. Drosera-rotundpfoli'a; s h o r t - l i v e d , . chasmogamous , s e l f - p o l l i n a t i n g f l o w e r . Two cleis togamous f lowers can be seen j u s t above the open f l o w e r . 76 nec ta r and p o l l e n . This seems p a r t i c u l a r l y t rue i n B l a c k w a l l Meadow, which has a l a rge number of entomophilous species and an extremely h i g h d e n s i t y of s imul taneous ly-b looming f lowers (see F i g s . 6 8 7 ) . Mosquin (1971) concluded t h a t , at l e a s t dur ing midsummer, the f l o r a o f a mountain v a l l e y near Banff competed s t r o n g l y f o r p o l l i n a t o r s , and some spec ies were unsuccess fu l i n the c o m p e t i t i o n . Hocking (19 68) and Kevan (1970, 1972) have noted a r e l i a n c e of f lowers on p o l l i n a t i n g i n s e c t s i n h i g h a r c t i c v e g e t a t i o n , w h i c h , i n d e n s i t y and showiness of f lowers and compressed growing season, i s somewhat s i m i l a r to suba lp ine meadow v e g e t a t i o n . Undoubtedly, i n t e r s p e c i f i c compe t i t i on has p layed a major r o l e i n the e v o l u t i o n o f showy f l o w e r s , as w e l l as p r e s su r ing some species i n t o the escapism of autogamy ( L e v i n 197 2 c ) . Another p o s s i b l e advantage of the showy f lowers of some subalp ine spec ies has been suggested by Hocking and S h a r p l i n (1965) and Kevan (1970). They have found tha t the i n t r a f l o r a l temperatures of l a r g e showy f lowers are g e n e r a l l y above the ambient i n the a r c t i c , and under optimum c o n d i t i o n s the temperatures i n f lowers shaped l i k e p a r a b o l i c r e f l e c t o r s can exceed the ambient by 5 \u00b0 ' t o 10\u00b0 C ( B l i s s 1971). In B l a c k w a l l Meadow, bowl-shaped f lowers l i k e those of Potentilla diversi-f o l i a , P. f l a b e l l i f o l i a ( F i g . 114)., R a n u n c u l u s e s c h s c h o l t z i i 3 and Anemone occidentalis serve as basking s i t e s f o r the va r ious d ip t e rans tha t are the p r i n c i p a l p o l l i n a t o r s of these s p e c i e s . Th i s f l ower bask ing i s probably most important t o the i n s e c t s du r ing e a r l y morning i n the mountains, when the sun i s up but the a i r temperature s t i l l low. Kevan (197 0) has suggested tha t 77 F i g . 114. Potentilla flabellifolia. S l i g h t l y protogynous f l o w e r ; nec ta r sec re ted as a t h i n , s h i n i n g f i l m on the b l a c k i s h - p u r p l e d i s c j u s t i n s i d e the f i l amen t bases . Bowl-shaped blossomrserves as bask ing s i t e f o r i n s e c t s . F i g . 115. Plantago maritima. S t rong ly protogynous, anemo-p h i l o u s flowers, borne i n a dense, a c r o p e t a l l y -f l o w e r i n g s p i k e . Note the e longate stigmas and w e l l - e x s e r t e d , v e r s a t i l e an thers . 78 the h igher i n t r a f l o r a l temperatures i n gene ra l may a l s o be important fo r p o l l e n tube growth, f e r t i l i z a t i o n , and p o s s i b l y seed development. A t h e o r e t i c a l aspect o f i n t e r s p e c i f i c compe t i t i on f o r p o l l i n a t o r s has been exp lored i n models by L e v i n and Anderson (1970) and Straw (1972). Both models have e s s e n t i a l l y the same i n i t i a l assumptions: (1) two randomly i n t e r m i x e d , s imul taneous ly f l o w e r i n g , s e l f - i n c o m p a t i b l e spec ies w i t h f lowers s u f f i c i e n t l y s i m i l a r t o a t t r a c t the same p o l l i n a t o r s ; (2) a s i n g l e p o l l i n a t o r spec ies to s e r v i c e both p l an t s p e c i e s ; (3) a d e f i c i t o f p o l l i n a t o r s r e l a t i v e to the number of f l o w e r s . Then, i f the r a t e of r e p r o d u c t i o n o f each f lower species 'depends on i t s success i n the compe t i t i on fo r p o l l i n a t o r s , the frequency o f a spec ies i n subsequent generat ions w i l l be p r o p o r t i o n a l to the degree of p o l l i n a t o r constancy to tha t spec ies , and the l e s s favored species e v e n t u a l l y should be e l i m i n a t e d . I f p o l l i n a t o r constancy i s d e n s i t y dependent ( c f . L e v i n and K e r s t e r 1969a), the m i n o r i t y spec ies w i l l be at an inc reased disadvantage and i t s compe t i t i ve e x c l u s i o n should be a c c e l e r a t e d (Straw 1972). The f i r s t two requirements f o r t h i s type of compe t i t i ve e x c l u s i o n r a r e l y , i f a t a l l , e x i s t i n nature (Crosswhi te and Crosswhi te 19 7 0 ; ;Macior\" 1971; Straw 1972). In the study communit ies, assumption (3) seems to be v a l i d , e s p e c i a l l y f o r the suba lp ine meadow. In the meadow, there are two spec ies p a i r s , A r n i c a l a t i f o l i a v s . A. m o l l i s and S e n e c i o i n t e g e r r i m u s v s . S. triangularis, t ha t have s i m i l a r f lowers and p o l l i n a t o r s and are a l l s e l f - i n c o m p a t i b l e . However, they do not r e l y on a s i n g l e p o l l i n a t o r spec ies but r a t h e r are s e r v i c e d by a v a r i e t y of 79 Hymenoptera, D i p t e r a , and L e p i d o p t e r a ; they are \"cornucopian\" f lowers i n the sense of Mosquin (1971). Nor are these spec ies randomly i n t e rmixed A r n i c a m o l l i s and S e n e c i o i n t e g e r r i m u s g e n e r a l l y occur on d r i e r s i t e s w i t h i n the meadow than t h e i r congeners. Fur thermore, S. integerrimus f lowers much e a r l i e r i n the season than the o ther three s p e c i e s , which are more or l e s s s imul taneous ly l a t e - f l o w e r i n g . In the study communit ies, these spec ies p a i r s are the c l o s e s t approaches to the t h e o r e t i c a l assumptions, and c l e a r l y the approaches are q u i t e remote. N e v e r t h e l e s s , i t i s safe t o say tha t avoidance of i n t e r s p e c i f i c compe t i t i on f o r p o l l i n a t o r s , by s h i f t i n g of f l o w e r i n g p e r i o d and\/or h a b i t a t preference and\/or p o l l i n a t i n g agent , has been i n s t rumen ta l i n the e v o l u t i o n of p l a n t species i n g e n e r a l . In p a r t i c u l a r , i t i s r e f l e c t e d i n the s p a t i a l d i s t r i b u t i o n s and phenology o f the spec ies o f the study a reas , and moreover i s much more apparent and presumably p lays a much more important r o l e i n the suba lp ine meadow than i n the s a l t marsh or bogs. Types o f p o l l i n a t i o n Gross f l o r a l morphology should be regarded as r e s u l t i n g from adapta t ions fo r f e r t i l i z a t i o n from f lowers of d i f f e r e n t i n d i v i d u a l s of the same spec ies (Darwin 1876, 1877; Whitehouse 1959). The except ions to t h i s gene ra l r u l e ( e g . , as occur i n s e l f - f e r t i l i z i n g spec ies ) are g e n e r a l l y regarded as secondary adapta t ions f o r the sake of g rea te r c e r t a i n t y o f seed p roduc t ion of a more or l e s s uni form genotype (Stebbins 19 50, 1957a; Grant 19 71)...The .adaptive nature .of t h e ' f l o w e r must be 80 apprec ia t ed i n a d i s c u s s i o n of p o l l i n a t i o n eco logy , but the t empta t ion to specu la te on the adapt ive func t ions of va r ious f l o r a l s t r u c t u r e s tha t defy l o g i c a l 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 should be r e s i s t e d . The i n t e r p r e t a t i o n of the f lower as a f u n c t i o n a l r e p r o d u c t i v e u n i t has l e d to the d e s c r i p t i o n of morpho log ica l c a t e g o r i e s f o r f lower forms r e l a t e d to t h e i r p o l l i n a t i o n ecology t hus , we have \"bee f l o w e r s \" , \" f l y f l o w e r s \" , \"bee t le f l o w e r s \" , and the l i k e . Crosswhite and Crosswhi te (1966) and Macior (1971) have po in t ed out the danger o f such a t y p o l o g i c a l approach, f o r many p l a n t spec ies u t i l i z e a wide v a r i e t y o f p o l l i n a t o r s and are not r e s t r i c t e d to one p a r t i c u l a r v e c t o r . This danger should be kept i n mind dur ing the f o l l o w i n g d i s c u s s i o n , s i nce I s h a l l be r e f e r r i n g to \"bumble bee f l o w e r s \" , e t c . , w h i l e o u t l i n i n g the p o l l i n a t i o n mechanisms c h a r a c t e r i s t i c o f c e r t a i n types of blossoms. C e r t a i n combinat ions of p o s i t i v e l y c o r r e l a t e d charac te r s or fea tures make up a p a r t i c u l a r syndrome ( F a e g r i and van der P i j l 19 71) corresponding to each of the f o l l o w i n g p o l l i n a t i o n s t r a t e g i e s . The o u t l i n e i s tha t o f F a e g r i and van der P i j l (1971); the examples are from the study communities. Anemophily Wind p o l l i n a t i o n or anemophily i s the dominant type o f a b i o t i c p o l l i n a t i o n i n p l a n t s , and i s the only type t ha t occurs i n the study communit ies , there being no w a t e r - p o l l i n a t e d s p e c i e s . The syndrome of anemophily i n c l u d e s the f o l l o w i n g 81 fea tures (Whitehead 19 69; F a e g r i and van der P i j l 19 71; P roc to r and Yeo 1973): the f lowers tend to be o f s m a l l s i z e , reduced and inconsp i cuous , to l a c k nectar and odor , and to be c l u s t e r e d i n dense i n f l o r e s c e n c e s ; the anthers have abundant, l i g h t , dry p o l l e n , and the stigmas have an expanded surface a rea . Anemo-p h i l o u s spec ies are f r equen t ly u n i s e x u a l , perhaps because s e l f -p o l l i n a t i o n i n an he rmaphrod i t i c , w i n d - p o l l i n a t e d f lower would be i n e v i t a b l e w i t h such a h i g h i nc idence of p o l l e n per u n i t area i n the immediate v i c i n i t y of the anthers ( F a e g r i and van der P i j l 1971). Then t o o , when p o l l e n i s the i n s e c t a t t r a c t a n t , the e v o l u t i o n o f u n i s e x u a l f lowers should tend toward anemo-p h i l y , s i nce entomophilous p l an t s would su f f e r a r e d u c t i o n i n the number of i n s e c t v i s i t s to female f lowers (Kaplan and Mulcahy 1971). U n i s e x u a l f lowers are found i n such wind-p o l l i n a t e d spec ies as M y r i c a g a l e , E m p e t r u m n i g r u m , C a r e x spp., and Thalictrum occidentale. I f an anemophilous spec ies has hermaphrodi t ic f l o w e r s , these are f r equen t ly s t r o n g l y dichogamous e g . , P l a n t a g o m a r i t i m a ( F i g . 115) and P. m a c r o c a r p a , S a l i c o r n i a v i r g i n i c a ( F i g . 116a S b ) , T r i g l o c h i n m a r i t i m u m , J u n c u s spp. ( F i g . 117) , L u z u l a s p p . , and S c i r p u s c e s p i t o s u s ( F i g . 118) . The genus Thalictrum p rov ides e x c e l l e n t examples of these t r ends . Kaplan and Mulcahy (19 71) showed tha t the d i o e c i o u s and p o l y -gamous spec ies of Thalictrum are i n gene ra l the most anemo-p h i l o u s , w h i l e hermaphrodite species are the most entomophilous. Fur thermore, the hermaphrodite Thalictrum species are d i c h o -gamous (Faeg r i and van der P i j l 1971). F i n a l l y , as an e c o l o g i c a l c o r o l l a r y , wind p o l l i n a t i o n most o f t en predominates i n open v e g e t a t i o n w i t h clumped species d i s t r i b u t i o n s (Whitehead 19 69) . C O F i g . 116a. S a l i c o r n i a v i r g i n i c a ; protogynous F i g - 116b. S a l i c o r n i a v i r g i n i c a ; f lowers f l o w e r s , here i n female stage i n male stage (anthers (st igmas r e c e p t i v e ) . ex se r t ed , shedding p o l l e n ) . F i g . 117. J u n o u s b a l t i o u s ; s t r o n g l y F i g . 118 protogynous , anemophilous f l o w e r s . CO CO 8 Seivpus oespitosus ; s t r o n g l y protogynous , anemophilous f lowers u n i t e d i n a s e v e r a l -f lowered , s o l i t a r y , t e r m i n a l s p i k e l e t . Male stage i n mid-foreground; female stage behind and to the l e f t . 84 Most of the s a l t marsh spec ies are e x c e l l e n t examples o f anemophiles , and the marsh i t s e l f i s a predominant ly wind-p o l l i n a t e d community. However-, as mentioned be fo re , s i x : t y p i c a l l y anemophilous s a l t marsh species are a l s o v i s i t e d by p o l l e n - f o r a g i n g bumble bees. Pojar (1973b) has d i scussed the p o s s i b l e s i g n i f i c a n c e o f t h i s f o r t u i t o u s entomophi ly . The s i t u a t i o n i s advantageous to the p l a n t s , s i n c e : t h e e f f i c i e n c y of wind p o l l i n a t i o n must be c o n s i d e r a b l y reduced i n the humid oceanic c l i m a t e o f the Tof ino a rea . The bumble bees seem to have responded to a p a u c i t y of p o l l e n - r i c h entomophilous f lowers by s h i f t i n g t h e i r p o l l e n - g a t h e r i n g a c t i v i t i e s to spec ies t h a t , i n most o ther c i r cums tances , would be ignored . Anemophily i s now g e n e r a l l y cons idered to be a d e r i v e d c o n d i t i o n i n angiosperms (Whitehead 1969; Kugle r 1970; Stebbins 1970a; F a e g r i and van der P i j l 1971). Apparent r e v e r s i o n s from wind p o l l i n a t i o n to secondary entomophily have occur red i n the t r o p i c a l sedge Dichromena oiliata (Leppik 1955; Baker 1963), i n c e r t a i n t r o p i c a l r a i n f o r e s t grasses (Soderstrom and Calderon 1971), inr ' the geniis -.Salix, and e x t e n s i v e l y i n the f a m i l y Moraceae (Stebbins 1970a). Other observers have noted p o l l e n -fo rag ing v i s i t s by honey bees (Bogdan 19 62) and hover f l i e s ( C l i f f o r d 1964) to grass f l o w e r s , and by honey bees to Plantago lanceolata ( C l i f f o r d 1962). In c o n t r a s t , adapt ive s h i f t s from i n s e c t to wind p o l l i n a t i o n are a l s o apparent i n some groups. The genera Plantago and Thalictrum both c o n t a i n i n s e c t - and w i n d - p o l l i n a t e d s p e c i e s , and the anemophily e v i d e n t l y i s de r i ved and of f a i r l y recen t o r i g i n (S tebbins-1970a; F a e g r i and van der P i j l 1971; Kaplan 85 and Mulcahy 1971). In the study communties, Plantago maritima, P. m a c r o c a r p a , and T h a l i c t r u m o o c i d e n t a l e ( F i g . 119a & b) are a l l predominant ly w i n d - p o l l i n a t e d . Thalictrum (Ranunculaceae) and Sanguisorba (Rosaceae) are two genera tha t belong to over -whelmingly entomophilous f a m i l i e s , but never the less c o n t a i n some anemophilous species (Knuth 1906-1909; Stebbins 1970a). Sanguisorba officinalis ( F i g . 120) , a t a l l p e r e n n i a l herb o f the sphagnum bogs, has apetalous f lowers w i t h lobed stigmas and the f lowers are aggregated i n a dense i n f l o r e s c e n c e . But the deep maroon-purple f lowers are markedly n e c t a r i f e r o u s , have s t i c k y (not powdery) orange p o l l e n , and are p o l l i n a t e d by d i p t e r a n s . S. officinalis represents a c o n d i t i o n in te rmedia te between anemophily and entomophi ly , and the d i r e c t i o n of the adap t ive s h i f t can o n ly be presumed, a l though the m a j o r i t y o f Sanguisorba spec ies are w i n d - p o l l i n a t e d (Stebbins 1970a). F l y p o l l i n a t i o n (myophily) There i s great v a r i a t i o n i n the p o l l i n a t i o n methods employed by D i p t e r a . There are many u n s p e c i a l i z e d f l i e s , o f sma l l s i z e and w i t h shor t probosces , t ha t are g e n e r a l l y r e s t r i c t e d - , to more \" p r i m i t i v e \" f lowers w i t h e a s i l y a c c e s s i b l e nec ta r . Blossoms c h a r a c t e r i s t i c a l l y p o l l i n a t e d by these s m a l l , u n s p e c i a l i z e d f l i e s form a f a i r l y d i s t i n c t type of f l y f l o w e r , a l though they are a l s o v i s i t e d by many sma l l hymenopterans (Kugle r 1955). The syndrome of t h i s type of myophily i n c l u d e s r e g u l a r , s i m p l e , g e n e r a l l y l i g h t but d u l l - c o l o r e d f lowers w i t h l i t t l e or no depth e f f e c t . The nec ta r i s very a c c e s s i b l e , the oo CD F i g . 119a. Thalictvum occidentale . Male F i g . 119b. Thaliotrum o c c i d e n t a l e . Female f l o w e r s . f lowers (stigmas a l ready w i t h e r e d ) . Th i s i s a d i o e c i o u s , anemophilous spec i e s . Note the elongate s t igmas , the e longa te , pendent stamens, and the reduced, inconspicuous f lowers which are ape t a lous , w i t h g r een i sh -wh i t e , caducous sepals ( f a l l e n o f f i n female f l o w e r s ) . 87a, Fig. 120. S a n g u i s o r b a . o f f i o i n a l i s ; flowers apetalous, sepals maroon to deep maroon purple; the s t i c k y , orange pollen i s often shed i n bud. Fig. 121. T r i e n t a l i s a r c t i o a ; the white petals form a shallowly bowl-shaped star; the thick, fleshy, stamen-bearing r i n g i s a t t r a c t i v e to f l i e s . 0 .05 ) . A l s o , Poa cusickii i s o v e r a l l the l e a s t v a r i a b l e s p e c i e s . Low w i t h i n -p o p u l a t i o n v a r i a b i l i t y i s to be expected i n agamospermous species (Gustafsson 1946-47) . H. Niche d i f f e r e n t i a t i o n . 136 Numerica l taxonomy o f f e r s an approach to the study of n iche d i f f e r e n t i a t i o n w i t h i n p l an t communities. Assuming tha t q u a n t i t a t i v e or q u a l i t a t i v e morpho log ica l and b e h a v i o r a l charac te r s are o f e c o l o g i c a l s i g n i f i c a n c e i n the l i f e o f p l a n t s , i t should be p o s s i b l e to quan t i fy the e c o l o g i c a l s i m i l a r i t i e s or d i s s i m i l a r i t i e s of a group of spec ies tha t c h a r a c t e r i s t i c a l l y grow toge the r . For example, a Festuca spec ies i s p h e n e t i c a l l y more l i k e a Deschampsia than a P o t e n t i l l a , but i s a Potentilla more l i k e a R a n u n c u l u s , an A n e m o n e , or an A r n i c a , e c o l o g i c a l l y speaking? Taxonomica l ly , these ques t ions are easy to answer, and the e c o l o g i c a l answers should be s i m i l a r , s i n c e a spec ies should be an e c o l o g i c a l u n i t as w e l l as a t axon . But an e c o l o g i c a l s i m i l a r i t y a n a l y s i s cou ld o f f e r i n t e r e s t i n g and perhaps s u r p r i s i n g i n s i g h t s . What spec ies i n the suba lp ine meadow i s e c o l o g i c a l l y most s i m i l a r t o Thalictrum occidentalel Does Gentiana douglasiana f u n c t i o n more l i k e Gentiana sceptrum or Trientalis arctica i n the sphagnum bogs? I w i l l t r y to answer these and s i m i l a r ques t ions by what amounts to a p a r t i a l d e s c r i p t i o n of each s p e c i e s ' n i c h e , n e c e s s a r i l y us ing on ly those charac te r s tha t are e a s i l y measured or observed and un fo r tuna t e ly o m i t t i n g p h y s i o l o g i c a l n iche components. The approach has been neo-Adansonian whereby equal weight i s g iven to a l l cha rac te r s used i n the a n a l y s i s . A s i m i l a r i t y measure fo r mixed q u a n t i t a t i v e and q u a l i t a t i v e a t t r i b u t e s (Anderson 1971) has been used. Anderson 's measure of s i m i l a r i t y i s as f o l l o w s . 137 I f a t t r i b u t e k i s q u a n t i t a t i v e and sca led to have approxi -mately u n i t range , l e t . f e = x i k x j k I f a t t r i b u t e A: i s q u a l i t a t i v e , l e t s = 0 w h e n *ih * * j k 13 k 0.5 when x ^ = x T h e n 1 m S . . = - Z S . , 7 13 m 13k where S . . i s the s i m i l a r i t y between o p e r a t i o n a l taxonomic 13 u n i t s (OTU's) . . and m the t o t a l number of a t t r i b u t e s . The d i s t ance d . . between OTU's . . i s then computed as 13 13 ^ d . . = (s . . + s . . - 2s . . ) 1 \/ 2 . 13 i i 33 13 The OTU's f o r each community are i t s c o n s t i t u e n t s p e c i e s . A t o t a l of 32 u n i t charac te r s were used to measure s i m i l a r i t y between OTU's. A u n i t cha rac te r i s def ined by Soka l and Sneath (19 63) as a \"taxonomic cha rac te r o f two or more s t a t e s , which w i t h i n the study at hand cannot be subd iv ided l o g i c a l l y , except f o r s u b d i v i s i o n brought about by changes i n the method of c o d i n g \" . Each u n i t charac te r has at l e a s t two s t a t e s . I have t r i e d to choose independent charac te r s but fo r the sake of completeness some redundancy was unavo idab le . The u n i t charac te r s are o u t l i n e d i n Table 17 and ( i f necessary) exp l a ined i n the f o l l o w i n g paragraphs. The l i f e - f o r m c l a s s i f i c a t i o n i s adopted from Raunkiaer (.1934). Therophytes are annuals i n which the renewal bud i s w i t h i n the seed coa t . Geophytes have perennat ing buds covered by s o i l , whereas the buds of hemicryptophytes are at or s l i g h t l y embedded i n the s o i l su r f ace . Chamaephytes are TABLE 17. E c o l o g i c a l u n i t c h a r a c t e r s . 138 U n i t charac te r Character s t a t e s Number o f s t a tes 1. l i f e - f o r m therophyte geophyte hemicryptophyte chamaephyte nanophanerophyte 5 2. organ of perennat ion seed only corm, t u b e r , or bulb b a s a l bud caudex rhizome bud stem bud 6 3. r oo t system taproot f a s c i c l e d f i b r o u s 3 4. stem form e rec t decumbent or matted acaulescent 3 5. stem cove r ing glabrous or s l i g h t l y pubescent moderately to densely pubescent , tomentose, v i l l o u s , e t c . g landular -pubescent g l andu la r 4 6. l e a f ( l e a f l e t ) s i z e l e p t o p h y l l nanophy l l m i c r o p h y l l mesophyl l macrophyl l 5 7. g r a s s , sedge, or rush l e a f w id th (at wides t p o i n t ) l e s s than 4 mm grea te r than 4 mm does not apply 3 8. l e a f form more or l e s s p lanar t e r e t e or i n v o l u t e 2 TABLE 17. (Continued) 139 U n i t charac te r Charac ter s t a t e s Number of s t a t e s 9. l e a f d i s p l a y b a s a l r o s e t t e h o r i z o n t a l (0-15 ) i n one plane h o r i z o n t a l i n s e v e r a l planes angled Q v e r t i c a l (70 or more) 5 10. l e a f margin e n t i r e - s e r r a t e l o b e d - i n c i s e d dissected-compound 3 11. l e a f p e r s i s t e n c e deciduous evergreen 2 12. l e a f t ex tu re membranous cor iaceous ( l e a t h e r y ) c a r t i l a g i n o u s f l e s h y or succu len t 4 13. l e a f cove r ing glabrous or s l i g h t l y pubescent moderately to densely pubescent , tomentose, v i l l o u s , e t c . g landular -pubescent g l andu la r 4 14. growth form s i n g l e , unbranched f r e e l y branched, rhizomatous or spreading ce sp i to se or clump-forming 3 15. vege t a t i ve propagat ion rhizome s t o l o n v i v i p a r y none 4 16. temporal s e p a r a t i o n homogamy 6 of sexua l func t ions weak dichogamy moderate dichogamy s t rong dichogamy monoecy d ioecy TABLE 17. (Continued) 140 U n i t cha rac t e r Charac ter s t a tes Number of s t a tes 17. p o l l i n a t i o n predominant ly s e l f - 12 mechanism p o l l i n a t i o n or c le i s togamy anemophily by muscid f l i e s by s y r p h i d f l i e s by bee f l i e s by s m a l l , s h o r t -tongued bees by bumble bees by b u t t e r f l i e s by hawk moths by ants by bee t l e s by hummingbirds 18. f lower c o l o r f lowers inconspicuous 10 g reen i sh whi te p ink orange red lavender purp le b lue y e l l o w 19. f lower n e c t a r i f e r y f a i n t 4 moderate s t rong none 20. f lower odor f a i n t 4 moderate s t rong none 21. f lower type amorphic 6 haplomorphic ac t inomorphic pleomorphic s tereomorphic zygomorpaic TABLE 17. (Concluded) 141 U n i t cha rac te r Character s t a t e s Number of s t a t e s 22* propagule.-type auxochore cyc lochore p terochore pogonochore desmochore sarcochore sporochore mic rosc l e rochore megasclerochore b a l l o c h o r e 10 23. d i s p e r s a l mechanism anemochory hydrochory epizoochory endozoochory autochory 5 24. p l a n t s ca rn ivorous +\u2022\/- 2 25. p l a n t s root\" \u2022\u201e p a r a s i t e s . . \u2022_.. - . + \/-i 2 26. p l a n t s ^ - f i x i n g + \/- 2 27. es t imate of degree of vege t a t i ve p ropaga t ion 0.2 0.4 0.6 0.8 1.0 5 28. average he igh t of f.lower or i n f l . n l 29. average wid th of f l o w e r o r i n f l . n 2 30. average l eng th of f l ower or i n f l . n 3 31. t ime of f l o w e r i n g n 4 32. l eng th of f l o w e r i n g n 5 5 T o t a l 124 + In 1 142 p r o s t r a t e p l a n t s or low shrubs w i t h buds above the s o i l , but not over 2 5 cm above. Nanophanerophytes bear t h e i r buds between 0.25 and 2 m above the su r f ace . The l i f e - f o r m g e n e r a l l y f o l l o w s from the organ of pe renna t ion , but not n e c e s s a r i l y . For i n s t a n c e , Junous balticus i s rhizomatous but i s c l a s s e d as a geophyte (Clapham, T u t i n and Warburg 1962). Hemicryptophytes may perennate by means o f a b a s a l bud, a rhizome bud, or a caudex. The caudex i s understood here as the woody, p e r s i s t e n t base of an o therwise annual herbaceous stem, as i n Castitleja m i n i a t a . The l e a f or l e a f l e t s i z e c l a s s e s are a l s o taken from Raunkiaer (1934). H i s c l a s s i f i c a t i o n i s as f o l l o w s : 2 l e p t o p h y l l s - area s m a l l e r than 2 5 mm 2 nanophyl l s - 25 - 225 mm m i c r o p h y l l s - 225 - 2,025 mm2 mesophylls - 2,025 - 18,225 mm2 macrophyl l s - 18,2 25 - 164,025 mm2 The f u r t h e r d i s t i n c t i o n between graminoid leaves wider or narrower than 4 mm f o l l o w s Knight (1965). The t y p o l o g i c a l c l a s s i f i c a t i o n of f lowers i s tha t of Leppik (1957, 1968a S b ) , who desc r ibed them as succes s ive l e v e l s i n the genera l t r end of f l o r a l e v o l u t i o n . Whether L e p p i k ' s phylogeny i s v a l i d or not i s not p e r t i n e n t he re ; the f l o r a l types seem to be e c o l o g i c a l types as w e l l , i n genera l e x p l o i t i n g d i f f e r e n t groups of animal p o l l i n a t o r s . The s i x types a re : amorphic , haplomorphic , ac t i nomorph ic , p leomorphic , s te reomorphic , and zygomorphic. Amorphic f lowers are supposedly p r i m i t i v e c l u s t e r s of d i s c o l o r e d leaves wi thout 143 p a r t i c u l a r form. Haplomorphic types are c h a r a c t e r i z e d by many separate f lower pa r t s f r equen t ly s p i r a l l y arranged. A c t i n o -morphic types c o n s i s t of po lype ta lous f lowers w i t h t h e i r par t s f l a t t e n e d i n t o more or l e s s one p lane . Pleomorphic f lowers are s i m i l a r to ac t inomorphic f lowers but have c e r t a i n symmetr ica l numerate pa t t e rns of p a r t s ; e g . , t r i - , t e t r a - , or pentamery. Stereomorphic f lowers are t h r ee -d imens iona l and u s u a l l y have some s o r t of f l o r a l tube . Zygomorphic types are the h i g h l y s p e c i a l i z e d f lowers w i t h b i l a t e r a l symmetry and f r equen t ly concealed n e c t a r . Species of Compositae, subfamily T u b i f l o r a e must be scored p o s i t i v e fo r both zygomorphic and s tereomorphic f l o w e r s , and, as the cap i tu lum i s probably the u n i t o f i n s e c t p e r c e p t i o n , a l s o scored p o s i t i v e f o r , say , actinomorphy i n the case of A r n i c a spp. , or pleomorphy i n A c h i l l e a m i l l e f o l i u m . Observat ions of f l ower c o l o r and odor are n e c e s s a r i l y human and s u b j e c t i v e . However, animals (most i m p o r t a n t l y i n s e c t s ) seem to respond to the same smel ls as man, but t h e i r sense of s m e l l i s g e n e r a l l y keener , e s p e c i a l l y f o r scents tha t s i g n a l the presence of food or s exua l pa r tners (Free 19 7 0a; Kugler. 1970.; :Faegr i and van der P i j l 1971; F r i s c h 1971). S i m i l a r l y , the v i s u a l range of the spectrum i n the most important animal p o l l i n a t o r s does not d i f f e r much from man's , a l though there i s a s h i f t towards shor t e r wavelengths ( F a e g r i and van der P i j l 1971). Bees can d i s t i n g u i s h four c o l o r groups: y e l l o w , b l u e - g r e e n , b l u e , and u l t r a v i o l e t , but not red ( F a e g r i and van der P i j l 1971). However,- f lowers could not be scored f o r the presence o f u l t r a v i o l e t nec ta r guides , which are i n v i s i b l e to humans but appear to be f a i r l y common i n some angiospermous 144 groups (Daumer 1958; Lindauer 1967; E i s n e r et al. 1969; Ornduff and Mosquin 1970). The scheme of propagule types i s a simple m o r p h o l o g i c a l c l a s s i f i c a t i o n adopted from F r e n k e l (1970) and o u t l i n e d p r e v i o u s l y i n Table 13. Undoubtedly, separate c a t e g o r i e s f o r propagule types and d i s p e r s a l mechanisms i n t r o d u c e some redundancy i n t o the a n a l y s i s , but f r e q u e n t l y the a c t u a l mode of d i s p e r s a l cannot be i n f e r r e d from d i a s p o r e morphology. The only c a r n i v o r o u s s p e c i e s i n t h i s study i s Drosera rotundif'olia. Pedioularis braoteosa, Castilleja miniata, and C. parviflora are the only documented r o o t p a r a s i t e s ( K u i j t 1969). Trifolium wormskjoldiiLupinus latifolius, and Myrioa gale are the three s p e c i e s w i t h n i t r o g e n - f i x i n g r o o t nodules. Character s t a t e s were assigned to each OTU ( s p e c i e s ) of each community, and the e c o l o g i c a l d i s t a n c e (.d) c a l c u l a t e d f o r a l l p o s s i b l e species p a i r s w i t h i n a community. These d's are a l l presented as p a r t of Appendix 3, but some p o i n t s of g e n e r a l i n t e r e s t are d i s c u s s e d below. Ranges of values f o r d (0<_d<_l) a r e : S a l t Marsh 0.25 (DCSxppu) 0.62 (TW0*SCA) Wade's Bog 0.15 (RALxSCS) 0.63 (COBxVVI) Ogg's Bog 0.14 (RALxSCS) 0.63 (COBxVVI) B l a c k w a l l 0.19 (VSCxVDE) 0.62 (PDIxVVD) Meadow The most e c o l o g i c a l l y d i s t i n c t i v e s p e c i e s ( i . e . , the s p e c i e s with the h i g h e s t average e c o l o g i c a l d i s t a n c e , d) are Spergularia canadensis i n the s a l t marsh, Vaooinium vitis-idaea i n both bogs, and Veratrum viride i n the s u b a l p i n e meadow. 145 The average e c o l o g i c a l d i s t a n c e (d) b e tween\u2022a i i - s p e c i e s . o f a g i v e n community i s 0 .46 , 0 .54, 0 .52 , and 0.49 fo r the S a l t Marsh, Wade's Bog, Ogg's Bog, and B l a c k w a l l Meadow, r e s p e c t i v e l y . That i s to\" s ay , s a l t marsh spec ies a r e , on the average, the most e c o l o g i c a l l y equ iva l en t and sphagnum bog spec ies the most e c o l o g i c a l l y d i s t i n c t i v e , accord ing to t h i s a n a l y s i s . I have t r i e d to f i t these community d's i n t o the genera l scheme of t h i s i n v e s t i g a t i o n i n F i g . 151. To get some i n d i c a t i o n o f the r e l a t i v e s t rengths o f i n t e r s p e c i f i c compe t i t i on w i t h i n the d i f f e r e n t communit ies , I have c a l c u l a t e d f o r each the product of average vascu l a r p l a n t cover (C) t imes the absolu te value of the average community c o r r e l a t i o n c o e f f i c i e n t ( | r | ) t imes the community species d i v e r s i t y (# ' ) (see Sec t . I I I - I ) . I have p l o t t e d t h i s product aga ins t the average community e c o l o g i c a l d i f f e r e n c e or d i s t i n c t i v e n e s s (d ) . My i n t e r p r e t a t i o n of F i g . 151 i s tha t i t r epresen ts a p o r t i o n o f a p o t e n t i a l area upon which a l l v e g e t a t i o n types (or s t r a t a the reof ) i n the wor ld cou ld be p l o t t e d , i f subjec ted to s i m i l a r a n a l y s i s . The upper l e f t of t h i s f i g u r e ( s t rong e c o l o g i c a l s i m i l a r i t y , s t rong i n t e r s p e c i f i c compe t i t ion ) might represen t the s i t u a t i o n i n a s t ra tum of t r o p i c a l r a i n f o r e s t v e g e t a t i o n or a c h a p a r r a l shrub community; the lower r i g h t ( e c o l o g i c a l d i s s i m i l a r i t y , weak compet i t ion) by , f o r example, sphagnum bogs and Sonoran d e s e r t ; the lower l e f t ( e c o l o g i c a l s i m i l a r i t y , weak compet i t ion) by h i g h s a l i n i t y s a l t marshes, c o l d deser t s ( e g . , Great Bas in) and a r c t i c and a l p i n e t und ra ; the upper r i g h t ( e c o l o g i c a l d i s s i m i l a r i t y , s t rong compet i t ion) by t a l l grass p r a i r i e . 146 cu F i g . 151. Strength of i n t e r s p e c i f i c competition vs. average e c o l o g i c a l d i s t i n c t i v e n e s s . 100 95 90 85 80 4-75 70 65 60 55 50 45 40 35 30 chaparral shrubs? rain forest stratum? \u00bb Blackwall Meadow t a l l grass prairie? M Salt Marsh Great Basin desert? Sonoran desert? Wade's Bog Ogg's Bog H h H h 0.44 0.45 .0.46 0.47 0.48 0.49 0.50 0.51 0.52 0.53 0.54 0.55 0.56 0.57 0.58 ^\u00bb d (AVERAGE ECOLOGICAL DISTINCTIVENESS) 147 Fur ther a n a l y s i s o f the d's i n Appendix 2 cor robora tes some o f the s p e c u l a t i o n s made i n the d i s c u s s i o n ( i n Sec t . I I I - B ) on i n t e r s p e c i f i c c o r r e l a t i o n . In the bogs, d between a l l grasses and g r a s s - l i k e spec ies i s about 0 .34; between a l l shrubs , 0 .45. In the suba lp ine meadow, d f o r a l l graminoids i s 0 .35 ; f o r Compositae, 0 .39. Thus, as suggested p r e v i o u s l y , there i s g rea te r niche d i f f e r e n t i a t i o n among the shrubs i n the bogs and the meadow composites than among the graminoid s p e c i e s , and t h i s apparen t ly r e s u l t s i n s t ronger compe t i t i on among grasses and g r a s s - l i k e p l a n t s . One f i n a l p o i n t : i f v i s a measure of i n t e r s p e c i f i c c o m p e t i t i o n , and d of e c o l o g i c a l s i m i l a r i t y , and, f u r t h e r , i f the most d i s s i m i l a r spec ies compete the l e a s t (and i f the converse i s t r u e ) ; then there should be a p o s i t i v e c o r r e l a t i o n between r and d, f o r a g iven community. Such i s the case fo r a l l four communities. For a l l quadrat s i z e s , t h i s new r i s p o s i t i v e , though u s u a l l y weakly so. I t s s t r eng th inc reases i f the communities are subd iv ided i n t o a s s o c i a t i o n s ; e g . , i n the s a l t marsh, i f the spec ies are put i n t o two groups (h igh marsh v s ; mud f l a t s ) and the r computed w i t h i n these two groups, the v i s more p o s i t i v e than i f a l l spec ies are grouped toge the r . Th i s makes sense, s ince the e f f e c t s of i n t e r s p e c i f i c compe t i t i on w i l l be r e f l e c t e d more r o b u s t l y by c o r r e l a t i o n c o e f f i c i e n t s between spec ies t ha t c h a r a c t e r i s t i c a l l y grow together i n a s s o c i a t i o n s , than between spec ies of d i f f e r e n t a s s o c i a t i o n s . 147 \u00ab-I . Dominance, d i v e r s i t y , and s t a b i l i t y . P l a n t communities are u s u a l l y mixtures of unequa l ly abundant species tha t form a p r o g r e s s i o n from dominant through in te rmed ia te to r a r e s p e c i e s . W i t h i n a g iven community i t i s p o s s i b l e to es t imate the s u b d i v i s i o n o f n iche space by assuming tha t the importance values of the species c o n s t i t u t i n g the community are express ions of t h e i r r e l a t i v e n iche s i z e s . I f the species are then arranged i n a sequence from most to l e a s t important and the s p e c i e s ' p o s i t i o n s i n t h i s sequence are then p l o t t e d aga ins t the l o g of t h e i r r e s p e c t i v e importance v a l u e s , a curve of c h a r a c t e r i s t i c genera l form r e s u l t s (Whi t taker 1965, 1969, 1970a). This has been done f o r each o f the four study communities and the curves are presented i n F i g . 15 2A-D. Note tha t the curves are roughly s h a l l o w l y s igmoid and tha t the s lope decreases i n order from A to D i n the f i g u r e . The s igmoid shape r e f l e c t s the f ac t t ha t each community has a few dominant spec ies and a few r a re s p e c i e s , p lus a l a r g e number of spec ies of in te rmedia te abundance. Compet i t ion theory holds tha t species tend to evolve toward n iche d i f f e r e n t i a t i o n and r e d u c t i o n of i n t e r s p e c i f i c c o m p e t i t i o n , and the r e s u l t a n t - d i v i s i o n of n iche space i s manifested i n the forms of such dominance -d ive r s i t y cu rves . The steepness of the curve f o r the f l o r i s t i c a l l y poor s a l t marsh agrees w i t h W h i t t a k e r ' s obse rva t ions tha t steep s lopes are d e r i v e d from samples o f low d i v e r s i t y communities of r i g o r o u s p h y s i c a l environments. The gradua l moderation of the s lope found i n the curves f o r the bogs and the suba lp ine meadow i s c o n s i s t e n t w i t h the ' i g . 15 2. Dominance-diversity c u r v e s , f o r the four study communities. 149 i n c r e a s i n g d i v e r s i t y of these communit ies. The more equal s u b d i v i s i o n of n iche space i n the communi-t i e s of h ighe r d i v e r s i t y i s a l s o i n d i c a t e d by a community index of dominance, e i t h e r the DI of McNaughton ( i n McNaughton and Wolf 1970) or the A of Simpson (1949) . DI = 100(y \u201e\/Y) where y-^ 2 i - s sum of the importance values o f the two most abundant spec ies in . the community and Y i s the t o t a l importance n 2 value of a l l spec ies i n the community; A. = ETT where TT i s the ^ 1 n n importance value of the nth spec ies i n the community. In e i t h e r case , the h ighe r the dominance i n d e x , the g rea t e r the c o n c e n t r a t i o n of abundance i n the fewer s p e c i e s . Ind ices of dominance fo r the four communities are l i s t e d i n Table 18. The two i n d i c e s g ive very s i m i l a r r e s u l t s and are l i n e a r l y r e l a t e d i n t h i s ins tance ( F i g . 153) . TABLE 18. Number of s p e c i e s , two i n d i c e s of dominance, and an index of d i v e r s i t y fo r the four study communities. N = number of spec ies sampled; DI = McNaughton's dominance i n d e x ; X = Simpson's dominance i n d e x ; H ' = the Shannon-Wiener index of d i v e r s i t y . Community N DI XxlO H ' S a l t Marsh 18 31.6 10.1 2.50 Wade's Bog 28 28. 7 8.2 2. 71 Ogg's Bog 41 27.2 7.7 3.03 B l a c k w a l l 45 24. 6 6.3 3.13 Meadow 150 a, Fig . 153. DI (McNaughton1s index of dominance), N (number of species i n a community) vs. H' (an index of d i v e r s i t y ) , A. (Simpson's dominance index). I 1 1 1 1 1 } 1 \\ 1 1 1 1.0 1.4 1.8 2.2 2.6 3.0 3.2 H' 0.0 0.2 0.4 0.6 0.8 1.0 1.1 X 151 The matter of d i v e r s i t y r e q u i r e s f u r t he r comment. Species d i v e r s i t y i s a concept vague enough and of s u f f i c i e n t l y broad a p p l i c a t i o n tha t i t can be (and has been) v a r i o u s l y def ined ( c f . H u r l b e r t 1971). D i v e r s i t y i s o f ten equated w i t h s imple spec ies r i c h n e s s ; i . e . , the number of spec ies present i n a g iven a rea . However, as s t r i c t l y d e f i n e d , species d i v e r s i t y i s a f u n c t i o n of the numbers of species present ( spec ies r i c h n e s s ) and the evenness w i t h which the t o t a l number of i n d i v i d u a l s (or t o t a l biomass) i n a community i s d i s t r i b u t e d among these spec ies ( spec ies e q u i t a b i l i t y ) . D i v e r s i t y and r i c h n e s s are u s u a l l y p o s i t i v e l y c o r r e l a t e d , but not n e c e s s a r i l y so (Johnson and Raven 1970; H u r l b e r t 1971). In the four study communit ies, the number of spec ies per community i s l i n e a r l y r e l a t e d to the spec ies d i v e r s i t y o f these communities ( F i g . 153) . D i v e r s i t y can be c a l c u l a t e d by a number of formulas (Mcintosh 1967a; L loyd et al. 1968; H u r l b e r t 1971; Whi t t aker 1972). The most commonly used i s the Shannon-Wiener i n f o r m a t i o n i n d e x , H ' = - p ^ l o g p ^ , where H ' i s the species d i v e r s i t y and p^ i s the p r o p o r t i o n of i n d i v i d u a l s (or biomass) i n the i - t h s p e c i e s . Th i s f u n c t i o n i s d e r i v e d from in fo rma t ion theory accord ing to the l o g i c t h a t , s i nce d i v e r s i t y i s a f u n c t i o n of the number of p o s s i b l e i n t e r a c t i o n s i n a system and the degree to which they are s t r u c t u r e d (Johnson and Raven 1970), d i v e r s i t y should inc rease as the number of spec ies inc reases and as the d i s t r i b u t i o n o f i n d i v i d u a l s i n t o spec ies becomes more even. The concept of d i v e r s i t y has proven most u s e f u l i n i n t e r p r e t a t i o n s of l a r g e - s c a l e p o p u l a t i o n a l pa t te rns i n na ture . For example, spec ies d i v e r s i t y o f both f l o r a s and faunas has 152 repea ted ly been observed to inc rease i n a g rad ien t from c o l d c l ima te s toward the warm t r o p i c s and from h i g h e l e v a t i o n s downward ( F i s c h e r 1960; MacArthur 1965; P ianka 1966; Baker 1970). Rough p o s i t i v e c o r r e l a t i o n s have been made between the area of i s l a n d s and the d i v e r s i t y of t h e i r b i o t a (MacArthur and Wi l son 1967; Johnson et al. 1968; Johnson and Raven 1970, 1973; S i m b e r l o f f 1970). Pa t te rns of d i v e r s i t y have been s tud i ed i n r e l a t i o n to w i t h i n - and between-habi ta t environmenta l he t e ro -gene i ty (Johnson et al. 1968; Whi t t aker 1970a, 1972) and as adjuncts to vege t a t i on c l a s s i f i c a t i o n s w i t h i n c e r t a i n geograph ica l areas (Whi t taker 1965; Monk 1967; A u c l a i r and Goff 1971; d e l Mora l 1973). More s p e c i a l i z e d i n v e s t i g a t i o n s have revea led both d i r e c t . (ednnell and Or ias 1964; MacArthur 1969; Singh and M i s r a 1969) and i nve r se (McNaughton 1968; Margalef 1969; Whi t taker 1972) c o r r e l a t i o n s between d i v e r s i t y and p r o d u c t i v i t y . In p l an t communit ies , attempts to produce c o n s i s t e n t c o r r e l a t i o n s between d i v e r s i t y and a number of environmental f a c t o r s such as mois ture (Monk 1967; A u c l a i r and Goff 1971; Whi t taker 1972) , a l t i t u d e (Whi t taker and N i e r i n g 1965; Daubenmire 1970), s o i l f e r t i l i t y and pH (Loucks 1962; Monk 1967), have not been s u c c e s s f u l . D i v e r s i t y i s g e n e r a l l y reduced by a i r p o l l u t i o n (Gordon and Gorham 1963; G i l b e r t 1965; Skye 1968; Nash 1972), gamma i r r a d i a t i o n (Woodwell 1967; Woodwell and Whi t t ake r 1968) , and ove rg raz ing (Whi t taker 1972), but Harper (19 69) has demonstrated tha t l i g h t or moderate g r a z i n g may a c t u a l l y inc rease d i v e r s i t y . D i v e r s i t y o f t en inc reases w i t h success ion (Margalef 1963; Monk 1967; Odum 1969; Reiners et al. 1971) , but i n some cases 153 decreases from l a t e s u c c e s s i o n a l stages to the c l imax (Loucks 1970; A u c l a i r and Goff 1971). Reiners et al. (1971) fo l lowed changes i n d i v e r s i t y dur ing success ion on g l a c i a l moraines of known age i n G l a c i e r Bay, A l a s k a . They found tha t an i n i t i a l r a p i d inc rease i n d i v e r s i t y dur ing the f i r s t 10 0 years was fo l l owed by a more g radua l inc rease to a maximum i n the muskeg steady s ta te (c l imax) at 200 years p lus (up to 1500 y e a r s ? ) . Success ion at G l a c i e r Bay proceeds from bare ground to shrub t h i c k e t s to a l d e r and cottonwood f o r e s t s t o S i t k a spruce and hemlock f o r e s t s and e v e n t u a l l y v i a p a l u d i f i c a t i o n to muskeg. Inasmuch as I i m p l i c a t e d p a l u d i f i c a t i o n of c o a s t a l f o r e s t i n the format ion o f the Tof ino area sphagnum bogs, one would expect tha t Wade's Bog, the most s u c c e s s i o n a l l y advanced, would have a h i g h e r d i v e r s i t y than Ogg's Bog. However, such i s not the case (Table 18 ) . There i s l i t t l e agreement as to the causes of d i f f e r e n c e s and g rad ien t s i n species d i v e r s i t y . Attempts have been made to e x p l a i n pa t te rns of d i v e r s i t y i n terms o f t ime a v a i l a b l e f o r e v o l u t i o n (Sanders 1969; Baker 1970), s p a t i a l he te rogene i ty of the p h y s i c a l environment (Ashton 1969; MacArthur 1969; Richards 1969; Whi t t aker 1972), i n t e r s p e c i f i c compe t i t i on (Dobzhansky 1950; M i l l e r 1969; Whi t t aker 1969) i n c l u d i n g a l l e l o p a t h y '.(Whittaker 1970b; Whi t t aker and Feeny 1971) , p r eda t i on pressure (Paine 1966; Spight 1967; Harper 1969; Janzen 1970) , hos t -p a r a s i t e i n t e r a c t i o n s (Barbehenn 1969), c l i m a t i c s t a b i l i t y or environmental p r e d i c t a b i l i t y (Sanders 19 68; S lobodk in and Sanders 1969), p r o d u c t i v i t y (Conne l l and Or ias 1964; McNaughton 1968; Singh and M i s r a 1969) , r e l a t i v e r a t e s of 154 of s p e c i a t i o n (Simpson 1964, 1969; S t e h l i et al. 1969), and va r ious combinations of the above (see P ianka 1966; Baker 1970; and Johnson and Raven 1970 fo r genera l d i s c u s s i o n s ) . This b r ings us to the ques t ion of the r e l a t i o n s h i p between d i v e r s i t y and s t a b i l i t y . As used he re , s t a b i l i t y does not r e f e r to the envi ronmenta l s t a b i l i t y o f the p rev ious paragraph, but i s to be understood r a t h e r as the s t a b i l i t y through time of a p l a n t community type . More p r e c i s e l y , the s t a b i l i t y of a community may be def ined as i t s r e s i s t a n c e to e x t e r n a l p e r t u r b -a t ions and consequent p e r s i s t e n c e i n time (Odum 1969; Pres ton 1969). Communities of g rea te r d i v e r s i t y are then s a i d to be more s t ab l e (Hutchinson 1959; Odum 1969; Whi t taker and Woodwell 1972; see Woodwell and Smith 1969). Given t h i s , and the f a c t tha t each o f the four study communities has a d i f f e r e n t species d i v e r s i t y , can I v a l i d l y c l a i m t h a t , f o r example, the s a l t marsh i s the l e a s t s t a b l e , and the subalpine meadow the most s t ab l e o f these communities? I t i s important to s p e c i f y the time s ca l e i n v o l v e d . In most d i s c u s s i o n s of d i v e r s i t y and s t a b i l i t y , a long- te rm ( e v o l u t i o n a r y ) view r a the r than a sho r t - t e rm ( s u c c e s s i o n a l ) view i s at l e a s t i m p l i c i t . Thus, we are d e a l i n g w i t h the p e r s i s t e n c e of a community type i n e v o l u t i o n a r y t i m e ; s u c c e s s i o n a l t rends are subl imated (McCullough 1970). When one speaks of the s t a b i l i t y of a community, one must i n c l u d e the s u c c e s s i o n a l phases of tha t community. For example, the e v o l u t i o n a r y future of the s a l t marsh community type i n v o l v e s both the p ioneer mud f l a t stages and the advanced h igh marsh, as w e l l as the degene ra t ive , eroded stages of the marsh ( c f . 155 R e d f i e l d 1972); heather meadows i n England ( B a r c l a y - E s t r u p 1970, 1971; B a r c l a y - E s t r u p and Gimingham 1969) and sugar maple f o r e s t s i n Wiscons in (Loucks 1970) evolve as systems o f seres c y c l e d by p e r i o d i c , random pe r tu rba t i ons ( c f . Watt 1947). The e v o l u t i o n a r y consequence of the proposed d i r e c t r e l a t i o n s h i p between d i v e r s i t y and s t a b i l i t y should be a p o t e n t i a l l y se l f -augment ing inc rease i n d i v e r s i t y (Hutchinson 1959; Whi t t aker 1969; Whi t t ake r and Woodwell 1972); i . e . , i f d i v e r s i t y inc reases s t a b i l i t y , there should be s e l e c t i o n f o r i n c r e a s e d d i v e r s i t y , which i n t u r n confers g rea te r s t a b i l i t y , e t c . Such a s e l e c t i v e mechanism r e q u i r e s tha t the p l a n t community i t s e l f be a u n i t of e v o l u t i o n (Whi t taker and Woodwell 1972). Th i s i s t h e o r e t i c a l l y f e a s i b l e , as to be a s e l e c t i v e uni t ' an e n t i t y need on ly have v a r i a t i o n , r e p r o d u c t i o n , and h e r i t a b i l i t y (Lewontin 1970). Genes, c e l l s , organs , organisms, p o p u l a t i o n s , spec ies , communit ies , and ecosystems f u l f i l l these requirements and thus a l l can evolve (Lewontin 1970). H e r i t a b i l i t y w i l l be g r e a t e r , and the r a t e of e v o l u t i o n f a s t e r , i n those e n t i t i e s w i t h the t i g h t e s t i n t e r n a l o r g a n i -z a t i o n (Lewontin 1970; Stebbins and Lewontin 1972). The c loseness of the o r g a n i z a t i o n decreases con t inuous ly i n the p r o g r e s s i o n : gene, c e l l , o rgan , organism, p o p u l a t i o n , s p e c i e s , community, ecosystem (Baker 1966a); or \" . . . c h a n c e p lays a grea ter r o l e at the ecosystem end of the p rog re s s ion\" (Langford and B u e l l 1969). A p l an t community may be viewed as a more or l e s s i n t e g r a t e d assemblage of spec ies r e s u l t i n g from \" n a t u r a l s e l e c t i o n ope ra t i ng between groups o f organisms and ad jus t i ng t h e i r mutual s t r a t e g i e s to permit i n c r e a s i n g d i v e r s i t y and more e f f i c i e n t environmenta l e x p l o i t a t i o n . . . \" (Harper 19 69) . Thus, a community has c h a r a c t e r i s t i c s such as growth, m a t u r i t y , homeostas is , and energy f l u x tha t correspond to those o f o ther b i o l o g i c a l e n t i t i e s . R e l a t i o n s h i p s such as h e r b i v o r e -p l a n t ( E h r l i c h and Raven 1964; Breedlove and E h r l i c h 1968; Janzen 1969, 1970; B e a t t i e , Breed love , and E h r l i c h 1973) , p a r a s i t e - h o s t (Person 1959; Chabora and P imen te l 1970), p reda tor -p rey ( H o l l i n g 196 5 ) , and p o l l i n a t o r - p l a n t i n t e r a c t i o n s ( F a e g r i and van der P i j l 1971) , symbioses (Janzen 1966; Henry 19 1967; Sco t t 1969), and p o s s i b l e synergisms (Baker , Cruden, and Baker 1971; Po ja r 1973b) w i l l by gene t ic feedback and p o p u l a t i o n r e g u l a t i o n inc rease the complexi ty and i n t e g r a t i o n o f the ecosystem, i n c r e a s i n g i t s homeostasis and presumably i t s s t a b i l i t y (Brock 1967; P imente l 1968; c f . H a i r s t o n et al. 1960 ; Murdoch 1966 ;\u2022* S lobodk in et al. 1967 , and E h r l i c h and B i r c h 1967). Note tha t t h i s does not r e q u i r e an organismic (Clements 19 36) concept of the p l an t community, nor does i t s t r i c t l y conform to the n o t i o n of a community as merely a f o r t u i t o u s assemblage of i n d i v i d u a l spec ies (Gleason 1926, 1939). Ra ther , t h i s approach appropr ia t e s elements of both t r a d i t i o n a l l y opposed ( c f . Whi t t ake r 1962, 19 67; Daubenmire 19 66; Mcin tosh 1967b, 1970, 1972; Langford and B u e l l 1969) schools of synecology. Species do evolve toward n iche and h a b i t a t d i f f e r -e n t i a t i o n , r e d u c t i o n o f c o m p e t i t i o n , and i n d i v i d u a l i t y o f d i s t r i b u t i o n (Whi t taker 1967, 1969, inter alia). However, superimposed on the spec ies e v o l u t i o n i s the h igher order 157 e v o l u t i o n of the community toward inc reased homeostasis (Odum 1969). I n t e r s p e c i f i c compe t i t i on c o n t i n u a l l y a p p l i e s a d i v i s i v e force to aggregat ions of s p e c i e s ; t h i s i s countered by the cohesive or c e n t r i p e t a l fo rce of s e l e c t i o n f o r community homeostasis and s t a b i l i t y . Now, a l l t h i s begs the q u e s t i o n , Does inc reased d i v e r s i t y really i nc rease s t a b i l i t y ? I can accept the argument tha t g rea te r d i v e r s i t y w i l l inc rease community s t a b i l i t y i n the face of a b i o l o g i c a l p e r t u r b a t i o n . For example, a monoculture (say a Midwestern c o r n f i e l d ) would be most s u s c e p t i b l e to an outbreak of a v i r u l e n t pathogen or p a r a s i t e or an h e r b i v o r e epidemic . Fur thermore, the more d i v e r s e a community i s , the b e t t e r i t should accommodate l e s s c a t a s t r o p h i c , commonplace occurrences such as death and replacement o f i n d i v i d u a l s , windthrow, d i s tu rbance by g r a z i n g or burrowing a n i m a l s , and en t ry of new t a x a . On the other hand, I t h i n k tha t p h y s i c a l f a c t o r s tha t cou ld d i s r u p t a community ( f i r e , g l a c i a t i o n , drought , r i s e and f a l l of the water t a b l e , c l e a r - c u t t i n g , p o l l u t i o n , b u l l d o z e r s , e t c . ) would act more i n d i s c r i m i n a t e l y , r e g a r d l e s s of community d i v e r s i t i e s . P l a n t communities have been shown to be s e n s i t i v e to the e f f ec t s of i o n i z i n g r a d i a t i o n (Woodwell 1967, 1970; Woodwell and Whi t t aker 1968), p e s t i c i d e s (Cant lon 1969; Mosser et al. 1972) , and e u t r o p h i c a t i o n (Hurd et al. 1971), as w e l l as to more heavy-handed treatments such as c l e a r - c u t t i n g and h e r b i c i d e a p p l i c a t i o n (Bormann et al. 1968; L ikens et al. 1970). The mechanisms of r ecovery from such d i s tu rbances have on ly begun to be s t ud i ed (L ikens et al. 1969; L ikens and Bormann 1972 158 Marks and Bormann 19 72) . The s tud ies have been done e i t h e r wi thou t the r e l a t i o n s h i p between d i v e r s i t y and s t a b i l i t y i n mind, or they are too e q u i v o c a l ( c f . Rosenzweig 19 71 and M c A l l i s t e r et al. 1972) fo r g e n e r a l i z a t i o n . Most o ther data o f fe red as evidence are l a r g e l y phy togeograph ica l and n e c e s s a r i l y anecdo ta l (see MacArthur and Wi l son 1967; S i m b e r l o f f 19 70; MacArthur 19 72; Diamond 19 73) . A recen t experiment (Hurd et al. 1971) more d i r e c t l y apropos the problem i n d i c a t e d t h a t , at l e a s t a t the l e v e l of pr imary producers ( p l a n t s ) , s t a b i l i t y was p o s i t i v e l y c o r r e l a t e d w i t h d i v e r s i t y . However, these r e s u l t s have been quest ioned ( c f . Harger et al. 1972; H o l t et al. 1972) , and a n o t - a l t o g e t h e r -comparable experiment by H a i r s t o n et al. (19 68) concluded tha t \" . . . m u c h more exper imenta l and o b s e r v a t i o n a l work i s necessary before the nature of any f u n c t i o n a l r e l a t i o n s h i p between d i v e r s i t y and s t a b i l i t y can be c la imed w i t h conf idence\" I agree. 159 J . Index o f p o t e n t i a l r ecombina t ion . The v a r i e t y of p l a n t gene t i c systems i n nature r e f l e c t s t h e i r s u s c e p t i b i l i t y to h e r e d i t a b l e v a r i a t i o n , a d a p t a t i o n , and n a t u r a l s e l e c t i o n ( D a r l i n g t o n 19 39; Grant 1958). That the d i v e r s i t y of these systems i s not random suggests tha t c o r r e l a t i o n s e x i s t between c e r t a i n k inds o f r ep roduc t ive s t r a t e g i e s and p a r t i c u l a r k inds o f gene t ic systems. The nonrandom v a r i a t i o n manifests i t s e l f i n a number of r ep roduc t i ve methods tha t embody a c h a r a c t e r i s t i c combinat ion or syndrome o f p o s i t i v e l y c o r r e l a t e d c h a r a c t e r s , m o r p h o l o g i c a l , p h y s i o -l o g i c a l , c y t o l o g i c a l , and e c o l o g i c a l (Ornduff 1969). For i n s t a n c e , herbaceous spec ies tha t are w i n d - p o l l i n a t e d and wind-d i spe r sed tend to have: reduced, inconspicuous f l o w e r s , o f ten u n i s e x u a l o r , - ' i f h e r maphrod i t i c , s t r o n g l y dichogamous ; f lowers aggregated i n t i g h t , many-flowered i n f l o r e s c e n c e s ; anthers exser ted and w i t h c o p i o u s , d r y , l i g h t ' p o l l e n ; exser ted stigmas w i t h a l a rge surface a r ea ; s m a l l , numerous propagules of ten beset w i t h winged, h a i r l i k e , or plumose appendages; and l a r g e , clumped popu la t ions occupying open h a b i t a t s ( P i j l 19 69; Whitehead 1969; F a e g r i and van der P i j l 1971). As another , l e s s i n c l u s i v e example, morpho log ica l dimorphism of f l o r a l pa r t s ( i . e . , d i s t y l y ) and d i a l l e l i c s e l f - i n c o m p a t i b i l i t y are two independent fea tures o f breeding systems g e n e r a l l y found toge ther presumably becauise they produce, i n combina t ion , an inc reased e f f i c i e n c y of c r o s s - p o l l i n a t i o n and c r o s s -f e r t i l i z a t i o n ( V u i l l e u m i e r 1967). I t i s p o s s i b l e to i n v e s t i g a t e the gene t ic systems o f 160 p l a n t s i n an e v o l u t i o n a r y c o n t e x t , keeping i n mind tha t these systems f u n c t i o n p r i m a r i l y i n ba l anc ing constancy and v a r i a -b i l i t y i n r ep roduc t ion (Stebbins 19 50; Grant 19 50) . The e f f i c i e n c y of t h i s compromise between immediate f i t n e s s and long-range f l e x i b i l i t y i s best eva lua ted by c o n s i d e r a t i o n of the c o n t r o l of r ecombina t ion . R e l a t i v e m u t a t i o n a l r a t e s o f h ighe r p l a n t spec ies are too p o o r l y known to assess m u t a t i o n a l c o n t r i b u t i o n s to gene t i c v a r i a b i l i t y , so i t i s necessary i n t h i s i n v e s t i g a t i o n to assume tha t the c h a r a c t e r i s t i c mutat ion ra tes of s e x u a l l y reproducing h ighe r p l a n t s are approximate ly the same, or at l e a s t of the same order of magnitude. The p o t e n t i a l performance o f d i f f e r e n t recombinat ion systems i n the genera t ion and maintenance of gene t ic v a r i a b l i t y can then be compared. In the f o l l o w i n g d i s c u s s i o n I have o u t l i n e d a scheme to evalua te the recombinat ion p o t e n t i a l of each species i n the study communities. Many o f the ideas are modi f ied from papers by F r y x e l l (1957) , Grant (1958) , Baker (1959) , C a r l q u i s t (1966) , Mosquin (1966) , and Ornduff (1969). The approach i n v o l v e s n u m e r i c a l l y s c a l i n g va r ious c y t o l o g i c a l , m o r p h o l o g i c a l , and e c o l o g i c a l components of the gene t i c system and combining values f o r a l l components i n a f i n a l Index of P o t e n t i a l Recombination ( I . P . R . ) fo r each s p e c i e s . W i t h i n a g iven component ( e g . , l e v e l of p l o i d y ) , there i s u s u a l l y a number of s t a t e s ( e g . , o c t o p l o i d and above, h e x a p l o i d , t e t r a p l o i d , d i p l o i d ) tha t have^been.- arranged ' in,.order;.6.f' i n c r e a s i n g p o t e n t i a l r ecombina t ion . 1. Length of gene ra t i on . 161 \"The recombinat ion system regu la t e s the genera t ion of v a r i a b i l i t y by r e s t r i c t i n g the types o f gametes produced and the types o f zygotes formed\" (Grant 1958). This r e g u l a t i o n i s imposed dur ing each success ive sexua l gene ra t ion . The genera t ion l eng th o f d i f f e r e n t p l a n t spec ies i s h i g h l y v a r i a b l e , so the amount of recombinat ion generated per u n i t t ime per spec ies w i l l be a f u n c t i o n of the genera t ion l e n g t h . A l l o ther t h ings be ing e q u a l , annuals or s h o r t - l i v e d p e r e n n i a l s should r e a l i z e more recombinat ion per u n i t time than l o n g - l i v e d p e r e n n i a l s and woody p l a n t s . A l s o , l engthy l i f e c y c l e s cou ld r e s u l t i n c r o s s i n g between parents and o f f s p r i n g and thus inc rease inbreed ing (Mosquin 1966). 1 - woody p l an t s 2 - l o n g - l i v e d herbaceous pe renn ia l s 3 - s h o r t - l i v e d herbaceous pe renn i a l s 4 - annuals 2. B a s i c chromosome number. Independent assortment of the chromosomes at meios i s w i l l recombine genes borne on separate chromosomes. The grea te r the number of chromosomes, the g rea te r the p o t e n t i a l recombinat ion (Grant 1958, 19 71; Stebbins 19 71b) . Al though the number of p o s s i b l e chromosome combinations inc reases e x p o n e n t i a l l y w i t h h a p l o i d chromosome number, I have chosen to reduce the p o s s i b i l i t i e s to a much s m a l l e r s c a l e , namely tha t def ined by 162 x (bas ic chromosome number)\/2. The range of the sca l e i n t h i s study would be 3 (x = 6 f o r T r i g l o o h i n m a r i t i m u m and P l a n t a g o m a o r o o a r p a and P. m a r i t i m a ) to 8.5 (x = 17 fo r N e p h r o -p h y l l i d i u m o r i s t a - g a l l i ) . 3. R e l a t i v e chromosome l e n g t h . Recombination o f l i n k e d genes i s promoted by a h igh chiasma frequency. I l a c k the data to c a l c u l a t e a recombina t ion index (Stebbins 1950) f o r each s p e c i e s , but w i l l use r e l a t i v e chromosome l eng th (based on camera l u c i d a drawings ( F i g s . 14-110) of microsporocyte m e i o t i c chromosomes) as an express ion of c rossover potential::,.--since the. number of chiasmata tends to inc rease w i t h chromosome l eng th (Swanson 1957; Stebbins 1971b). Q u a l i t a t i v e es t imates of chromosome leng th have been sca led from 1 to 7. 1 - e g . , J u n o u s spp., C a r e x spp. 3 L u p i n u s l a t i f o l i u s . 2 - e g . , V a o o i n i u m s p p . , G a u l t h e r i a s h a l l o n , V a l e r i a n a s i t c h e n s i s 3 V e r o n i c a c u s i c k i i , E p i l o b i u m a l p i n u m . 3 - e g . , Ledum g r o e n l a n d i c u m 3 L i n n a e a b o r e a l i s , G l a u x m a r i t i m a , S p e r g u l a r i a c a n a d e n s i s 3 C a s t i l l e j a m i n i a t a . 4 - e g . , S a l i o o r n i a v i r g i n i c a 3 S a n g u i s o r b a o f f i c i n a l i s 3 A r n i c a m o l l i s 3 P h l o x d i f f u s a . 5 - e g . , P l a n t a g o m a r i t i m a 3 P u o c i n e l l i a p u m i l a 3 A p a r g i d i u m b o r e a l e 3 G e n t i a n a d o u g l a s i a n a 3 S i l e n e p a r r y i . 6 - e g . , D e s o h a m p s i a o e s p i t o s a , G e n t i a n a s c e p t r u m 3 F e s t u o a v i r i d u l a 3 C l a y t o n i a l a n c e o l a t a , S e n e c i o t r i a n g u l a r i s . 7 - e g . , M a i a n t h e m u m d i l a t a t u m , E l y m u s g l a u o u s . 4 . L e v e l of p l o i d y . 16 3 P o l y p l o i d y i s cons idered to promote and preserve pheno-t y p i c u n i f o r m i t y i n popu la t ions wh i l e r e t a r d i n g species e v o l u t i o n a r y change (Stebbins 1950, 1971b; Mosquin 1966; De Wet 1971a). The d i s r u p t i v e e f f e c t s of muta t ion , r ecombina t ion , and s e l e c t i o n tend to be buf fe red by polysomic i n h e r i t a n c e (Dawson 1962; Stebbins 1971b). Presumably the b u f f e r i n g e f f e c t inc reases w i t h h ighe r p l o i d y l e v e l s . P o l y h a p l o i d y ( f u n c t i o n a l hap lo idy r e s u l t i n g from a r e v e r s a l o f p o l y p l o i d y ) i s qu i t e p o s s i b l e i n f l o w e r i n g p l a n t s ( c f . Raven and Thompson 1964; Jones 1970; Ornduff 1970a; Stebbins 1970b; De Wet 1971b; Anderson 1972), but i s not r e a l l y important i n t h i s a n a l y s i s , s ince the p l o i d y s ta tus quo i s under c o n s i d e r a t i o n , not i t s mode of o r i g i n . 1 - o c t o p l o i d and up 2 - h e x a p l o i d 3 - t e t r a p l o i d 5 - d i p l o i d The breeding system of f l o w e r i n g p l a n t s , through a v a r i e t y o f morpho log ica l and p h y s i o l o g i c a l mechanisms, c o n t r o l s the amount of o u t c r o s s i n g i n spec ies popu la t ions and consequently e x e r c i s e s a powerful c o n t r o l >0.05). There are some other i n t e r e s t i n g o b s e r v a t i o n s . Species tha t appear to be predominant ly autogamous have an average I . P . R . of 4 0 . 3 , compared w i t h an average fo r a l l spec ies of 47 .0 . The inbreeders thus have more r e s t r i c t e d recombina t ion systems than the ou tbreeders , a cco rd ing to t h i s a n a l y s i s . Autogamous s p e c i e s , because of t h e i r r e s t r i c t e d recombina t ion 175 TABLE 19. Index of P o t e n t i a l Recombination ( I . P . R . ) . f e n e r a t i o n l eng th . . 2 B a s i c chromosome number\/2. 3 R e l a t i v e chromo-some l e n g t h . ' 'Leve l of p l o i d y . 5Dichogamy. 6 S p a t i a l s epa ra t i on of s exua l organs. 7 H e t e r o s t y l y . 8 S e x u a l r e p r o d u c t i o n . 9Mode o f p o l l i n a t i o n . 1 0 C l e i s t o g a m y . 1 f o m p a t i b l i t y . 1 2 Gei tonogamy. 1 3 D i s p e r s i b i l i t y . 1 ^ P o p u l a t i o n s i z e and s t r u c t u r e . 1 5 E c o t y p i c v a r i a t i o n . 1 6 H y b r i d i z a t i o n . 0 \u2022 a Species I 2 3 5 6 - . \u2022 - - -\u2022 \u2022. \u2022 - S a l t Marsh Des champ si d~ cesp it os.d 2 3.5 6 3 2 2 F e s t u c a r u b r a 2 3.5 6 2 2 2 T r i g l o c h i n - m a r i t i m u m 2 3 3 1 3 1 S a l i c o r n i a virgih'icd 2 4.5 4 3 3 1 P l a n t a g o m a r i t i m a 2 3 5 5 2 . 5 2 J u n c u s b a l t i c u s 2 5 1 1 3 2 C a r e x lyngbyei 2 5 1 1 3 3 G l a u x m a r i t i m a 2 7 . 5 3 5 1 1 P o t e n t i l l a p a c i f i c a 2 3.5 4 3 2 1.5 A g r o s t i s e x a r a t a 2 3.5 6 3 1.5 1.5 S t e l l a r i a h u m i f u s a 2.5 6 . 5 4 5 1.5 1.5 T r i f o l i u m w o r m s k j o l d i i 2 4 3 3 3 1.5 S c i r p u s c e r n u u s 3.5 5 2 2 2.5 1 P u c c i n e l l i a p u m i l a 2.5 3.5 5 2 1 1 S p e r g u l a r i a c a n a d e n s i s 4 4.5 3 3 1 1 D i s t i c h l i s s p i c a t a 2 2.5 3 1 3 5 H o r d e u m b r a c h y a n t h e r u m 2 3.5 5 3 1 1 L i l a e o p s i s o c c i d e n t a l i s 2 5.5 4 3 1 1 Wade * s Bog C a r e x p l u r i f l o r a 2 5 1 3 3 3 Apargidium'' b o r e a l e 2 4.5 5 5 3 1.5 A g r o s t i s a e q u i v a l v i s 2 3.5 5 5 1.5 1 C a r e x o b n u p t a 2 5 1 1.5 3 3 S a n g u i s o r b a o f f i c i n a l i s 2 3.5 4 3 1 1 K a l m i a p o l i f o l i a 1 6 3 5 1.5 2 V a c c i n i u m oxy c o c c u s 1.5 6 2 3 1.5 1.5 C a r e x c a n e s c e n s 2 5 1 2 3 3 D r o s e r a r o t u n d i f o l i a 3 5 3 5 1 1 Ledum g r o e n l a n d i c u m 1 6.5 3 5 1.5 2 T r i e n t a l i s a r c t i c a 2.5 11 1 1 1 2 E m p e t r u m n i g r u m 1 6 . 5 3 5 3 5 M y r i c a g a l e 1 4 2 1 3 5 T o f i e l d i a g l u t i n o s a 2 7.5 4 5 1.5 2 G e n t i a n a d o u g l a s i a n a 4 6.5 5 5 2 1.5 L i n n a e a b o r e a l i s 1.5 4 3 3 1.5 1 TABLE 19. (Continued) 176 1 6 7 8 9 10 1 1 1 2 13 Ik 15 16 E=IPR-1 S a l t Marsh :\u2022.. ~? 5 -~ a -s6 \u2022 . -P3 7 3 3 4 5 3 58.5 r.. \u2022 -. 2;5 6 3 7 3 2 3 4 2 51.0 I 3.5 . 7 3 2 2 3 5 2 1 42. 5 I 4 6 3 2 2 3 4 1.5 1 45.0 l 5 6 3 1 2 3 4 2 1 47.5 I 2 6 3 1 2 4 3 3 3 42.0 I 2.5 7 3 1 2 2 3 1.5 2 40. 0 l 3 2.5 3 1 2 3 2 3 1 41.0 i 3 4 4 5 2.5 2 2 3 2 44 . 5 l 3 6 3 7 3 3 2 4 2 51.5 l 3 3 3 1 2 3 3 2 1 42.0 i 3 5 4 7 3 2 2 4 1 48. 5 I 5 4 2 . 5 1 2 2 3 3 1 40 . 5 l 5 3 2 2 2 2 2 2 2 38 . 0 I 5 1 1.5 1 2 4 1 2 1 36.0 I 2 7 3 7 3 2 1 2 1 45 . 5 I 3.5 3.5 2 1 2 2 1 4 5 40 . 5 I 3 3 3 1 2 2 1 2 1 35.5 range: 35.5_- 58.5 unweighted x = 4 3.8 weighted x = 47.2 Wade's Bog 1 2.5 7 3 1 2 2 4 1 3 43. 5 1 5 4 4 7 3 5 5 2 1 58.0 1 4 7 3 3 2 3 5 1 2 49. 0 1 2.5 7 3 1 2 2 4 1 2 41. 0 1 4.5 4 3.5 2 1 3.5 5 3;5 4 46 . 5 1 4 5 4 3 2 5 3 4 2 51.5 1 2.5 5 4 2 2 4 3 2.5 3 44. 5 1 3.5 7 3 1 2 2 3 3 2 43.5 1 5 1.5 2 1 2.5 4 5 2 4 43.5 1 . 5 5 4 2 1 5 4 3 3 52.0 1 3 3 4 7 3 2 5 2 2.5 51. 0 1 5 7 3 7 3 4 3 4 1 61. 5 1 5 7 3 7 3 3.5 2 3.5 1 52.0 1 4 4 4 2 1 4 4 4 1 51. 0 1 5 5 4 2 1.5 3 4 1 1 51.5 1 2.5 4.5 4 7 3 4 2 3.5 1 46.5 177 TABLE 19 (cont inued) Species 1 2 3 4 5 6 Wade's Bog (cont inued) M a i a n t h e m u m d i l a t a t u m 2 4. 5 7 3 2 2 R h y n c h o s p o r a a l b a 2 6.5 1 5 3 1 V a o o i n i u m v i t i s - i d a e a 1.5 6 2 5 1 1 S o i v p u s o e s p i t o s u s 2 5 1 1 3 1 J u n o u s s u p i n i f o r m i s 2 5 1 1 3 1. 5 V a c c i n i u m u l i g i n o s u m 1 6 2 3 1 1 G a u l t h e r i a s h a l l o n 1 5 . 5 2 1 1. 5 1 G e n t i a n a s c e p t r u m 2 6.5 6 5 3 2 C o p t i s a s p l e n i f o l i a 2 4.5 3 5 2 1.5 Ogg' 1 s Bog M y r i c a g a l e 1 4 2 1 3 5 A p a r g i d i u m b o r e a l e 2 4.5 5 5 3 1.5 C a r e x o b n u p t a 2 5 1 1.5 3 3 C a r e x p l u r i f l o r a 2 5 1 3 3 3 A g r o s t i s a e q u i v a l v i s 2 3.5 5 5 1.5 1 S a n g u i s o r b a o f f i c i n a l i s 2 3.5 4 3 1 1 Ledum g r o e n l a n d i c u m 1 6.5 3 5 1.5 2 V a c c i n i u m oxy c o c c u s 1.5 6 2 3 1.5 1.5 D r o s e r a r o t u n d i f o l i a 3 5 3 5 1 1 K a l m i a p o l i f o l i a 1 6 3 5 1.5 2 E m p e t r u m n i g r u m 1 6.5 3 5 3 5 T r i e n t a l i s a r c t i c a 2 . 5 11 1 1 1 2 C a r e x c a n e s c e n s 2 5 1 2 3 3 T o f i e l d i a g l u t i n o s a 2 7 . 5 4 5 1.5 2 L i n n a e a b o r e a l i s 1.5 4 3 3 1.5 1 R h y n a h o s p o r a a l b a 2 6.5 1 5 3 1 P l a n t a g o m a c r o c a r p a 2 3 5 3 3 2 G a u l t h e r i a s h a l l o n 1 5.5 2 1 1.5 1 G e n t i a n a s c e p t r u m 2 6.5 6 5 3 2 C a l a m a g r o s t i s n u t k a e n s i s 2 3.5 6 3 2 2 M a i a n t h e m u m d i l a t a t u m 2 4.5 7 3 2 2 S c i r p u s c e s p i t o s u s 2 5 1 1 3 1 C o p t i s a s p l e n i f o l i a 2 4.5 3 5 2 1.5 C o p t i s t r i f o l i a 2 4.5 3 5 2 1. 5 J u n c u s s u p i n i f o r m i s 2 5 1 1 3 1.5 TABLE 19. (Continued) 178 1 o I I 1 2 1 3 1 it 1 5 1 6 1 6 S = IPR l Wade's Bog (continued) 1 3 3.5 4 7 3 4 1 3 1 51.0 1 3 7 3 1 2 2 2 2 1 42.5 1 2.5 5 4 2 2 4 2 3 1 43 . 0 1 4.5 7 3 1 1.5 2 2 3.5 1 39. 5 1 2 7 3 1 2 4 1 1 2 37.5 1 5 5 4 2 2 4 1 4 2 44. 0 1 5 5 4 1 1 4 1 2 1 37.0 1 5 5 4 2 2 4 3 1 1 52.5 1 3 3 3.5 3 1.5 2.5 1 2 1 39 . 5 range : 36. 0 - 61.5 unweighted x = 47. 0 weighted x = 48 . 5 Ogg ' s Bog 1 5 7 3 7 3 3.5 4 3.5 1 54.0 1 5 4 4 7 3 5 5 2 1 58 . 0 1 2 . 5 7 3 1 2 2 4 1 2 41. 0 1 2.5 7 3 1 2 2 4 1 3 43.5 1 4 7 3 3 2 3 5 1 2 49. 0 1 4.5 4 3.5 2 1 3.5 5 3.5 4 46.5 1 5 5 4 2 1 5 4 3 3 52.0 1 2.5 5 4 2 2 4 3 2.5 3 44. 5 1 5 1.5 2 1 2.5 4 5 2 4 43 . 5 1 4 5 4 3 2 5 3 4 2 51.5 1 5 7 3 7 3 4 3 4 1 61. 5 1 3 3 4 7 3 2 4 2 2.5 50 . 0 1 3.5 7 3 1 2 2 3 3 2 43. 5 1 4 4 4 2 1 4 3 4 1 50.0 1 2.5 4.5 4 7 3 4 3 3 . 5 1 47.5 1 3 7 3 1 2 2 2 2 1 42.5 1 5 7 3 1 2 3.5 3 1 1 45.5 1 5 5 4 1 1 4 2 2 1 38.0 1 5 5 4 2 2 4 3 1 1 52.5 1 4 7 3 7 3 2.5 2 2 2 52.0 1 3 3 . 5 4 7 3 4 2 3 1 52.0 1 4.5 7 3 1 1.5 2 2 3.5 1 39 . 5 1 3 3. 3.5 3 1.5 2.5 2 2 1 40.5 1 3 3 4 3 2.5 2 2 3 1 42.5 1 2 7 3 1 2 4 2 1 2 37.5 TABLE 19. (cont inued) 179 Species 1 2 3 5 6 Ogg's Bog (continued) D e s c h a m p s i a ce s p i t o s a 2 3.5 6 3 2 2 G e n t i a n a d o u g l a s i a n a 4 6.5 5 5 2 1.5 V a c c i n i u m o v a t u m 1 6 2 5 1 1 V a c c i n i u m v i t i s - i d a e a 1.5 6 2 5 1 1 C o r n u s u n a l a s c h k e n s i s 1.5 5.5 3 3 1.5 1.5 V a c c i n i u m u l i g i n o s u m 1 6 2 3 1 1 E e p h r o p h y l l i d i u m c r i s t a - g a H i 2 8.5 1 2 2 2 E r i o p h o r u m p o l y s t a c h i o n 2 5 2 2 3 3 C a r e x p a u c i f l o r a 2 5 1 1.5 2 3 B l a c k w a l l Meadow V a l e r i a n a s i t c h e n s i s 2 4 2 1 2 2 F e s t u c a v i r i d u l a 2 3 . 5 6 3 1.5 1.5 L u p i n u s l a t i f o l i u s 2 6 1 1 1 1 E r i g e r o n p e r e g r i n u s 2 4.5 4 5 3 1.5 Anemone o c c i d e n t a l i s 2 4 4 5 2 2 P o t e n t i l l a f l a b e l l i f o l i a 2 3 . 5 2 3 2 1.5 E r y t h r o n i u m g r a n d i f l o r u m 3 6 7 5 1 1.5 V a c c i n i u m s c o p a r i u m 1 6 2 5 1 1 C l a y t o n i a l a n c e o l a t a 3 4 6 5 2 2 A r e n a r i a c a p i l l a r i s 2 5.5 4 5 2 2 A n t e n n a r i a l a n a t a 2 3.5\" 5 3 3 5 V e r o n i c a c u s i c k i i 2 4.5 2 1 2 2 A g o s e r i s a u r a n t i a c a 2 4.5 4 3 3 1 P h l e u m a l p i n u m 2 3.5 5 3 2 2 A r n i c a l a t i f o l i a 2 5 4 3 3 1.5 L u z u l a h i t c h c o c k i i 2 3 3 3 3 2 T h a l i c t r u m o c c i d e n t a l e 2 3.5 2 1 3 4 A c h i l l e a m i l l e f o l i u m 2 4.5 4 2 3 1.5 T r i s e t u m s p i c a t u m 2 3 . 5 5 3 2 2 C a r e x r o s s i i 2 5 1 2 2 3 E l y m u s g l a u c u s 2 3.5 7 3 1 1 S i l e n e p a r r y i 2 6 5 3 3 2 A r n i c a m o l l i s 2 5 4 1 3 1. 5 P o a e p i l i s 2 3.5 3 3 3 5 S e n e c i o i n t e g e r r i m u s 2 5 5 3 3 1.5 180 TABLE 19. (cont inued) 1 6 7 8 9 1 0 I I 1 2 1 3 l >t 1 5 1 6 Z = IPR l Ogg's Bog (cont inued) 1 5 6 3 7 3 2 . 5 1 5 3 55.0 1 5 5 4 2 1.5 3 1 1 1 49. 5 1 5 5 4 2 1 4 2 1 1 41. 0 1 2.5 5 4 2 2 4 2 3 1 43.0 1 2.5 4 4 7 3 4 1 2.5 3 48.0 1 5 5 4 2 2 4 1 4 2 44.0 3 3.5' 3.5 4 2 2 2 1 2 1 41. 5 1 2 7 3 1 2 5 1 2 3 44. 0 1 3 7 3 1 2 3 1 2 1 . 38.5 range: 37.0 _ 61.5 unweight ed x = 46.6 weij jhted x = 49 . 8 B l a c k w a l l Meadow 1 4 4 4 2 1 5 4 3 2 43. 0 1 5 7 3 3 2 2 5 1 3 49 . 5 1 5 5 4 7 3 3 5 4 5 54. 0 1 4 4 4 7 3 5 5 4 3 60.0 1 5 3 4 2 3 5 5 2 1 50.0 1 4 4 4 6 2 2 5 1 2 45. 0 1 5 5 4 2 3 2 5 3 2 55 . 5 1 5 5 4 3 2 4 4 1 1 46.0 1 5 4 4 2 2.5 2 5 5 1 53.5 1 5 4 4 7 3 4 4 4 4 60.5 1 4 4 3 7 3 5 4 1 3 56.5 1 4 4 4 3 2 3 4 2 1 41. 5 1 5 4 4 3 5 3 3 1 5 3 . 5 1 4 7 3 1 1 4 3 5 2 48.5 1 4 4 4 7 3 5 2 4 3 55.5 1 2 7 3 6 2 2 2 1 1 43.0 1 3 7 3 5 3 2 1 3 2 45.5 1 4 4 4 7 3 5 2 5 4 56.0 1 5 7 3 7 3 4 2 5 3 57. 5 1 5 7 3 1 2 2 2 3 2 43.0 1 5 3 2 1 2 2 1 3 5 42.5 1 5 6 4 2 2 3 2 1 1 48 . 0 1 4 4 4 7 3 5 2 2 3 51.5 1 1 7 3 1 3 2 1 2 4 44. 5 1 5 4 4 7 3 5 2 3.5 2 56.0 TABLE 19. (Continued) 181 Species B l a c k w a l l Meadow (cont inued) H i e r a o i u m g r a c i l e 2 4.5 5 5 3 1 L u z u l a s p i a a t a 2 3 2 3 3 2 Sedum l a n c e o l a t u m 2 4 4 5 1 2 C a s t i l l e j a m i n i a t a 2 6 3 5 1 2 P e n s t e m o n p r o c e r u s 1.5 4 3 5 2 1 P e d i o u l a r i s b r a c t e o s a 2 4 4 . 5 1 2 P h l o x d i f f u s a 1.5 3. 5 4 5 1 1 P o t e n t i l l a d i v e r s i f o l i a 2 3.5 2 1 2 1.5 C a r e x s p e c t a b i l i s 2 5 2 1 3 3 E p i l o b i u m a l p i n u m 2 4.5 2 3 1 1 D e l p h i n i u m n u t t a l l i a n u m 2 4 4 3 2.5 1 C a s t i l l e j a p a r v i f l o r a 2 6 3 5 1 2 R a n u n c u l u s e s c h s c h o l t z i i 2 4 5 3 2 1.5 S i b b a l d i a p r o c u m b e n s 2 3.5 2 5 1 1. 5 M i c r o s t e r i s g r a c i l i s 4 4 4 5 1 1 J u n o u s d r u m m o n d i i 2 5 1 1 2. 5 2 E y d r o p h y l l u m f e n d l e r i 2 4.5 4 3 2 2 S e n e c i o t r i a n g u l a r i s 2 5 6 3 3 1. 5 V a c c i n i u m d e l i c i o s u m 1 6 2 3 1 1 TABLE 19. (Concluded) 182 1 o I l 1 2 1 3 1 \"t 1 5 1 6 1 6 Z = I P R l 1 4 4 3 7 3 5 1 4 3 55.5 1 5 7 3 1 2 2 2 3 1 42. 0 1 4 5 4 7 3 4 1 3 3 53.0 1 5 6 4 7 3 5 2 4 4 60 . 0 1 4 4 4 2 1 4 1 2 3 42. 5 1 5 5 4 7 3 3 2 3 . 5 1 52.5 1 5 5 4 7 3 2 1 3 1 48.0 1 5 4 4 5 2 2 1 3.5 4 43.5 1 3 7 3 1 2 1 1 3 3 40 . 0 1 3 2.5 3 1 2.5 5 1 4 3 39 . 5 1 5 5 4 3 2.5 4 1 3 4 49 . 0 1 5 5 4 7 3 5 2 3 3 \u202257.0 1 5 3 4 2 2 2 1 4 2 43.5 1 3 2 3 2 2 2 1 3 1 35. 0 1 4 2.5 3.5 1 2.5 4 1 3 1 43.5 1 4 7 3 1 2-..5 4 1 2.5 1 40.5 1 4 5 4 3 '2 2 1 2 1 42. 5 1 5 4 4 7 3 5 1 3 . 5 1 55.0 1 5 5 4 3 2 4 1 1 1 41. 0 range: 3 5.0_- 6 0.5 unweighted x = 48.6 weighted x = 50.6 'Arranged i n order of dec reas ing importance v a l u e . 183 systems, might be expected to have l e s s v a r i a b l e popu la t ions than comparable o u t c r o s s i n g spec ies (Stebbins 1957a; Grant 1958). However, a l a rge body of recent evidence suggests tha t some inb reed ing spec ies have mainta ined a h i g h l e v e l o f p o p u l a t i o n v a r i a b i l i t y presumably through o c c a s i o n a l o u t c r o s s i n g between s t r o n g l y homozygous b io types ( A l l a r d 1965; Kannenberg and A l l a r d 1967; J a i n and M a r s h a l l 1967; R o l l i n s 1967; A l l a r d , J a i n and Workman 1968; J a i n 1969; Kondo 1972, inter alia). Species of Cyperaceae, Gramineae, Juncaceae, and L i l i a c e a e have average I . P . R . ' s of 41 .4 , 4 8 . 3 , 4 0 . 2 , and 5 2 . 5 , r e s p e c t i v e l y . Average I . P . R . ' s fo r r e p r e s e n t a t i v e s of some common nor th temperate d ico ty ledonous f a m i l i e s a re : Caryo-phy l l aceae ( 4 5 . 4 ) ; Compositae ( 5 5 . 1 ) ; E r i c a c e a e , i n c l u d i n g Empetrum nigrum ( 4 6 . 2 ) ; Ranunculaceae ( 4 5 . 1 ) ; Rosaceae ( 4 2 . 9 ) ; Sc rophula r iaceae ( 5 0 . 7 ) . The phy logene t i c s i g n i f i c a n c e o f these f i g u r e s w i l l not be specula ted on. TABLE 20. Average community I . P . R . ' s . Community x , unweighted x , weighted 1 S a l t Marsh Wade's Bog Ogg's Bog B l a c k w a l l Meadow 43.8 47.0 46 . 6 48.6 47.2 48.5 49.8 50.6 Weighted by m u l t i p l y i n g the unweighted I . P . R . f o r each species by i t s corresponding I . V . and d i v i d i n g the community sum o f these products by the average I . V . fo r tha t community. 184 The most s t r i k i n g g e n e r a l i z a t i o n tha t emerges from t h i s a n a l y s i s i s tha t the d i f f e r e n t f a c t o r s i n a s p e c i e s ' recombina t ion system f r equen t ly work i n o p p o s i t i o n to one another but a r r i v e at an e v o l u t i o n a r y compromise between f i t n e s s and f l e x i b i l i t y . This compromise i s u s u a l l y d i f f e r e n t f o r d i f f e r e n t species and tends to be c o r r e l a t e d w i t h the s p e c i e s ' e c o l o g i c a l s t r a t egy and the type o f vege t a t i on of which i t i s a c o n s t i t u e n t . Some examples of the i n t e r p l a y o f opposing f a c t o r s are g iven i n Table 21. The s i t u a t i o n i n d i f f e r e n t species i s i n s t r u c t i v e . For example, the e r icaceous shrub Kalmia polifolia has a long genera t ion t ime , moderate vege ta t ive r ep roduc t ion v i a shor t rhizomes and t i l l e r i n g , and i s p a r t i a l l y s e l f - c o m p a t i b l e ; but i t i s d i p l o i d , has a h igh b a s i c chromosome number, showy f lowers tha t are h i g h l y ou tc ros sed , and minute , l i g h t seeds w i t h good d i s p e r s i b i l i t y . Spergularia canadensis , though autogamous (most o f i t s f lowers are c le is togamous) and a t e t r a p l o i d w i t h f a i r l y s m a l l chromosomes and clumped p o p u l a t i o n s , i s an annual tha t u s u a l l y has a few chasmogamous f lowers on each p l a n t , and has s m a l l , g l andu la r -pubescen t , s l i g h t l y -winged seeds and l a rge p o p u l a t i o n s . Even Poa cusickii v a r . epilis , though i t appears to reproduce p r i m a r i l y by agamospermy, has l a rge chromosomes and i s d ioec ious and wind-p o l l i n a t e d , so tha t whenever t h i s Poa happens to reproduce s e x u a l l y , i t i s l i k e l y to r e l ea se a l a rge amount o f r ecombina t iona l v a r i a b i l i t y (Clausen 1954, 1961). 185 TABLE. 21. Combinations of opposing r e g u l a t o r y f a c t o r s . Fac tors promoting v s . f a c t o r s r e s t r i c t i n g recombinat ion Examples shor t gene ra t ions , good seed d i s -p e r s a l , l a rge popu la t ions v s . autogamy, clumped popu la t ions d ioecy or s t rong dichogamy, anemo-p h i l y v s . h igh p o l y p l o i d y , s m a l l chromosomes l a rge chromosomes, s e l f - i n c o m p a t i -b i l i t y , o u t c r o s s i n g vs.. p o l y p l o i d y , vege t a t i ve r ep roduc t ion Spergularia canadensis, M i c r o s t e r i s g r a c i l i s T r i g l o c h i n maritimum, Juncus b a l t i c u s , Myrica gale, Thalictrum occiden-t a l e many Gramineae and Compositae showy f l o w e r s , s e l f - i n c o m p a t i b i l i t y , Lupinus l a t i f o l i u s , Arnica h igh b a s i c numbers, f r e q u e n t ' i n t e r -s p e c i f i c h y b r i d i z a t i o n , e c o t y p i c d i f f e r e n t i a t i o n v s . p o l y p l o i d y , sma l l chromosomes d i p l o i d y , h i g h b a s i c chromosome number v s . s e l f - c o m p a t i b i l i t y r e l a t i v e l y inconspicuous f lowers ' h igh b a s i c number, s e l f - i n c o m p a t i -b i l i t y v s . h i g h p o l y p l o i d y , s m a l l chromosomes h i g h b a s i c number v s . long genera t ion time l a rge chromosomes, anemophily, frequent h y b r i d i z a t i o n v s . p o l y p l o i d y , i nb reed ing d i o e c y , wind p o l l i n a t i o n v s . low b a s i c numbers, p o l y p l o i d y , ex tens ive vege ta t ive r ep roduc t ion mon^eey,, ^ dieta^gagrjf,;; wind; - p p l l i r t : ^nafiLOBi HPS . . . s p a l l ' Ghrompsome^Sr, good d i s p e r s a l , h y b r i d i z a t i o n v s . autogamy, p o l y p l o i d y , sma l l chromosomes l a t i f o l i a , Arnica mollis Glaux maritima, S t e l l a r i a humifusa, T o f i e l d i a g l u t i -nosa, Drosera r o t u n d i f o l i a T r i e n t a l i s a r c t i c a , Lupinus l a t i f o l i u s E r i c a c e a e , Empetrum nigrum Elymus glaucus, Hordeum brachy an the rum D i s t i c h l i s spicata Carex lyngbyei, C. obnupta, C. p l u r i f l o r a Epilobium alpinum TABLE 21. (Continued) 186 Fac tors promoting v s . f a c to r s r e s t r i c t i n g recombinat ion , Examples l a rge chromosomes, d i p l o i d y , dichogamy, anemophily v s . low b a s i c number, s e l f -c o m p a t i b i l i t y good d i s p e r s a l , h y b r i d i z a t i o n v s . autogamy, p o l y p l o i d y , s m a l l chromosomes showy f l o w e r s , w e l l - d e v e l o p e d entomophi ly , good d i s p e r s a l v s . s e l f - c o m p a t i b i l i t y l a rge chromosomes, showy, out -crossed f l o w e r s , e c o t y p i c d i f f e r e n t i a t i o n v s . s e l f - c o m p a t i -b i l i t y , poor d i s p e r s a l P l a n t a g o m a r i t i m a E p i l o b i u m a l p i n u m V a l e r i a n a s i t o h e n s i s 3 D e l p h i n i u m n u t t a l l i a n u m , Er icaceae C l a y t o n i a l a n c e o l a t a , E r y t h r o n i u m g r a n d i f l o r u m l a rge chromosomes, d i p l o i d y , showy, Gentiana d o u g l a s i a n a 3 entomophilous f l o w e r s , good \\G. soeptrum, Anemone d i s p e r s a l v s . s e l f - c o m p a t i b i l i t y , ocoidentalis no h y b r i d i z a t i o n , l i t t l e or no e c o t y p i c d i f f e r e n t i a t i o n In summary, t h i s a n a l y s i s has shown tha t there are d i f f e r ences i n the recombinat ion p o t e n t i a l s of d i f f e r e n t spec ies but tha t these d i f f e r e n c e s are apparent ly not g rea t . The g e n e r a l i z a t i o n tha t autogamous species have more r e s t r i c t e d recombina t ion systems than comparable xenogamous or o u t c r o s s i n g spec ies seems v a l i d . However, spec ies w i t h r e s t r i c t e d recombi -n a t i o n p o t e n t i a l do not dominate the vege t a t i on of any o f the study communit ies; indeed , they are u s u a l l y very minor elements ( c f . Tables 3, 4 , 5, 6 w i t h Table Table 19) . Furthermore, no 187 s i g n i f i c a n t d i f f e r e n c e was found between l e v e l s of p o t e n t i a l recombinat ion w i t h i n any of the communities. This f i n d i n g i s at odds w i t h Mosquin 's (1966) t h e s i s tha t species of harsh p h y s i c a l environments have r ep roduc t i ve s p e c i a l i z a t i o n s fo r reduc ing gene t i c v a r i a b i l i t y ; i . e . , c l o sed recombina t ion systems. Of course , i t i s d i f f i c u l t t o decide which of the four communities has the harshest p h y s i c a l environment. The c l ima te of the Tof ino area may be c l a s s e d as warm, temperate , and r a i n y ; tha t of the subalp ine meadow as c o n t i n e n t a l , c o l d , and humid but summer dry ( a f t e r K r a j i n a 19 65) . The o v e r a l l c l i m a t e of the subalp ine meadow might thus be cons idered harsher . But there are l o c a l microenvi ronmenta l c o m p l i c a t i o n s , The s a l t marsh i s subjec ted to great ranges i n d a i l y temperature , and the p l a n t s are p e r i o d i c a l l y baked by the sun and f looded by s a l t water (Chapman 1960, 19 64) . Dur ing the growing season, p l an t s i n the suba lp ine meadow are exposed to great d i u r n a l temperature f l u c t u a t i o n bo'thaabpveaand below ground (Kuramoto and B l i s s 197 0; B a l l a r d 1972),and may a l s o experience mois ture s t r e s s on s h a l l o w , w e l l - d r a i n e d s o i l s (Eady 1971). Only the c o a s t a l sphagnum bog has a t r u l y equable c l i m a t e , but i t s o v e r a l l p h y s i c a l environment, c h a r a c t e r i z e d by a low n u t r i e n t s t a tus and a wate r logged , p o o r l y oxygenated subs t ra te (Gorham 1957 ; Smal l 19 7 2a .. 8 b),,. .must .be. . .considered .compara t ive ly harsh Furthermore, as was po in ted out i n the s e c t i o n on l e v e l s of p o l y p l o i d y , environmenta l r i g o r i n c l u d e s r e l a t i v e ampli tude of environmental f l u c t u a t i o n s and the i r r e g u l a r i t y of these f l u c t u a t i o n s , as w e l l as average c o n d i t i o n s of the e n v i r o n -188 mental complex such as c l i m a t i c means and n u t r i e n t l e v e l s . The d i f f e r e n t l e v e l s of p o l y p l o i d y i n the three vege t a t i on types seemed to be best exp l a ined by d i f f e r e n t degrees of e n v i r o n -mental i n s t a b i l i t y . However, the l o g i c of Sec t . I I I - C does not apply to the f i n d i n g s of the I . P . R . , of which p o l y p l o i d y i s but one aspec t . D i f f e r e n t s e l e c t i o n pressures have been ope ra t i ng i n s a l t marshes, sphagnum bogs, and suba lp ine meadows, and d i f f e r e n t types of spec ies s t r a t e g i e s and community fea tures have evolved as a r e s u l t ('see s ec t . I V ) . A l l four communities have harsh environments , i n at l e a s t some r e s p e c t s . But there i s no i n d i c a t i o n tha t the d i f f e r e n t types of s e l e c t i o n d i f f e r i n o v e r a l l s e v e r i t y , a l though one ( tha t o f the suba lp ine meadow) may be caused more by b i o t i c f a c t o r s than another ( tha t of the s a l t marsh). I t i s not s u r p r i s i n g t ha t the I . P . R . ' s f o r the four communities are not s i g n i f i c a n t l y d i f f e r e n t . What i s more, t h i s a n a l y s i s says no th ing d i r e c t l y about the a c t u a l l e v e l s of gene t ic v a r i a b i l i t y w i t h i n the spec ies p o p u l a t i o n s . To get some measure of t h i s i t would be necessary to study enzyme polymorphisms or seed p r o t e i n v a r i a b i l i t y or some such molecu la r approach. But even i f one community had more spec ies w i t h more h e t e r o z y g o s i t y than the other communit ies , one s t i l l cou ld not ma in ta in tha t tha t community had b e t t e r adapted spec ies than the o t h e r s , s ince spec ies adaptedness can be achieved e i t h e r by i n d i v i d u a l a d a p t a b i l i t y (a homeostat ic genotype) or by gene t i c a d a p t a b i l i t y ( d i v e r s i t y of genotypes) (Dobzhansky 1968; c f . Gooch and Schopf 1972; L e v i n t o n 1973). Bradshaw (1971) puts the problem i n proper p e r s p e c t i v e : \"There i s l i t t l e d i f f e r e n c e i n essence between 189 normal s i t u a t i o n s and extreme environments from an e v o l u t i o n a r y po in t of v i ew\" . N e v e r t h e l e s s , i t would be extremely i n t e r e s t i n g to subjec t other predominant ly herbaceous v e g e t a t i o n types to s i m i l a r a n a l y s i s . Would, say , the extremely d i v e r s e , t a l l g r a s s p r a i r i e s of the Midwest or the c o a s t a l p r a i r i e s of Texas have h ighe r average I . P . R . ' s than the study communities? Would a r c t i c tundra or the d i s t u r b e d , weedy C e n t r a l V a l l e y g r a s s l and of C a l i f o r n i a have lower a v e r a g e . I . P . R . ' s ? What about o ther community comparisons; e g . , Great B a s i n v s . Sonoran d e s e r t s , C a l i f o r n i a c h a p a r r a l v s . A u s t r a l i a n thorn scrub? Sure ly here i s a tremendous f i e l d fo r fu ture i n v e s t i g a t i o n . Summary, Discussion, and Conclusions. 190 The f i n d i n g s of the present i n v e s t i g a t i o n are best summarized and con t ras t ed i n a balance sheet (Table 22) . General i n d i c a t i o n s are tha t i n t e r s p e c i f i c compe t i t ion increases from sphagnum bogs to s a l t marsh to suba lp ine meadow, w h i l e the s t r e s s of the p h y s i c a l environment inc reases from sphagnum bogs to subalp ine meadow to s a l t marsh. These i n d i c a t i o n s are not w h o l l y c o n s i s t e n t , but s e v e r a l genera l t rends are ev ident i n Table 22. A more s p e c i f i c summary f o l l o w s : 1. The vege t a t i on o f a s a l t marsh, two sphagnum bogs, and a suba lp ine meadow were desc r ibed by means o f t ab l e s of importance values and by modi f ied species c o n s t e l l a t i o n s 2. The p o p u l a t i o n s t r u c t u r e of the i n d i v i d u a l spec ies has been assessed w i t h a D\/d index of aggrega t ion . Aggrega-t i o n increases as environmental he te rogene i ty and p h y s i c a l s t r e s s i n c r e a s e , and decreases as success ion proceeds and i n t e r s p e c i f i c compe t i t i on i n c r e a s e s . 3. Ind ices of a s s o c i a t i o n and c o r r e l a t i o n have been used to s t a t i s t i c a l l y assess i n t e r s p e c i f i c r e l a t i o n s h i p s . I n t e r s p e c i f i c a s s o c i a t i o n and c o r r e l a t i o n , both p o s i t i v e and n e g a t i v e , inc rease w i t h i n c r e a s i n g environmenta l he te rogene i ty and c o m p e t i t i o n . 4. Chromosome counts were made fo r most of the s p e c i e s , and l e v e l s of p o l y p l o i d y fo r both the f l o r a and vege t a t i on of a l l four communities were e s t a b l i s h e d . Leve l s of po ly p l o i d y appear to be c o r r e l a t e d w i t h environmenta l r i g o r TABLE 22. Resu l t s and i n d i c a t i o n s from var ious aspects of the i n v e s t i g a t i o n ; a l l i n d i c a t i o n s are r e l a t i v e . Community Spec i e s - a r ea curves Community d e s c r i p t i o n s Popu la t i on s t r uc tu r e (D\/d index) S a l t Marsh Wade's Bog Ogg's Bog B l a c k w a l l Meadow co >. cu M-l CO \u2022H cu CO Cl) & U CU O > \u2022H \u2022H TJ >i >, +-> rG \u2022H ft O cu rtj ft bO rtf o O ft o bO -H a o \u2022H >, o u \u2022H u S rd O >, rrj +-> +-> cu +-> \u2022H g o cu r -4 h igh s a l i n i t y c o a s t a l s a l t marsh, mainly i n r a p i d expansion and a c c r e t i o n c o a s t a l sphagnum bog at or approaching c l imax c o a s t a l sphagnum bog, f a i r l y e a r l y i n success ion l u s h , more or l e s s mes ic , Cascadian suba lp ine meadow; c l imax (?) species most aggregated; s t rong h a b i t a t zona t ion ; harsh p h y s i c a l environment; moderate compe t i t i on l e a s t aggrega t ion; weak i n t e r -s p e c i f i c compe t i t i on ; success-i o n a l ma tur i ty s t rong aggrega t ion ; micro topo-graphic mosaicism; weak compet-i t i o n ; s u c c e s s i o n a l youth moderate aggrega t ion ; m ic ro -topographic mosaicism; s t rong compe t i t i on ; s u c c e s s i o n a l ma tu r i ty Community A s s o c i a t i o n C o r r e l a t i o n Sa l t . Marsh Wade's Bog Ogg's Bog B l a c k w a l l Meadow heterogeneous moderate environment; compe t i t i on t i g h t m i c r o s i t e s p e c i f i c i t y g rea te s t p h y s i c a l s t r e s s +\/- homogeneous weak environment; compe t i t i on loose m i c r o s i t e s p e c i f i c i t y i n t e rmed ia t e s t r e s s heterogeneous weak environment; compe t i t i on . t igh t m i c r o s i t e s p e c i f i c i t y - i n t e rmed ia te s t r e s s heterogeneous s t rong environment; compe t i t i on t i g h t m i c r o s i t e s p e c i f i c i t y lowest p h y s i c a l s t r e s s Levels of F lower ing P o l l i n a t i o n p o l y p l o i d y phenology ecology most p o l y -p l o i d y ; g rea tes t \" r i g o r \" ( h i s t o r i c a l and p h y s i c a l ) l e a s t p o l y -p l o i d y ; l e a s t r i g o r l e a s t p o l y -p l o i d y ; l e a s t r i g o r much p o l y -p l o i d y ; moderate-strong r i g o r CD CO rd CD P . O CD T 3 T 3 O \u2022r-i U CD ft hO u o I\u2014I 4 H M H O -H M | CD rH a) ca0| -rd & CD > weak i n t e r s p e c i f i c compet i t ion f o r p o l l i n a t i o n v e c t o r s ; p r i m a r i l y a b i o t i c p o l l i n a t i o n moderate compe t i t i on both b i o t i c and a b i o t i c p o l l i n a t i o n moderate compe t i t i on both b i o t i c and a b i o t i c p o l l i n a t i o n in tense c o m p e t i t i o n ; predominantly b i o t i c p o l l i n a t i o n DO o an 3 M OP pj pj fD pu OP ft H P) O -\u2022 CD rt ft x cn -\u2022 o t, cn s; pj tD pj H O to H OP So sh o o rt ft p f l cn ft cn tr ex, H H i cn O H - O 4 CD H ' O 4 rt H - O 4 rt H - H - O O rt H -CO 3 H' fu cn 3 H - 4 cn 3 H - 4 0 cn 3 4 4 cn >d OP 3 r V T j OP 3 O *d OP 3 0 rtW OP O \u2022d CD 1 P) CD 1 pj 3 CD 1 pJ 3 H - CD 1 ft 3 CD 4 ft 4 O 4 ft 4 OP 4 O J 4 OP O 4 ft H.rjq 4 cn p . p- O cn H - H - cn H - H - cn H - cn cn P) cn H 3 pj cn H O pi cn H o pj cn >d o P) H rt ^ 0 H <+ \u2022< O 1\u20141 rt CD O H P) fD PJ 3 p. 3 pj 4 3 3 PJ rt 3 t r u 3 tr>d CD o tr H - O H - CD O H - (D 4 O P) fD o (D H - r+ (C 0 rt fD O rt H - fD h-1 rt O O H - rt H - rt H - 3 H - H rt O H- rt H- rt PJ P) rt O H - 3 O H - O H - 4 OP H - OP O v . . O O H - fD 0 \u2022< 3 3 H 3 3 ^ rt cn average n iche s i z e decreases + to i n t e r s p e c i f i c compe t i t i on inc reases average species v a r i a b i l i t y decreases cn cn cD 3 rt p- o o 3 O ft H ' H (D H O 4 OP P) OP PJ 4 H - rt O H - O fl) O rt P) I 3 \"< H cn 4 o 3 CD o CD O 3 H *d ^ \u2022 CD rt H > rt H -O 3 rt 4 O 3 OP O O 3 \u2022d fD fD O rt cn OP cn H -o pj s: H fD PJ ft X - H -cn rt H -3 O rt I rt -H -O 3 \u2022d O 3 CD O O H PJ < CD 3 CD CD O rt cn OP H - 01 H rt O P-s: CD PJ ft W H -cn rt H -3 O rt I cn s! cn cn >0 CD H- rt CD PJ 3 4 O X H - O H - I H 3 H i 3 P) OP H - O 4 O ft H - (D CD rt O 0 4 ^ 0 O P ) H 3 rt O O \u2022d CD H i O P CD H-rt H - Cn O H - 3 *d P) rt rt fD H \u2022 CD O 4 H -I CD cn O 3 index of dominance decreases spec ies d i v e r s i t y inc reases ^ C s t r e s s of p h y s i c a l environment decreases s t a b i l i t y inc reases (?????) 2 H -O tr CD rt tr CD o 4 S i H -O tr CD ft H -H i H I CD 4 CD 3 rt H -PJ rt H -O 3 PJ O 3 0> ft 3 H -cn 3-rt P) pj 3 tr o H - CD rt \u00a361 ft < ft) 4 cn H -rt 194 s e n s u l a t o . 5. The f o l l o w i n g aspects of species r ep roduc t ive b i o l o g y were s t u d i e d : a . f l o w e r i n g phenology b . f l o r a l b i o l o g y and p o l l i n a t i o n ecology c . d i s p e r s a l ecology d. c o m p a t i b i l i t y A l l four communities are dominated by o u t c r o s s i n g s p e c i e s . 6. The theory of the niche and va r ious c o r o l l a r i e s concern-i n g c o m p e t i t i o n , dominance, and v a r i a b i l i t y were d i s cus sed . The most abundant species (at l e a s t of grasses) are the most v a r i a b l e and presumably have the l a r g e s t n i c h e s . Niche s i z e and p o p u l a t i o n v a r i a b i l i t y decrease as i n t e r s p e c i f i c compe t i t i on i n c r e a s e s . 7. A numer ica l taxonomic approximat ion of each s p e c i e s ' n iche was at tempted. E c o l o g i c a l d i s s i m i l a r i t y reduces i n t e r s p e c i f i c c o m p e t i t i o n . 8. The not ions of d i v e r s i t y and s t a b i l i t y were i n v e s t i g a t e d . 9. An index of p o t e n t i a l recombina t ion was d e v i s e d , accord ing to which 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 p o t e n t i a l r e combina t i on , on the average, between spec ies of the four d i f f e r e n t communities. Fur ther b r i e f summaries of the f i n d i n g s and i n t e r p r e t a t i o n s o f t h i s i n v e s t i g a t i o n as they apply to each community type would be u s e f u l and are o u t l i n e d - b e l o w . 195 S a l t marsh Only a few species can s u r v i v e the s t r i n g e n t p h y s i c a l and chemica l c o n d i t i o n s of a h igh s a l i n i t y s a l t marsh. The marsh vege t a t i on i s dominated by the s t i l l 'fewer species tha t can not on ly s u r v i v e but a l so t h r i v e . Tide l e v e l s e x e r c i s e s t rong c o n t r o l over the v e r t i c a l d i s t r i b u t i o n s of s a l t marsh p l a n t s . The h a b i t a t zona t ion and m i c r o s i t e s p e c i f i c i t y promote aggregated popu la t ions and inc rease the s t r eng th o f i n t e r s p e c i f i c a s s o c i a -t i o n . Strong a s s o c i a t i o n and gross morpho log ica l and f u n c t i o n a l s i m i l a r i t i e s among marsh species r e s u l t i n s t rong i n t e r s p e c i f i c c o m p e t i t i o n . However, the harsh p h y s i c o - c h e m i c a l environment, by p l a c i n g a premium on s u r v i v a l and l i m i t i n g spec ies d i v e r s i t y , tends to counterac t the e f f e c t s o f compe t i t i on and d imin i shes the s t reng th o f i n t e r s p e c i f i c c o r r e l a t i o n . The phys iog raph ic i n s t a b i l i t y inheren t i n the c o a s t a l s a l t marsh environment has s e l e c t e d fo r a l l o p o l y p l o i d y i n most of the marsh s p e c i e s . The long growing season and low number of species i n the community i s r e f l e c t e d i n the long average f l o w e r i n g per iods of s a l t marsh s p e c i e s . The marsh i s dominated by p e r e n n i a l , hemicrypto-p h y t i c g rasses , sedges, rushes , and g r a s s - l i k e f o r b s ; t h i s growth form dominance i s r e f l e c t e d i n the predominant wind p o l l i n a t i o n . Vegetatiy . e . j a sexua l r ep roduc t ion i s common and e x t e n s i v e , and s e l f - c o m p a t i b i l i t y i s common, but the f l o r a l and p o l l i n a t i o n b i o l o g y of the spec ies i s such tha t the m a j o r i t y are outcrossed to some ex ten t . E f f e c t i v e long d i s t ance seed d i s p e r s a l i s p robably p r i m a r i l y b i o t i c . D i s p e r s a l by wind and water i s more e f f e c t i v e over shor t d i s t a n c e s . The dominant s a l t marsh 196 s p e c i e s have l a r g e niches and v a r i a b l e p o p u l a t i o n s . Most of the s p e c i e s have wide (though l o c a l ) d i s t r i b u t i o n s ; many probably have ecotypes and a few h y b r i d i z e with c l o s e l y r e l a t e d s p e c i e s or s u b s p e c i f i c taxa. Sphagnum bogs The low n u t r i e n t l e v e l s and c o l d , waterlogged, p o o r l y oxygenated s u b s t r a t e of a c o a s t a l sphagnum bog c o n s t i t u t e a severe p h y s i c o - c h e m i c a l environment, but the equable macro-c l i m a t e and the long term p h y s i o g r a p h i c s t a b i l i t y have c o n t r i b u t e d to the r e l a t i v e l y h i g h s p e c i e s d i v e r s i t y and r e l a t i v e l y low p o l y p l o i d y l e v e l s of the two study bogs. The nature of a sphagnum bog, which i s e s s e n t i a l l y a community of v a s c u l a r p l a n t s e p i p h y t i c upon mosses, s i g n i f i e s a mosaic of microtopography, water, and n u t r i e n t c o n d i t i o n s , a l l c o n t r o l l e d p r i m a r i l y by the moss s u b s t r a t e . T h i s p a t t e r n i n g r e s u l t s i n aggregated p o p u l a t i o n s and marked i n t e r s p e c i f i c a s s o c i a t i o n i n e a r l y s u c c e s s i o n a l stages. However, s i n c e a v a s c u l a r p l a n t i n a sphagnum bog e x i s t s more i n terms of Sphagnum spp. than i n terms of other v a s c u l a r p l a n t s , i n t e r s p e c i f i c c o m p e t i t i o n i s much l e s s of a f a c t o r than i n e i t h e r the s a l t marsh or the s u b a l p i n e meadow, and c o r r e l a t i o n tends to be weak. The great e c o l o g i c a l d i f f e r e n c e s among the bog s p e c i e s f u r t h e r reduce co m p e t i t i o n . The s p e c i e s are more or l e s s e q u a l l y anemophilous or entomophilous. As with the s a l t marsh, long d i s t a n c e seed d i s p e r s a l i s probably p r i m a r i l y zoochorous, with anemochory and hydrochory more important over short d i s t a n c e s . V e g e t a t i v e 197 r ep roduc t ion i s common, but n e i t h e r as common nor ex tens ive as i n the s a l t marsh. More than 7 0% of the spec ies o f both bogs are at l e a s t p a r t i a l l y s e l f - c o m p a t i b l e , but t h e i r f l o r a l and p o l l i n a t i o n b i o l o g y promote o u t c r o s s i n g . The dominant bog spec ies probably have f a i r l y l a rge n iches and v a r i a b l e p o p u l a t i o n s , but apparen t ly are r e l a t i v e l y , poor competi tors . ' Many of the spec ies range throughout the b o r e a l and s u b a r c t i c nor the rn hemisphere, but there i s a conspicuous P a c i f i c Northwestern American element i n the bog f l o r a . Ecotypels p r o b a b l y \" a r e common, w h i l e i n t e r s p e c i f i c h y b r i d i z a t i o n i s apparent ly r e l a t i v e l y uncommon. Subalpine meadow The suba lp ine meadow has the most f avorab le o v e r a l l p h y s i c a l environment dur ing the growing season, and probably even dur ing the w i n t e r . Fac tors f a v o r i n g the e v o l u t i o n o f a l u s h , d i v e r s e p l a n t community i n an presumably unfavorable environment have been deep, w e l l - a e r a t e d s o i l , abundant mo i s tu r e , adequate n u t r i e n t s , and the h igh s o l a r r a d i a t i o n input and warm daytime temperatures dur ing the growing season, at l e a s t i n the mesic s i t e s tha t predominate i n the study meadow. H y d r i c and x e r i c s i t e s a l s o occur f r e q u e n t l y ; t h e i r d i s t r i b u t i o n i s s t r o n g l y c o r r e l a t e d w i t h micro topographic pa t te rns tha t c o n t r o l snow accumula t ion , water and n u t r i e n t r e l a t i o n s , and so- i l depth and development. The mosaic of m ic rohab i t a t s has l e d to moderately aggregated popu la t ions of s i g n i f i c a n t l y a s s o c i a t e d s p e c i e s . St rong a s s o c i a t i o n combined w i t h l u s h , t h i c k l y - g r o w i n g v e g e t a t i o n , h i g h spec ies d i v e r s i t y , 198 a shor t growing season, and a preponderance of showy-f lowered, entomophilous forbs produces in tense i n t e r s p e c i f i c c o m p e t i t i o n , which r e s u l t s i n very s h o r t , s taggered f l o w e r i n g per iods and s t rong c o r r e l a t i o n . Al though most of the spec ies are i n s e c t -p o l l i n a t e d , seed d i s p e r s a l (both s h o r t - and l o n g - d i s t a n c e ) i s most l i k e l y p r i m a r i l y by wind.' As i n the s a l t marsh, long- te rm phys iograph ic i n s t a b i l i t y has been i n s t r u m e n t a l i n s e l e c t i o n f o r a l l o p o l y p l o i d y i n a h igh percentage of the meadow f l o r a . Vege ta t ive r e p r o d u c t i o n i s f a i r l y common but not e x t e n s i v e . Almost 50% of the species are predominant ly s e l f - i n c o m p a t i b l e , and the m a j o r i t y are ou tc rossed . Subalpine meadow species probably have r e l a t i v e l y s m a l l , s p e c i a l i z e d n iches and popu la t ions tha t are not as v a r i a b l e as but have b e t t e r compe t i t i ve a b i l i t y than those of comparable s a l t marsh and sphagnum bog s p e c i e s . There are a few w i d e - r a n g i n g , a r c t i c -a l p i n e spec ies but the b u l k r o f t h e meadow f l o r a i s . widespread i n the c o r d i l l e r a of western North Amer ica . Both e c o t y p i c v a r i a t i o n and i n t e r s p e c i f i c h y b r i d i z a t i o n ^probably are r e l a t i v e l y common. My conc lus ions are best framed as answers to the three b a s i c ques t ions posed i n the I n t r o d u c t i o n . (1) Do p l an t communities of harsh p h y s i c a l environments e x h i b i t any c h a r a c t e r i s t i c p h y t o s o c i o l o g i c a l fea tures? Y e s , i f they have heterogeneous microtopographies (most d o ) , such communities w i l l have clumped spec ies popu la t ions and e x h i b i t s t rong i n t e r s p e c i f i c a s s o c i a t i o n and c o r r e l a t i o n . A l s o , l i f e - f o r m d i v e r s i t y w i l l be reduced i n communities where c o n d i t i o n s fo r growth are unfavorable f o r a major p o r t i o n of 199 each year ( e g . , a r c t i c and a l p i n e t und ra ) . An environment tha t i nc ludes c o n d i t i o n s tha t are severe but never the less w i t h i n the e c o l o g i c a l amplitude o f adapted species throughout most o f the year w i l l evolve many l i f e - f o r m s ( e g . , the Sonoran d e s e r t ) . (2) Are there any c o r r e l a t i o n s between environmenta l harshness and c e r t a i n s y n e c o l o g i c a l p r o p e r t i e s of such communities? Yes: (a) l e v e l s of p o l y p l o i d y w i t h i n the p l a n t communities i n t h i s i n v e s t i g a t i o n seem to be c o r r e l a t e d w i t h the degree of long- te rm phys iog raph ic i n s t a b i l i t y of the environment . (b) i f envi ronmenta l r i g o r i nc ludes a shortened growing season and i f the community i s p r i m a r i l y b i o t i c a l l y p o l l i n a t e d , i n t e r s p e c i f i c compe t i t i on f o r p o l l i n a t o r s w i l l i n c r e a s e , f l o w e r i n g times w i l l evolve d i f f e r e n t peaks , and there w i l l be s e l e c t i o n f o r e i t h e r showy, s p e c i a l i z e d f lowers or reduced, autogamous f l o w e r s . P ropo r t i ons of b i o t i c v s . a b i o t i c p o l l i n a t i o n and seed d i s p e r s a l are r e l a t e d only s e c o n d a r i l y to envi ronmenta l harshness ; they are e s s e n t i a l l y s ide e f f ec t s of s e l e c t i o n for an optimum growth form. I f hemic ryp tophy t i c g ra s ses , sedges , and rushes are best adapted f o r a c e r t a i n environment, wind p o l l i n a t i o n w i l l p r e v a i l ; i f herbaceous vege t a t i on i s f avo red , b i o t i c p o l l i n a t i o n w i l l predomi-nate . Seed d i s p e r s a l w i l l c l o s e l y depend on the r e l a t i v e a v a i l a b i l i t y o f b i o t i c and a b i o t i c vec to r s such as mammals, migra tory and r e s i d e n t b i r d s , w ind , and water . There i s no a priori reason f o r one or the other mode o f p o l l i n a t i o n or d i s p e r s a l to p r e v a i l i n severe environments. 200 (c) n iche s i z e (at l e a s t w i t h respec t to p h y s i c a l parameters) and p o p u l a t i o n v a r i a b i l i t y inc rease as envi ronmenta l harshness inc reases and i n t e r s p e c i f i c compe t i t i on decreases . (d) dominance o f the community by r e l a t i v e l y few species inc reases as harshness i n c r e a s e s . (e) spec ies d i v e r s i t y i s a good i n d i c a t o r of o v e r a l l e n v i r o n -mental s e v e r i t y and the s e l e c t i v e m i l i e u w i t h i n a p l a n t community. However, t h i s i s t rue on ly i n e x t e r n a l p e r s p e c t i v e . A s a l t marsh has a more r i g o r o u s e n v i r o n -ment than a maple-basswood f o r e s t , but i n which community i s Salioornia v irg m i c a more l i k e l y to su rv ive? L o w - d i v e r s i t y communities of severe p h y s i c a l environments are not n e c e s s a r i l y l e s s s t a b l e than any other communities (3) Are spec ies of such communities s e l e c t e d fo r r e p r o d u c t i v e s p e c i a l i z a t i o n s tha t ( accord ing to e s t a b l i s h e d e v o l u t i o n a r y theory) tend to reduce gene t i c v a r i a b i l i t y ? No, there i s l i t t l e evidence tha t t h i s i s so i n the four study communities. A l l four are dominated by o u t c r o s s i n g s p e c i e s ; there i s no pronounced t rend toward autogamy or apo-m i x i s (one might p r e d i c t tha t autogamous species w i l l evolve i n any undis turbed n a t u r a l p l a n t community, but w i l l not dominate such a community). Fur thermore, 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 p o t e n t i a l r e combina t i on , on the average, between the spec ies of the four d i f f e r e n t communities. The reason f o r the absence of d i f f e r e n c e s i s r e l a t e d to the l a r g e r problem of the genera t ion and maintenance o f v a r i a b i l i t y i n . ' popu la t ions and i t s r e l a t i o n to developmental 201 homeostasis and e v o l u t i o n a r y f l e x i b i l i t y . Why are 25 t o 50% of a l lozyme l o c i polymorphic i n a v a r i e t y of organisms, why are 10 to 15% of a i n d i v i d u a l ' s l o c i he te rozygous , and why i s there not a g rea te r range of v a r i a b i l i t y from organism to organism] ( c f . inter alia Gooch and Schopf 1972 ; Stebbins and Lewontin 1972; Lev in ton 1973)? Presumably, these l e v e l s of gene t ic v a r i a b i l i t y represent an e q u i l i b r i u m value adjusted by n a t u r a l s e l e c t i o n . The e q u i l i b r i u m can be mainta ined by a v a r i e t y of adap ta t ions . The d i f f e r e n t s t r a t e g i e s and adapta t ions of the species of the study communities r e f l e c t d i f f e r e n t types of s e l e c t i o n inheren t i n t h e i r community types . From an e v o l u t i o n a r y po in t of v i ew, there i s l i t t l e or no i n d i c a t i o n tha t the d i f f e r e n t types of s e l e c t i o n d i f f e r i n o v e r a l l s e v e r i t y , and the average l e v e l s of p o t e n t i a l recombina t ion probably represent some so r t o f e q u i l i b r i u m v a l u e . However, t h i s g e n e r a l i z a t i o n can be v a l i d a t e d on ly by s tudy ing o ther comparable communities a long s i m i l a r l i n e s . V. LITERATURE CITED. 202 Adams, D .A. 1963. Fac tors i n f l u e n c i n g v a s c u l a r p l a n t zona t ion i n Nor th C a r o l i n a s a l t marshes. Ecology 44: 445-456. A l l a r d , R.W. 1965. 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D i v e r s i t y and s t a b i l i t y i n e c o l o g i c a l systems. Brookhaven Symp. B i o l . 22: 1-264. 226 , and R.H. Whittaker. 1968. Ef f e c t s of chronic gamma i r r a d i a t i o n on plant communities. Quart. Rev. B i o l . 43: 42-55. Appendix 1. Species l i s t s . S a l t Marsh 227 Species Acronym A g r o s t i s e x a r a t a T r i n . AEX C a r e x tyngbyei Hornem. CLY D e s o h a m p s i a o e s p i t o s a ( L . ) Beauv. v a r . longi f l o r a DCS Bea l D i s t i o h t i s s p i c a t a ( L . ) Greene v a r . b o r e a l i s DSP ( P r e s l ) Bee t le F e s t u o a r u b r a L . v a r . l i t t o r a l i s Vasey FRU G l a u x m a r i t i m a L . GMA H o r d e u m b r a o h y a n t h e r u m Nevsk i HBR J u n o u s b a l t i o u s W i l l d . JBA L i l a e o p s i s o o o i d e n t a l i s C o u l t . \u00a3 Rose LOC * * O e n a n t h e s a r m e n t o s a P r e s l * P l a n t a g o m a o r o o a r p a Cham. S S c h l e c h t . P l a n t a g o m a r i t i m a L . ssp . j u n o o i d e s (Lam.) Hul ten PMA P o t e n t i l l a p a o i f i o a Howel l PPA P u c c i n e l l i a p u m i l a (Vasey) A . S . Hi t chcock PPU S a l i o o r n i a v i r g i n i o a L . SPA S o i r p u s o e r n u u s Vah l SCE S p e r g u l a r i a c a n a d e n s i s (Pe r s . ) G. Don v a r . SCA o o o i d e n t a l i s Rossbach S t e l l a r i a h u m i f u s a Ro t tb . SHU 2 2 8 Trifolium wormskjoldii Lehm. TWO Triglochin maritimum L. TMA Wade's Bog Agrostis aequivalvis ( T r i n . ) T r i n . AAE Apargidium boreale (Bong.) T. S G. ABO Blechnum spicant (L.) Roth BSP Carex canescens L. CCA Carex obnupta L.H. B a i l e y COB Carex pluriflora H u l t e n CPL Coptis asplenifolia S a l i s b . CAS *Coptis trifolia (L.) S a l i s b . CTR Drosera rotundifolia L. DRO Empetrum nigrum L. ENI Gaultheria shallon Pursh GSH Gentiana douglasiana Bong. GDO Gentiana sceptrum G r i s e b . GSC Juncus supiniformis Engelm. JOR Kalmia polifolia Wang. KPO Ledum groenlandicum Oeder LGR Linnaea borealis L. LBO Maianthemum dilatatum (Wood) N e l s . S Macbr. MDI Myrica gale L. MGA Pinus contorta Dougl. PCO Rhynchospora alba (L.) V a h l RAL Sanguisorba officinalis L. SMI Scirpus cespitosus L. SCS Thuja plicata D. Don TPL 229 T o f i e l d i a glutinosa (Michx . ) Pe r s . v a r . b r e v i s t y l a TGL ( H i t c h e . ) C . L . H i t c h c . T r i e n t a l i s a r c t i c a F i s c h . TAR * *Vaccinium ovatum Pursh VOV Vaccinium oxy coccus L . VOX Vaccinium uliginosum L . VUL Vaccinium v i t i s - i d a e a L . ssp . minus (Lodd.) Hu l t en VVI Ogg's Bog Agrostis aequivalvis ( T r i n . ) T r i n . AAE Apargidium boreale (Bong.) T. \u00a3 G. ABO Blechnum spicant ( L . ) Roth BSP Calamagrostis nutkaensis ( P r e s l ) Steud. CNU Carex canescens L . CCA Carex obnupta L . H . B a i l e y COB Carex p a u c i f l o r a L i g h t f . CPA Carex p l u r i f l o r a H u l t e n CPL Coptis a s p l e n i f o l i a S a l i s b . CAS Coptis t r i f o l i a ( L . ) S a l i s b . CTR Cornus unalaschkensis^ Ledeb. CUN Desehampsia cespitosa ( L . ) Beauv. v a r . DCS l o n g i f l o r a Bea l Drosera r o t u n d i f o l i a L . DRO Empetrum nigrum L . ENI Eriophorum polystachion L . EPO Nephrophyllidium o r i s t a - g a l l i (Menzies) G i l g FCG Gaultheria shallon Pursh GSH Gentiana douglasiana Bong. GDO Gentiana sceptrum G r i s e b . GSC 230 *Goodyera o b l o n g i f o l i a Raf . *Habenaria sacoata Greene **Juncus effusus L . v a r . paoifious Fern.. S Wieg. Junous supiniformis Engelm. JOR Juniperus communis L . v a r . montana A i t . JCO Kalmia p o l i f o l i a Wang. KPO Ledum groenlandioum Oeder LGR Linnaea borealis L . LBO Lyoopodium olavatum L . LCL Lyoopodium inundatum L . LIN Lysichitum amerioanum Hu l ten \u00a3 S t . John LAM Maianthemum dilatatum (Wood) N e l s . \u00a3 Macbr. MDI Myrioa gale L . MGA Pinus oontorta Dougl . PCO Plantago maorooarpa Cham. \u00a3 S c h l e c h t . PMC Rhynohospora alba ( L . ) Vah l RAL Sanguisorba o f f i c i n a l i s L . SMI Scirpus cespitosus L . SCS Thuja p l i c a t a D. Don TPL T o f i e l d i a glutinosa (Michx . ) Pe r s . v a r . TGL b r e v i s t y l a ( H i t c h c . ) C L . H i t c h c . T r i e n t a l i s a r o t i c a F i s c h . TAR Vaooinium ovatum Pursh VOV Vaccinium oxyoooous L . VOX Vaccinium uliginosum L . VUL Vaccinium v i t i s - i d a e a L . ssp . minus (Lodd.) Hul ten VVI B l a c k w a l l Meadow A c h i l l e a m i l l e f o l i u m L . AMI 2 3 1 A g o s e r i s a u r a n t i a c a (Hook.) Greene AAU Anemone o o o i d e n t a l i s Wats. AOC A n t e n n a r i a l a n a t a (Hook.) Greene ALA A r e n a r i a oaipillaris P o i r . v a r . a m e r i o a n a (Maguire) ACA Davis A r n i c a l a t i f o l i a Bong. ALF A r n i c a m o l l i s Hook. AMO * B r o m u s v u l g a r i s (Hook.) Shear * * C a r e x n i g r i c a n s C . A . Meyer C a r e x r o s s i i Boot t CRO C a r e x sipectabilis Dewey CSP C a s t i l l e j a m i n i a t a Dougl . CMI C a s t i l l e j a p a r v i f l o r a Bong. v a r . CAL a l b i d a (Penne l l ) Ownbey C l a y t o n i a l a n c e o l a t a Pursh CLA D e l p h i n i u m n u t t a l l i a n u m P r i t z . DNU ^ D e s o h a m p s i a a t r o p u r p u r e a (Wahl.) Scheele l a t i f o l i a (Hook.) S c r i b n . * D r a b a s t e n o l o b a Ledeb. v a r . n a n a (Schulz) C L . H i t c h c . E l y m u s g l a u o u s B u c k l . v a r . b r e v i a r i s t a t u s Davy EGL E p i l o b i u m a l p i n u m L . v a r . l a c t i f l o r u m EAL (Hausskn. ) C L . H i t chcock E r i g e r o n p e r e g r i n u s (Pursh) Greene ssp t EPE c a l l i a n t h e m u s (Greene) Cronq. E r y t h r o n i u m g r a n d i f l o r u m Pursh EGR F e s t u o a v i r i d u l a Vasey FVI H i e r a o i u m g r a o i l e Hook. HGR 232 E y d r o p h y l l u m f e n d l e r i (Gray) H e l l e r v a r . a l b i f r o n s HFE ( H e l l e r ) Macbr. J u n o u s d r u m m o n d i i E . Meyer v a r . s u b t r i f l o r u s JDR (E. Meyer) C L . H i t chcock * * L o m a t i u m g e y e r i (Wats.) C o u l t . \u00a3 Rose * * L u e t k e a p e o t i n a t a (Pursh) Kuntze L u p i n u s l a t i f o l i u s Agardh v a r . * s u b a l p i n u s LLA ( P i p e r \u00a3 Robbins) C P . Smith 2 L u z u l a h i t o h c o c k i i Hamet-Ahti i LHI L u z u l a s p i c a t a ( L . ) DC. LSP M i c r o s t e r i s g r a c i l i s (Hook.) Greene MGR * M i t e l l a p e n t a n d r a Hook. * O s m o r h i z a c h i l e n s i s H . \u00a3 A . P e d i o u l a r i s b r a o t e o s a Benth . v a r . l a t i f o l i a PBR (Penn.) Cronq. * ^ P e d i o u l a r i s r a o e m o s a Dougl . P e n s t e m o n p r o c e r u s Dougl . va r . t o l m i e i (Hook.) Cronq. PPR P h l e u m a l p i n u m L . PAL P h l o x d i f f u s a Benth . v a r . l o n g i s t y l i s (Wherry) Peck PDI * * P h y l l o d o o e empetrif'ormis (Swartz) D. Don P o a c u s i c k i i Vasey v a r . e p i l i s CSc r ibn . ) C L . H i t c h c . PEP P o t e n t i l l a d i v e r s i f o l i a Lehm. PDV P o t e n t i l la flab e l l i f o l i d ^ K o o ' k . PFL R a n u n c u l u s e s c h s c h o l t z i i S c h l e c h t . RES * * S a x i f r a g a o o o i d e n t a l i s Wats. Sedum l a n c e o l a t u m T o r r . SLA S e l a g i n e l l a w a l l a c e i H i e r . SWA S e n e c i o i n t e g e r r i m u s N u t t . v a r . e x a l t a t u s (Nu t t . ) Cronq. SIN 233 S e n e c i o t r i a n g u l a r i s Hook. STR S i b b a l d i a p r o c u m b e n s L . SPR Silene parryi (Wats.) H i t c h c . S Maguire SPA T h a l i c t r u m Occident ale Gray TOC T r i s e t u m s p i c a t u m L . TSP * * T r o l l i u s laxus S a l i s b . v a r . a l b i f l o r u s Gray V a c c i n i u m d e l i c i o s u m P i p e r VDE V a c c i n i u m s c o p a r i u m L e i b . VSC V a l e r i a n a s i t c h e n s i s Bong. VSI * V e r a t r u m v i r i d e A i t . VVD V e r o n i c a c u s i c k i i Gray VCU V e r o n i c a w o r m s k j o l d i i Roem. \u00a3 S c h u l t . *Species present i n t r a n s e c t s but not i n random quadra ts . -\" 'Species present i n sampling area but l o c a t e d n e i t h e r by t r ansec t s nor random quadra ts . \"\"\"after Ca lder and T a y l o r (19 68 ) . 2 a f t e r Hamet-Ahti (1971). Appendix 2. Some s t a t i s t i c s . 234 The arrangement of the 15 columns of i n fo rma t ion i s as f o l l o w s . The s i x l e t t e r s of the f i r s t column are acronyms of a g iven species p a i r ; e g . , DCSFRU r e f e r s to the species p a i r D e s c h a m p si a c e s p i t o s a x F e s t u c a r u b r a . Refer to Appendix 1 f o r the i d e n t i t y o f the acronyms. W i t h i n each community, the spec ies are arranged i n order of decreas ing importance v a l u e . Next , there f o l l o w f i v e columns of + ' s or - ' s tha t i n d i c a t e s i g n i f i c a n t 2 p o s i t i v e or negat ive X values of a s s o c i a t i o n . Each column r e f e r s to a d i f f e r e n t quadrat s i z e ; i n o r d e r , the s i z e s are 1, 0 .25 , 0 . 0 1 , 0 .45 , and 0.09 m . In a g iven column, there are three + ' s or - ' s i f P = 0 .005; two i f P = 0 .01 ; one i f P = 0 .05 ; none i f P i s g rea te r than 0 .05. Columns 6-11 g ive the s i g n and va lue of C o l e ' s c o e f f i c i e n t of i n t e r s p e c i f i c ' a s s o c i a t i o n f o r each of the 2 f i v e quadrat s i z e s ; aga in 1, 0 .25 , 0 . 0 1 , 0 .45 , and 0.09 m , r e s p e c t i v e l y . Columns 12, 13 , and 14 g ive the s i gn and va lue of 2 r, the c o e f f i c i e n t of c o r r e l a t i o n , f o r the 1, 0 .25 , and 0.01 m quadra ts , r e s p e c t i v e l y . An a s t e r i s k f o l l o w i n g an r value i n d i c a t e s P = 0 .10. The l a s t column l i s t s the value of d, the e c o l o g i c a l d i s t ance between members of a g iven spec ies p a i r (see Sec t . I I I - F ) . S a l t 1 2 3 4 5 6 7 8 DCSFRU +++ +++ +++ +++ + . 39 + \u2022 40 DCSTMA +++ +++ DCSSVI mm - \u2014 - mm . 78 DGSPMA + \u2014 + \u2022 09 06 DCSJBA ++ +++ + +++ \u2022+\u2022++ + \u2022 13 + \u2022 18 DCSCLY +++ +++ + , 03 0. 0 DCSGMA +++ +++ +++ +++ +++ + \u2022 28 22 DCSPPA + + +++ +++ + . 06 + \u00bb 07 DCSAEX + +++ +++ 05 + \u2022 07 DCSSHU ++ +++ +++ + . 10 06 DCSTWO +++ 03 + , 03 DCSSCE +++ +++ 05 + , 01 DCSPPU 14 mm j 06 DCSSCA - \u2014 km . 18 mu m 40 DCSDSP mm mm - \u2014 \u2014 - 50 - \u2022 65 DCSHBR + \u2022 01 02 DCSLOC ++ +++ FRUTMA + ++\u2022+ + \u2022 02 FRUSVI \u2014 \u2014 99 88 FRUPMA 26 - . 20 FRUJBA +++ + ++ + +++ +++ 35 + , 33 FRUCLY + \u2014 M m. 44 45 FRUGMA ++ + +++ +++ +++ +++ + \u2022 56 + , 43 FRUPPA ++ + + ++ +++ +++ +++ + \u2022 16 + \u2022 15 FRUAEX +++ + ++ +++ +++ + \u2022 14 + \u2022 15 FRUSHU +++ + + + +++ +++ + . 20 + t 14 FRUTWO + + +++ +++ + \u2022 08 + \u2022 06 FRUSCE +++ - \u2022 16 35 FRUPPU 67 84 FRUSGA - \u2022 71 51 FRUDSP - \u2014 mm ^ 80 mm m 79 FRUHBR +++ + + + + , 03 + , 03 FRULOC +++ TMASVI 05 TMAPMA +++ + . 03 TMAJBA + TMACLY + + + + 06 TMAGMA + + + + 00 TMAPPA Marsh, 9 10 1 1 12 13 14 15 + \u202207 + \u2022 98 + < .49 -.125 -.157 -.354 .265 + \u202208 - i .20 + .098 + \u2022123 -.157 .399 -\u2022 15 -\u2022 57 -, .38 -\u2022568* -.121 -.054 \u2022 508 -\u202209 -\u2022 14 - i \u2022 50 -.539* -\u2022274* -.750 .426 + . 17++# 72 + i .49 +.364* + .192 -.409 \u2022 406 + .07 + . 10 .24 -\u2022094 + .012 -.078 .465 + .37 + . 98 + . 38 -.115 -\u2022350 + .041 \u2022 544 -.20 + \u2022 99 + \u2022 05 + .001 -.038 \u2022 591 + .02 96 + i l l -\u2022353 -.091 .294 + .02 + \u2022 96 \u2022 11 + .068 -.145 -.141 \u2022 591 + .01 68 + \u2022 11 + .263 -.041 .549 -.33 + \u2022 94 \u2022 03 -.119 + .353 .405 MM * 62 \u2022 68 -.420 -.522 .247 -\u2022 33 - .63 -.127 .518 - , 60 - \u2022 69 .393 + \u2022 73 + \u00bb01 .337 .506 + .08 + .07 +.305* +.337* + .115 .410 -.29 67 .64 -.299 -.277 -.325 .482 -.41 03 - \u2022 31 -.162 -.259 .445 + . 17 + , 82 .69 -.036 -\u2022166 -.425 .354 + .06 + \u00bb 32 .08 -\u2022211 -.270 + .016 .382 + .49 + \u2022 83 + \u00ab .56 + \u2022129 + .093 -.115 .483 + .18 + \u2022 95 + \u2022 09 -\u2022231 -.276 -.570 .558 + .02 + . 87 + \u2022 20 -\u2022328 -#563* .353 + .23 + \u2022 78 + .14 + .284 + \u2022088 .551 + .04 + \u2022 99 + i .20 -.213 -\u2022359 \u2022 499 -.57 + \u2022 38 + i \u2022 01 + .383 .397 -\u2022 73 - i .98 .319 - t 91 mm .98 .529 \u2014 | 80 + .88 .04 .270 .342 + \u2022 74 .01 .423 -.08 - t 58 -, .60 -.065 -\u2022103 -.125 .373 -.27 + .01 -\u2022052 + .066 -.041 .432 + .02 .01 -.400* -.519* -.245 .288 -.07 + . 17 + , .08 -.065 + .059 -.063 .362 + .17 + .04 + .249 -.071 + .048 .453 -.10 .00 -.551* -.593* + .375 .569 TMAAEX + + + -.34 \u2022 00 + i \u2022 01 -\u2022046 -.047 .411 TMASHU + + .01 + .05 + < .01 -\u2022082 + .145 + .119 \u2022 540 TMATWO -.43 + .01 + i .01 + \u2022029 -.438 \u2022 542 TMASCE \u2022 00 -.40 .00 + .220 -.253 .391 TMAPPU \u2022 00 + . 52 -, .10 +\u2022617* +.644* \u2022 354 TMASGA \u2022 00 + \u202256 - i .08 +\u2022557* + .387 \u2022 541 TMADSP \u2014 \u2022 00 -.28 -\u2022 66 - i .41 \u2022 375 TMAHBR - + -.46 .00 .443 TMALOC .00 \u2022 460 SVIPMA + + + + + + ++ + + + + + + + + \u202239 + .50 + .38 + . 28 + i .19 +\u2022576* +\u00ab529* + .193 \u2022 479 SVUBA -\u202283 -.78 -.87 -\u00ab 63 - i .68 -.349 + \u00ab022 \u2022 403 SVICLY + + + + + + ++ + + \u202241 + .48 + .35 + \u2022 26 - i .11 + .056 -.074 + .329 .432 SVIGMA \u2014 -.77 -.48 -.41 -\u00bb 49 - .47 -.422 -.162 + .218 \u2022 462 SVIPPA -.53 -.63 -.79 -\u2022 29 - i .93 \u2022 544 SVIAEX -.91 -.91 -.76 -\u2022 21 - i .72 \u2022 498 SVISHU + *20 + .19 + .11 + * 12 -. .09 -\u2022130 .076 -.269 \u2022 480 SVITWO - \u2014 -. 19 -.20 - . 74 - .86 -\u2022455 \u2022 532 SVISCE ++ + ++ + . 13 + .09 + .10 -\u2022 04 - .01 + .163 + .245 + .154 \u2022 432 SVIPPU + + + + + + + + + + .11 + .09 + \u00ab 97 + .11 +\u2022494* + .578 \u2022 491 SVISCA + + + + + + + + + .08 + .05 + \u2022 61 + (.09 +.734* + .578 \u2022 556 SVIDSP + + + + + + .05 + .0 7 -.15 + \u2022 42 + .04 \u2022 461 SVIHBR am a * \u2014 -\u2022 83 -< .99 \u2022 507 SVILOC \u2014 -\u00bb 89 -< .93 \u2022 497 PMAJBA ...... ammmmm - -.41 -.41 -.81 10 - .29 -.418 + .218 \u2022 446 PMACLY \u2022+\u2022 + + +++ + + + \u2014 + .30 + .35 -.01 + \u2022 45 - .47 -.086 -.123 + .284 \u2022 468 PMAGMA -.20 -.15 -.67 -\u2022 05 mm .04 -.379 -\u2022149 \u2022 480 PMAPPA - -.23 -\u202235 08 - .48 -.121 + .492 \u2022 529 PMAAEX - -\u202262 -\u202213 12 - .37 \u2022 376 PMASHU + + + +++ + + + + .28 + .25 + . 11 -* 12 + \u2022 10 -\u2022 164 -\u2022195 -.450 \u2022 533 PMATWO + .01 + .02 -.36 -1 44 - .48 -.320 \u2022 532 PMASCE + + + +++ ++ + + + + + + + + .20 + .12 + .20 + \u00bb 78 + t .07 + \u2022336 + .270 + .028 \u2022 488 PMAPPU + + + + + + + .09 + .09 + \u2022 74 + i .04 +.769* +.711* .430 PMASCA + + + + + + + + + .08 + .08 + \u2022 49 + ( .09 +\u2022594* + \u2022597 .522 PMADSP - -.07 + \u202201 + .02 -\u2022 29 mm t .30 \u2022 447 PMAHBR - 64 .99 .410 PMALOC + \u00ab 20 + .01 .461 JBACLY ++ + -.67 -.67 -.57 + \u00ab 27 + .04 + .051 -.198 .354 JBAGMA + + + + + + + + + + + + + + \u202241 + .37 + .18 + \u00ab 73 + .43 + .205 + .270 + .115 \u2022 480 JBAPPA + + + + + + + + + + + + + + + + .25 + .24 + .24 89 + .08 -\u2022103 + .110 -.557 \u2022 555 JBAAEX +++ +++ + +++ + + + + 1 24 + .28 + .19 + \u202280 + \u2022 17 -\u2022444* -.364 .425 JBASHU - +++ + + + + , 02 -\u202209 + .46 + \u00bb07 + \u2022001 -\u2022063 .532 JBATWO +++ + + + 05 + .01 + \u202281 + .17 -.127 .531 JBASCE - \u2014 + + + + 53 + .29 + .06 .420 JBAPPU --- \u2014 85 -.99 -.99 .396 JBASCA - - rn.rn.rn. - 4 60 -.99 - i .99 \u2022 562 JBADSP mm mm mm mm rn.rn.rn. .rn.rn. -\u202295 - i .99 \u2022 328 JBAHBR + + + + + + + 1 .05 + .05 + .03 \u2022 476 JBALOC + + + + + + + \u2022 03 \u2022 482 CLYGMA + + + - \u00ab 40 -.12 \u2022 00 + \u202229 + i .04 -.141 -.252 +.430 .444 CLYPPA mm mm, mm \u2014 - | 47 -.65 -.76 -\u202211 -( .28 -.413 \u2022 490 CLYAEX \u2014... - 45 -.47 -.73 + .19 + ( .01 -.442 -.117 \u2022 430 CLYSHU ++ + + + + + + + | 15 + .15 + .15 + .51 + .03 + \u2022001 + \u2022010 -.166 .517 CLYTWO m. + . 02 -.07 -.03 -\u202208 - .26 +.799* .517 GLYSCE + + + + + + + + + \u00ab 11 + .07 -. 19 + \u00ab .02 + \u2022061 +\u2022649* \u2022 358 GLYPPU + + + \u00ab 08 + .07 -\u202206 -< .12 + \u2022411 -\u2022364 .420 CLYSCA + + + + + < 06 + .05 + \u202228 + < .03 +.690* \u2022 567 CLYDSP - - I 04 -.15 -\u2022 16 mm .17 .337 CLYHBR - \u2014 - 1 73 -.70 -.09 - i .41 .421 CLYLOC + + + \u2022 00 + i .01 \u2022 450 GMAPPA + + + + + + + + \u00ab 13 + .05 + .03 + \u2022 90 + t .08 -.070 -.043 .525 GMAAEX + + + + + + + + + \u00ab 13 + .12 + .05 + .42 + i .14 + .084 + \u2022082 .501 GMASHU + + + + + + + + ( 11 + .12 + . 15 + \u202236 + ( .08 + .162 + .091 -.645* .439 GMATWO + + + + ++ + + | 08 + .04 + .07 + .81 + i .15 +\u2022667* + .636 \u2022 480 GMASCE - - + + + -\u2022 16 -.60 + \u202242 + i .01 -\u2022280 .526 GMAPPU rn.rn.rn. - 63 -.60 -\u202289 -< .79 \u2022 492 GMASCA rn.rn.rn. mm | 36 -.40 -. 87 - i .72 \u2022 526 GMADSP + rn.rn.rn. rn.rn.rn. 54 -.49 -.53 -< .48 \u2022 443 GMAHBR + + i 04 + .06 + .24 -< .29 .511 GMALOC + + + + + + + \u202275 .01 .379 PPAAEX +++ + ++ +++ + + + + + + + t 44 + .46 + .54 + .26 + .76 +\u2022543* +,596* -.272 \u00bb545 PPASHU -, 04 -.22 -.46 + \u202202 - .25 -\u2022024 \u2022 453 PPATWO + + + + + + + + + \u2022 13 + .20 + \u202245 + .50 \u2022 523 PPASCE -1 78 + \u202204 -.30 .556 PPAPPU - mi -.99 - i .99 \u2022 569 PPASCA - \u2014 -.99 -< .99 \u2022 586 PPADSP \u2014 -< .99 .525 PPAHBR ++ +++ + + + + + + 15 + .18 + \u202262 + i .06 \u2022 539 PPALOG + + + + + + + .35 + i .05 .473 AEXSHU \u2014 - -.69 -.80 AEXTWO + + + +++ + .09 \u2022 14 AEXSCE \u2014 +++ AEXPPU - \u00ab\u2022\u2022\u00ab\u2022\u00ab\u2022 AEXSCA \u2014 mmmmmm AEXDSP \u2014 \u2014 AEXHBR +++ +++ + + + ++ + + . 18 + .18 AEXLOC + SHUTWO + + + \u2022 16 + .09 SHUSGE + + + + +++ + . 12 + .07 SHUPPU \u2014 \u2014 -.60 -.63 SHUSCA M M W* -.48 -\u202244 SHUDSP mm SHUHBR SHULOC TWOSCE -TWOPPU \u2014 TWOSGA \u2014 TWODSP mm mm TWOHBR + + ++ + + .02 + .06 TWOLOC ++ SCEPPU ++ + + .28 + .23 SCESCA +++ + \u2022\u00a5 + + .24 + .27 SCEDSP - + .08 + .09 SCEHBR SCELOC + + PPUSCA +++ +++ ++ + + .46 + .15 PPUDSP ++ + + .06 + .01 PPUHBR - -PPULOC - -SCADSP + + + + + + +++ + .33 + .63 SCAHBR Mi SCALOC - -DSPHBR -DSPLOG HBRLOC + .03 + \u202204 .536 + .79 + .19 .498 + \u202205 + .11 .412 -.99 -.99 .283 -.99 -.99 .472 -\u202299 -.99 .404 + .71 + .12 \u2022 248 + \u202235 + .01 \u2022 461 09 + .18 + .05 -.076 .532 16 + .31 + .01 -.048 \u2022 544 -.75 -.90 .544 -.65 -.62 .510 -.20 -.08 .498 .512 + .14 + .01 \u2022 482 -. 17 -.55 .558 -.99 -.99 .540 -.99 -.99 .617 -.99 -.99 + .13 .499 .508 + .27 \u2022 00 .480 -.41 + .08 -.227 .328 -.09 + \u202206 -.072 .495 10 -.67 -.64 .396 .401 + .26 + \u202203 .470 + .60 + .32 +.912* .461 + .16 + .18 .350 .298 .452 + .09 + .13 .561 .462 .470 .395 \u2022 422 .452 Wade's Bog CPLABO -.63 + .05 + .01 -.236* -\u2022436* -.600* .530 CPLAAE + \u202228 -.13 -.14 + .168 + .152 + .149 .375 CPLCOB -.39 -.41 -.274* -.191 -.302 .290 CPLSMI mm 16 -.28 +.318* + .154 +.509* .478 CPLKPO + .27 -.18 + .235 + .223 +.508* \u2022 506 CPLVOX -.23 -.29 + .058 -\u2022041 -.270 .534 CPLCCA _. -\u202255 -.51 + .01 -.442* -.413 \u2022 328 CPLDRO +++ + .11 + .07 + .08 + .14 + .231 + .204 -.066 \u2022 508 CPLLGR + .03 + .03 -.09 -. 18 -.07 -.002 + .059 .585 CPLTAR - + -.41 + .06 + .05 -.022 -.168 + .210 .505 CPLENI + \u202202 + .01 -.39 .00 -.04 -.448 -.306 .527 CPLMGA + .02 -.17 -.05 \u2022 00 -.037 + .004 .463 CPLTGL -.54 .\/\/ + .01 + .218 + .499 \u2022 464 CPLTPL + \u202201 -.19 -.04 + .348 CPLPCO + .01 + .01 .00 -.313 CPLGDO + \u202201 + .01 -.05 -.513 .499 CPLLBO + \u202201 -.08 \u2022 00 -.446 .544 CPLMDI + .01 .470 CPLRAL \u2022 00 \u2022 00 \u2022 365 CPLVVI \u2022 602 CPLSCS \u2022 331 CPLJOR \u2022 00 \u2022 00 \u2022 403 CPLBSP GPLVUL \u2022 530 CPLGSH \u2022 565 GPLGSC \u2022 509 GPLCAS \u2022 441 ABOAAE + + + +++ *.07 + .43 + .31 + .211 + .071 + .121 \u2022 505 ABOCOB + .02 + .11 + .08 + .05 \u2022 00 + .182 + .175 + .356 .526 ABOSMI -.11 + .07 + .04 -.122 + .213 -.535* \u2022 499 ABOKPO + + + +++ + .01 -.08 + .18 + .06 -\u2022081 + .013 + .112 .519 ABOVOX + + + +++ + .05 -.18 + .33 + .17 -.071 + .031 -.249 \u2022 546 ABOCCA -.04 + .03 + .01 -.06 \u2022 00 -.305 -.307 -.164 \u2022 510 ABODRO + + + +++ -.16 + .24 + .09 -.129 -.097 + .524 .495 ABOLGR -.17 -.11 -.12 + .02 -.01 -.110 -\u2022400 .527 ABOTAR + + + ++ + -.17 + .19 + .13 + .097 + .149 + .013 \u2022 489 ABOENI + .03 + .03 + .02 + .03 .00 -.371 -.600 \u2022 581 ABOMGA -.31 -.29 -.29 -\u202236 + .235 -.029 .561 ABOTGL + -.52 -.41 + .04 + .02 -.018 -.289 .507 ABOTPL - -\u202236 ABOPCO . . . -.38 ABOGDO + + *+ + + \u202201 ABOLBO -.15 ABOMDI + .01 ABORAL - . . . ABOVVI ABOSCS ABCUQR \u2014 . . . ABOBSP ABOVUL ABOGSH ABOGSC ABOCAS AAECOB +++ ++ + .11 \u2014 . AAESMI ++ +++ + .01 \u2014 . AAEKPO +++ +++ + .11 \u2014 . AAEVOX 4-++ + + + + .13 + \u2022 AAECCA mm + \u202202 -.19 + . AAEDRO + + + +++ + .19 + \u2022 AAELGR + .05 + \u2022 AAETAR + + + + +++ + .28 \u2014 . AAEENI + \u202201 + .02 \u2014 . AAEMGA - + \u202201 -\u202206 AAETGL + .02 + .03 AAETPL mm mm mm* + \u202201 AAEPCO + \u202201 AAEGDO - ++ AAELBO hmm - \u2014 \u2022 00 AAEMDI \u2022 00 AAERAL \u2014 . . . AAEVVI AAESCS AAEJOR . . . . . . AAEBSP AAEVUL AAEGSH AAEGSC AAECAS .00 -.15 -.008 -\u202205 -\u202227 +.722* + .06 + .03 \u2022 480 + .01 -.14 \u2022 524 \u2022 488 -.21 -.19 \u2022 510 \u2022 536 .524 -.54 \u2022 531 .480 \u2022 495 .435 \u2022 546 13 + \u202210 + .06 -.168 -\u2022133 \u2022 393 01 + .16 + .15 + \u2022192 -.011 -\u2022418 .487 05 + \u202209 + .20 -.146 + \u2022082 + .283 \u2022 520 31 + .35 + .18 + \u2022213 + .083 \u2022 553 08 + .03 -.10 - \u2022 4 1 9 * -\u2022078 + \u2022656 \u2022 325 16 + .20 + .13 + .118 + \u2022097 + \u2022 194 \u2022 516 03 + .02 + .01 -\u2022141 -\u2022043 \u2022 593 02 + .26 + .23 +.297* + \u2022111 + .085 .424 13 + .01 -.04 -\u2022359 -.522 \u2022 510 -.17 -.23 + .176 + .167 .478 + .02 -\u202207 -\u2022304 -.172 .476 -.01 -.29 -\u2022352 .00 -\u202210 -\u2022340 + .03 + \u202203 -\u2022547 \u2022 456 -.19 -\u202246 + \u2022069 \u2022 574 .471 -\u202246 -.46 \u2022 379 .587 \u2022 356 -\u202240 -.99 \u2022 341 \u2022 .542 \u2022 568 \u2022 474 \u2022 517 COBSMI - . 3 8 - . 2 0 + . 0 9 + . 0 5 + . 0 3 - . 3 8 1 * - . 1 9 6 + . 6 4 6 . 5 1 3 COBKPO +++ + . 16 - . 3 8 + . 2 6 + . 0 4 + .112 + . 1 5 3 . 5 1 5 COBVOX +++ + - . 4 1 - . 3 6 + . 1 0 + . 05 - . 0 9 5 - . 1 6 8 .572 COBCCA - . 0 4 + . 0 7 + . 0 6 - . 1 4 - . 2 6 1 - . 2 6 7 \u2022 300 COBDRO + - . 1 7 - . 2 7 - . 2 9 + . 1 0 - . 0 3 - . 4 7 1 * - . 2 4 9 . 5 4 2 COBLGR +++ + + \u2022 17 + . 0 5 + .11 + . 4 7 + . 0 5 + .203 + \u2022525* \u2022 592 GOBTAR + ++ + . 0 6 + . 12 + . 0 8 - . 0 3 7 - . 1 0 2 \u2022 541 GOBENI + + . 0 2 + . 1 0 + .11 - . 1 0 - . 0 3 + .361 - . 2 0 5 \u2022 514 COBMGA + . 0 6 + . 0 2 - . 0 1 \u2022 00 - . 1 4 7 + . 1 5 3 \u2022 453 COBTGL - . 4 2 - . 0 3 + . 0 4 - . 1 4 - . 2 1 5 - . 4 6 5 \u2022 516 GOBTPL + . 01 - . 0 6 - . 0 7 - . 1 0 7 COBPCO \u2022 00 - . 0 1 + \u202202 + . 197 GOBGDO + \u202203 - . 1 5 - \u2022 0 7 - . 5 7 5 \u2022 516 COBLBO - \u2022 1 7 + .01 + \u202201 . 5 9 4 COBMDI \u2022 00 \u2022 489 COBRAL - . 8 0 - . 9 9 \u2022 369 COBVVI \u2022 633 COBSCS \u2022 337 COBJOR \u2014 \u2014 - . 8 2 - \u2022 9 9 \u2022 432 COBBSP COBVUL \u2022 549 COBGSH . 5 3 4 GOBGSC . 5 1 7 COBCAS \u2022 516 SMIKPO + +++ + .11 + . 0 3 - . 0 1 + . 0 3 + .12 + .346* + .542* + . 161 .521 SMIVOX +++ - . 0 9 - . 0 8 + . 0 9 + . 1 5 + .406* + .098 + . 6 9 9 * \u2022 562 SMI CCA +++ + . 0 4 + . 0 1 + . 0 9 + . 0 3 + . 0 4 + .127 + . 2 2 0 + . 6 2 9 . 5 2 4 SMIDRO +++ + .01 - . 2 4 + .11 + . 0 3 + . 15 + . 3 1 3 * + .0 34 + . 1 2 0 \u2022 410 SMILGR + . 13 + . 0 9 - . 0 9 - . 0 5 \u2022 00 + .121 + . 0 9 3 . 5 6 0 SMITAR - . 2 4 - . 2 0 + .01 + . 0 3 + .130 - . 0 3 8 + . 001 \u2022 471 SMIENI - . 1 6 - . 0 4 - . 5 5 + .02 + .01 - . 1 2 1 + . 496 . 5 8 2 SMIMGA - . 1 3 - . 1 0 - . 0 4 - . 4 6 + . 7 4 1 * + . 9 6 6 * \u2022 477 SMITGL - . 0 4 + . 0 4 - . 2 7 - . 0 3 + .191 + . 0 2 5 \u2022 430 SMITPL - . 2 0 - . 0 2 - . 1 1 - . 3 4 4 SMIPCO - . 0 6 \u2022 00 - . 0 1 - . 4 1 3 SMIGDO +++ \u2022 00 + . 02 + . 0 3 + .673 \u2022 486 SMILBO - - - . 4 1 \u2022 00 - . 2 8 . 5 6 7 SMIMDI \u2014 - - . 7 6 .522 SMIRAL - . 0 7 - . 1 2 . 5 0 5 SMIVVI SMISGS SMIJOR SMIBSP SMIVUL SMIGSH SMIGSC SMICAS KPOVOX KPOCCA KPODRO KPOLGR KPOTAR K.POENI KPOMGA KPOTGL KPOTPL KPOPCO KPOGDO KPOLBO KPOMDI KPORAl KPOVVI KPOSCS KPOJOR KPOBSP K.POVUL KPOGSH KPOGSC KPOCAS VOXGCA VOXDRO VOXLGR VOXTAR VOXENI VOXMGA VOXTGL VOXTPL VOXPCO - . 2 1 - . 6 4 +++ . 5 9 6 \u2022 526 \u2022 560 \u2022 498 \u2022 561 \u2022 452 \u2022 439 +++ +++ + . 2 9 + .51 + . 2 6 + \u202224 + . 3 9 8 * + \u2022390* - . 3 7 8 \u2022 461 + \u202202 \u2022 00 + . 0 6 \u2022 00 + .02 - . 1 9 7 - \u2022 3 0 9 \u2022 524 +++ +++ + . 0 1 + . 31 + . 3 4 + . 1 6 + . 2 2 4 + . 3 8 2 * - \u2022 0 0 7 \u2022 533 +++ +++ + \u202205 + . 1 6 + . 0 3 + . 12 + . 15 + . 4 2 6 * + \u2022473* \u2022 367 +++ +++ + .21 + . 0 6 + . 2 7 + . 1 9 - \u2022 0 0 9 - \u2022 0 6 9 + . 2 8 4 \u2022 514 + . 01 + . 0 4 - . 3 8 - \u2022 0 3 + . 02 - . 2 6 1 - . 1 7 0 \u2022 503 + \u202201 + . 0 5 + .01 - \u2022 0 6 - . 3 3 5 - \u2022 2 1 7 \u2022 486 + \u202202 + . 0 1 + . 0 3 + .02 - . 0 9 9 + \u2022302 \u2022 491 + \u202201 + .01 - . 0 4 - . 2 5 1 + \u202201 \u2022 00 \u2022 00 - . 2 5 3 \u2022 00 - . 1 9 - \u2022 1 5 + \u2022211 \u2022 454 \u2022 00 \u2022 00 - . 1 6 - . 2 1 7 \u2022 471 \u2022 00 . 5 1 9 - . 3 1 - . 3 7 \u2022 548 \u2022 484 \u2022 526 \u2014 - . 8 4 - . 9 9 \u2022 528 \u2022 387 \u2022 415 \u2022 422 \u2022 509 + . 0 2 - . 6 8 + . 0 7 + .01 + .01 - \u2022 0 6 1 - . 0 2 6 + . 6 2 4 * \u2022 583 +++ +++ + . 1 7 + . 2 4 + . 2 0 + . 1 7 + . 3 6 1 * + \u2022237 - . 0 6 4 \u2022 556 +++ +++ + . 0 3 + . 0 3 + . 0 8 + . 12 + . 3 9 0 * + \u2022007 . 5 1 3 +++ +++ + . 2 6 + . 14 + .22 + . 2 0 + \u2022073 - \u2022 0 8 9 - . 2 2 4 \u2022 568 +++ + . 0 1 - . 0 5 - . 3 6 + .02 + .03 - . 2 8 7 - . 4 2 8 \u2022 438 + . 0 1 . 0 0 + .01 \u2022 00 - \u2022 4 2 2 - \u2022 2 2 9 \u2022 579 +++ ++ + . 0 2 + . 0 4 + . 42 + .02 + \u2022088 + \u2022465* \u2022 535 + + \u202201 + .01 + \u202201 - \u2022 1 5 3 + + .01 + .01 + .01 - . 1 7 0 VOXGDO + + \u2022 00 + .04 -.04 + .492 \u2022 530 VOXLBO \u2022 00 \u2022 00 + \u202201 +\u2022903* .390 VOXMDI \u2022 00 .511 VOXRAL > -.38 -.51 .538 VOXVVI \u2022 364 VOXSCS \u2022 552 VOXJOR -.55 -\u202299 \u2022 538 VOXBSP VOXVUL \u2022 457 VOXGSH \u2022 473 VOXGSC \u2022 543 VOXCAS \u2022 500 CCADRO + + -.04 -.06 -.29 + \u202205 + .03 -.314 -\u2022489* \u2022 534 CCALGR -\u202236 -.19 -.08 + \u202202 +\u2022522* + .553 \u2022 596 CCATAR -.04 + .05 + .13 .00 + \u202202 -.017 + .039 -.245 .483 CCAENI -\u202268 -.67 -.19 \u2022 00 \u2022 524 CCAMGA + .07 + .08 \u2022 00 \u2022 00 -\u2022170 .479 CCATGL + .20 + .12 + .09 -.27 -.481* -\u2022744* \u2022 514 CCATPL -\u202243 + .03 -.09 +\u2022591* CCAPCO - -.38 -.13 -.74 CCAGDO + \u202208 \u2022 498 CCALBO -.65 + .01 + .01 \u2022 590 CCAMDI -.58 .507 CCARAL - -.45 -.84 \u2022 330 CCAVVI \u2022 631 CCASCS \u2022 300 CCAJQR - -\u202232 \u2022 381 CCABSP CCAVUL \u2022 558 CCAGSH \u2022 565 CCAGSC \u2022 504 CCACAS \u2022 549 DROLGR + .02 + .07 + .13 + .212 -.098 \u2022 508 DROTAR ++ + . 13 + .24 + .10 +\u2022506* + \u2022160 -.240 \u2022 463 DROENI - -.55 -.51 -.057 \u2022 544 DROMGA -.31 + .01 -.175 -.271 .494 DROTGL + .08 + .05 + .137 + .135 \u2022 442 DRGTPL + .04 -\u2022123 DROPCO -.17 -.362 DROGDO + \u202202 + .184 \u2022 500 DROLBO -\u202262 -.08 \u2022 00 \u2022 559 DROMDI -.77 \u2022 513 DRORAL -.40 -.37 \u2022 513 DROVVI \u2022 604 DROSCS \u2022 536 DROJOR \u2014- -.68 -.99 .537 DROBSP \u2022 541 DROVUL DROGSH \u2022 568 DROGSC \u2022 \u2022516 DROCAS \u2022 505 LGRTAR +++ + .03 + .01 -.50 + .04 + \u202209 +\u2022350* + .149 \u2022 507 LGRENI + + +++ + .04 + .04 + .33 + \u202209 + \u202213 +\u2022708* \u2022 514 LGRMGA + + .09 -.04 + .05 + \u202201 -.261 \u2022 509 LGRTGL + + -.10 + .09 + .11 + .04 + .017 +.770* .518 LGRTPL + + \u202208 + .05 + \u202202 -\u2022137 LGRPCO + .04 + .03 \u2022 00 -\u2022403 LGRGDO + .04 -.19 -.40 +\u2022906* \u2022 501 LGRLBO + +++ + .06 + .04 + .02 + .404 \u2022 486 L.GRMD I + .02 \u2022 483 LGRRAL -.64 -.77 \u2022 571 LGRVVI \u2022 499 LGRSGS \u2022 584 LGRJOR - \u2014 -.56 -\u202299 \u2022 586 LGRBSP LGRVUL \u2022 461 LGRGSH \u2022 388 LGRGSC \u2022 469 LGRCAS \u2022 558 TARENI - +++ -.21 \u2022 00 + .01 + .04 -\u2022252 + .314 \u2022 523 TARMGA mmmm -.31 + .03 -.29 -.35 -\u2022288 -\u2022322 .473 TARTGL -\u202204 + .02 + \u202204 \u2022 00 + \u2022042 -.118 \u2022 442 TARTPL - + .01 -.16 -\u202228 -\u2022425 TARPCO + \u202201 -.07 -.12 -\u2022077 TARGDO +++ + \u202201 + .08 + \u202202 + .260 \u2022 412 TARLBO - - + .01 -.27 -.32 - \u2022 7 5 0 * \u2022 562 TARMDI -\u202243 .457 TARRAL -.38 -.61 \u2022 480 TARVVI \u2022 588 TARSCS .497 TARJOR - \u2014 -.99 -\u202299 .488 TARBSP TARVUL \u2022 504 TARGSH \u2022 536 TARGSC \u2022 466 TARCAS \u2022 484 ENIMGA -.18 -.38 + .04 + \u202202 .416 ENITGL - - - -.52 -.55 -.25 -\u202239 .572 ENITPL +++ + .01 + .06 + \u202212 ENIPCO +++ + .14 + .08 + \u202206 EN IGDO \u2014 - - -\u202245 -.54 -\u202233 \u2022 519 ENILBO + +++ +++ + .21 + \u202211 + .09 \u2022 506 ENIMDI +++ + .23 .549 ENIRAL + .03 -.20 .473 ENIVVI \u2022 444 ENISCS .487 ENIJOR - \u2022 -.57 \u2022 497 ENIBSP ENIVUL .479 ENIGSH .490 ENIGSC .564 ENICAS \u2022 541 MGATGL + .04 -\u202203 -\u202210 -.494 \u2022 501 MGATPL +++ + .08 + .12 + .12 MGAPCO + .20 + \u202204 + .03 MGAGDO - - -.49 -.85 \u2022 486 MGALBO + .03 \u2022 588 MGAMDI + .13 \u2022 546 MGARAL \u2022 472 MGAVVI \u2022 611 MGASCS \u2022 482 MGAJOR \u2022 521 MGABSP MGAVUL \u2022 480 M6AGSH .532 MGAGSC \u2022 510 MGAGAS .540 TGLTPL + \u202204 +\u202205 -.12 +.207 TGLPCO -\u202238 -\u202209 -.24 TGLGDO -.11 -\u202210 -.06 .475 TGLLBO + \u202201 \u2022 00 .491 TGLMDI -.17 .370 TGLRAL -\u202232 -.10 \u2022 514 TGLVVI .587 TGLSCS \u2022 498 TGLJOR -\u202268 -.67 \u2022 499 TGLBSP TGLVUL .523 TGLGSH \u2022 534 TGLGSC \u2022 481 TGLCAS \u2022 445 TPLPCO + +.09 +.12 +.07 TPLGDO - + -.25 -.85 TPLLBO +++ +++ + .17 +\u202227 +.30 TPLMOI -\u202209 TPLRAL TPLVVI TPLSCS TPLJOR TPLBSP TPLVUL TPLGSH TPLGSC TPLCAS PCOGDO \u2014 -.29 -.08 -.87 PCOLBO + \u202209 +.02 + \u00bb04 PCOMDI + .11 \u2022 . PCORAL PCOVVI PCOSCS PCOJOR PCOBSP PCOVUL PCOGSH PCOGSC PCOCAS GDOLBO + .03 -.26 -.76 .527 GDOMDI + .05 .492 GDORAL -.22 -.08 .496 GDOVVI \u2022 538 GDOSCS .509 GDOJOR - -.42 \u2022 534 GDOBSP GDOVUL \u2022 443 GDOGSH \u2022 479 GDOGSC \u2022 399 GDOCAS .517 LBOMDI +++ + .44 \u2022 486 LBORAL \u2022 574 LBOVVI \u2022 353 LBOSCS \u2022 593 LBOJOR \u2022 575 LBOBSP LBOVUL \u2022 408 LBQGSH \u2022 445 LBOGSC \u2022 478 LBOCAS \u2022 493 MDIRAL \u2022 529 MDIVVI \u2022 533 MDISCS \u2022 508 MDIJOR \u2022 511 MDIBSP MDIVUL \u2022 485 MDIGSH \u2022 448 MDIGSC \u2022 484 MDI GAS \u2022 487 RALVVI .573 RALSCS \u2022 149 RALJQR +++ +++ +.36 +.15 \u2022 336 RALBSP RALVUL \u2022 540 RALGSH .553 RALGSC .517 RALGAS .537 VVISCS \u2022 588 VVIJOR \u2022 574 VVIBSP VVIVUL \u2022 403 VVIGSH .417 VVIGSC \u2022 513 VVICAS \u2022 539 SCSJOR \u2022 311 SCSBSP SCSVUL \u2022 555 SCSGSH \u2022 565 SCSGSC \u2022 539 SCSCAS \u2022 516 JORBSP JORVUL \u2022 578 JORGSH \u2022 591 JORGSC \u2022 551 JORCAS \u2022 521 BSPVUL BSPGSH BSPGSC BSPCAS VULGSH \u2022 371 VULGSC \u2022 347 VULCAS \u2022 493 GSHGSC \u2022 447 6SHCAS \u2022 518 GSCCAS \u2022 510 Ogg's Bog MGAABO + ++ + + , 07 + , 09 + MGATPL \u2014 .26 - i 08 MGACOB + + + + + , .12 + i 15 \u2014 MGACPL + + +++ + + + ++ + + , 28 + , 35 + MGAAAE + + + + 4 08 .10 ** MGASMI + + + + + , 32 + , 14 + MGALGR - + - - 4 05 + , 10 MGAVOX + + 1 .03 07 *\u2022 MGADRO + 4 .06 + , .07 MGAKPO *\u00bb4 .48 .40 -MGAENI *\u2022 \u2014 , .47 - , .41 \u2014 MGATAR + 4 .04 - 4 .08 + MGACCA + + + , 04 + 4 07 + MGATGL + , .02 + 4 08 -MGALBO \u2014\u2014 .00 \u2014 , 10 \u2014 MGARAL -| , 13 - 1 .26 -MGAPMC \u2014\u2014 + ( .03 .00 MGAGSH \u2014 1 .29 - .88 MGABSP - -< .42 - .47 MGAG5C - .16 -( .11 MGACNU -- \u2014 i .46 -( .31 MGAMDI .24 -< .14 MGASCS mm, mm mm .47 \u2014 .41 MGACAS + .01 .00 MGACTR + , .01 + .01 MGAJOR _ \u2014 > - .06 MGAPCO \u2014 .62 MGADCS - .03 - 01 MGALCL _ - \u2014 ,55 MGAGDO \u2014 .25 MGAVOV \u2014\u2014 + .01 MGAVVI -MGACUN - \u2014 \u2014 MGAVUL MGAJCO MGAFCG MGALIN MGAEPO MGACPA 01 + \u00ab 15 + . 08 + 4 199 + < i076 + .039 .554 22 13 -, .22 + 4 ,011 + < ,035 01 -| 15 + 4 .11 + 4 146 + ! ,224 -.237 .443 18 + \u00ab 36 + 4 .11 + * .200 + | ,003 -.361 .457 01 + 4 06 + 4 .10 + 4 188 -< ,175 + .474 .471 02 07 + 4 .07 + 4 403* + i ,100 -.282 .472 65 + , 07 - , .09 \u00bb 4 .208 *\u2022 , ,275* .509 07 + 4 03 + 4 .05 + \u00ab ,285 ~ l .073 + .039 \u2022 566 21 + , 02 -\u00ab .07 .262* -1 .229 + .004 .486 58 + 4 03 - , 14 .048 - I ,100 .483 84 -, 23 - 4 .44 + < ,102 + | .227 .402 11 + 4 03 - 4 ,03 - I .185 mm .134 + .001 .457 10 + , 02 - 4 .48 + 4 .354* + i .265 -.167 .474 22 + 4 02 - 4 ,04 - 4 ,167 + < .009 .494 48 12 -, ,16 - I ,155 - \u2022 090 \u2022 580 43 .17 ,31 - I ,067 + .140 .461 08 + ) .02 ,62 + ( .229 + . 188 .473 ,09 .45 + ( .349* - .209 .523 mm . , 18 - ,61 + .699* ,08 - .18 mm . .072 * .207 \u2022 506 ,00 - .23 + .121 \u2022 215 .471 .07 - .06 .141 + .078 .538 ,00 mm .45 -< .054 .473 .00 - .27 + .371 - .926* .530 ,07 - .50 + .185 - \u2022 260 .553 .25 - .63 .564 .03 - .63 \u2022 .00 - .27 + .427 \u2022 468 mm . .20 - .91 .15 - .55 .472 - .07 - .51 .470 \u2014 .30 \u2014 .57 .601 .30 .00 .00 -.79 .27 .51 \u2022 560 .467 545 486 495 MGALAM ABOTPL +++ + , .28 .07 ABOCOB -, .25 .16 ABOCPL ++ +++ + +++ + i .42 .37 ABOAAE +++ + ++ + +++ + , .36 .23 ABOSMI +++ +++ +++ .07 ABOLGR +++ +++ +++ +++ .50 .28 ABOVOX ++ +++ + +++ +++ + i .36 + < .56 ABODRO +++ + +++ +++ .50 .20 ABOENI +++ .06 - i 03 ABOKPO +++ +++ +++ .22 .09 ABOTAR +++ +++ +++ +++ + , .49 ,32 ABOCCA - + , .03 ,01 ABOTGL +++ +++ +++ +++ .27 ,36 ABOLBO + + + .18 ,08 ABORAL \u2014 . - \u2014 - - , .62 ,65 ABOPMC \u2014 - .81 ,37 ABOGSH .09 ,02 ABOBSP ++ .09 + , 01 ABOGSC - \u2014 .38 ,30 ABOCNU +++ .04 + , 03 ABOMDI + 1 06 -, ,11 ABOSCS .57 - I ,65 ABOCAS + i .03 + , 02 ABOGTR + mm mm mm + | .04 ,05 ABOJOR .79 ABODCS -< .82 77 ABOPCO + , .02 ABOLCL - .00 ABOGDO -1 .36 ABOVOV + ( .02 ABOVVI \u2014 ABOCUN ABOVUL ABOJCO ABOFCG ABOLIN ABOEPO ABOCPA 549 -< ,26 ,03 ,07 -\u2022396* -.292 \u2022 -, ,17 ,32 ,22 -\u2022170 -.052 - \u2022 477 .519 + 1 01 + , 16 .23 + .128 -.140 + .307 .530 + 1 ,12 + , 38 ,21 + .155 + \u2022098 - .159 .503 + \u00ab ,03 35 ,17 -.204 \u00ab.217 + .325 .502 \u2014 1 ,34 + , 24 ,13 -.189 -\u2022074 .525 + 1 , 18 26 .12 + .065 +\u2022323* + \u2022 013 .547 + 1 05 + , 15 ,18 -.055 + .237 - .230 .499 Mm { ,09 + i 08 mm , ,51 -.157 -\u2022062 + 473 .583 + , 04 + , 13 + i .06 + \u2022049 + .033 .517 ,03 ,14 ,13 -\u2022047 -.058 .000 .490 ,75 ,01 - , .45 \u2022 000 -.238 .511 ,03 ,15 + | ,09 -\u2022013 + .102 .508 .26 + , 04 + ( .03 -\u2022378* -.344 \u2022 528 82 27 ,26 -\u2022684 .510 ,78 15 - , 53 -.237 -.377 \u2022 436 + , 02 - , 42 + \u2022051 -.293 \u2022 488 +, ,01 ,43 + .258 + \u2022148 \u2022 02 ,17 + \u2022378 -.128 .439 + , .02 ,02 + .084 -\u2022065 .530 01 - , 30 -\u2022480* -\u2022313 .484 + \u00ab .02 ,39 .524 + t .03 ,37 -.221 -\u2022202 .547 + i .03 .40 + \u2022051 + \u2022257 \u2022 553 mm* M .52 .70 .529 + , .01 .00 ,00 .00 mm , .11 .63 .35 .66 -.355 .523 .478 ,01 .00 .00 - i .63 .73 .52 \u2022 510 .538 .469 .07 .00 mm .00 .38 .482 475 471 545 ABOLAM ,432 TPLCOB \u2014 44 -.27 -.49 mm - 34 - 4 ,35 - 4 231 -.338 TPLCPL 09 + .01 -.67 00 - 4 ,09 mm j ,109 -.338 TPLAAE + + , 11 - .12 -.49 + 4 09 - 4 ,11 ,199 -.265 TPLSMI +++ ++ + + .02 -. 56 + \u00ab 10 + 4 ,02 mm | ,108 -.155 TPLL6R +++ + +++ + + + + , 68 + .39 + .26 + 4 19 + 4 ,08 - i ,105 -.323 TPLVOX + i 29 + .42 -.34 + 4 05 - < .14 \u2014 4 ,208 -.063 TPLDRO + + + + + , 29 + .29 -.08 + 4 15 ,06 - 4 ,037 -.246 TPLKPO ++ + ++ + + 4 22 + .28 + , 09 ,09 \u2014 4 ,101 -.178 TPLENI + 4 11 .13 + .25 - 4 05 mm , ,09 - 4 ,115 -.137 TPLTAR + + + + +++ + 4 29 + .06 + 4 18 + 4 ,12 - 4 ,044 -.155 TPLCCA \u2014 \u2014 11 -.30 -, 30 - 4 ,55 - 4 182 + .070 TPLTGL + + + + 1 19 - .04 + 4 18 - 4 ,13 - ( , 199 -.209 TPLLBO + + ++ ++ + + + ++ + + 4 15 + .28 + .30 + , 32 + 4 ,21 + \u00ab ,329* + .126 TPLRAL - \u2014 - | .29 .64 - 4 39 - 1 ,78 mm . ,446 TPLPMC - \" I \u2022 23 -.43 mm 65 - ( ,94 -\u00ab ,067 TPLGSH ++ + + ++ + + 4 ,17 + .01 + 4 17 + ( ,10 + , .310 + .078 TPLBSP + ++ +++ + | ,11 + .08 + 4 15 + ( .11 + , 351 + .422 TPLGSC mmtmU \u2014 - , 34 -.68 + | ,06 -< .42 - i .184 TPLCNU + ++ + 1 ,05 .04 + \u00ab ,09 + ,06 -< .167 -.513 TPLMDI +++ ++ + + i ,08 + .15 + 4 ,09 + .11 + ( .541* + .091 TPLSCS - -4 ,16 -\u2022 52 + 4 ,08 ,54 \u2014 i .497 TPLCAS + + + ++ + - 4 ,09 -.18 + 4 ,20 + 1 .14 *\u2022> .126 TPLCTR + + + + + + ,01 .36 + < ,16 + \u00ab ,07 - i .563* TPLJOR - \u2014\u2014 ,53 - 4 ,79 \u2014 ( .93 TPLDCS -t , 19 .36 -( ,05 TPLPCO + , 02 + ( .04 a w ,16 TPLLCL + + + + ( .03 + < ,02 + .03 TPLGDO - + , .02 .58 -.99 TPLVOV + ++ .01 + ,06 + .03 TPLVVI +++ + .06 + .09 TPLCUN .06 + .01 TPLVUL + ( .02 .00 TPLJGO + , .04 - .16 TPLFCG TPLLIN TPLEP0 TPLCPA TPLLAM COBCPL -.22 + i .19 + .07 - \u00ab .32 00 + .133 + .103 -.420 .283 GOBAAE - -.73 .26 -.57 + 4 ,03 - 4 ,10 -.032 -\u2022137 .391 COBSMI . . . + < .10 + .12 - 4 ,59 mm . 12 +.284* + \u2022229 + .341 .506 GOBLGR -.53 - ( .15 -.66 - I ,24 - 4 ,07 +.339* +.547* \u2022 587 COBVOX ++ -.39 mm .19 + .24 ,00 + \u00bb ,08 + .202 + .098 + .250 \u2022 562 COBDRO . . . . . . . . - .34 -.04 - , 70 - . ,48 -.163 -\u2022233 \u2022 536 COBKPO . . . . . . \u2014 . . . -.87 - .57 -.19 ,38 - , ,24 -.194 -.205 .510 GOBENI - \u2014 -.29 - .29 -.38 - , . 12 - i .39 -.369 -\u2022438 .507 COBTAR \u2014 . -. 14 - .03 + .07 \u2022 62 -( .36 -.042 -.109 .531 COBCCA + i \u2014 - + . 13 + .06 -.49 + ,04 4M> , .31 -.019 -.136 .291 GOBTGL . . . -.43 - .25 -.49 mm . .85 - .40 -.006 + \u2022035 .509 COBLBO - + .04 - .08 .00 - .08 - .18 -.058 + \u2022169 \u2022 586 COBRAL + .01 + .01 + .14 - .66 - .33 -.216 -.455 .355 COBPMC +++ + .08 + .04 -.55 + .11 - .92 + .017 + .143 .439 GOBGSH -.11 - .09 - .16 mm .64 -.213 -.023 .533 GOBGSC - -.08 + .07 - . 19 - \u2022 12 -.111 - 284 .513 COBBSP . . . - -.34 -.51 mm .33 4Kt .89 -\u2022427 .297 COBCNU \u2014 -. 16 - .38 - .38 - .54 -.460 COBMDI -.16 - .19 - .19 + .02 + \u2022044 -.090 \u2022 482 COBSCS -.02 -.08 mm .34 - .70 -.459 .325 COBCAS - -.29 -.37 - .43 - .23 -.196 .508 COBCTR \u2014 . -.30 - .51 mm .37 - .72 + .237 .519 COBJOR + + .03 + \u2022 06 - \u2022 48 -.397 .429 COBDCS -.25 -\u2022 26 - .15 - .39 \u2022 410 GOBPCO -. 13 \u2014 .19 - .75 \u2022 COBLCL \u2014 - \u2022 00 .84 - .99 \u2022 506 GOBGDO \u2014 - -.35 - \u2022 42 - .27 COBVOV + .02 - .05 - .32 .526 COBVVI -\u2022 11 - .49 .625 COBCUN . . . .56 \u2014 .78 . 501 COBVUL - \u2022 15 - .39 .540 COBJCO + \u2022 01 \u2014 .32 .489 COBFCG COBLIN .329 GOBEPO COBCPA \u2022 372 GOBLAM -\u2022024 -.632 .555 CPLAAE +++ ++ + \u202227 + .21 -. 16 + .05 - \u2022 01 + .199 \u2022 371 CPLSMI + \u202207 + \u2022 08 -.03 -\u2022 24 + .03 + .122 + .186 + .404 .478 GPLLGR -\u202222 + .07 -.34 + .08 + .02 -.305 -.252 .583 GPLVOX + + \u2022 00 + .15 .00 + .07 + .06 -\u2022130 -\u2022064 + .139 .531 CPLDRO + + . 16 + .08 + .05 + .06 + .04 + .044 -.054 -.368 .508 GPLKPO -.36 -.19 -.47 -.05 -.03 -.159 + .095 \u2022 506 CPLENI -.27 -.36 -.39 + .02 -.56 + \u2022057 + .092 .527 CPLTAR + + + + \u202213 + .04 + .04 + .03 + .07 -.028 -\u2022038 -.070 .502 CPLCCA + + .02 + .07 -.67 + .04 -.49 -.117 -.166 .328 GPLTGL - + \u202203 + .01 + .02 .00 -.018 + .077 \u2022 462 CPLUBO .00 -.08 - 34 -.18 + .01 -.153 + .376 .544 CPLRAL -.29 -.41 - 28 -.03 -.39 -.089 -\u2022239 \u2022 362 CPLPMC -.27 -.12 - 41 -\u202221 -.61 -.020 -.529 \u2022 465 GPLGSH \u2014 -.42 -.17 -.22 -.41 + .008 -\u2022029 \u2022 562 CPLBSP _ -.17 -.10 -.04 -.56 -.157 GPLGSC -.18 -.21 -.31 -.34 -.259 -\u2022625 .510 CPLCNU - -.11 -.10 + .01 -.18 -.229 + \u2022046 \u2022 402 GPLMDI - -.13 -.02 -.25 -.20 -.196 +\u2022900* \u2022 469 CPLSCS -.57 -.83 -.27 -.36 .330 CPLCAS \u2014 + \u202203 + .04 + \u202203 -.36 -.404 + .467 \u2022 440 CPLCTR -.02 + .05 -.09 -.43 + .472 -\u2022073 .470 CPLJOR -.03 \u2022 00 -.62 \u2022 400 CPLDCS -.50 -.78 -.11 \u2022 00 .420 CPLPCO - -\u202222 -.20 -.56 \u2022 GPLLCL -.30 -\u202201 -\u202243 CPLGDO -.61 -.47 -.54 \u2022 497 CPLVOV - + .01 -\u202225 -.62 .519 CPLVVI + .01 -.59 \u2022 600 CPLCUN \u2014 + \u202201 -.62 \u2022 489 CPLVUL - -.33 \u2022 00 .528 CPLJCO + \u202201 -.37 CPLFCG .515 C P L L I N CPLEPO \u2022 311 GPLCPA .326 CPLLAM .560 AAESMI +++ + + + + + + + + .75 + .38 -.45 + .29 + .17 + .127 +\u2022319* \u2022 484 AAELGR + + .09 + .06 + .13 + .18 -.04 -.116 + .122 .589 AAEVOX + + + + + + + .28 + .33 + .07 + \u00ab 16 + .12 +.629* + \u2022152 -.324 .547 AAEDRO +++ + + + + + + + .30 + .08 -.23 + .20 + .19 + .090 -\u2022049 .513 AAEKPO + .01 -.14 -.19 + .06 \u2022 00 + .198 + .117 .516 AAEENI - -< l l O -.16 AAETAR + + i .24 + .04 AAECCA + , 02 + .10 AAETGL + + +++ + 4 ,16 + .04 AAELBO \u2014 -4 ,08 -.11 AAERAL -1 ,18 -.40 AAEPMC \u2014\u2014 -4 ,10 + .02 AAEGSH - -4 ,10 -.01 AAEBSP \u2014 - - , 10 -.29 AAEGSC \u2014 -t , 15 -.06 AAECNU + 4 ,02 -.20 AAEMDI -4 .14 -.42 AAESCS - -4 ,47 -.40 AAECAS - + 4 ,03 -.08 AAECTR - + , 03 + .01 AAEJOR -1 .37 AAEDCS - -1 ,10 -.47 AAEPCO - ,00 AAELCL .00 AAEGDO -1 , 16 AAEVOV + 4 ,02 AAEVVI \u2014 AAECUN AAEVUL + AAEJCO AAEFCG AAELIN AAEEPO AAECPA AAELAM SMILGR +++ + + \u2022 13 + .10 + SMIVOX + + +++ +++ + . 16 + .24 + SMIDRO +++ ++ +++ + i 13 + .02 SMIKPO ++ + .09 -.11 SMIENI - \u2014 + .04 + .06 + SMI TAR +++ +++ +++ + . 14 + .07 + SMIGCA + .05 + .05 + SMITGL +++ +++ +++ + .11 + .01 + SMILBO + + .06 -.01 -+ \u2022 04 -.50 -.207 -\u2022488 .506 .24 + \u2022 11 mm .11 + .057 \u2022 000 .417 \u2022 00 + \u2022 01 -.34 -.113 -\u2022238 .320 \u2022 00 + . 10 + .09 -.228 -.058 .473 \u2022 49 + .04 -.22 -.168 + .231 .570 \u2022 26 - .14 -.22 -\u2022281 + .323 .373 \u2022 11 .00 -.48 -.126 + .300 \u2022 458 -\u2022 01 -.26 -.266 -.405 .564 \u2022 00 -.39 -.356 + \u2022 04 -\u2022 22 -.212 + .031 \u2022 474 \u2022 00 + \u2022 01 + .079 + .086 \u2022 329 -\u2022 06 -.10 -.244 \u2022 464 + \u2022 04 -.33 + .084 \u2022 350 + .03 -\u2022 23 + .407 .512 + .03 -.31 + .031 + .121 .503 -.37 -.68 \u2022 341 \u2022 00 -.78 .319 -\u2022 15 mm .55 -.27 -.19 + \u2022 02 mm .89 .449 -.27 -\u2022 46 \u2022 543 -\u2022 10 -.75 .583 -.47 -\u2022 61 \u2022 484 \u2022 00 + \u2022 01 .537 + \u2022 02 -.46 .525 \u2022 336 .322 \u2022 512 .09 + .15 -\u2022 08 - . 2 5 6 * -.193 -.740* .557 \u2022 13 + \u2022 23 + .10 + .295 + .062 + .204 \u2022 561 \u2022 00 + \u2022 13 + \u2022 16 + .031 -\u2022176 + . 8 8 3 * \u2022 410 + \u2022 05 + .04 + .168 + .255 .520 .13 + \u2022 04 -\u2022 42 -.011 -.028 -\u2022 286 .584 \u2022 08 + \u2022 14 + .12 + .091 + \u2022044 + \u2022 316 .471 . 04 -.07 -.64 +.505* + .369 \u2022 524 \u2022 14 + \u2022 19 + \u2022 11 + .052 +.486* \u2022 430 \u2022 56 + .04 + .01 -.002 + .153 .567 SMIRAL . . . - - - . 2 7 - . 6 8 - . 2 6 - . 0 9 - . 3 8 6 - . 4 5 9 \u2022 504 SMI PMC - \u2014 - - \u2022 6 8 - . 0 2 + . 0 7 . 0 0 - . 5 3 + . 5 0 1 * + . 1 7 3 \u2022 493 SMIGSH +\u2022++ + \u202204 . 0 0 + . 0 9 - . 3 9 + \u2022239 + \u2022146 .551 SMIBSP + \u2014 . + \u202202 . 0 0 + . 0 4 - . 4 0 - . 1 1 2 - \u2022 0 0 9 \u2022 SMIGSC - + - \u2022 7 0 - . 0 2 + . 0 6 - . 0 4 - \u2022 1 4 2 - \u2022 5 5 8 .451 SMICNU mm mm + \u202202 - . 1 3 + \u202202 - . 3 5 + .146 - . 2 0 1 . 5 0 4 SMIMDI + \u2022 03 - . 0 5 + . 0 3 - . 6 3 + \u2022162 + \u2022868* \u2022 516 SMISGS + - - . 7 6 - . 1 8 + \u2022 0 4 - . 2 6 + \u2022627* \u2022 526 SMICAS + + . 0 1 + . 01 + . 0 5 - . 5 4 + . 8 5 1 * + . 7 3 9 * . 4 4 0 SMICTR + - + . 0 1 \u2022 00 + . 0 4 - . 3 3 + .385 + \u2022279 \u2022 466 SMIJOR \u2014 \u2014 - \u2014 - . 3 5 - . 5 3 - . 6 3 \u2022 558 SMIDCS - - . 3 4 + .01 + . 0 1 - . 2 2 . 5 0 0 SMIPCO - \u2014 + . 0 1 \u2022 00 - . 6 7 + . 2 2 7 \u2022 SMILCL - + . 0 1 + . 0 1 - . 5 1 \u2022 SMIGDO - . 1 3 - . 1 4 - . 7 0 . 4 8 2 SMIVOV - + . 0 1 + . 01 - . 6 7 . 5 0 9 SM-IVVI - \u2014 + . 0 2 - . 4 7 . 5 9 7 SMICUN - + . 0 2 - . 3 7 .501 SMIVUL + . 01 \u2022 00 . 4 9 8 SMIJCO \u2022 00 - . 5 1 SMIFCG . 4 9 0 SMI L IN SMIEPO \u2022 522 SMI CPA \u2022 526 SMI LAM \u2022 541 LGRVOX ++ +++ + +++ + . 3 5 + \u202255 + . 4 7 + . 2 3 + . 0 6 + .066 +\u2022081 - . 1 9 4 . 5 0 6 LGRDRO +++ +++ +++ +++ + . 4 2 + . 4 8 + . 0 8 + . 2 0 + .18 - . 1 8 5 + \u2022013 . 5 0 4 LGRKPO +++ + ++ +++ +++ + . 3 3 + . 2 5 + . 11 + . 19 + \u202215 - . 0 0 2 - , 0 3 7 \u2022 365 LGRENI +++ \u2014 + . 0 8 + . 02 + . 2 0 + \u202212 - . 3 5 + . 074 - \u2022 3 6 6 \u2022 511 LGRTAR ++ ++ +++ +++ + . 2 8 + . 2 7 - . 6 0 + . 2 6 + . 1 7 + . 103 + \u2022130 . 5 0 0 LGRCCA - . 0 5 - . 2 7 - . 2 0 - . 2 8 - . 4 0 + . 227 - \u2022 3 3 9 . 5 9 5 LGRTGL ++ + +++ ++ + . 3 0 + . 2 9 + . 1 6 + . 1 6 + . 0 4 + .105 - . 1 4 2 . 5 1 5 LGRLBO ++ + + +++ +++ + . 19 + . 1 7 + . 16 + . 21 + .18 + .156 - \u2022 0 8 7 \u2022 481 LGRRAL \u2014 - - \u2022 \u2014 \u2014 - . 4 2 - . 6 0 - . 2 4 - . 5 0 - . 5 9 5 \u2022 566 LGRPMC - - \u2014 - \u2022 6 4 - . 4 4 - . 2 9 - . 2 2 - . 7 4 + .240 + . 1 2 9 . 5 9 7 LGRGSH + + +++ + \u2022 13 - . 0 1 + . 12 - . 6 0 - . 3 6 2 * - . 1 7 0 \u2022 372 LGRBSP + + \u2014 . + \u202207 + . 0 6 + . 0 7 - . 5 7 - . 0 8 8 + . 5 0 5 \u2022 LGRGSC . . . . . . - . 5 0 - . 3 3 - . 0 8 - . 3 2 + .176 \u2022 466 LGRCNU + + . 0 5 + .01 + . 0 5 + .02 + . 354 + \u2022041 \u2022 589 LGRMDI + + + + + + \u2022 0 9 + . 1 0 + . 0 7 - . 2 2 + .070 + . 0 3 2 . 4 7 4 LGRSCS - \u2022 13 - \u2022 4 0 + . 02 - . 5 3 - . 3 4 1 .581 LGRCAS + M M i M - \u2022 0 5 + \u202202 + . 0 8 - . 5 2 - . 7 1 6 * . 5 5 3 LGRCTR mm mm mm + . 02 + \u2022 0 6 + . 0 4 - . 7 1 - . 2 1 1 + . 178 . 5 4 4 LGRJOR mm mm \u2014 \u2014 - \u2022 5 7 - . 6 4 - . 8 7 \u2022 582 LGRDCS - . 2 5 - . 2 0 - . 1 6 \u2022 00 \u2022 555 LGRPCO + \u2022 03 + .01 - . 4 0 + .678 \u2022 LGRLCL + \u202202 + .02 - . 1 1 \u2022 LGRGDO - . 1 3 - . 1 0 - . 5 7 . 4 9 3 LGRVOV + \u202202 + \u202201 - . 4 0 . 3 7 7 LGRVVI + \u202202 - . 6 2 . 4 9 4 LGRCUN + + . 0 4 - . 3 2 . 5 1 2 LGRVUL \u2022 00 \u2022 00 . 4 5 4 LGRJCO + . 02 - . 6 0 \u2022 LGRFCG . 5 4 9 LGRLIN \u2022 LGREPO \u2022 561 LGRCPA \u2022 598 LGRLAM \u2022 560 VOXDRO + + + + ++ + + + \u2022 2 5 + . 2 4 + . 1 5 + . 2 3 + . 0 8 + . 229 + . 3 3 5 * + . 6 0 7 * . 5 5 6 VOXKPO + + + + + . 1 5 + . 1 4 + . 0 4 + . 1 5 + . 0 6 - . 1 7 1 - . 0 8 4 . 4 5 5 VOXENI + . 0 6 + . 0 7 + . 0 5 - . 0 1 - \u2022 4 0 - . 0 2 4 + . 3 1 9 . 4 3 8 VOXTAR ++ + + ++ + + + . 2 7 + . 2 4 + . 1 3 + . 16 + . 0 7 - . 0 4 9 + .054 - . 5 0 0 . 5 6 8 VOXCCA + . 0 2 + . 0 1 - . 2 5 + .01 - . 5 2 - . 0 5 7 + .204 . 5 8 0 VOXTGL + + + + ++ + . 14 + . 0 9 + . 0 5 + . 1 8 + . 0 7 + . 3 3 9 * + .600* . 5 3 3 VOXLBO + + + + . 0 7 + . 1 2 - . 6 3 + . 11 + .01 - . 0 5 7 + . 0 4 7 .391 VOXRAL ...... - \u2014 - . 6 3 - . 8 3 - . 2 7 - . 4 9 - . 2 8 9 . 5 3 6 VOXPMC - - . 3 1 - . 2 2 . 0 0 - . 3 0 - . 6 8 + .111 + . 0 1 7 \u2022 569 VOXGSH + + . 0 8 + . 0 3 + .01 - . 6 4 - . 1 4 7 - \u2022 0 1 2 \u2022 462 VOXBSP + . 0 3 + . 0 2 - . 0 3 - . 6 6 - . 2 2 8 - . 0 5 8 \u2022 VOXGSC - - . 1 4 - \u2022 3 4 + \u2022 03 + .02 - . 2 7 7 + . 5 0 0 \u2022 543 VOXCNU + + . 0 3 + . 0 6 + . 0 3 + .01 + .225 - . 2 4 6 \u2022 602 VOXMDI - + . 0 4 + \u2022 0 3 - . 2 5 - . 4 5 - . 2 0 5 + . 2 0 6 . 5 0 5 VOXSCS - . 0 8 - \u2022 3 4 + . 0 5 - . 4 7 + .578 . 5 5 0 VOXCAS \u2014 + . 0 4 + . 0 3 + . 0 5 - . 3 2 + .421 + . 5 9 0 . 4 9 8 VOXCTR - + . 0 3 \u2022 00 + . 0 4 - . 3 6 + . 7 7 2 * + .702* . 4 8 9 VOXJOR - . 8 0 - . 6 1 - . 6 7 . 5 3 3 VOXDCS mm + .01 - . 5 6 + . 02 - . 0 1 - . 3 7 0 . 5 8 5 VOXPCO + . 02 - . 2 5 - . 7 9 - . 1 0 2 \u2022 VOXLCL .\u2014 + . 02 17 -.85 . VOXGDO ... \u2022 00 \u2014 \u2022 21 -.49 .530 VOXVOV \u00ab... \u2014 \u2022 02 \u2014 . 67 -.59 .471 VOXVVI ... \u2014 a 25 -.55 .364 VOXCUN \u2014 . 16 -.38 .430 VOXVUL - 70 + .01 \u2022 457 VOXJCO - \u2014 . 36 -.79 VOXFCG \u2022 565 VOXLIN \u2022 VOXEPO \u2022 540 VOXCPA \u2022 531 VOXLAM .552 DROKPO + + + + + + + + + + 21 + .16 27 + \u2022 39 + \u202224 -.124 -.048 \u2022 533 DROENI + + . . . + 4 10 + .07 + \u2022 03 + , 12 -.50 -.196 -.109 .548 DROTAR + + + + + + + + + + . 28 + .19 + . 04 + . 31 + .25 + .057 +.353* .464 DROCCA + + -\u2014 + \u2022 07 + .14 \u2014 . 54 mm m 17 -.59 -.301 -.302 .534 DROTGL + + + + + + + + + + + + + \u2022 26 + .30 - \u2022 08 40 + .21 + .086 +.340* .443 DROLBO + + \u2022 04 + .03 + . 04 + \u00ab 12 + .03 + .258 -.330 .559 DRORAL -\u2014 - - , 35 -.31 \u2014 . 67 06 -.30 -.285 -.298 .514 DROPMC \u2014_ ... 55 -.45 -\u00ab 27 -.72 -.083 -.096 \u2022 486 DROGSH - i 02 -.03 + , 05 -.23 -.326 -.075 .559 DROBSP + t 03 + .03 + . 04 -.05 -.387 +.709* \u2022 DROGSC ... - + + ... mm | 50 -.48 + . 12 -.34 -.013 \u2022 516 DROCNU ++ ++ + \u00bb 03 + .06 + . 08 + .04 -.229 + .009 .531 DROMDI - \u00bb 01 -.14 + . 04 -.70 + .059 -.467 \u2022 510 DROSCS ... + + + 46 -.41 + . 10 -.26 \u2022 538 DROCAS + + + + \u2022 01 -.33 + . 14 -.57 -.523 .506 DROCTR + + + \u00ab 06 + .05 + i 07 -.58 -.051 \u2022 494 DROJOR \u2014. ... -, 78 -. 57 -.88 .536 DRODCS - i 25 \u2022 00 + \u2022 01 + .01 \u2022 510 DROPCO + i 02 + \u00ab 02 -.50 DROLCL 22 + . 02 -.06 \u2022 DROGDO 01 + , 05 -.92 .497 DROVOV 22 mm _, 08 -.50 \u2022 538 DROVVI -, 17 -.73 .605 DROCUN + + t 06 -.36 .494 DROVUL + , 01 + .02 .541 DROJCO -\u00ab 06 .00 DROFCG .534 DROLIN DROEPO DROGPA DROLAM KPOENI + \u2022 + + \u202216 + .20 KPOTAR + +++ +++ + .21 + .29 mm K.POCCA \u2022 t o n -.07 -.28 + KPOTGL +++ + +++ +++ + \u202259 + .25 + KPOLBO + ++ + +++ + .08 + .23 + K.PORAL \u2014- - - -.51 -.58 KPOPMC \u2014 -.40 -.66 KP06SH + ++ \u2014 + . 16 + .15 KPOBSP + +++ + . 14 + .13 KPOGSC -.06 -.43 KPOCNU +++ +++ + + . 14 +.16 KPOMDI + - + .09 + .07 KPOSCS ++ + - + .03 + .05 K.POCAS ++ + .05 + .06 KPOCTR + + - + .04 + .07 KPOJOR - \u2014 \u2014- -.71 KPODCS -.25 -.25 KPOPCO + .04 K.POL.CL + .03 KPOGDO - + .04 KPOVOV -.65 KPOVVI -KPOCUN KPOVUL KPOJCO K.POFCG K.POLIN KPOEPO K.POCPA KPOLAM ENITAR + .07 + .14 EN I CCA mm - \u2014 \u2022 00 -.54 ENITGL + . 12 + .03 -ENILBO + + - + .31 + .33 + ENIRAL -.41 -.58 532 519 517 + .06 -.53 -.148 + .015 \u2022 502 .09 + .27 + .23 + .184 +.593* \u2022 510 \u2022 11 -.09 - .62 -.005 -.569 .524 \u2022 11 + .34 + .15 -.065 + \u2022383 .489 .11 + .23 + .15 + .011 -\u2022373 \u2022 469 -.26 -.49 \u2022 544 -.19 -.67 -.043 \u2022 575 + .09 -\u2022 68 +.406* + .235 \u2022 408 + .07 -\u2022 41 -.233 -\u2022091 \u2022 + .06 an .05 -.127 .421 + .07 + \u2022 02 +\u2022598* +.920* .523 + .07 -.39 + .398 +\u2022736* .516 + .12 -\u2022 41 + .123 .525 + .11 -\u2022 60 + .335 .507 + .10 MM \u2022 50 -.532 -\u2022107 .499 -\u202270 -\u2022 81 \u2022 525 -.31 + \u2022 01 \u2022 518 + .03 \u2022 00 \u2022 + .04 \u2022 00 \u00ab + .04 -\u2022 63 \u2022 448 + .01 + \u2022 01 .344 + .01 -\u2022 51 \u2022 481 + .02 + \u2022 01 .482 \u2022 00 -\u2022 28 \u2022 383 -.01 -.70 \u2022 \u2022 524 \u2022 481 \u2022 541 .567 \u2022 26 + .13 -.59 -.171 -\u2022122 .523 -.35 -\u2022 52 -.350 +\u2022796* .524 .26 + .05 - .64 + .099 +\u2022538* .573 .06 + ,09 mm \u2022 38 + .009 + \u2022450 .510 -.67 -.37 .474 EN I PMC - -\u202245 + .12 + .05 -.78 +.833* \u2022 487 ENIGSH +++ + + \u202245 + .16 + .03 -.19 -.124 + .145 \u2022 487 ENIBSP + + +++ + .15 + .16 + .13 + .19 -.177 -.081 \u2022 ENIGSC ... -.69 -.72 + .05 -.12 +.798* \u2022 568 ENICNU + .12 + .12 + .03 + .03 + .286 \u2022 534 ENIMDI + +++ + . 19 + .04 -.25 + .09 -.308 -.226 \u2022 551 ENISCS + .02 + .03 -.23 + .05 \u2022 487 EN I CAS + .08 + .04 -.11 \u2022 00 -.479 \u2022 542 ENICTR + .05 -.63 -. 16 + .03 +.844* .531 ENIJOR - \u2014. -.51 -.99 .496 ENIDCS -.24 + .01 \u2022 00 .510 ENIPCO +++ + .03 + .13 + .02 \u2022 ENILCL + ++ + .08 + .08 + .01 .521 ENIGDO + + + .07 + .09 + .05 ENIVOV + + .04 + .06 + .02 .454 ENIVVI ++ + .05 + .06 .445 ENICUN + + .10 + .02 .522 ENIVUL + + .05 \u2022 00 .479 ENIJCO + .03 \u2022 ENIFCG .573 ENILIN \u2022 500 ENIEPO ENICPA .525 EN I LAM \u2022 550 TARCCA ... -.14 -.17 40 -.11 -.74 + .038 -.153 .480 TARTGL +++ +++ + .05 + .14 + . 02 + .33 + .13 + .075 +.638* \u2022 441 TARLBO +++ +++ + .04 + .08 + .19 + .15 + .084 -.055 \u2022 563 TARRAL *\u2022>*\u00bb\u00ab\u00bb ... \u2014 -.40 -.66 -\u2022 56 -.34 -.09 -.623 \u2022 479 TARPMC ... ... \u2014 -.60 -.73 + . 01 -.53 -.57 -.192 \u2022 496 TARGSH + + +++ + . 13 + .11 + .18 -.74 + .143 -.074 .527 TARBSP + +++ + .06 + .08 + .15 -.42 -.138 -.333 \u2022 TARGSC + \u2014. -.09 -.24 + .11 -.38 + .266 .467 TARCNU + + +++ +++ + .08 + .10 + .17 + .10 -.233 -.155 \u2022 487 TARMDI + ... + .04 + .08 + .08 -.61 + .157 + .034 .453 TARSCS \u2014 - -.44 -.77 + .07 -.33 .496 TARCAS +++ \u2014 + .04 + .03 + .20 -.77 + .166 -\u2022708* \u2022 483 TARCTR +++ +++ + .03 + .11 + . 14 -.24 -.244 .471 TARJOR ... mm mm* mm ... -.78 -.85 -.77 \u2022 484 TARDCS - -.42 -.39 + .02 + .01 .440 TARPCO +++ + \u202203 + .07 -.39 -.493 a TARLCL + + .02 + .04 -.27 . TARGDO -\u2022 10 + .01 -.70 .412 TARVOV + + \u202202 + .04 -.27 .500 TARVVI +++ + .08 -.67 \u2022 588 TARCUN + + .06 -.27 .475 TARVUL + .02 -.13 \u2022 504 TARJCO -.24 -.27 TARFCG \u2022 480 TARlilN \u2022 TAREPO \u2022 494 TARGPA .499 TARLAM \u2022 457 CCATGL. + .20 -.13 +.02 -.07 -.62 -.129 +.368 \u2022 514 CCALBO - \u2014-. + \u202203 -.48 -\u202258 -.75 -.117 \u2022 589 CCARAL \u2014 * -.47 -.34 + .06 -.93 \u2022 328 CCAPMC + -T+ + +++ + \u202210 + .08 +\u202243 +.16 -.99 +.119 -.169 .442 CCAGSH -\u2022 17 -\u202246 -.64 -.11 +.632* .559 CCABSP mm - -\u202220 + .02 -.53 -.48 -.685 GCAGSC -*26 -.13 + .12 -.49 + .171 .504 CCAMDI -.08 -.68 -.31 + .12 + .336 .505 CCASCS -.40 -.56 -.50 + .01 .299 CCACAS + \u202202 + .05 .00 -.24 \u2022 548 CCACTR - mm mm -.26 -.61 -.52 -.77 .522 CCAJOR - + .08 + .03 .378 CCADCS -.66 -\u202242 -.35 .321 CCAPCO -\u202220 -.08 -.52 CCALCL - -.52 CCAGDO - -\u202260 -.81 -.41 .494 CCAVOV - + \u202203 .549 CCAVVI - -.74 .629 CCACUN - -.80 -.59 .524 CCAVUL -.35 .555 CCAJCO + .02 CCAFCG .513 CCALIN CCAEPO \u2022 314 CCACPA .297 GCALAM .559 TGLLBO + \u202206 + .04 + .08 + .02 -.175 -.390 .492 TGLRAL \u2014 -\u202236 -.40 -.13 \u2022 00 -.50 -.056 .513 TGUPMC - -.27 -.64 + .01 -.07 -.53 -.237 \u2022 499 TGLGSH -\u2022OA -.26 + .04 -.53 + .152 .526 TGLBSP + .06 + .01 + .07 -.65 -.440 \u2022 TGLGSC + + -.18 -.20 + . 14 -.25 -.010 .482 TGLCNU + + .04 .00 + .09 + .04 + .136 .530 TGLMDI + .06 + .02 + .03 -.71 -.345 \u2022 364 TGLSCS + + + -. 18 -\u202210 + .15 -.58 + .492 .497 TGLCAS + + + ' + .03 + .04 + . 16 -.53 + .443 .444 TGLCTR + + + - + .08 + .08 + .11 -.57 + .264 \u2022 452 TGkJOR \u2014- -.59 -.60 .497 TGLDCS -\u202230 -.20 \u2022 00 + .01 \u2022 542 TGLPCO + .01 + .02 -.65 \u2022 TGLLCL -.02 + .02 -.20 \u2022 TGLGDO + + + - + .03 + .09 -.67 .472 TGLVOV -\u202235 + .02 -.65 \u2022 516 T G L W I - + .02 -.61 \u2022 586 TGLCUN + + .07 -.54 .389 TGLVUL + .02 + .01 \u2022 523 TGLJCO -.20 -.65 \u2022 TGLFCG \u2022 440 T GL L I N \u2022 TGLEPO .491 TGLCPA \u2022 441 TGLLAM \u2022 495 LBORAL -.78 -.99 -.67 -.37 -.465 .575 LBOPMC -\u202259 -.57 -.59 -.67 \u2022 535 LBOGSH + + + + + + + + + + \u202237 + .24 + .40 -.80 +.052 +.277 .432 LBOBSP + + + + + + \u202219 + .08 + .25 -.56 -.107 \u2022 LBOGSC -\u202262 + .01 -.19 .478 LBOCNU + + + + + + + \u202204 + .12 + . 16 + .07 +.573 +.112 \u2022 589 LBOMDI +++ +++ + + + + .27 + .22 + . 18 -.70 +.563* +.376 \u2022 481 LBOSCS - -\u202267 -.29 -. 10 -.42 .594 LBOCAS + + + + + .05 + .10 + .20 -.66 + .098 .493 LBOGTR -\u202223 -.68 + .05 -.21 .484 LBOJOR - -.70 -.99 .573 LBODCS \u2014 -.07 + .02 \u2022 573 LBOPCO + .03 + .04 + .02 \u2022 LBOLCL + +++ + \u202208 + .09 + .01 LBOGDO - -.67 -.04 -.66 .526 LBOVOV + \u202205 + .04 + .01 .446 LBOVVI +++ + .13 + .01 .354 LBOCUN +++ + .13 + .02 .440 LBOVUL. + .01 -.12 .409 LBOJCO -.33 \u2022 LBOFCG .509 LBOL.IN LBOEPO .567 LBOCPA .557 LBOLAM \u2022 531 RALPMC +++ + +++ + .64 + .33 +\u202218 +.17 + .20 -.239 +.533 \u2022 424 RALGSH - - - MP -\u202256 -.55 -.59 .543 RALBSP mm mm - -\u202264 + .22 \u2022 RALGSC + + + .33 + .19 + \u202205 + .06 -.450 .516 RALCNU - -.69 -.51 -.84 .379 RALMDI - \u2022 -.59 -.71 -.99 .524 RALSCS +++ +++ + .28 + .23 + .02 + .06 -.618 \u2022 145 RALGAS -\u202261 -.42 -.90 .537 RALCTR ++ + -.34 -.47 -. 10 + .25 .542 RALJOR +++ ++ + + .28 + .32 -.14 -.031 .332 RALDCS +++ +++ + .33 + \u202232 -.217 -.039 \u2022 348 RALPCO -.42 + .02 RALLCL -.55 + .02 \u2022 RALGDO ++ + * 16 + .09 -.05 .490 RALVOV -.55 -.23 \u2022 534 RALVVI - -.58 .573 RALCUN + .10 + .01 \u2022 542 RALVUL -.17 .539 RALJCO + .04 -.23 RALFCG \u2022 516 RALLIN RALEPO .374 RALCPA .355 RALLAM .532 PMCGSH - -.45 -.70 -.72 -.85 -.421 .528 PMCBSP - -.85 -\u202288 + .09 PMC6SC ++ + + .27 + .17 + .03 + .01 +.473* -.397 .508 PMCCNU \u2014 -.99 -.83 \u2022 450 PMCMDI -.37 -.65 -.55 -.99 .491 PMCSCS ++ + +++ + . 18 + .22 + .03 + .05 -.271 -.811* \u2022 440 PMCCAS - - \u2014 -.76 -.99 -.99 \u2022 517 PMCCTR mm -.59 -.07 + .30 \u2022 540 PMCJOR + + + + +++ + .10 + .23 -.12 .503 PMCDCS +++ + + .17 + .13 + .02 -.149 .479 PMCPCO - -.70 + .06 \u2022 PMCkCL + .01 \u2022 PMCGDO + .03 + \u2022 10 + .04 .494 PMCVOV -.54 + .06 \u2022 533 PMCVVI - - + .05 .543 PMCCUN - + .04 \u2022 513 PMCVUL + .02 .522 PMCJCO + .01 .00 PMCFCG \u2022 443 PMCLIN PMCF.PO \u2022 474 PMGCPA \u2022 443 .470 PMCLAM GSHBSP +++ + + + + + .28 + .13 + .39 + .09 + .266 \u2022 GSHGSC -.39 -. 18 -.19 -.425 .435 GSHCNU + + + + + .12 + .27 + .22 -.604 .547 GSHMDI +++ + + + + .34 + .19 + .31 -.11 -.498* .445 GSHSCS + .02 + .06 -.39 -.35 \u2022 561 GSHCAS + + + + . 12 + .07 + .26 + .07 +.971* .514 GSHGTR + \u2022 00 -.34 + .12 -.68 .503 GSHJOR -.86 + .01 .587 GSHDCS -.24 + .03 + .01 .559 GSHPCO + .03 + .04 -.67 GSHLCL + + + + + + + .12 + .15 -.51 GSHGDO + .03 + .02 .00 .469 GSHVOV + + + + + .08 + .12 .219 GSHYVI + + + + .25 -.28 .407 GSHCUN + + + + .17 + .01 .450 GSHVUL -.37 + .01 .358 GSHJCO + .01 GSHFCG GSHLIN GSHEPO 6SHCPA GSHLAM BSPGSC BSPCNU ++ + BSPMDI +++ ++ BSPSCS BSPCAS +++ BSPCTR BSPJOR BSPDCS BSPPCO +++ BSPLCL + BSPGDO BSPVOV B S P W I BSPCUN BSPVUL BSPJCO BSPFCG B S P L I N BSPEPO BSPCPA BSPLAM GSCCNU GSCMDI 6SCSCS GSCCAS GSCCTR GSCJOR GSCDCS + + GSCPCO GSCLCL GSCGDO GSCVOV GSCVVI GSCCUN GSCVUL GSCJCO -.51 -.05 .00 -.29 \u2022 ++ + .25 + .34 + .18 + .02 -.259 +++ +++ + .39 + .35 + .34 -.93 + .370 -.29 -.04 + .07 * + + .29 + .14 + .17 -.14 -.133 \u2022 ++ + -.01 + .12 + .20 + .06 \u2022 \u2014 \u2014 -.99 -.99 -.35 \u2022 + .22 + .09 + .02 +++ +++ + .11 + .15 + .04 + + .04 -.02 + .05 \u2022 + + .05 + .09 + .01 \u2022 +++ + .27 + .03 +++ +++ + .27 + .06 +++ + .02 \u2022 00 -,40 -.46 + .01 + .01 -.01 -.54 .510 -.30 -,42 -.23 -.30 .477 +++ + \u2022 13 + .16 + .18 + .02 -.328 .540 + .01 + . 11 -.21 .511 ++ + .03 -.29 + .11 + .06 .499 + .08 + .01 -.30 .552 + + .11 + .19 + .05 .00 \u2022 468 + -.01 + .08 -.78 \u2022 \u2014 -.35 \u2022 + .04 + .06 - . 46 .395 + .03 -.13 \u2022 386 \u2014 -.81 + .02 .514 -.04 -.07 .495 +++ + .08 .348 -.12 -.13 \u2022 GSCFCG .447 GSCLIN GSCEPO .503 GSCCPA .506 GSCLAM .482 GNUMDI +++ + \u202221 +.32 + . 13 -.16 -.705 .506 CNUSCS +.07 -.20 + .07 + .04 \u2022 363 CNUCAS +++ +++ +.17 +.08 + .41 + .23 .555 GNUCTR +++ +.14 +.06 + .28 -.75 .562 CNUJOR \u2014 -.71 -.99 .413 CNUDCS + .05 .302 CNUPCO +++ +++ + .23 + .04 + .11 CNULCL +++ +++ +++ + .24 + .20 + .12 GNUGDO + +++ + .15 + .11 + .09 .531 CNUVOV +++ + .07 + .05 .541 CNUVVI +++ +++ + .20 + .10 .633 CNUCUN +++ +++ + .20 + .09 .528 CNUVUL .561 CNUJCO -.05 \u2022 00 GNUFCG .515 CNULIN CNUEPO .371 CNUCPA .405 CNUkAM .549 MDISCS + .04 -.22 -.08 .507 MDICAS +++ .00 + .11 + .23 .489 MDICTR +++ + .08 + .03 -.84 .497 MDIJOR \u2014 - -.68 .505 MDIDCS -.48 + .01 .553 MDIPCO + + .07 + . 13 -.67 MDILCL +++ + .03 + .16 MDIGDO - + .07 + .07 .484 MDIVOV + .03 + .08 .492 MDIVVI +++ - + .24 -.82 .530 MDICUN +++ + .20 -.72 .326 MDIVUL + .04 .481 MDUCO -.28 -.33 MDIFCG .388 MDILIN \u2022 . -+ \u202209 - . 6 2 +++ - . 5 1 + \u202232 + .08 + . 1 9 - . 5 3 + .01 +.27 +.32 + . 0 4 + .02 + . 01 + . 02 - . 0 3 + . 0 2 +++ + . 2 8 + . 2 6 - . 3 9 - . 4 0 - . 0 3 - . 3 2 + . 0 4 - . 2 2 + .02 + . 0 3 + .04 + .02 MDIEPO MDICPA MDILAM S C S C A S SCSCTR SCSJOR + SCSDCS +++ +++ SCSPCO S C S L C L SCSGDO +++ SCSVOV SCSVVI SCSCUN SGSVUL S C S J C O SCSFCG S C S L I N SCSEPO SCSCPA SCSLAM CASCTR GASJOR CASDCS CASPCO +.11 - \u00ab 1 3 - . 3 4 GASLCL + + \u00ab 1 3 +.08 \u2022 CASGDO + \u00ab 0 3 CASVOV + + \u00ab 0 3 +.08 \u00ab ^ 8 6 CASVVI + - 1 3 - . 6 4 . 5 3 9 +.13 - . 4 4 . 3 9 0 . 4 9 2 . 4 2 0 .521 CASLAM CTRJOR CTRDCS + . 461 . 468 . 4 2 7 . 5 1 7 . 5 2 3 . 3 0 6 . 3 7 9 \u2022 508 . 5 4 9 . 5 8 8 . 5 2 7 . 5 5 5 !498 . 3 4 4 . 3 2 8 . 5 4 4 +.02 +.06 +.15 - . 8 4 .211 - . 8 2 - . 5 8 \"517 - . 2 1 . 581 CASCUN + CASVUL CASJCO CASFCG C A S L I N CASEPO CASCPA *ttZ \u00b0> . 5 2 7 Cn - . 0 1 - . 6 2 . 5 0 8 - \u2022 1 5 +.08 \u00ab 5 5 7 GTRPCO + + + + + + * 1 7 +.14 +.09 * CTRLCL + .01 + .05 \u2022 CTR6D0 + + .08 +.16 .504 GTRVOV + + + + .05 +.06 .475 CTRVVI +.04 +.04 .530 CTRCUN + +++ +.12 +.08 .401 CTRVUL .00 .498 GTRJCO -.25 \u2022 CTRFCG \u00bb 4 5 0 CTRLIN \u2022 CTREPO \u00ab 5 1 4 CTRCPA \u00bb 4 8 7 CTRLAM \u00ab 5 3 5 JORDCS +.08 .410 JORPCO JORkCL \u2022 JORGDO *\u2022 ^.27 .529 JORVOV \u00bb 5 8 2 JORVVI - , 5 7 2 JORCUN * 5 0 7 JORVUL * 5 7 5 JORJCO \u2022 JORFCG * 5 4 8 JORLIN * JOREPO \u00ab 3 4 3 JORCPA \u00bb 3 5 4 JORUAM * 5 4 2 DCSPCO +.08 \u2022 DCSLCL \u2022 DCSGDO +++ +.38 +.02 +.01 .507 DCSVOV +.0 8 \u00bb 5 2 2 DCSVVI \u00ab 6 1 6 DCSCUN +.03 .572 DCSVUL * 5 3 8 DCSJCO \u2022 DCSFCG \u00bb 5 7 4 DCSLIN \u2022 DCSEPO \u00bb 3 6 7 DCSGPA \u00bb400 DCSLAM \u2022 5 Z f 0 PCOkCL ++ + \u202229 + \u202205 PCOGDO + \u202208 + \u202202 PCOVOV +++ +++ + .24 + .15 PCOVVI + .03 -.33 PCOCUN +++ +++ + .24 + .11 PCOVUL +++ +++ + .19 + .07 PCOJCO + .07 PCOFCG PCOLIN PCOEPO PCOCPA PCOLAM LCLGDO +++ +++ +.13 + .18 + .08 LCLVOV + + .17 + .06 LCLVVI +++ +++ + .40 + .14 LCLGUN +++ +++ + .33 + .09 LCLVUL LCLJCO LCLFCG LCLLIN LCLEPO LCLGPA LCLLAM GDOVOV -.07 +.01 .437 GDOVVI +++ +.14 +.18 .538 GDOGUN + +.01 +.07 .468 GDOVUL + +.09 +.01 .444 GDOJCO \u2022 6D0FCG .494 GDOLIN \u2022 GDOEPO .491 GDOCPA .477 GDOLAM .527 VOVVVI +++ ++ +.20 +.05 .418 VOVCUN +++ + .27 .479 VOVVUL .288 VOVJCO VOVFCG VOVLIN VOVEPO VOVCPA VOVLAM VVICUN +++ +++ +.44 +.31 VVIVUL V V U C O VVIFCG V V I L I N VVIEPO VVICPA W I L A M CUNVUL + ++ +\u00ab09 +.06 CUNJCO +.01 CUNFCG CUNLIN CUNEPO CUNCPA CUNLAM VULJCO +\u00ab11 VULFCG VULLIN VULEPO VULGPA VULLAM JCOFCG JC O L I N JCOEPO JCOCPA JCOLAM FCGLIN FCGEPO FCGGPA FCGLAM LINEPO LINGPA LINLAM EPOCPA EPOLAM CPALAM Blackwall Meadow PDVSPR +++ +++ + .16 + .06 .465 PDVMGR .465 PDVSTR - .436 PDVVDE +++ + .06 + .07 .475 PDVVVI .475 EALRES +++ +++ + .48 + .17 .507 EALSPR . 364 EALMGR + .20 .507 EALSTR +++ +++ + .29 + .22 .529 EALVDE .482 EALVVI +++ +++ + .22 + .13 .537 RESSPR .479 RESMGR + .27 + .12 .439 RESSTR +++ +++ + .43 + .19 .483 RESVDE .451 RESVVI +++ +++ .548 SPRMGR .479 SPRSTR .521 SPRVDE + .08 + .03 .490 SPRVVI .564 MGRSTR + .25 + .10 .502 MGRVDE .410 MGRVVI + . 19 + .12 .595 STRVDE .436 STRVVI +++ +++ + .38 + .23 .522 VDEVVI .573 VSIFVI -.06 -.19 -.195 + .007 + .331 .559 VSILLA +++ +++ + .14 + .12 + .29 + .28 + .080 -.107 -.018 .503 VSIEPE + .06 - .05 + .07 -.05 + .01 -.232 -.226 + .062 .314 VSIAOC + + + + + + + + +++ + .34 + .36 + .51 + .22 + .132 + .049 .476 VSIPFL + +++ + .47 + .14 + .08 + .17 + .02 \"-\u2022213 -.236 -.334 .418 VSIEGR + .03 - .26 -.64 -.36 -.12 + .200 -.213 .544 VSIVSC + + + + .05 + .12 -.25 + .11 + .02 -.440* -.540* .542 VSICLA + . 12 + .14 + .12 + .02 -.03 -.071 -.243 -.066 .532 VSIACA - - .81 -.85 -.60 -.631* -.544* .527 VSIALA - -.64 _ .33 -.69 -.50 -.53 -.580* -.485* .435 VSIVCU \u2014 + .03 - .03 -.37 -.50 -.21 -.530* -.398 .468 VSIAAU +++ - + .01 - .18 + . 17 -.17 -. 157 +.652* .482 VSIPAL + -.07 + .05 .00 + .02 + .04 -.095 . -.163 .554 VSIALF + + + + + + - . 33 - . 46 . 00 + . 12 .04 - .407 + .064 .356 VSILHI + + ++ + + + ++ + + .22 + .18 + .19 + .34 .13 - .243 - .483* + * 108 .503 VSITOC + + + + + .05 + .03 + .11 + .04 .03 + .261 .523 VSIAMI - . 1 4 - . 46 - . 64 - . 61 .33 - . 6 4 1 * - .825* .354 VSITSP \u2014 - . 4 0 - . 19 - . 2 6 .41 - .262 + .265 .616 VSICRO ' - - . 4 0 - .48 - . 65 .71 - .387 - .530 .583 VSIEGL + + + + .03 + .02 + .03 .02 .594 VSISPA - . 0 7 - .21 + .01 .14 - .301 .537 VSIAMO - .28 - . 52 - . 25 .41 + .257 .400 VSIPEP - . 10 .00 + .01 +\u00ab .01 - .192 - .815* .599 VSISIN \u2014 - . 12 - . 29 - . 3 0 .49 + .163 .407 VSIHGR \u2014 + .01 + .01 + .02 ,34 - .193 + .181 .471 VSILSP - - . 6 0 - .71 - .48 ,61 .558 VSISLA - . 5 6 - . 59 .71 .509 VSICMI - - . 18 - . 39 - . 2 2 ,21 .525 VSIPPR - . 2 3 - . 29 - . 6 7 .74 - .878* .467 VSIPBR + .01 + .02 .00 .476 VSIPDV - .51 .73 .452 VSIEAL + + + + + .04 + \u2022 .02 .524 VSIRES + +++ + .03 .02 .481 VSISPR \u2014 - . 5 0 ,80 .547 VSIMGR + .02 .01 .550 VSISTR + + ++ + .05 + ,02 .356 VSIVDE .524 VSIVVI + ++ + .03 + .02 . 528 FVILLA + - .36 + ,02 + .236 + .180 .000 .509 FVIEPE ++ + - .12 + ,04 + .091 + .045 +.587* .551 FVIAOC - .16 + .01 - .026 - .059 .521 FVIPFL +++ - . 02 + .07 + .234 +.270* + .548 .463 FVIEGR ++ + + .04 + .02 +.337* + .014 + .243 .486 FVIVSC - . 12 + .01 - .354 - .173 + .476 .494 FVICLA - .08 - .04 + .007 - .072 + .330 .518 FVIAGA ++ + .02 + .02 - .181 - .012 + .138 .481 FVIALA ++ + + .05 + .01 - .453* - .350* + .226 .502 FVIVCU + .05 + .01 + .229 - .182 - .633 .454 FVIAAU .00 + .187 + .093 .490 FVIPAL + - .12 + .01 + .124 + .152 .252 FVIALF + .02 .00 + .059 + .096 . 544 FVILHI + .04 + .01 + .064 + .172 - .194 .417 FVITOC -.41 -.28 - . 7 0 1 * - . 8 8 0 * .426 FVIAMI + + .08 + .01 + .128 + .326 +.702* .559 FVITSP .00 + .020 + .425 .313 FVICRO .00 -.279 -.019 .275 FVIEGL -.25 .326 FVISPA .00 + .425 + .586 .488 FVIAMO .00 + .098 + .336 . 544 F V I P E P .00 + .122 + .397 .251 F V I S I N .00 193 -.340 .530 FVIHGR .00 -.456 -.435 .510 F V I L S P .00 -.251 .256 F V I S L A .00 + .340 .446 FVICM I + .00 .000 + .102 .461 FVIPPR + .01 + .014 + .356 .508 FVIPBR .00 + .269 .459 FVIPDV .00 . 500 FVIEAL -.04 .435 FVIRES .00 .464 FVISPR -.01 .484 FVIMGR .00 .483 FVISTR -.39 .492 FVIVDE .00 .455 F V I V V I -.06 .552 LLAEPE + .11 + .05 + .02 + .06 -.01 -.084 -.234 .530 LLAAOC ++ + ++ + + .06 + .07 + .05 + .19 + .08 -.076 + .058 .399 LLAPFL + + + ++ + + .17 + .00 + .01 + .35 + .17 -.207 + .185 .475 LLAEGR -\u202244 -.11 -.33 -.15 -.252 -.025 .546 LLAVSC ++ + ++ + + .03 + .07 + . 14 + .07 + .07 + .154 + .197 .522 LLACLA + .17 + .04 + .03 -.023 -.027 .000 .562 LLAACA + .03 + .01 -.08 -. 84 -.29 -.035 -.036 .590 LLAALA + .05 + .02 + .11 -.24 -.02 -.082 + .225 .437 LLAVCU - + .05 + .07 -.62 -.04 -.157 + .167 .524 LLAAAU - + .06 + .01 -.12 -.17 + .032 + .446 .500 LLAPAL + .02 + .01 -.37 -.15 .00 + .251 -.005 .502 LLAALF + + + .04 -.03 -.37 + .05 + .02 -.063 + .356 .540 LLALHI +++ + + + + .05 + .04 + . 13 + .16 + .08 + .031 -.020 .523 LLATOC - - -.32 -.37 . 00 -.20 .477 LLAAMI \u2014 -. 14 -.35 -.83 -.31 - . 4 2 8 * -.243 .505 LLATSP + .03 -.39 -. 19 -.14 + .252 + .250 .520 LLACRO + .03 -.37 -.48 -.60 - < .37 -.066 + .305 .550 LLAEGL - -.44 -.11 .00 .22 .496 LLASPA -.30 -.02 .00 .00 + .380 -.025 .501 LLAAMO - \u2014 -.36 -.51 -.38 39 + .381 + .184 .489 LLAPEP \u2014 + .01 + .03 -.10 43 + .204 + .445 .551 LLAS IN + .01 -.05 -.57 24 - . 5 8 1 * -.654 .484 LLAHGR \u2014 + .01 -.05 + .01 +. .01 + .204 + .179 .509 LLALSP - \u2014 + .01 + .01 -.32 43 + .008 .542 LLASLA - + .01 -.36 .26 -.493 .548 LLACMI \u2014 - + .01 -.53 -.10 -, 22 + .443 .453 LLAPPR -.34 -.25 -.72 .59 -.076 -*324 .505 LLAPBR + .01 + .01 +, .01 -.010 .454 LLAPDV -. 18 .54 .454 LLAEAL + .01 .01 .542 LLARES + .01 +, .01 .532 LLASPR -.03 - 1 .63 .516 LLAMGR .00 +, .01 .532 LLASTR + .01 .02 .504 LLAVDE -.37 .50 .504 LLAVVI + .01 + , .01 .522 EPEAOC -.29 -.2 1 -.05 + .00 .02 + .016 + .147 .520 EPEPFL + + .30 + .19 + .03 + .17 .07 +.424* + .085 .444 EPEEGR ++ +++ +++ + .22 + .29 -.05 + .08 . 10 -.157 -.167 .536 EPEVSC \u2014 \u2014 -.51 -.54 .00 -.08 .10 + .197 + .087 .495 EPECLA - -.21 -.29 + .02 .09 + .069 -.088 .502 EPEACA - .00 -.2 1 -.61 -.08 .02 + .063 + .070 .531 EPEALA -.38 -.43 -.58 -.13 -, .14 -. 105 -.210 .461 EPEVCU + + +++ + .09 + .13 + .05 + .06 .08 + .320 -.110 .471 EPEAAU + .07 + .06 + .02 + \u00ab .01 + .188 -.232 .412 EPEPAL + ++ + +++ + . 16 + .10 + .04 + . 13 + .04 -.031 +.533* .542 EPEALF + + + .07 + .06 -.33 + .05 + .02 + .264 + .177 .248 EPELHI -.35 -.23 -.05 -.09 + .02 -.164 -.367 .452 EPETOC .00 -.03 -.33 -.02 - .09 - . 9 0 5 * .535 EPEAMI + + +++ ++ + + .12 + .10 + .07 + .08 + .08 -.080 -.191 .338 EPETSP .00 -.13 + .01 .00 + .024 -.297 .605 EPECRO -.23 -.11 -.08 - .07 + .230 -.047 .553 EPEEGL - + .02 + .02 -.08 - . 18 .571 EPESPA + + .06 + .04 + .01 + .01 + .292 .526 EPEAMO + .05 + .04 + .02 + .01 + .178 -.174 .314 EPEPEP + .03 + .01 -.05 -.09 + .107 + .015 .589 EPESIN + .05 + .03 + .03 .00 -.250 -.651* .380 EPEHGR -.46 -.49 -.12 -.51 +.985* .433 EPELSP + .04 + .02 .00 .00 -.098 .545 EPESLA +++ + .04 + .01 + .01 + .158 .496 EPECMI ++ + i03 + .03 + .01 + .01 + .133 .000 .516 EPEPPR + + .05 + .04 + .02 + .02 + .291 + .345 .451 EPEPBR + .02 + .01 -.09 -.052 .465 EPEPDV -.05 + .01 .461 EPEEAL + .01 .00 .525 EPERES + + .01 + .01 .501 EPESPR + .00 + .01 .565 EPEMGR . 00 + .01 .519 EPESTR -.03 -.11 .325 EPEVDE .00 .00 .475 EPEVVI + .01 -.10 .566 AOCPFL +++ +++ + .25 + .06 + .40 + . 14 + .10 + .100 + .081 .404 AOCEGR - -.33 -.07 .00 -.49 -.02 + .088 -.057 .577 AOCVSC + + +++ +++ + . 14 + .19 + .11 + .22 + .17 -.048 -.059 .562 AOCCLA + .22 -.11 -.05 -.05 -.364* -.295 .517 AOCACA + .02 -.07 + . 13 -.27 -.33 -.031 + .114 .502 AOCALA + .08 + .19 -.11 -.01 -.01 -.070 + .110 .402 AOCVCU + .02 + .09 -.11 -.03 + .01 -.355 -.255 .514 AOCAAU .00 + .04 + .02 -.01 -.314 + .104 .526 AOCPAL + . 10 + .02 + .05 -.08 + .05 -.116 + .101 .529 AOCALF + + .00 -.19 + .05 + .09 + .04 + .111 +.784* .546 AOCLHI + + +++ + .20 + .11 \u00ab 00 + .17 + .23 + .040 -.339 .512 AOCTOC - -.11 -.22 + .05 -.15 -.32 .467 AOCAMI \u2014 -.31 -.37 . 00 -.49 -.21 + .048 -.155 .457 AOCTSP + .02 + .01 -.04 -.08 -.122 + .151 .532 AOCCRO + .02 -.04 + .03 -.36 -.50 + .284 -.017 .538 AOCEGL .00 -.06 -.22 -.76 .511 AOCSPA - \u2014 -.44 -.65 + .02 .00 -.350 .512 AOCAMO - -.23 -.48 -.26 -.23 -.298 .493 AOCPEP ++ + .02 + .14 .00 + .01 -.040 -.300. .527 AOCSIN - + .01 -.22 -.18 -.13 -.122 .494 AOCHGR + + + ++ + .06 + .07 + .06 + .03 -.562 -.397 .480 AOCLSP - -. 12 -.37 -.52 -.66 -.299 .512 AOCSLA - -.31 -.50 -.64 .502 AOCCMI -.06 -.33 AOCPPR - -.34 -.22 AOCPBR + + + .03 AOCPDV AOCEAL AOCRES +++ AOCSPR -AOCMGR AOCSTR AOCVDE AOCVVI PFLEGR +++ -.33 -.17 PFLVSC + +++ .00 -.03 PFLCLA ++ + . 14 .00 PFLACA - -.29 + .03 PFLALA -.56 .00 PFLVCU + .08 + .06 PFLAAU +++ + .17 + .03 PFLPAL + .07 -.07 PFLALF + .07 + .07 PFLLHI +++ +++ + .04 + .06 PFLTOC - -.07 -.37 PFLAM I .00 + .01 PFLTSP + .05 + .09 PFLCRO -.64 -.06 PFLEGL + .01 -.25 PFLSPA + .04 + .05 PFLAMO -.23 -.03 PFLPEP + .04 + .03 PFLSIN - - -.47 -.22 PFLHGR .00 + .02 PFLLSP - - - -.50 -.25 PFLSLA - - -.27 PFLCMI + .02 + .04 PFLPPR -.21 .00 PFLPBR + + .01 PFLPDV PFLEAL + PFLRES + -.20 -.20 +.013 .500 -.45 -.65 -.465 .530 +.04 +.01 +.221 .521 -.42 -.61 .378 .00 +.01 .515 +.03 +.03 .465 -.46 -.85 .519 +.01 +.01 .536 .00 .00 .515 -.99 .546 +.01 -.13 .516 +.03 +.18 -.115 +.146 .483 +.06 +.04 +.05 +.001 +.373 .469 +.14 +.03 +.06 -.204 -.234 .440 -.51 -.80 -.27 +.005 +.171 .399 -.48 -.19 +.01 +.151 +.323 .441 +.01 -.30 -.05 +.159 +.035 .390 -.04 -.03 +.170 +.397 .451 +.07 +.03 +.01 -.026 +.535* .451 -.17 +.02 +.01 +.125 +.353 .436 +.03 +.16 +.10 -.204 -.229 .433 .00 -.15 -.799* .453 +.03 -.58 -.07 -.177 -.117 .395 -.12 .00 +.126 -.359 .525 -.75 -.26 -.042 +.366 .480 -.09 -.43 .485 +.01 +.01 -.116 +.640* .491 -.12 -.18 -.525 -.234 .440 -.07 .00 -.006 +.246 .505 -.20 -.11 +.128 .447 .00 +.01 +.547 -.644 .454 -.25 -.30 +.463 .452 -.30 -.24 +.326 .405 .00 -.08 -.488 -.076 .461 -.65 -.25 +.007 +.103 .431 +.01 +.01 +.307 .356 +.49 +.34 .238 +.01 +.01 .472 +.01 +.01 .376 PFLSPR PFLMGR PFLSTR PFLVDE PFLVVI EGRVSC -.31 -.28 EGRCLA + .05 + .06 EGRACA +++ + .08 -.07 EGRALA + .01 -.11 EGRVCU ++ +++ + . 13 + .15 EGRAAU .00 + .09 EGRPAL + + + .29 + .25 EGRALF + .03 + .04 EGRLHI -.24 -.01 EGRTOC + .04 + .01 EGRAMI + -.08 + .09 EGRTSP + + . 13 + .10 EGRCRO -.29 -.04 EGREGL + .02 + .03 EGRSPA + .01 + .08 EGRAMO + + .08 + .08 EGRPEP + + .11 + .06 EGRSIN + .02 + .07 EGRHGR - - -.29 -.41 EGRLSP + + + .07 .00 EGRSLA + .02 EGRCMI + -.11 + .02 EGRPPR + +++ + .06 + .10 EGRPBR + .03 EGRPDV + + EGREAL EGRRES + EGRSPR EGRMGR EGRSTR + EGRVDE EGRVVI VSCCLA - -.23 -.19 VSCACA + + .66 + .41 -.05 -, 10 .399 . 00 00 .460 .00 -\u00ab 07 .403 -.38 -, 49 .449 .01 00 .492 -.29 -.57 -, 34 -.692* -.728* .439 + .37 -.34 + 4 01 + .251 + .067 .402 -.16 + .07 + I 14 + .231 -.071 .519 + . 15 + .03 + . 03 + .113 + .346 .533 -.11 + . 14 + \u00ab 12 +.735* -.004 .531 -.23 + , 01 -.044 -.172 .434 + .21 + .07 + t 02 + .175 -.179 .495 + .05 -.05 - i 04 + .072 -.216 .513 .00 -.63 03 + .212 + .091 .545 . 00 - , 06 -.865* .553 .00 + .02 ,05 + .371 -.047 .561 + .06 ,02 -.181 -.664* .564 + .19 + .04 ,01 + .382 + .259 .522 .00 .26 .544 + .01 - , .22 + .141 + .564 .477 + .03 + .03 -.175 + .248 .528 + .01 .00 -.414 + .236 .527 -.05 ,00 + .459 + .557 .456 -.25 - .31 .481 + .03 + .01 -.267 .513 + .03 .00 -.126 .485 + .03 + .02 .517 + .06 + .06 -.135 -.030 . 546 -. 17 - .14 .468 + .05 + .02 .497 + .01 .00 .498 + .01 + .01 .501 + .01 + .01 .484 + .01 + .01 .503 + .03 + .02 .462 + .01 + .01 .439 + .02 .00 .580 -.37 -.28 - .18 -.138 -.393 .505 -.12 + .12 + .02 +.585* +.746* .452 VSCALA +++ + + +++ +++ + .59 + .37 + .29 + .22 + .22 + .274 .491 VSCVCU + .27 + .12 + .06 + .03 - .111 + .258 .483 VSCAAU + . 17 - . 25 + .07 + .01 - .332 .482 VSCPAL + .21 - .08 .00 - .01 + .02 - .256 .461 VSCALF + - . 05 - .31 .00 . 00 + .05 + .425 .488 VSCLHI + +++ +++ + .35 + .34 + . 19 + .29 + .22 + .074 +.611* .519 VSCTOC - - .539 VSCAMI \u2014 - - . 5 5 - . 7 2 - . 1 7 - .32 .540 VSCTSP + . 15 + .04 - .01 + .01 + .094 - .133 .550 VSCCRO + .23 + .09 .00 + .02 + .444 + .194 .489 VSCEGL - .496 VSCSPA - . 15 - . 12 + .03 + .02 .502 VSCAMO - . 5 7 - .41 - . 5 0 - . 8 0 .510 VSCPEP + . 16 + .08 + .01 + .11 + .573 .531 VSCSIN + - . 4 0 - . 6 7 + .17 - .01 .500 VSCHGR ++ + +++ +++ +++ + .29 + .25 + .17 + .11 + .669 + .528 .519 VSCLSP \u2014 - - . 0 5 - .47 - .73 - .81 .516 VSCSLA - . 4 7 - . 16 - .32 .460 VSCCMI - - . 4 0 - . 62 - .58 - .63 .473 VSCPPR - . 4 0 - . 4 7 - .48 - .48 .462 VSCPBR +++ +++ - . 2 1 + .11 + .06 .456 VSCPDV - - - .43 - .52 .494 VSCEAL \u2014 - . 65 - . 89 .482 VSCRES - \u2014 - . 63 .488 VSCSPR - . 0 9 .00 .489 VSCMGR - . 7 0 .410 VSCSTR - .77 - .92 .459 VSCVDE - .31 - .45 .187 VSCVVI - - .81 - . 99 .591 CLAACA - .21 - . 44 - .45 - . 5 0 - .05 + .278 - .007 .515 CLAALA - - . 0 3 - . 2 0 - .61 - .28 - . 06 - .257 - .161 .498 CLAVCU - - . 0 3 - . 1 7 - .22 - .48 - . 14 + .285 - .033 .542 CLAAAU ++ + .02 - . 03 + .07 + .03 - .142 + .194 .472 CLAPAL - . 0 7 - . 0 7 + .03 - . 24 + .01 - .135 - .328 .524 CLAALF - .11 + .03 - . 33 - . 27 - .161 - .271 .513 CLALHI - . 4 9 - .11 - . 0 6 - .13 + .014 + .073 .542 CLATOC + .04 + .03 + .03 .00 + .140 + .824 .559 C L A A M I - - . 4 2 - . 19 - . 55 - .41 - .08 + .366 + .434 .496 CLATSP - . 4 0 - . 39 - .01 + .01 + .108 - .102 .558 CLACRO -.40 -.22 -.09 -.01 -.137 -.343 .531 CLAEGL + .02 + .03 .00 -.45 .538 CLASPA + .02 + .02 -.05 .00 + .412 + .020 .455 CLAAMO - -.29 -.31 -.02 + .02 -.435 .532 CLAPEP + .01 .00 + .01 .00 + .558 + .227 .539 CLASIN -. 12 -.06 -.11 + .01 + .183 + .075 .502 CLAHGR -.03 -.22 + .02 -.04 -.568 .526 CLALSP \u2014 -. 17 -.50 + .02 + .01 + .548 .506 CLASLA - -.46 -.24 .00 .520 CLACMI -.24 -.11 + .01 + .01 .549 CLAPPR -.27 -.12 .00 + .02 + .210 + .176 .524 CLAPBR -. 19 -.21 -.36 .518 CLAPDV + .02 + .01 .464 CLAEAL + + .04 + .01 . 508 CLARES + .02 .00 .496 CLASPR + + .01 + .01 .563 CLAMGR + .02 + .01 .530 CLASTR + .03 + .01 .502 CLAVDE -.99 .488 CLAVVI .00 -.21 .606 ACAALA ++ + +++ +++ +++ + .40 +.44 +.09 + .37 + .19 + .229 + .103 .514 ACAVCU + + + .21 +.15 -.26 + .33 + .24 + .223 -.124 .409 ACAAAU +++ + . 17 .00 -.02 + .05 - . 4 0 3 * - . 6 8 1 * .488 ACAPAL + + + +++ -.21 -.28 +.02 -.23 -.09 -.090 + .154 .505 ACAALF + + -o 14 +.04 +.02 -.20 -.28 -.160 -.324 .523 ACALHI - -.35 -.2 1 -. 16 -.10 + .226 + .404 .507 ACATOC -.80 -.61 -.73 .555 ACAAMI + +++ -.26 -.10 +.04 + . 18 + .32 +.561* -.081 .489 ACATSP ++ + +++ +++ + .21 + .19 + .25 + .07 -.057 + .107 .540 ACACRO +++ +++ +++ +++ + .31 +.32 +.13 + .32 + .12 +.452* + .064 .498 ACAEGL \u2014 \u2014 -.72 -.60 -.76 -.025 .519 ACASPA -.01 -.13 + .02 .00 -.124 -.442 .506 ACAAMO ++ + + .04 + .05 + .05 + .06 + .248 + .129 .509 ACAPEP + + - + . 10 + .14 -.09 -.66 .505 ACASIN + + + + +++ + . 12 + .11 + .14 + .11 + .307 + .019 .517 ACAHGR + + + + + . 10 + .11 + .05 + .02 -.167 -.323 .488 ACALSP + + + . 10 + .06 + .04 + .02 + .385 .471 ACASLA +++ ++ + + .05 + .13 + .05 - . 7 5 6 * .421 ACACMI + .04 + .02 + .03 + .02 -.359 .479 ACAPPR + + + +++ +++ + . 12 + .14 + .30 + .22 + .458 + .378 .449 ACAPBR -.07 -.08 -.47 .462 ACAPDV +++ +++ + .17 + .10 .442 ACAEAL -.84 -.93 .470 ACARES -.99 -.99 .497 ACASPR + +++ .441 ACAMGR .479 ACASTR .480 ACAVDE + +++ .489 ACAVVI .553 ALAVCU + +++ + .36 + .25 -.22 + .14 + .03 -.032 + .105 .433 ALAAAU + + . 17 + .06 + .02 + .09 -.041 -.200 .489 ALAPAL - -*03 -.22 -. 19 -.39 + .095 -.516 .491 ALAALF - -.04 + .10 + .17 -.19 -.39 +.677* + .303 .490 ALALHI + .07 + .07 -.11 -.01 + .02 -.319 -.252 .522 ALATOC -- -.76 -.92 .537 ALAAMI -.29 -.16 -.11 -.04 -.17 + .521 +.712* .453 ALATSP ++ + + + + + + .34 + .2 1 + .28 + .04 -.297 -.277 .512 ALACRO + + + +++ +++ +++ + .40 + .44 + .01 + .34 + .13 +.451* +.663* .535 ALAEGL - - -.60 -.99 .557 ALASPA -.21 -.33 + .02 -.36 .416 ALAAMO .00 + .02 + .05 -.13 + .463 .436 ALAPEP + + * 15 + .15 -.23 -.66 + .371 + .413 .524 ALAS IN + + + + . 17 + .14 -.08 + .06 +.723* +.783* .422 ALAHGR +++ +++ +++ +++ + 4.21 + .19 + . 16 + .17 +.577* + .441 .435 ALALSP + + . 12 + .11 + .00 + .01 -.429 .510 ALASLA + .05 + .06 -.03 +.923* .481 ALACMI + .03 + .07 -.09 -.19 .442 ALAPPR + +++ +++ + .07 + .15 + .21 + .10 + .249 + .136 .470 ALAPBR + .03 + .06 + .01 .502 ALAPDV +++ +++ + .17 + .09 .400 ALAEAL \u2014 -.75 -.86 .491 ALARES -.60 -.77 .463 ALASPR +++ + .03 + .04 .531 ALAMGR .500 ALASTR -.87 -.90 .444 ALAVDE + +++ + .06 + .04 .453 ALAVVI - - -.80 -.82 .487 VCUAAU + + .35 + .14 + .02 -.13 -.062 -.046 .500 VCUPAL + + . 18 + .07 + .08 + .10 -.370 + . 142 .442 VCUALF + . 18 + .01 + i 03 -.08 + .03 -.218 + .031 .461 VCULHI + + + -.01 -.04 + .07 + .12 + .046 + .353 .422 VCUTOC - \u2014 -.73 -.43 - .56 .481 VCUAMI + + ++ + -.12 -.19 + < 27 + .15 + .14 -.248 - .261 .443 VCUTSP + + + + . 14 + .01 + .15 + .02 + .140 + .121 .499 VCUCRO + + + .07 -.04 , 15 + .11 - .02 -.176 - .057 .489 VCUEGL - - -.28 - .59 .511 VCUSPA + + + + + + + .13 + .01 + .06 + .05 -.249 .462 VCUAMO - -.33 -.57 + .04 + .01 .426 VCUPEP + + .10 + .13 + .01 - .42 -.329 - .559 .499 VCUSIIM + + + + .07 + .08 + .30 + .03 -.213 - .121 .497 VCUHGR - + .07 -.28 -. 14 - .45 .439 VCULSP -.25 + .02 .00 - .34 .000 \u2022 442 VCUSLA + + + .01 + .06 + .03 .452 VCUClMI + .07 + .02 + .01 + .02 .435 VCUPPR + + + + .02 + .09 + .15 + .01 -.106 - .287 .351 VCUPBR + + .07 + .03 + .02 .434 VCUPDV + + .06 - .04 .416 VCUEAL -.11 - .22 .459 VCURES -.16 - .21 .452 VCUSPR + .02 .15 .469 VCUMGR -.61 - .49 .432 VCUSTR -.56 - .66 .455 VCUVDE + + .01 + .02 .444 VCUVVI - - -.40 -.50 .483 AAUPAL + . 17 + .21 + .04 + .02 +.457* + .628* .479 AAUALF + .20 + .31 -.06 - .38 +.620* + .472 .403 AAULHI + .09 -.19 .00 .00 + .037 .466 AAUTOC -.29 + .03 -.76 - .92 .512 AAUAMI + + + +++ -.05 -.06 + . 17 + .06 + .127 .443 AAUTSP + + . 17 + .2 1 + .06 + .28 -.069 - .216 .549 AAUCRO + + .11 + .11 + .10 + .08 -.312 - .301 .492 AAUEGL -.67 -.29 -.22 - .21 .510 AAUSPA + . 12 + .13 -.01 +.738* .461 AAUAMO + + + -.09 + .13' + .03 + .12 + .201 .424 AAUPEP + .05 + .02 -.40 .00 -.546 .515 AAUS IN + .03 -.11 + . 14 + .07 -.472 .430 AAUHGR .00 -.55 .00 - .40 .000 .403 AAULSP + \u00bb00 + .13 +.07 +.01 -.396 .482 AAUSLA +.02 -.02 +.02 +.830* .508 AAUCMI + +.05 +.09 +.07 +.05 -.306 .486 AAUPPR + +++ +.03 +.02 +.14 +.16 +.258 .480 AAUPBR -.17 +.04 .00 .432 AAUPDV + +++ +.12 +.10 .487 AAUEAL - -.39 .484 AAURES -.22 .507 AAUSPR +++ +++ +.06 +.06 .491 AAUMGR -.36 .525 AAUSTR - \u2014 -.64 -.99 .382 AAUVDE - .481 AAUVVI - - -.80 .572 PALALF +++ +.31 -.08 +.07 +.02 +.211 +.544 .501 PALLHI -.24 -.11 +.07 -.05 -.02 \".281 -.253 .383 PALTOC -.11 +.01 +.11 -.21 -.27 .402 PALAM I + -.04 +.09 -.05 +.09 -.047 -.402 .552 PALTSP +.05 -.10 +.01 -.34 +.278 -.255 .351 PALCRO -.21 -.40 +.09 -.13 -.16 -.248 .284 PALEGL -.37 -.13 .00 -.33 .315 PALSPA +++ +.09 +.10 +.03 +.03 +.208 .497 PALAMO + -.01 +.05 +.05 +.05 +.406 .537 PALPEP +.11 -.13 +.01 -.47 -.432 .297 PALSIN -.09 -.18 +.07 +.03 -.695* .528 PALHGR - - -.53 -.73 -.20 -.52 .498 PALLSP +.06 -.13 +.02 +.01 -.309 .327 PALSLA +.01 +.04 +.02 -.605 .455 PALCMI +.03 +.04 +.04 +.02 -.242 .451 PALPPR -.35 -.35 -.01 +.01 .497 PALPBR +.04 +.03 .00 .451 PALPDV +.01 +.01 .491 PALEAL +.03 .00 .422 PALRES +.03 +.01 .431 PALSPR -.27 -.53 .472 PALMGR +.01 .00 .431 PALSTR +.02 -.14 .487 PALVDE -.17 -.27 .420 PALVVI +.01 -.03 .562 ALFLHI .00 +.14 +.17 +.05 +.02 +.247 -.050 .443 ALFTOC -. 14 -.12 + .11 -.22 - .55 .494 ALFAMI + + + + + . 24 + .41 + .07 . 00 - .04 + .323 + .173 .351 ALFTSP + + . 14 + .28 -.18 + .04 + . 175 -.263 .583 ALFCRO \u2014 -.09 + .18 + .09 -.49 - .67 + .560* +.767* .513 ALFEGL + -. 20 + .01 + .03 + .05 .531 ALFSPA + . 17 + .17 -.04 - .07 + .336 +.805* .535 ALFAMO - + .08 + .17 -.18 - .46 - .017 + .592 .268 ALFPEP + .08 + .08 .00 - .51 + .062 .552 ALFSIN .00 -.34 -.10 - .47 + .116 .346 ALFHGR -.40 -.34 -.26 - .82 .422 ALFLSP + . 10 + .01 -.32 .66 .000 .537 ALFSLA + * 12 -.05 .00 + .437 .487 ALFCMI + + + . 14 + .14 + .04 + .03 \u2014 .394 .526 ALFPPR - - + .05 + .14 -.37 - .55 + .553 + .208 .461 ALFPBR + + .09 + .07 + .03 .437 ALFPDV \u2014 -.51 - .56 .473 ALFEAL + .04 + .01 .533 ALFRES + .03 + .03 .475 ALFSPR . 542 ALFMGR + .08 + .01 .494 ALFSTR + .06 + .01 .283 ALFVDE -.30 \u2014 .33 .468 ALFVVI + .01 - .49 .575 LHITOC -.49 + .02 .00 \u2014 .17 .375 LHI AM I + .01 + .05 -.12 - .11 + .707* + .219 .444 LHITSP + \u202211 + .10 -.02 .00 - .434 -.147 .468 LHICRO -.11 -.24 -.29 - .46 - . 150 -.231 .416 LHIEGL - + .01 -.09 -.07 - .48 .423 LHISPA + + + + .01 + .11 + .06 + .01 + .247 -.415 .553 LHIAMO - -.35 .00 -. 19 - .29 .485 LHIPEP -.04 + .09 + .03 + .01 \u2014 .307 + .600 .464 LHISIN -.11 -.43 -.08 - .16 + .556 .512 LHIHGR + + + + + .11 + .13 + .05 + .03 - .391 -.026 .464 LHILSP -.29 -.54 -.50 - .87 .360 LHI SLA - -.40 -.42 - .48 .473 LHICMI - \u2014 -. 11 -.07 -.27 - .50 .510 LHIPPR - -.40 -.13 -.29 .28 .480 LHIPBR + + + + + + + .07 + .05 + .02 .438 LHIPDV -.34 .62 .475 LH I HAL- -.28 -.33 .461 L-HIRES \u2014 -.07 -.75 .474 LHISPR -.24 -.19 .488 LHIMGR -.06 -.51 .474 LHISTR -.21 -.73 .495 LHIVDE -.78 -.87 .501 LHIVVI -. 30 -.71 .547 TOCAMI + .26 + .46 + .07 -.17 -.56 .509 TOCTSP \u2014 - - -.60 -.71 .518 TOCCRO - -.99 -.99 .398 TOCEGL ++ + +++ ++ + ++ + + .52 + .63 + .36 + .30 .368 TOCSPA + .01 -.07 + .03 .00 .546 TOCAMO + .08 -.07 + .08 + .03 .525 TOCPEP -.45 -. 57 .400 TOCSIN -.76 -.99 .557 TOCHGR - .535 TOCLSP -. 19 .447 TOCSLA - -.72 .520 TOCCMI - -.75 .520 TOCPPR - \u2014 -.29 -.69 -.79 .532 TOCPBR .467 TOCPDV -.76 -.76 .501 TOCEAL + .06 -.02 .483 TOCRES -.44 .456 TOCSPR .473 TOCMGR .512 TOCSTR + ++ + + .16 +. 14 .526 TOCVDE .521 TOCVVI -.44 .508 AMITSP -.03 + .12 + .06 -.01 186 -.309 .569 AMICRO + + + +++ -. 13 + .01 + .19 + . 16 + .05 + \u2022 866* + .479 .563 AMIEGL - + .05 + .06 -.23 -.47 .579 AM I SPA + + .21 + .10 + .04 + .01 053 .517 AM IAMO + + + + + + + .27 + .23 + .06 + .04 + . 090 +.697* .375 AMIPEP -.08 -.21 -.21 .00 .583 AM I SIN +++ +++ + . 12 + .06 + .17 + .06 + . 857* .431 AMIHGR - -.76 -.67 -.67 -.92 .444 AMILSP + + +++ + . 17 + .06 + .05 + .03 +. O i l .538 AM I SLA + +++ +++ + . 19 + .09 + .07 + .087 .505 AM I CM I + + + + + . 15 + .14 + .07 + .03 - .194 .525 AMIPPR + +++ +++ + . 18 + .14 + . 19 + .17 +.583 +.515 .462 AM IPBR + .03 - .11 + .01 .436 AMIPDV +++ +++ + .13 + .05 .411 AMIEAL - - .21 - .47 .535 AM I RES - .23 - .41 .512 AMISPR + + .03 + .01 .526 AMIMGR - . 1 7 - \u2022 62 .513 AMISTR \u2014 - . 44 - . 94 .407 AMIVDE + + + .03 + .02 .522 AMIVVI - . 8 7 - . 99 .538 TSPCRO ++ + + .11 + .17 + .31 + .03 +.104 +.199 .390 TSPEGL - - - . 77 - .77 .409 TSPSPA + .05 - . 0 2 - . 3 4 + .377 .547 TSPAMO +++ + ++ + . 13 + .28 + . 14 + .09 - .404 - .452 .570 TSPPEP ++ + .26 + .19 .00 - .41 - .126 - .039 .376 TSPSIN + + . 10 + .09 + .07 + .07 - .265 .545 TSPHGR + . 15 + .16 + .03 + .04 + .390 .532 TSPLSP + .09 + .08 + .04 + .03 +.901* .328 TSPSLA + + .17 + .07 + .04 - .551 .527 TSPCMI + + .08 + .18 + .09 .00 .507 TSPPPR +++ +++ + . 10 + .19 + . 34 + .13 - .039 - .139 .564 TSPPBR + .07 + .02 - .21 .505 TSPPDV +++ + .22 + .03 .528 TSPEAL - - .58 - . 49 .516 TSPRES - . 32 - .57 .507 TSPSPR + + .06 + .03 .543 TSPMGR - .63 .489 TSPSTR - - .58 - .26 .571 TSPVDE - .58 - .57 .516 TSPVVI - . 3 2 .557 CROEGL - - - . 77 - . 79 .360 CROSPA - . 3 5 - . 07 - . 5 0 .525 CROAMO + + .07 + .14 + .08 + .07 - .337 . 548 CROPEP - - . 13 + .03 - . 64 .284 CROSIN + + + + +++ ++ + + .28 + .21 + .25 + .14 +.313 +.139 .541 CROHGR + +++ + .20 + .14 + .03 + .09 - .061 .523 CROLSP +++ ++ + . 15 + .20 + .11 + .02 + .252 .338 CROSLA + + + +++ + * 11 + .23 + .09 - .393 .519 CROCMI ++ + .03 + .10 + .15 + .01 .518 CROPPR + +++ ++ + +++ + .22 + .3 1 + .52 +.24 +.563 +.428 .540 CROPBR - - . 6 2 - . 80 - .63 .500 CROPDV + + + +++ + .28 + .12 .521 CROEAL - - . 9 9 .489 CRORES \u2014 .459 CROSPR +++ ++ + + .11 + .07 .531 CROMGR .459 GROSTR \u2014 - . 99 - .99 . 504 CROVDE +++ +++ + .09 + .06 .489 GROW I \u2014 - .582 EGLSPA - . 2 9 - . 32 .540 EGLAMO + .23 + .05 - . 05 - .53 .564 EGLPEP -* 23 .343 EGLSIN - - - . 79 - .83 .569 EGLHGR . 544 EGLLSP - .32 .414 EGLSLA .505 EGLCMI .518 EGLPPR - \u2014 - . 17 - .99 .560 EGLPBR .479 EGLPDV - . 5 9 .522 EGLEAL - .41 .475 EGLRES .485 EGLSPR .487 EGLMGR .466 EGLSTR +++ ++ + + .22 + .21 .529 EGLVDE .476 EGLVVI .573 SPAAMO - . 0 3 + .05 + .05 - .19 .483 SPAPEP - .18 - . 4 4 - .48 .497 SPASIN + + .06 + .08 + .01 .505 SPAHGR - . 64 - .55 .499 SPALSP + .02 + .06 + .01 .499 SPASLA + . 13 - .32 + .02 .541 SPACMI + + .23 + .22 + .11 + .01 .451 SPAPPR - .11 - . 4 4 + .02 .00 .481 SPAPBR ++ + .03 + .02 + .10 .522 SPAPDV + .02 + .03 .472 SPAEAL + .09 + .02 .500 SPARES + .09 .490 SPASPR . 543 SPAMGR .508 SPASTR -.59 -.35 .485 SPAVDE .464 SPAVVI + .03 .551 A M O P E P + .07 -.44 -.03 + .02 .585 AMOS IN + .16 + .07 -.04 + .03 .311 AMOHGR - -.54 -.31 -.79 -.78 .447 AMOLSP + + + .32 + .26 + .02 + .02 .555 AMOSLA + + + +++ + + .56 + .48 + . 14 +.10 +.186 .487 AMOCMI + + + +++ +++ + + .35 + .3 1 + .18 + .06 .474 AMOPPR + + + + ++ + + .40 + . 15 +.17 +.005 .465 AMOPBR + .07 -\u2022 18 -.59 .457 AMOPDV + +++ + .15 + .10 .452 AMOEAL + .08 .00 .538 AMORES + .02 + .01 .530 AMOSPR -.49 -.43 .545 AMOMGR + .03 -.43 .514 AMOSTR + .03 -.04 .276 AMOVDE + .01 + .02 .491 AMOVVI + .02 .00 .517 PEPSIN + .09 + .17 + .05 -.54 .573 PEPHGR + .04 -.44 -.06 .00 .533 PEPLSP + . 13 + .11 .00 + .03 .326 PEPSLA -.14 + .05 .00 .493 PEPCMI -.04 -.29 -.35 .00 .523 PEPPPR + .09 + .12 .00 -.07 .548 PEPPBR -.36 -.37 + .04 .522 PEPPDV -.37 .00 .542 PEPEAL + .04 + .01 .481 PEPRES -.28 .469 PEPSPR .524 PEPMGR + .01 + .05 .505 PEPSTR + .04 .539 PEPVDE + .01 .497 PEPVVI + . 10 .604 SINHGR -.17 -.22 -.20 -.17 .443 SINLSP +++ +++ + +++ + . 37 + .31 + .08 +.06 -.427 .532 SINSLA + . 19 + .04 -.01 .462 SINCMI + .21 -.11 + .04 + .01 .477 SINPPR + + +++ +++ + .34 + .37 + .22 +.22 +.048 .495 SINPBR + .05 + .02 .00 .440 SINPDV ++ + .06 + .06 .456 SINEAL -.24 -.25 .565 SINRES -.18 -.68 .504 SINSPR +++ +++ + .07 + .07 .555 SINMGR .00 + .01 .523 SINSTR -.49 -.99 .247 SINVDE + .04 + .02 .480 SINVVI -.59 -.76 .494 HGRLSP .498 HGRSLA -.69 .452 HGRCMI - -.37 -.11 .474 HGRPPR + .08 + .22 -.49 + .01 .459 HGRPBR -. 17 + .06 -.21 .473 HGRPDV -.47 -.63 .417 HGREAL .478 HGRRES .432 HGRSPR .00 .00 .451 HGRMGR .489 HGRSTR .436 HGRVDE .500 HGRVVI .571 LSPSLA ++ + ++ ++ + .39 + .18 + .07 .456 LSPCMI + +++ +++ + .29 + .07 + .22 + .09 .492 LSPPPR +++ +++ +++ + + + + .47 + .75 + .16 +.07 -.092 .498 LSPPBR + .09 .453 LSPPDV + .06 + .03 .494 LSPEAL -.58 -.02 .461 LSPRES -.32 .450 LSPSPR + + .06 + .07 .489 LSPMGR + . 12 + .02 .468 LSPSTR + .09 .510 LSPVDE + .07 + .02 .481 LSPVVI + .03 + .01 .586 SLACMI +++ + + + + + .47 + .18 + .14 .482 SLAPPR + +++ +++ + .41 + .29 + .10 .489 SLAPBR + .12 .446 SLAPDV +++ +++ + .24 + .11 .443 SLAEAL -.71 .434 SLARES .425 SLASPR + .03 .00 .443 SLAMGR .501 SLASTR - .440 SLAVDE + + + + .11 + .02 .459 SLAVVI .567 CMIPPR + +++ +++ + .34 + .22 + .06 .432 CM IPBR + .29 .377 CMIPDV ++ + +++ -.31 + .23 + .11 .440 CM IEAL + + .17 + .05 .510 CMIRES +++ + .17 + .04 .423 CMISPR + .02 + .02 .500 CMIMGR + .20 + .16 .482 CMISTR + + . 13 + .08 .474 CMIVDE -.21 + .04 .432 CM IVVI + .06 -.37 .528 PPRPBR \u2014 - .453 PPRPDV +++ +++ + .40 + .11 .434 PPREAL - - -.67 -.99 .496 PPRRES - .451 PPRSPR +++ +++ + .10 + .07 .487 PPRMGR .506 PPRSTR . -.99 -.99 .453 PPRVDE +++ +++ + .09 + .09 .442 PPRVVI - - .535 PBRPDV -.64 .424 PBREAL -.49 .509 PBRRES .444 PBRSPR .426 PBRMGR .463 PBRSTR .416 PBRVDE .435 PBRVVI .522 PDVEAL - .513 PDVRES .357 288 Appendix 3. The scheme tha t has been fo l l owed i n Appendix 3 i s a much condensed and l i m i t e d v e r s i o n of tha t used i n the B i o l o g i c a l F l o r a of the B r i t i s h I s l e s , p u b l i s h e d r e g u l a r l y i n the J o u r n a l of Eco logy . The format i s o u t l i n e d below: Name o f species Vege ta t ive h a b i t ; vege t a t i ve r e p r o d u c t i o n Flower type (see Sec t . I I I - H ) ; f l o r a l b i o l o g y ; blooming p e r i o d Mode of p o l l i n a t i o n and types of p o l l i n a t o r s C o m p a t i b i l i t y Propagule type (see Sec t . I I I - F ) ; probable mode of d i s p e r s a l E c o t y p i c v a r i a t i o n H y b r i d i z a t i o n D i s t r i b u t i o n A d d i t i o n a l comments, i f any C o m p a t i b i l i t y r e l a t i o n s were t e s t e d by s e l f i n g experiments performed r\\in i n s e c t exc losures o f f i n e mesh c h e e s e c l o t h , both i n the f i e l d and i n the greenhouse. Plump f r u i t - s e t and matura t ion of r i p e , f u l l seeds were cons idered evidence o f s e l f - c o m p a t i b i l i t y . With respect to e c o t y p i c v a r i a t i o n , the view adopted here was tha t ecotypes are l i k e l y to evolve i n any wide- rang ing ( e i t h e r a r e a l l y or a l t i t u d i n a l l y , or b o t h ) , s e x u a l l y r e p r o d u c i n g , angiosperm s p e c i e s . 289 A. S a l t Marsh Agrostis exarata T r i n . Tufted p e r e n n i a l , short-rhizomatous; p l a n t s o f t e n some-what decumbent and n o d a l l y r o o t i n g . Flowers amorphic; p e r f e c t , weakly protandrous, reduced, inconspicuous, o d o r l e s s , n e c t a r l e s s ; blooming (1971) J u l y 5-10 t o J u l y 30-Aug. 5. P o l l i n a t i o n anemophilous and f r e q u e n t l y entomophilous, by Bombus t e v r i c o l a occiden t a l i s Grne. ( P o j a r 1973b). 5\/5 i n d i v i d u a l s s e l f - i n c o m p a t i b l e . Propagule a m i c r o s c l e r o c h o r e ( s m a l l , unawned, w i t h a s l i g h t l y bearded c a l l u s ) ; d i s p e r s a l anemochorous and epizoochorous (mud). E c o t y p i c v a r i a t i o n probably s t r o n g , i n view of s p e c i e s ' d i s t r i b u t i o n . H y b r i d i z a t i o n l i k e l y , as A. exarata i s a polymorphic species c o n s i s t i n g of s e v e r a l i n t e r g r a d i e n t races (Hitchcock et al. 1969). D i s t r i b u t i o n : \"Moist areas, from near sea l e v e l t o midmontane; n. A l a s , t o C a l i f . , e. t o A l t a . , S.D., Neb., Tex., and n. Mex.\" (Hitchcock et al. 1969). Carex lyngbyei Hornem. P e r e n n i a l sedge, somewhat clumped along w e l l - d e v e l o p e d , creeping rhizomes. Flowers amorphic; u n i s e x u a l , reduced, i n c o n s p i c u o u s , o d o r l e s s , n e c t a r l e s s , aggregated i n t o 3-7 t e r m i n a l s p i k e s ; p l a n t s monoecious, f l o w e r i n g (1971) A p r i l 20-25 to .2,3-28. P o l l i n a t i o n anemophilous. 7\/7 i n d i v i d u a l s s e l f -compatible; East (1940) thought t h a t the genus was s e l f - c o m p a t i b l e , but t h a t s e l f - f e r t i l i z a t i o n was l i m i t e d by dichogamy. In t h i s r e s p e c t , t h e r e are protogynous, protandrous, and homogamous i n d i v i d u a l s i n the same p o p u l a t i o n of C. l y n g b y e i , as w i t h the c l o s e l y r e l a t e d C. obnupta. As f a r as I know, t h i s i s a h e r e t o f o r e unreported phenomenon. Propagule -a m i c r o s c l e r o c h o r e (achene + f i r m , t h i c k - w a l l e d perigynium); d i s p e r s a l hydrochorous and epizoochorous (mud). E c o t y p i c v a r i a t i o n u n l i k e l y . H y b r i d i z a t i o n may occur w i t h the r e l a t e d and sometimes sympatric C. obnupta. D i s t r i b u t i o n : \" C o a s t a l marshes and t i d a l f l a t s ; i n t e r r u p t e d l y c i r c u m b o r e a l along the s e a c o a s t s , i n Am. s. to Que. and n. C a l i f . \" (Hitchcock et al. 1969). Deschampsia aespitosa (L.) Beauv. S t r o n g l y c e s p i t o s e p e r e n n i a l ; no v e g e t a t i v e r e p r o d u c t i o n i n the study a r e a , but o c c a s i o n a l v i v i p a r y and agamospermy have been r e p o r t e d from the D. aespitosa complex (Lawrence 1945; Nygren 1949 ; Clausen and Hiesey 1958; Kawano .1963). Flowers amorphic; p e r f e c t , moderately protandrous, reduced, inconspicuous, o d o r l e s s , n e c t a r l e s s ; blooming (1971) June 10-20 t o J u l y 20-24. P o l l i n a t i o n anemophilous and f r e q u e n t l y entomophilous, by Bombus t e r r i c o l a o c c i d e n t a l i s Grne. ( P o j a r 1973b). 7\/7 i n d i v i d u a l s s e l f - i n c o m p a t i b l e (0-5% f e r t i l e s p i k e l e t s ) ; S a i r f i e l d (1954) found 0-15% s e l f - c o m p a t i b i l i t y i n Deschampsia flexuosa. Propagule a sporochore and pogonochore, awned and beset w i t h c a l l u s h a i r s ; d i s p e r s a l anemochorous ( R i d l e y 1930) and epizoochorous (mud). E c o t y p i c v a r i a t i o n marked (Lawrence 1945; Kawano 1963; T i e s z e n and Bonde 1967; Ward 1969; Pearcy and Ward 1972). H y b r i d i z a t i o n r e p o r t e d by Kawano (1963). D i s t r i b u t i o n : \"From a l p i n e r i d g e s and t a l u s s l o p e s , montane meadows and moist areas i n the mountains to c o a s t a l marshes and moist ' p r a i r i e s 1 ; A l a s , t o Greenland, s. to C a l i f . , A r i z . , N.M. , Mich., W i s e , and Ga.; Mex. ; E u r a s i a \" (Hitchcock et al. 1969); \"modern d i s t r i b u t i o n area ...covers almost the e n t i r e area of P l e i s t o c e n e g l a c i a t i o n i n the northern hemisphere\" (Kawano 1963). D i s t i c h l i s spioata (L.) Greene. E x t e n s i v e l y rhizomatous p e r e n n i a l grass. Flowers amorphic; u n i s e x u a l , reduced, i n c o n s p i c u o u s , o d o r l e s s , n e c t a r l e s s ; p l a n t s d i o e c i o u s , f l o w e r i n g (1971) J u l y 20-30 to Aug. 15-25. P o l l i n a t i o n anemophilous. C o m p a t i b i l i t y i r r e l e v a n t , as p l a n t s are d i o e c i o u s and not agamospermous. Propagule a mega-s c l e r o c h o r e ; d i s p e r s a l anemochorous and epizoochorous (mud). E c o t y p i c v a r i a t i o n ( l a t i t u d i n a l ) p o s s i b l e . Apparently no h y b r i d i z a t i o n . D i s t r i b u t i o n : \" C o a s t a l beaches and s a l t marshes; Van. I . t o C a l i f . , and on the e. coast of N. Am. from Canada to F l a . , La., and Tex.; W. I n d i e s \" (Hitchcock et al. 1969). Note: D. spioata was present i n the study marsh only i n the p i s t i l l a t e c o n d i t i o n . Festuca rubra L. P e r e n n i a l , u s u a l l y rhizomatous, forms l a r g e but loose clumps; hemicryptophyte; v e g e t a t i v e r e p r o d u c t i o n moderate i n the study marsh, but i n B r i t a i n e x t e n s i v e (Harberd and Owen 1969), forming clones up t o 240 yds. i n diameter (Harberd 1961a). Flowers amorphic; p e r f e c t , moderately protandrous, reduced, inconspicuous, o d o r l e s s , n e c t a r l e s s ; blooming (1971) J u l y 1-5 to 20-30. P o l l i n a t i o n anemophilous and f r e q u e n t l y entomophilous, by Bombus terricola occi-dentalis Grne. (Pdjar 1973b). 5\/5 i n d i v i d u a l s s e l f - i n c o m p a t i b l e ; i n B r i t a i n the s p e c i e s i s a l s o s e l f - i n c o m p a t i b l e , though Harberd (1961a) obtained.a few s u c c e s s f u l s e l f i n g s ; East (1940) a l s o found F . rubra s t r o n g l y s e l f - i n c o m p a t i b l e . Propagule a sporochore and pogonochore, awned and v i l l o u s ; d i s p e r s a l anemochorous and e p i -zoochorous (mud). E c o t y p i c v a r i a t i o n moderate (Harberd 1961b) to strong (consider range and e c o l o g i c a l amplitude of the s p e c i e s ) . H y b r i d i z a t i o n probable, at l e a s t at the r a c i a l l e v e l w i t h i n t h i s polymorphic species. D i s t r i b u t i o n : \" C o a s t a l marshes and dunes to montane f o r e s t and meadows; A l a s , to Newf., s. to s. C a l i f . , N.M., Tex., and S.C.; Europe\" (Hitchcock et al 1969); a wide-ranging, c i r c u m p o l a r -c i r c u m b o r e a l complex (Hulten 1968). Comment: Harberd (1961a) maintained t h a t , due t o e x t e n s i v e v e g e t a t i v e apomixis, th e r e are r e l a t i v e l y few genotypes i n any p a r t i c u l a r a r e a , say, 100 yds. square (at l e a s t i n B r i t a i n ) . 291 G l a u x maritima L. F l e s h y , m o d e r a t e l y r h i z o m a t o u s , p e r e n n i a l h e r b ; h e m i c r y p t o p h y t e . F l o w e r s s t e r e o m o r p h i c ; p e r f e c t , homogamous, s l i g h t l y n e c t a r i f e r o u s b u t o d o r l e s s ; c a l y x ( p e t a l s l a c k i n g ) w h i t e t o p a l e p i n k i s h ; s t i g m a s b o r n e s l i g h t l y above t h e a n t h e r s , so t h a t autogamy, t h o u g h p o s s i b l e , i s n o t i n e v i t a b l e ; b l o o m i n g (1971) May 25-31 t o J u l y 15-20. P o l l i n a t i o n by s e l f i n g and e n t o m o p h i l y a n t s ( p e r s o n a l o b s e r v a t i o n s and r e p o r t e d i n F a e g r i and v a n d e r P i j l ( 1 9 7 1 ) ) and s m a l l m u s c i d f l i e s ( Clapham, T u t i n and Warburg 1 9 6 2 ) . 5\/5 i n d i v i d u a l s s e l f - c o m p a t i b l e . P r o p a g u l e a c y c l o c h o r e ( d i s p e r s a l u n i t a p p e a r s t o be t h e seeds + t h e c o h e r e n t p l a c e n t a , w h i c h i s spongy and b u o y a n t ) ; d i s p e r s a l h y d r o c h o r o u s and e p i z o o c h o r o u s (mud). E c o t y p i c v a r i a t i o n p r o b a b l e ; l a t i t u d i n a l , and i n l a n d vs. m a r i t i m e ? ( c f . C a l d e r and T a y l o r 1 968). H y b r i d i z a t i o n none ( m o n o t y p i c g e n u s ) . D i s t r i b u t i o n : \" . . . o v e r much o f A r c t i c and t e m p e r a t e N. Am.; E u r a s i a \" ; i n t e r r u p t e d l y c i r c u m b o r e a l - s u b a r c t i c ( H u l t e n 1 9 6 8 ) . Hordeum brachyantherum N e v s k i . T u f t e d p e r e n n i a l ; h e m i c r y p t o p h y t e ; v e g e t a t i v e r e p r o d u c t i o n none. F l o w e r s a m o r p h i c ; p e r f e c t , homogamous, r e d u c e d , i n c o n s p i c u o u s , o d o r l e s s , n e c t a r l e s s ; a n t h e r s o n l y p a r t i a l l y e x s e r t e d a t a n t h e s i s , some f l o r e t s seem t o be c l e i s t o g a m o u s ; b l o o m i n g (1971) J u l y 5-10 t o J u l y 30-Aug. 5. P o l l i n a t i o n by s e l f i n g and a n e m o p h i l y . 3\/3 i n d i v i d u a l s s e l f - c o m p a t i b l e ; t h i s a g r e e s w i t h t h e f i n d i n g s o f M i t c h e l l and W i l t o n ( 1 9 6 4 ) . P r o p a g u l e a m e g a s c l e r o c h o r e and desmochore (whole s p i k e o r awned s p i k e l e t s ( n o d a l g r o u p s o f 3 s p i k e l e t s ) ) ; d i s p e r s a l anemo-c h o r o u s and e p i z o o c h o r o u s ( s t i c k t i g h t ) . E c o t y p i c v a r i a t i o n a l m o s t c e r t a i n . H y b r i d -i z a t i o n r e p o r t e d by R a j h a t h y and M o r r i s o n ( 1 9 5 9 ) , M i t c h e l l and W i l t o n ( 1 9 6 4 ) , and C a l d e r and T a y l o r ( 1 9 6 8 ) . D i s t r i b u t i o n : \"Ocean b e a c h e s t o m o u n t a i n meadows, u s u a l l y where m o i s t , as a l o n g s t r e a m s o r a r o u n d m a r s h e s , b u t a l s o f r o m d r y s a g e b r u s h d e s e r t t o open k n o l l s and r o c k y r i d g e s ; A l a s , t o C a l i f . , and N.M.; r e p o r t e d a l s o f r o m n.e. A s i a \" ( H i t c h c o c k e t al. 1 9 6 9 ) . Juncus balticus W i l l d . P e r e n n i a l w i t h t h i c k , w i d e - s p r e a d i n g r h i z o m e s ; g e o p h y t e (Clapham, T u t i n and Warburg 1 9 6 2 ) . F l o w e r s a c t i n o m o r p h i c ; p e r f e c t , - s t r o n g l y p r o t o -g y n o u s , i n c o n s p i c u o u s , o d o r l e s s , n e c t a r l e s s ; ' 3 s t i g m a s b o r n e w e l l above t h e a n t h e r s f l o w e r s l o w e s t i n t h e i n f l o r e s c e n c e f l o w e r f i r s t , and f l o w e r i n g p r o c e e d s upwards and outwards t h r o u g h t h e e s s e n t i a l l y cymose i n f l o r e s c e n c e ; b l o o m i n g (1971) June 12-25 t o J u l y 20-30. B r i t i s h s p e c i e s o f Juncus a r e a l l p r o t o g y n o u s , a n e m o p h i l o u s ; some s p e c i e s f l o w e r i n \" p u l s e s \" ; i . e . , s i m u l t a n e o u s a n t h e s i s o f a l l f l o w e r s i n t h e i n f l o r e s c e n c e ( R i c h a r d s and Clapham 1941; R i c h a r d s 1943; J o n e s and R i c h a r d s 1954; Welch 1966) t h i s i s n o t t h e case w i t h t h e s t u d y p o p u l a t i o n o f J . balticus. P o l l i n a t i o n a n e m o p h i l o u s and e n t o m o p h i l o u s , by Bombus terricola occidentalis Grne. 29 2 (Pojar 1973b). 5\/5 i n d i v i d u a l s s e l f - c o m p a t i b l e , but o u t c r o s s i n g favored by protogyny s e l f - i n c o m p a t i b i l i t y i n Juncaceae unknown to East (1940). Propagule a sporochore (small, very f i n e l y s t r i a t e - r e t i c u l a t e , unappendaged seeds); d i s p e r s a l anemochorous and epizoochorous (mud (Ridley 1930; P i j l 1969)). Ecotypic v a r i a t i o n probable (species has broad d i s t r i b u t i o n , with montane, l i t t o r a l , and c o a s t a l r a c e s ) ; Waisel (1972) mentions that h a l o p h y t i c ecotypes are common in.Juncus. H y b r i d i z a t i o n probabli (see taxonomic treatments i n Calder and Taylor 1968; Hulten 1968; Hitchcock et al. 1969). D i s t r i b u t i o n : \"Wet places, o f t e n where s a l i n e or a l k a l i n e , ' over much of \u2022 temperate and s u b a r c t i c N. Am. and E u r a s i a ; S. Am.\" (Hitchcock et al. 1969). Lilaeopsis occidentalis Coult. \u00a3 Rose. Small, semi-aquatic, rhizomatous p e r e n n i a l ; hemicryptophyte; vegetative reproduction extensive ( c f . Stevenson 1947). Flowers pleomorphic; p e r f e c t , homogamous or s l i g h t l y protandrous, minute, greenish-yellow, odorless, stylopodium minutely beaded with nectar; blooming (1971) J u l y 9-15 to 25-30. P o l l i n a t i o n probably ( i n f r e q u e n t l y ) by small muscid f l i e s ; s e l f - p o l l i n a t i o n p o s s i b l e but u n l i k e l y because of herkogamy; hydrophily by high t i d e s a l s o a p o s s i b i l i t y . 3\/3 i n d i v i d u a l s self-compatible. Propagule a megasclerochore (corky-thickened, ribbed, glabrous, ovoid f r u i t ) ; d i s p e r s a l hydrochorous and epizoochorous (mud). E c o t y p i c v a r i a t i o n p o s s i b l e ( l a t i t u d i n a l ? ) ; c f . H i l l (1927). Apparently no h y b r i d i z a t i o n . D i s t r i b u t i o n : \"Marshes, sa l t , f l a t s , and sandy or muddy beaches and shores along and near the coast, ...s. Van. I. to c. C a l i f . \" (Hitchcock et al. 1961). Note: see B e l l (19 71) f o r a d i s c u s s i o n of breeding systems i n Umbelliferae. Plantago maritipia L. Scapose perennial from a stout taproot often surmounted by an erect caudex; hemicryptophyte; vegetative reproduction none. Flowers stereomorphic; p e r f e c t , strongly protogynous, borne i n a dense, a c r o p e t a l l y - f l o w e r i n g s p i k e , odorless, n e c t a r l e s s ; gamopetalous c o r o l l a s reduced, inconspicuous, s c a r i o u s ; anthers and elongate stigmas well-exserted during anthesis; blooming (1971) June 10-25 t o July 20-Aug. 5. P o l l i n a t i o n anemophilous and entomophilous, by Bombus terricola occidentalis Grne. (Pojar 1973b);' i n s e c t v i s i t o r s t o species of Plantago have a l s o been noted by C l i f f o r d (1962), Sagar and Harper (1964), and F a e g r i and van der P i j l (1971). 5\/5 i n d i v i d u a l s self-compatible, but strong protogyny and acropetal flowering favor outcrossing; Gregor (1939) noted that a l l N. Am. pop-u l a t i o n s were self-compatible, whereas the European were s e l f - i n c o m p a t i b l e ; East (1940) predicted s e l f - c o m p a t i b i l i t y f o r the Plantaginaceae; Sagar and Harper (1964) found s e l f - c o m p a t i b i l i t y i n P. major, media, and lanceolata. Propagule a sporochore and desmochore (smooth seeds that become mucilaginous upon wetting (Ridley 1930; P i j l 1969)); d i s p e r s a l anemochorous and epizoochorous (mud and mucilage). E c o t y p i c 293 v a r i a t i o n c l i n a l , w e l l - s t u d i e d (Gregor 1939; Gregor and Watson 1961). H y b r i d i z a t i o n not reported. D i s t r i b u t i o n : \"A widespread and v a r i a b l e maritime sp e c i e s . . . \" (Hitchcock et al. 1959); also i n S. Am. (Hulten 1968). Potentilla pacifica Howell. Strawberrylike p e r e n n i a l ; hemicryptophyte; vegetative reproduction extensive by long, p r o s t r a t e f r e e l y r o o t i n g stolons. Flowers pleomorphic; p e r f e c t , weakly protogynous, s l i g h t l y o d o r i f e r o u s , moderately n e c t a r i f e r o u s ; c o r o l l a bowl-shaped, b r i g h t yellow; e a r l y i n a n t h e s i s , the r i n g of stamens i s . f l u s h with the c o r o l l a , but as flowering progresses, the stamens arch up over the stigmas, making s e l f - p o l l i n a t i o n p o s s i b l e ; blooming (1971) June 13-20 to J u l y 3-20. P o l l i -nation by s e l f i n g , and entomophily (bumble bees, muscid and syrphid f l i e s , and bee f l i e s ) ; Clapham, T u t i n and Warburg (1962) (re P. anserina): \" V i s i t e d by various i n s e c t s , s e l f - p o l l i n a t i o n p o s s i b l e \" . Weakly self-compatible (3-5 r i p e achenes formed on 4\/7 s e l f e d f l o w e r s ) ; Rousi (1965) obtained s i m i l a r r e s u l t s ; Clausen and Hiesey (1958) noted that some species of Potentilla contain both self-compatible and s e l f - i n c o m p a t i b l e races. Propagule a megasclerochore; d i s p e r s a l epizoochorous (mud) and hydrochorous; buoyancy \u00b0f seeds noted by Rid l e y (1930), and Rousi (1965) stated that the \"hard wall of the achenes ...has a l a y e r of a i r - c o n t a i n i n g c e l l s enabling them to f l o a t f o r long periods i n water\". Ecotypic v a r i a t i o n l i k e l y , l a t i t u d i n a l and perhaps maritime vs. i n l a n d . H y b r i d i z a t i o n probable, at l e a s t with segregates of P. anserina sensu lato. D i s t r i b u t i o n : \"Coastal dunes, beaches, sand f l a t s , and marsh edges and stream banks; Ala's, to s. C a l i f . , perhaps very o c c a s i o n a l l y i n l a n d \" (Hitchcock et al. 1961). Puccinellia pumila (Vasey) A.S. Hitchcock. Cespitose p e r e n n i a l ; hemicryptophyte; vegetative reproduction none. Flowers amorphic; p e r f e c t , homogamous, reduced, inconspicuous, o d o r l e s s , n e c t a r l e s s ; p a r t i a l l y cleistogamous (anthers only part-i a l l y exserted at a n t h e s i s ) ; blooming (1971) June 13-25 to J u l y 20-24. 4\/4 i n d i v i d u a l s self-compatible. P o l l i n a t i o n by s e l f i n g and wind. Propagule a micro-sclerochore; d i s p e r s a l anemochorous, hydrochorous, and epizoochorous (mud). Eco-t y p i c v a r i a t i o n probable, l a t i t u d i n a l . H y b r i d i z a t i o n with c l o s e l y r e l a t e d species of the north P a c i f i c coast also probable ( c f . Calder and Taylor 1968; Hitchcock et al. 1969). D i s t r i b u t i o n : \" T i d e f l a t s and s a l t marshes; Alas, southward t o Mendocino Co., C a l i f . ; Lab., Newf., Que., to Conn.\" (Hitchcock et al. 1969). 294 Salicornia virginica L. M a t t e d , s u c c u l e n t p e r e n n i a l f r o m an e x t r e m e l y d e n s e , woody, p e r e n n i a l s t o c k capped d u r i n g t h e g r o w i n g s e a s o n by a d d i t i o n a l a n n u a l g r o w t h (Cameron 1 9 7 2 ) ; chamaephyte; moderate v e g e t a t i v e r e p r o d u c t i o n - b y f r e e l y b r a n c h i n g r h i z o m e s and p r o s t r a t e a e r i a l stems ( s t o l o n s ) . F l o w e r s a c t i n o m o r p h i c ; p e r f e c t o r i n p a r t i m p e r f e c t , s t r o n g l y p r o t o g y n o u s , h i g h l y r e d u c e d , . o d o r l e s s , n e c t a r l e s s , s e s s i l e and sunk i n d e p r e s s i o n s o f t h e f l e s h y , a c r o p e t a l l y - f l o w e r i n g s p i k e ; a t a n t h e s i s , f i r s t t h e s t y l e s and t h e n t h e a n t h e r s a r e w e l l - e x s e r t e d ; b l o o m i n g (1971) J u l y 20-30 t o Aug. 25-Sept. 5. P o l l i n a t i o n a n e m o p h i l o u s and e n t o m o p h i l o u s , by Bombus terricola occidentalis Grne. ( P o j a r 1 9 7 3 b ) ; D a l b y (1962) f o u n d t h e B r i t i s h a n n u a l s p e c i e s t o be h a b i t u a l l y s e l f i n g , and b o t h D a l b y (1962) and F e r g u s o n (1964) n o t e d a t r e n d t o w a r d homogamy i n t h e a n n u a l s . 3\/3 i n d i v i d u a l s s e l f - c o m p a t i b l e ; n e v e r t h e l e s s , o u t c r o s s i n g f a v o r e d by s t r o n g p r o t o g y n y and a c r o p e t a l f l o w e r i n g o f t h e s p i k e . P r o p a g u l e a desmochore ( p u b e r u l e n t s e e d ) and c y c l o c h o r e ( i n d e h i s c e n t u t r i c l e + spongy, a c c r e s c e n t p e r i a n t h , o r an e n t i r e stem segment t h a t c a n f l o a t i n s e a w a t e r up t o 3 months ( D a l b y 1 9 6 3 ) ) ; d i s p e r s a l e p i z o o c h o r o u s (mud and s t i c k - -t i g h t ) and h y d r o c h o r o u s v i a t h e c y c l o c h o r e and f l o a t i n g s e e d l i n g s ( R i d l e y 1 930; P i j l 1 9 6 9 ) . L a t i t u d i n a l e c o t y p i c v a r i a t i o n p o s s i b l e . A p p a r e n t l y no h y b r i d i z a t i o n . D i s t r i b u t i o n : \" S a l t marshes and b e a c h e s a l o n g t h e c o a s t ; A l a s , t o B a j a C a l i f . , A t l a n t i c c o a s t f r o m Mass. t o S.C., and a l o n g t h e G u l f C o a s t ; w. E u r o p e and t h e M e d i t e r r a n e a n r e g i o n \" ( H i t c h c o c k e t al. 1 9 6 4 ) . Comments: S a l i s b u r y ( 1 9 4 2 ) . n o t e d t h e h i g h f e c u n d i t y o f 2 B r i t i s h s p e c i e s o f Salicornia; D a l b y (1962) m e n t i o n e d t h a t t h e r e i s seed d i m o r p h i s m i n Salicornia i n t h a t t h e c e n t r a l f l o w e r o f e a c h cymule a l w a y s has a l a r g e r seed t h a n e i t h e r o f t h e l a t e r a l s . Scirpus cernuus V a h l . D i m u n i t i v e t u f t e d a n n u a l o r s h o r t - l i v e d p e r e n n i a l ; t h e r o -p h y t e o r h e m i c r y p t o p h y t e ; v e g e t a t i v e r e p r o d u c t i o n none. F l o w e r s a m o r p h i c ; p e r f e c t , m o d e r a t e l y p r o t o g y n o u s , r e d u c e d , i n c o n s p i c u o u s , o d o r l e s s , n e c t a r l e s s ; f l o w e r i n g p r o c e e d s a c r o p e t a l l y a l o n g t h e s o l i t a r y s p i k e l e t ; b l o o m i n g (1971) J u l y 1-5 t o 20-25. 5\/5 i n d i v i d u a l s s e l f - c o m p a t i b l e . P o l l i n a t i o n a n e m o p h i l o u s and by s e l f i n g , o f w h i c h t h e r e i s p r o b a b l y a f a i r amount\u2014 t h e p r o t o g y n y o f t h e f l o w e r s i s n o t c o m p l e t e , t h e y a r e s e l f - c o m p a t i b l e , and t h e p l a n t s o f t e n grow n e a r l y h i d d e n among t h e stems o f t h e l a r g e r s p e c i e s i n t h e marsh where t h e r e i s l i t t l e w i n d . P r o p a g u l e a m e g a s c l e r o c h o r e ; d i s p e r s a l h y d r o c h o r o u s ( a c h e n e s b u o y a n t f o r s h o r t d i s t a n c e s ( R i d l e y 19 30)) and e p i z o o c h o r o u s (mud). E c o t y p i c v a r i a t i o n p r o b a b l e , i n v i e w o f wide r a n g e . No h y b r i d i z a t i o n a p p a r e n t . D i s t r i b u t i o n : \" F r e s h t o more o f t e n b r a c k i s h o r s a l i n e m a r s h e s , b e a c h e s , a n d . s h o r e s ; n e a r t h e c o a s t (and a b o u t P u g e t Sound) f r o m s. B.C. t o n. Hex.; i r r e g u l a r l y s c a t t e r e d i n p a r t s o f S. Am., E u r o p e , A f r . , and Aus. and N.Z.\" ( H i t c h c o c k et al. 1 9 6 9 ) . 2 9 5 Spergularia canadensis (Pers.) G. Don. Small, glandular-pubescent annual; thero-phyte; vegetative reproduction none. Flowers pleomorphic; p e r f e c t , homogamous, small, o d o r l e s s , s l i g h t l y n e c t a r i f e r o u s ; p e t a l s pink; plants o f t e n bear both chasmogamous and cleistogamous flowers, the chasmogamous flowers when present are borne uppermost on the p l a n t ; blooming (1971) J u l y 3-10 to J u l y 20-Aug. 5. 7\/7 i n d i v i d u a l s f u l l y s elf-compatible; B r i t i s h species of Spergularia, i n c l u d i n g the c l o s e l y a l l i e d S. marina, are a l s o self-compatible (Clapham, T u t i n and Warburg 1962); East (1940) stated that no self-compatible species were known from the Caryophyllaceae. P o l l i n a t i o n p r i m a r i l y by s e l f i n g , although the chasmogamous flowers may be outcrossed by small muscid f l i e s . Propagule a sporochore and desmochore (small, glandular-pubescent, s l i g h t l y winged seeds) ; d i s p e r s a l anemo-. chorous, hydrochorous, and epizoochorous (mud). Ecotypic v a r i a t i o n probable; l a t i t u d i n a l (Rossbach 1940 recognized, northern and southern p o p u l a t i o n s ) . Hybrid-i z a t i o n u n l i k e l y , as Ratter (1964) found a l l attempts at i n t e r s p e c i f i c h y b r i d -i z a t i o n with S. marina blocked by F-L hybrid s t e r i l i t y . D i s t r i b u t i o n : \"Coastal t i d e l a n d s and mudflats; A l a s , to n. C a l i f , and'-from N.Y. to Newf.\" (Hitchcock et al . 1964) .. Stellaria humifusa Rottb. Spreading, somewhat f l e s h y p e r e n n i a l herb; chamaephyte; vegetative reproduction marked, by decumbent, nodally r o o t i n g stems ( s t o l o n s ) . Flowers pleomorphic; p e r f e c t , weakly protandrous, moderately n e c t a r i f e r o u s at the base of the stamens, with a d i s t i n c t , sweet odor; c o r o l l a small, white; anthers and stigmas remote at a n t h e s i s , so s e l f - p o l l i n a t i o n u n l i k e l y ; flowering (1971) May 31-June 13 to J u l y 20-Aug. 10. P o l l i n a t i o n by'small muscid f l i e s (personal observations and Knuth 1906-1909), and probably a small amount of s e l f i n g . 5\/5 i n d i v i d u a l s self-compatible (according to East (1940), s e l f - c o m p a t i b i l i t y was unknown i n the Caryophyllaceae). Propagule a megasclerochore; d i s p e r s a l anemo-chorous and epizoochorous (mud) (Ridley 1930). Ecotypic v a r i a t i o n probably e x i s t s i n t h i s species, although there i s great phenotypic p l a s t i c i t y (Calder \u00a3 Taylor 1968). H y b r i d i z a t i o n improbable. D i s t r i b u t i o n : \"Circumpolar; A l a s , to Lab. and s. along the w. coast, to a few known l o c a l i t i e s i n Wash, and Oreg.\" (Hitchcock et al. 1964). Trifolium wormskjoldii Lehm. Taprooted per e n n i a l with decumbent-based and o f t e n rhizomatous stems; hemicryptophyte. Flowers zygomorphic; i r r e g u l a r , t u b u l a r flowers aggregated into a showy head; flowers p e r f e c t , s t r o n g l y protandrous (the anthers dehisce i n the bud well before anthesis) , stigma extends s l i g h t l y beyond anthers; sweetly smelling of c l o v e r , strongly n e c t a r i f e r o u s ; p e t a l s deep pink to .296 r e d p u r p l e ; b l o o m i n g (1971) June 13-20 t o J u l y 20-31. P o l l i n a t i o n e n t o m o p h i l o u s , by s p e c i e s o f Bombus. The bumble bees o p e r a t e t h e v a l v u l a r mechanism o f t h e f l o w e r , whereby .the s t i g m a and a n t h e r s p r o t r u d e f r o m t h e c a r i n a so l o n g a s t h e p r e s s u r e o f t h e v i s i t o r i s exserted on t h e s t a n d a r d and w i n g s . P o l l e n i s t h u s d e p o s i t e d on t h e u n d e r s i d e o f t h e v i s i t i n g bumble bee's head ( K n u t h 1906-1909; F r e e 1 9 7 0 ) . 5\/5 i n d i v i d u a l s s e l f - i n c o m p a t i b l e . P r o p a g u l e a b a l l o c h o r e and desmo-c h o r e ; d i s p e r s a l a u t o c h o r o u s ( e x p l o s i v e d e h i s c e n c e o f l e g u m e ) , h y d r o c h o r o u s ( b u o y a n t l e g u m e ? ) , and e p i z o o c h o r o u s ( s t i c k t i g h t , by t h e s p i n y c a l y x ) . E c o t y p i c v a r i a t i o n p r o b a b l e ( c o n s i d e r d i s t r i b u t i o n ) . A p p a r e n t l y no h y b r i d i z a t i o n . D i s t r i -b u t i o n : \"Meadows and s t r e a m banks f r o m B.C. t o C a l i f , and Mex., e., a t e l e v a t i o n s up t o 8000 f t , t o I d a . , U t a h , C o l o . , and N.M.\" ( H i t c h c o c k e t al. 1 9 6 1 ) . Comments: T. wormskjoldii has n i t r o g e n - f i x i n g r o o t n o d u l e s . I n two summers I have o b s e r v e d no seed s e t i n t h i s s a l t marsh p o p u l a t i o n o f T. wormskjoldii. P r e s u m a b l y t h e s p e c i e s p e r s i s t s i n t h e marsh t h r o u g h i t s v e g e t a t i v e r e p r o d u c t i o n . However, i n v i e w o f t h e s e l f - i n c o m p a t i b i l i t y , f l o r a l b i o l o g y , and bumble bee p o l l i n a t i o n , when t h e s p e c i e s does s e t seed i t i s l i k e l y t h e r e s u l t o f o u t c r o s s i n g , a l t h o u g h i t c o u l d be due t o i n f r e q u e n t breakdown o f t h e i n c o m p a t i b i l i t y s y s t e m . Triglochin maritimum L. More o r l e s s s c a p o s e p e r e n n i a l w i t h h e a v y , o f t e n woody r h i z o m e s ; h e m i c r y p t o p h y t e ; v e g e t a t i v e r e p r o d u c t i o n more e x t e n s i v e i n t h e s a l t marsh meadow t h a n on t h e mud f l a t s , where t h e s p e c i e s t e n d s t o f o r m l a r g e d e n s e c l u m p s . F l o w e r s a c t i n o m o r p h i c ; p e r f e c t , s t r o n g l y p r o t o g y n o u s , i n c o n s p i c u o u s , o d o r l e s s , n e c t a r l e s s ; s t i g m a s l a v e n d e r , b r u s h l i k e ; a n t h e r s n e a r l y s e s s i l e i n a x i l s o f p e r i a n t h segments; f l o w e r s s e t i n a c l o s e l y f l o w e r e d raceme t h a t f l o w e r s a c r o p e t a l l y and u l t i m a t e l y e l o n g a t e s c o n s i d e r a b l y i n f r u i t ; b l o o m i n g (1971) A p r i l 15-20 t o June 1 5 - J u l y 20. P o l l i n a t i o n a n e m o p h i l o u s ( p e r s o n a l o b s e r v a t i o n s ; Clapham, T u t i n and Warburg 1962; K u g l e r 1970; F a e g r i and van d e r P i j l 1 9 7 1 ) ; a f t e r a n t h e r d e h i s c e n c e , t h e c o p i o u s d r y p o l l e n i s h e l d between t h e p e r i a n t h segments u n t i l t h e w i n d i s s t r o n g enough t o b l o w (and s h a k e ) i t o u t ( K u g l e r 197 0; F a e g r i and van d e r P i j l 1 9 7 1 ) . 7\/7 i n d i v i d u a l s s e l f - c o m p a t i b l e ; autogamy p r o b a b l y l i m i t e d by s t r o n g p r o t o g y n y and a c r o p e t a l f l o w e r i n g o f t h e raceme. P r o p a g u l e a s p o r o c h o r e ( s m a l l , n a r r o w l y f u s i f o r m s e e d ) ; d i s p e r s a l anemochorous, e p i z o o c h o r o u s (mud), and h y d r o c h o r o u s v i a f l o a t i n g s e e d l i n g s ( R i d l e y 1 9 3 0 ) . E c o t y p i c v a r i a t i o n c o u l d be i n f e r r e d f r o m wide r a n g e and e x i s t e n c e o f chromosomal r a c e s ( L o v e and Love 1958a 6 b ) . A p p a r e n t l y no h y b r i d i z a t i o n . D i s t r i b u t i o n : \"Marshy t o m o i s t and g r a v e l l y a r e a s , m o s t l y where s a l i n e o r a l k a l i n e ; Alas', t o L a b . , s. t o B a j a C a l i f . , Mex., Neb., I a . , 1 1 1 . , I n d . , and P a . , S. Am., E u r a s i a \" ( H i t c h c o c k e t a l . - 1 9 6 9 ) . 297 B. Sphagnum Bogs Agrostis aequivalvis ( T r i n . ) T r i n . P e r e nnial grass, forming small clumps, though moderately rhizomatous. Flowers amorphic; p e r f e c t , s l i g h t l y protandrous, reduced, inconspicuous, o d o r l e s s , n e c t a r l e s s ; blooming (1971) June 10-14 to J u l y 14-19 (Wade's Bog), June 9-14 to June 29-July 4 (Ogg's Bog). 7\/7 i n d i v i d u a l s at l e a s t p a r t i a l l y s e l f-compatible. P o l l i n a t i o n anemophilous. Propagule a microsclerochore; d i s p e r s a l anemochorous and epizoochorous (mud). Ec o t y p i c v a r i a t i o n u n l i k e l y . H y b r i d i z a t i o n may occur with A. thurheviana. D i s t r i b u t i o n : \"In bogs and wet places near the coast; Alaska southward to the south end of Van. I. and the n. end of the Olympic Peninsula\" (Hitchcock et al. 1969). Apargidium boreale (Bong.) T. S G. Usually scapose, +\/- taprooted, p e r e n n i a l herb; vegetative reproduction none. Flowers zygomorphic, c a p i t u l a c a p i t a t e ; flowers a l l l i g u l a t e and p e r f e c t s t r o n g l y protandrous, b r i g h t yellow, with a mild, t e a - l i k e fragrance, moderately n e c t a r i f e r o u s . The f l o r a l biology of Compositae i s w e l l known, and a b r i e f d e s c r i p t i o n here w i l l s u f f i c e f o r the other asteraceous species i n t h i s study. In the markedly protandrous, hermaphrod-i t i c flowers p o l l e n i s discharged i n t o the anther c y l i n d e r that surrounds the s t y l e . As the s t y l e elongates i t pumps the p o l l e n out of the c y l i n d e r , sweeping the grains along with the help of h a i r s or p a p i l l a e on the s t y l e surface. L a t e r , the s t y l a r branches diverge, then r e f l e x , and the now-receptive stigmas can pick up p o l l e n s t i l l adhering to the sweeping'hairs. Nectar i s secreted by a r i d g e surrounding the base of the s t y l e , and i s so abundant as to r i s e i n the c o r o l l a tube. I t i s thus a c c e s s i b l e to both long- and short-tongued i n s e c t s ( a f t e r Knuth 1906-1909). Flowers blooming (1971) May 30-June 8 to J u l y 10-30 (Wade's Bog), May 18-31 to June 29-July 19 (Ogg's Bog). P o l l i n a t i o n entomophilous, p r i m a r i l y by syrphid f l i e s and Bombus spp. 7\/7 i n d i v i d u a l s s e l f - i n c o m p a t i b l e . Propagule a pogonochore (pappose achene); d i s p e r s a l anemochorous. L a t i t u d i n a l ecotypic v a r i a t i o n p o s s i b l e . H y b r i d i z a t i o n apparently none. D i s t r i b u t i o n : \"Sphagnum bogs and wet meadows i n the mts.; coast of s Alas, (beyond the panhandle), s. to n.w. C a l i f . , not occurring east of the Cascade range\" (Hitchcock et al. 195S). -2 9 8 Calamagrostis nutkaensis ( P r e s l . ) S t e u d . S t r o n g l y t u f t e d p e r e n n i a l g r a s s w i t h s h o r t r h i z o m e s . F l o w e r s a m o r p h i c ; p e r f e c t , m o d e r a t e l y p r o t a n d r o u s , r e d u c e d , i n c o n s p i c u o u s , o d o r l e s s , n e c t a r l e s s ; b l o o m i n g (1971) J u l y 28-Aug. 3 t o Aug. 10-14. P o l l i n a t i o n a n e m o p h i l o u s . 3\/3 i n d i v i d u a l s s e l f - i n c o m p a t i b l e ; p o l l e n good, m e i o s i s r e g u l a r , no i n d i c a t i o n o f agamospermy, w h i c h a g r e e s w i t h N y g r e n (19 5 4 ) . P r o p a g u l e a m e g a s c l e r o c h o r e and desmochore (awned f r o m t h e lemma and b e a r d e d f r o m t h e c a l l u s ) ; d i s p e r s a l anemochorous and e p i z o o c h o r o u s ( s t i c k t i g h t ) . L a t i t u d i n a l e c o t y p i c v a r i a t i o n p o s s i b l e . H y b r i d i z a t i o n w i t h C. canadensis i n f e r r e d by \u2022 H u l t e n (1968) b u t by C a l d e r and T a y l o r ( 1 9 6 8 ) . D i s t r i b u t i o n : \" C o a s t a l , o f t e n i n sand d u n e s ; A l a s , t o San L u i s O b i s p o Co., C a l i f . , f r o m a l o n g t h e b e a c h e s i n t o t h e a d j a c e n t low m o u n t a i n s \" ( H i t c h c o c k et al. 1 9 6 9 ) . Cavex oanesoens L. P e r e n n i a l sedge w i t h l o o s e l y t o d e n s e l y c l u s t e r e d c u l m s and s h o r t r h i z o m e s . F l o w e r s a m o r p h i c ; u n i s e x u a l , a g g r e g a t e d i n t o 4-8 s e s s i l e , g y n a e c a n d r o u s s p i k e s t h a t a r e s t r o n g l y p r o t a n d r o u s , f I s . . r e d u c e d , i n c o n s p i c u o u s , o d o r l e s s , n e c t a r l e s s ; p l a n t s m o n o e c i o u s , f l o w e r i n g (1971) June 1-4 t o J u n e 11-14 (Wade's B o g ) , May 22-27 t o June 2-8 (Ogg's Bog ) . P o l l i n a t i o n a n e m o p h i l o u s . 5\/5 i n d i v i d u a l s s e l f - c o m p a t i b l e . P r o p a g u l e a m i c r o s e l e r o c h o r e and desmochore (achene + s h o r t - b e a k e d p e r i g y n i u m ) ; d i s p e r s a l anemochorous, h y d r o c h o r o u s , and e p i z o o c h o r o u s (mud and s t i c k t i g h t ) . E c o t y p i c v a r i a t i o n h i g h l y p r o b a b l e ( s e e t a x o n o m i c d i s c u s s i o n i n C a l d e r and T a y l o r 1 9 6 8 ) . H y b r i d i z a t i o n ? D i s t r i b u t i o n : \"Wet meadows and o t h e r wet p l a c e s , sometimes i n sphagnum b o g s , f r o m t h e l o w l a n d s t o n e a r t i m b e r l i n e i n t h e m t s . ; c i r c u m b o r e a l , e x t e n d i n g s. t o Va., A r i z . , and C a l i f . \" ( H i t c h c o c k et al. 1 9 6 9 ) . Carex obnupta L.H. B a i l e y . D e n s e l y t u f t e d , p e r e n n i a l sedge w i t h e x t e n s i v e v e g e t a t i v e r e p r o d u c t i o n v i a l o n g , c r e e p i n g r h i z o m e s . F l o w e r s a m o r p h i c ; u n i s e x u a l , r e d u c e d , i n c o n s p i c u o u s , o d o r l e s s , n e c t a r l e s s ; p l a n t s m o n o e c i o u s , u p p e r 1-3 s p i k e s s t a m i n a t e , t h e o t h e r s p i s t i l l a t e o r a n d r o g y n o u s ; as i n C. lyngbyei, i n d i v i d u a l s i n t h e same p o p u l a t i o n can be e i t h e r s t r o n g l y p r o t a n d r o u s , s t r o n g l y p r o t o g y n o u s o r +\/-homogamous. P l a n t s f l o w e r i n g (1971) A p r i l 8-12 t o May-28-June 4 (Wade's Bog) May 6-19 t o June 1-7 (Ogg's Bog ) . P o l l i n a t i o n a n e m o p h i l o u s . 7\/7 i n d i v i d u a l s s e l f - c o m p a t i b l e . P r o p a g u l e a m e g a s c l e r o c h o r e (achene + t o u g h , c o r k y - t h i c k e n e d p e r i g y n i u m ) ; d i s p e r s a l h y d r o c h o r o u s and e p i z o o c h o r o u s (mud). E c o t y p i c v a r i a t i o n 299 u n l i k e l y . H y b r i d i z a t i o n may occur w i t h C. lyngbyei. D i s t r i b u t i o n : \"In water or very wet ground, e s p e c i a l l y along r i v e r s or i n c o a s t a l swamps, from the Cascade Hts. to the c o a s t ; s. B.C. (Van. I.) t o C a l i f . \" (Hitchcock et al. 1969). Carex p a u a i f l o r a L i g h t f . P e r e n n i a l sedge, stems a r i s i n g s i n g l y or few t o g e t h e r from rhizomes. Flowers amorphic; u n i s e x u a l , reduced, i n c o n s p i c u o u s , o d o r l e s s , n e c t a r l e s s ; p l a n t s monoecious; spikes s h o r t , s o l i t a r y , androgynous, moderately protandrous; f l o w e r i n g (1971) May 29-June 3 to June 10-15 (Ogg's Bog). P o l l i n a t i o n anemophilous. 3\/3 i n d i v i d u a l s s e l f - c o m p a t i b l e . Propagule a m i c r o s c l e r o c h o r e and desmochore (achene + beaked perigynium); d i s p e r s a l anemochorous, hydrochorous, and epizoochorous (mud, s t i c k t i g h t ) . E c o t y p i c v a r i a t i o n probable. H y b r i d i z a t i o n not apparent. D i s t r i b u t i o n : \"Sphagnum bogs; c i r c u m b o r e a l , extending s. near the coast i n w. N. Am. as f a r as Van. I . and Whatcom Co., Wash., and i n e. N. Am. t o the New England S t a t e s \" (Hitchcock et al. 1969). Carex p l u r i f l o r a ' H u l t e n . P e r e n n i a l sedge w i t h stems a r i s i n g s i n g l y or few t o g e t h e r from l o n g , creeping rhizomes. Flowers amorphic; u n i s e x u a l , reduced, inconspicuous, o d o r l e s s , n e c t a r l e s s ; p l a n t s monoecious, s t r o n g l y protogynous', t e r m i n a l spike staminate, p i s t i l l a t e s p i k e s l a t e r a l , 1-3; p l a n t s f l o w e r i n g (197.1) May 3-7 t o May 20-30 (Wade's Bog), May 2-8 t o May 25-30 (Ogg's Bog). P o l l i n a t i o n . ; : anemophilous. 7\/7 i n d i v i d u a l s s e l f - c o m p a t i b l e . Propagule a megasclerochore (achene + densely p a p i l l a t e perigynium); d i s p e r s a l hydrochorous and epizoochorous (mud). E c o t y p i c v a r i a t i o n u n l i k e l y . H y b r i d i z a t i o n w i t h C. r a v i f l o r a considered probable by Calder and T a y l o r (1968). D i s t r i b u t i o n : \"Marshes, streambanks, and boggy shores near the c o a s t , sometimes under the i n f l u e n c e of s a l t spray; n.w. Wash, n to s. A l a s . , and w. through the A l e u t i a n I s l a n d s t o the coast of S i b e r i a and K u r i l e I . \" (Hitchcock et al. 1969). Coptis a s p l e n i f o l i a S a l i s b . Scapose, evergreen, p e r e n n i a l herb, w i t h e x t e n s i v e v e g e t a t i v e r e p r o d u c t i o n v i a s l e n d e r , y e l l o w i s h rhizomes. Flowers act i n o m o r p h i c ; p e r f e c t or staminate, weakly protogynous, o d o r l e s s , s t r o n g l y n e c t a r i f e r o u s . Sepals 5-8, f i l i f o r m - l a n c e o l a t e , g r e e n i s h - w h i t e , p e t a l o i d , deciduous; p e t a l s 5-7, s l i g h t l y s h o r t e r than s e p a l s , f i l i f o r m - c l a w e d , blade narrowly l i g u l a t e above the broadened, nectar gland-bearing base. At a n t h e s i s , the anthers are at about the same l e v e l as the stigmas and some dehisce c l o s e t o the stigmas, so t h a t s e l f i n g i s p o s s i b l e . P l a n t s f l o w e r i n g (1971) A p r i l 25-28 to May 5-9 (Ogg's Bog). P o l l i -n a t i o n entomophilous, by muscid and ( o c c a s i o n a l l y ) s y r p h i d f l i e s . 3\/3 i n d i v i d u a l s 300 s e l f - c o m p a t i b l e . P r o p a g u l e a m e g a s c l e r o c h o r e ; d i s p e r s a l h y d r o c h o r o u s ( s p l a s h cup) and e p i z o o c h o r o u s (mud). A l t i t u d i n a l e c o t y p i c v a r i a t i o n p o s s i b l e . A p p a r e n t l y no h y b r i d i z a t i o n . D i s t r i b u t i o n : \" M o i s t woods and b o g s ; A l a s , t o Van. I . and s. B.C. on t h e w. s i d e o f t h e Cascade M t s . , n o t q u i t e r e a c h i n g Wash.\" ( H i t c h c o c k et al. 1964) Coptis t r i f o l i a ( L . ) S a l i s b . S c a p o s e , p e r e n n i a l , e v e r g r e e n h e r b w i t h m o d e r a t e v e g e t a t i v e r e p r o d u c t i o n v i a s l e n d e r y e l l o w i s h r h i z o m e s . F l o w e r s a c t i n o m o r p h i c ; p e r f e c t o r s t a m i n a t e , w e a k l y p r o t o g y n o u s , s t r o n g l y n e c t a r i f e r o u s , o d o r l e s s . S e p a l s w h i t e , p e t a l o i d , n a r r o w l y l a n c e o l a t e t o s p a t u l a t e ; ' p e t a l s a b o u t h a l f t h e l e n g t h o f t h e s e p a l s , f l e s h y , h o l l o w e d and n e c t a r i f e r o u s a t t h e t i p s . A t a n t h e s i s , t h e s t i g m a s p r o j e c t a b o u t 2 mm beyond t h e s t a m e n s ; a u t o m a t i c s e l f - p o l l i n a t i o n t h u s u n l i k e l y . P l a n t s f l o w e r i n g (1971) May .3-6 t o May 16-20 (Wade's B o g ) , May 2-6 t o 14-19 (Ogg's Bog) . P o l l i n a t i o n e n t o m o p h i l o u s , by m u s c i d f l i e s and o c c a s i o n a l l y s y r p h i d f l i e s . 5\/5 i n d i v i d u a l s a t l e a s t p a r t i a l l y s e l f - c o m p a t i b l e . P r o p a g u l e a m i c r o s c l e r o c h o r e ; d i s p e r s a l anemochorous, h y d r o c h o r o u s ( s p l a s h c u p ? ) , and e p i -z o o c h o r o u s (mud). E c o t y p i c v a r i a t i o n p r o b a b l e ( s e e t a x o n o m i c d i s c u s s i o n i n C a l d e r and T a y l o r 1 9 6 8 ) . A p p a r e n t l y no h y b r i d i z a t i o n . D i s t r i b u t i o n : \"Muskegs t o deep woods; A l a s , s . , a t s c a t t e r e d s t a t i o n s , t o Van. I . and s. B.C., w. t o J a p a n and S i b e r i a \" \" ( H i t c h c o c k et al. 1 9 6 4 ) . h e r b s . F l o w e r s p l e o m o r p h i c ; s e m i c a p i t a t e , racemose i n f l o r e s c e n c e w i t h i t s f o u r p e t a l o i d b r a c t s i s a l s o a p l e o m o r p h i c f l o r a l u n i t ; f I s . p e r f e c t , s l i g h t l y p r o t a n d r o u s f a i n t l y f r a g r a n t , m o d e r a t e l y n e c t a r i f e r o u s . B r a c t s w h i t e , p e t a l s s m a l l , g r e e n i s h -w h i t e w i t h a deep p u r p l i s h t i n g e . N e c t a r s e c r e t e d by a f l e s h y r i n g s u r r o u n d i n g t h e s t y l e . P l a n t s f l o w e r i n g (1971) June 19-29 t o J u l y 9-19 (Ogg's B o g ) . P o l l i n a t i o n e n t o m o p h i l o u s , by s y r p h i d and m u s c i d f l i e s . 3\/3 i n d i v i d u a l s s e l f - i n c o m p a t i b l e . P r o p a g u l e a s a r c o c h o r e ( b r i g h t r e d b e r r y ) ; d i s p e r s a l e n d o z o o c h o r o u s ( R i d l e y 1930; P i j l 1 9 6 9 ) . A l t i t u d i n a l e c o t y p i c v a r i a t i o n p r o b a b l e . H y b r i d i z a t i o n w i t h C. canadensis and C. suecica i s v e r y p r o b a b l e . . D i s t r i b u t i o n : Sphagnum b o g s , wet f o r e s t s , s u b a l p i n e meadows, f r o m s e a l e v e l t o s u b a l p i n e a l o n g t h e c o a s t ; s.e. A l a s , t o Van. I . , ( t o n. C a l i f . ? ) ( a f t e r C a l d e r and T a y l o r 1965, 1 9 6 8 ) . Low, w i d e l y r h i z o m a t o u s h a l f - s h r u b s t o p e r e n n i a l 301 Drosera votundifolia L. Scapose p e r e n n i a l with r o s u l a t e , g l a n d u l a r - v i s c i d leaves; vegetative reproduction l i t t l e ( S a l i s b u r y (1915) reported v i v i p a r y by leaf-budding i n Drosera) or none. Flowers pleomorphic; p e r f e c t , homogamous, odor l e s s , weakly n e c t a r i f e r o u s ; petals white. Some flowers are cleistogamous; the chasmogamous flowers open f o r only a day or two, and at anthesis the 5 s t y l e s spread wide so that the stigmas touch the 5 anthers. Plants flowering (1971) J u l y 22-23 to J u l y 25-27 (Wade's Bog), J u l y 24-26 to J u l y 27-29 (Ogg's Bog). P o l l i n a t i o n predominantly by s e l f i n g , o c c a s i o n a l l y entomophilous by small muscid f l i e s . 7\/7 i n d i v i d u a l s f u l l y self-compatible. Propagule a sporochore and pterochore (seed + loose, f l a t t e n e d t e s t a ) ; d i s p e r s a l anemochorous, hydrochorous ( p o s s i b i l i t i e s discussed by Ridley 1930), and epizoochorous (mud). Ec o t y p i c v a r i a t i o n p o s s i b l e , c o n s i d e r i n g d i s t r i b u t i o n . H y b r i d i z a t i o n reported with D. anglioa and D. linearis by Wood (1955) and Hitchcock et al. (1962). D i s t r i b u t i o n : \"Swamps and bogs; Alas, to Lab., s. to C a l i f . , Nev., Ida., Mont., Minn., 111., A l a . , F l a . , Va., and N. Eng.; E u r a s i a \" (Hitchcock et al. 1962). Note: the plants are i n s e c t i v o r o u s . Empetrum nigrum L. Procumbent, spreading, evergreen shrub; o c c a s i o n a l vegetative reproduction by l a y e r i n g ( B e l l and T a l l i s 1973). Flowers actinpmorphic; u n i s e x u a l , reduced, inconspicuous, odorless, n e c t a r l e s s . Plants d i o e c i o u s ; male flowers with 3 well-exserted stamens, female with lobed, p e l t a t e stigma. Plants flowering (1971) A p r i l 10-15 to A p r i l 25-30 (Wade's Bog), A p r i l 7-11 to April'22-28 (Ogg's Bog). P o l l i n a t i o n anemophilous. C o m p a t i b i l i t y i r r e l e v a n t , as plants are dioecious. Propagule a sarcochore (black b e r r y ) ; d i s p e r s a l endozoochorous (Ridley 1930; B e l l and T a l l i s 1973). Ecotypic v a r i a t i o n h i g h l y probable; the species i s very v a r i a b l e and has a broad e c o l o g i c a l amplitude (see d i s c u s s i o n i n B e l l and T a l l i s 1973). No h y b r i d i z a t i o n reported. D i s t r i b u t i o n : \"More or l e s s circumpolar, extending southward, i n the Cascade Mts. and along the coast, to Del Norte Co., C a l i f . , mostly on exposed rocky b l u f f s but also i n peat bogs\" (Hitchcock et al. 1962). Note: the study populations were dioecious and d i p l o i d (n = 13), i n o p p o s i t i o n to the conclusions of Love and Lbve (1959). Winter flower buds have mature p o l l e n . Eriophorum polijstachion L. P e r e n n i a l , e x t e n s i v e l y c o l o n i a l from creeping rhizomes. Flowers amorphic; p e r f e c t , strongly protogynous, reduced, inconspicuous, o d o r l e s s , n e c t a r l e s s ; blooming (1971) A p r i l 27-May 2 to May 8-12 (Ogg's Bog). P o l l i n a t i o n anemophilous; s e l f i n g can occur, as the o l d e s t (erect) s p i k e l e t produces mature anthers before the younger (pendulous) s p i k e l e t s , so that the p o l l e n of the o l d e s t s p i k e l e t f a l l s onto the stigmas of the younger ones ( P h i l l i p s 1954). 2\/2 i n d i v i d u a l s 302 self-compatible. Propagule a pogonochore (achenes + numerous, persistent, white bristles); dispersal anemochorous (Ridley 1930). Ecotypic variation probable. Hybridization reported by Raymond (1957), who also maintained that hybridization is frequent within the genus. Distribution: \"Cold swamps and bogs at moderate to high elevations in the mts., commoner northward; circumboreal at high latitudes, extending s. in Am. to Mex,...N. Mex.,...and c. Oreg.\" (Hitchcock e t al. 1969). G a u l t h e r i a s h a l l o n Pursh. Erect, evergreen shrub, occasionally reproducing by layering. Flowers stereomorphic, urn-shaped; perfect, faintly protandrous, mildly fragrant, strongly nectariferous; stigma slightly exserted from the white corolla tube, anthers included; blooming (1971) June 12-16 to June 29-July 4 (Ogg's Bog). Pollination entomophilous, primarily by Bombus spp., frequently by syrphid flies, occasionally by small muscid flies, and quite often ornithophilous by Rufous hummingbirds, which are probably ineffective pollinators. 3\/3 individuals at least partially self-compatible. Propagule a sarcochore (capsular fruit, appearing berry-like by the pulpy thickening of the enclosing calyx tube) ; dispersal endo-zoochorous. Altitudinal ecotypic,variation possible, but not probable. Apparently-no hybridization. Distribution: \"Moist to dry woods, lowl. to lower mts.; B.C. south, from e. base of Cascades to coast, to s. Calif.\" (Hitchcock e t al. 1959). G e n t i a n a d o u g l a s i a n a Bong. Freely-branching, annual herb. Flowers stereomorphic; perfect, moderately protandrous, faintly fragrant, moderately nectariferous. Corolla white, lobes dotted with blue nectar guides. Stigmas on a level with, but not adjacent to, the stamens. Nectar secreted at the base of the ovary. Plants flowering (1971) May 15-June 24 to July 24-Aug. 8 (Wade's Bog), June 9-July 1 to July 30-Aug. 13. Pollination entomophilous, primarily by Bombus spp., also by syrphid flies. 5\/5 individuals self-compatible. Propagule a microsclerochore dispersal anemochorous and hydrochorous (splash-cup). Ecotypic variation unlikely. Apparently no hybridization. Distribution: \"Bogs and tundra from Alas. s. to Van. I. and...two localities... in Wash.\" (Hitchcock e t al. 1959). Gentiana sceptrum Griseb. Caespitose perennial herb, without vegetative reproduction Flowers stereomorphic; perfect, strongly protandrous, very aromatic, smelling of vanilla or coumarin, strongly nectariferous; corolla deep-tubular, blue (often streaked or mottled with green); stigmas borne slightly above level of the anthers; nectar secreted at the base of the ovary. Pollination entomophilous, by Bombus spp. Plants flowering (1971) Aug. 3-8 to Sept. 3-12 (Wade's Bog), Aug. 10-17 to Aug. 29-Sept. 9 (Ogg's Bog). 4\/4 individuals self-compatible. Propagule a microsclero-chore; dispersal anemochorous. Ecotypic variation unlikely. Apparently no hybrid-ization. Distribution: \"Bogs and wet places, B.C. s., on the w. side of the Cascades, especially near the coast, to n.w. Calif.\" (Hitchcock e t al. 1959). 303 Junous supiniformis Engelm. P e r e n n i a l , o f t e n w i t h f l o a t i n g l e a v e s and s t e m s ; stems g e n e r a l l y t u f t e d a l o n g r h i z o m e s , o f t e n +\/- decumbent and f r e e l y r o o t i n g b e l o w ; v e g e t a t i v e r e p r o d u c t i o n a l s o by v i v i p a r y . F l o w e r s a c t i n o m o r p h i c ; p e r f e c t , s t r o n g l y p r o t o g y n o u s ( l o w e s t f l o w e r s i n t h e i n f l o r e s c e n c e f l o w e r f i r s t ) , r e d u c e d , i n c o n s p i c u o u s , o d o r l e s s , n e c t a r l e s s ; b l o o m i n g (1971) June 2 7 - J u l y 1 t o J u l y 19-24 (Wade's B o g ) , June 14-22 t o J u l y 9-14 (Ogg's B o g ) . 2\/3 i n d i v i d u a l s s e l f - c o m p a t i b l e . P r o p a g u l e a s p o r o c h o r e ; d i s p e r s a l anemochorous and e p i z o o c h o r o u s . ( m u d ) . E c o t y p i c v a r i a t i o n u n l i k e l y . H y b r i d i z a t i o n p o s s i b l e . D i s t r i b u t i o n : \" M a r s h e s , p o n d s , d i t c h e s , and r i v e r s . . . ; A l a s , s . , a l o n g t h e c o a s t w. o f t h e ; C a s c a d e s , t o M e n d o c i n o Co., C a l i f . \" ( H i t c h c o c k et al. 1 9 6 9 ) . Kalmia polifolia Wang. E r e c t - s t r a g g l i n g , e v e r g r e e n s h r u b ; m o d e r a t e v e g e t a t i v e r e p r o d u c t i o n v i a l a y e r i n g and s h o r t r h i z o m e s . F l o w e r s p l e o m o r p h i c - s t e r e o m o r p h i c ( c o r o l l a g a m o p e t a l o u s , b u t s h a l l o w l y c r a t e r i f o r m ) ; p e r f e c t , w e a k l y p r o t o g y n o u s , m o d e r a t e l y n e c t a r i f e r o u s , f a i n t l y f r a g r a n t ; p i n k . A t a n t h e s i s , t h e a n t h e r s a r e h e l d u n d e r t e n s i o n i n c o r o l l a p o u c h e s ; t h e e l a s t i c i t y o f t h e f i l a m e n t s c a u s e s them t o s p r i n g o u t and d u s t i n s e c t v i s i t o r s t h a t t r i p t h e mechanism. A t f i r s t , t h e s t y l e d i v e r g e s f r o m t h e a x i s o f t h e p e d i c e l and f l o w e r , b u t as t h e f l o w e r m a t u r e s , t h e s t y l e g r a d u a l l y s t r a i g h t e n s . The n e c t a r , s e c r e t e d a t t h e b a s e o f t h e o v a r y , i s p r o t e c t e d by dense h a i r s o f t h e f i l a m e n t b a s e s . P o l l i n a t i o n e n t o m o p h i l o u s by Bombus s p p . , a n d r e n i d b e e s , s y r p h i d and m u s c i d f l i e s . F l o w e r s b l o o m i n g (1971) May 10-15 t o June 1-7 (Wade's B o g ) , May 10-18 t o June 3-17 (Ogg's B o g ) . 5\/5 i n d i v i d u a l s p a r t i a l l y s e l f - c o m p a t i b l e ( c f . J a y n e s 1 9 6 8 a ) . P r o p a g u l e a s p o r o c h o r e ( m i n u t e , l i n e a r s e e d ) ; d i s p e r s a l anemochorous. E c o t y p i c v a r i a t i o n c e r t a i n , i f one a c c e p t s K. polifolia sensu lato. No h y b r i d i z a t i o n , a c c o r d i n g t o J a y n e s (196.8b). D i s t r i b u t i o n : \" L o w l a n d bogs t o m o u n t a i n meadows; A l a s . s. t o C a l i f . , C o l o . , and e. N. Am.\" ( H i t c h c o c k et al. 1 9 5 9 ) . N o t e : K. polifolia has f u l l y formed a n t h e r s w i t h mature p o l l e n i n t h e o v e r w i n t e r r i n g f l o w e r buds ( c f . B e l l and B u r c h i l l 1 9 5 5 ) . Ledum groenlandioum Oeder. E r e c t , e v e r g r e e n s h r u b , w i t h o u t v e g e t a t i v e r e p r o d u c t i o n . F l o w e r s p l e o m o r p h i c ; i n f l o r e s c e n c e t e r m i n a l , d e n s e , u m b e l - l i k e ; f l o w e r s p e r f e c t , s l i g h t l y p r o t o g y n o u s , s t r o n g l y s p i c y - f r a g r a n t , s l i g h t l y n e c t a r i f e r o u s , w h i t e ; n e c t a r s e c r e t e d a t t h e base o f t h e o v a r y , e a s i l y a c c e s s i b l e ; b l o o m i n g (1971) June 9-14 t o J u l y 9-19 (Wade's B o g ) , June 1-9 t o June 2 4 - J u l y 6 (Ogg's B o g ) . 5\/5 i n d i v i d u a l s s e l f - c o m p a t i b l e . P r o p a g u l e a s p o r o c h o r e ( m i n u t e , e l o n g a t e , w i n g e d s e e d s ) ; d i s p e r s a l anemochorous. E c o t y p i c v a r i a t i o n p r o b a b l e . H y b r i d i z a t i o n w i t h 304 L. palustre ( s e e d i s c u s s i o n i n C a l d e r and T a y l o r 1968) and L. glandulosum ( S a v i l e 1971) h i g h l y p r o b a b l e . P o l l i n a t i o n e n t o m o p h i l o u s , by Bombus s p p . , a n d r e n i d b e e s , s y r p h i d and m u s c i d f l i e s . . D i s t r i b u t i o n : \" G r e e n l a n d t o A l a s . , s. t o New E n g l a n d , i n t h e w. a l o n g t h e c o a s t , c h i e f l y i n swamps and bogs t o t h e Ol y m p i c P e n i n s u l a and n.w. Oreg. and i n B.C. t o p o s s i b l y n. I d a . \" ( H i t c h c o c k et al. 1 9 5 9 ) . N o t e : o v e r w i n t e r r i n g f l o w e r buds have f u l l y f o r m e d a n t h e r s w i t h mature p o l l e n ( c f . B e l l and B u r c h i l l 1 9 5 5 ) . Linnaea borealis L. S l e n d e r , c r e e p i n g and t r a i l i n g , e v e r g r e e n h a l f - s h r u b . F l o w e r s s t e r e o m o r p h i c ; p e r f e c t , w e a k l y p r o t o g y n o u s , v e r y f r a g r a n t , m o d e r a t e l y n e c t a r i f e r o u s ; c o r o l l a f u n n e l f o r m , c a m p a n u l a t e , w h i t e , t i n g e d w i t h r o s e - p u r p l e , h a i r y i n s i d e . F l o w e r s b l o o m i n g (1971) J u l y 16-19 t o Aug. 6-9 (Wade's B o g ) , J u l y 18-23 t o J u l y 30-Aug. 3 (Ogg's B o g ) . P o l l i n a t i o n e n t o m o p h i l o u s , by ( p r i m a r i l y ) s y r p h i d f l i e s , o c c a s i o n a l l y by Bombus spp. Though t h e f l o w e r s a r e p e n d e n t , t h e stamens a r e s h o r t e r t h a n t h e e l o n g a t e , s l i g h t l y e x s e r t e d s t y l e , so t h a t s e l f - p o l l i n a t i o n i s u n l i k e l y . 3\/3 i n d i v i d u a l s s e l f - i n c o m p a t i b l e , w h i c h a g r e e s w i t h E a s t ( 1 9 4 0 ) . P r o p a g u l e a desmochore ( g l a n d u l a r f r u i t ) ; d i s p e r s a l e p i z o o c h o r o u s ( s t i c k t i g h t ) . E c o t y p i c v a r i a t i o n p r o b a b l e , i n v i e w o f t h e s p e c i e s ' d i s t r i b u t i o n ( s e e d i s c u s s i o n i n H i t c h c o c k et al. 1 9 5 9 ) . No h y b r i d i z a t i o n . D i s t r i b u t i o n : \" I n open o r dense woods and b r u s h a t v a r i o u s e l e v a t i o n s t h r o u g h o u t o u r range;\" c i r c u m p o l a r , e x t e n d i n g s. i n A m e r i c a t o C a l i f . , A r i z . , N.M., S.D., I n d . and W.Va.\" ( H i t c h c o c k et al 1 9 5 9 ) . Maianthemum dilatatum (Wood) N e l s . S Macbr. H e r b a c e o u s p e r e n n i a l , w i d e l y r h i z o m -a t o u s . F l o w e r s p l e o m o r p h i c ; p e r f e c t , m o d e r a t e l y p r o t o g y n o u s , n e c t a r l e s s b u t v e r y f r a g r a n t , s m a l l , w i t h 4 w h i t e t e p a l s ; b l o o m i n g (1971) June 1-10 t o 23-29 (Ogg's Bog; t h e s p e c i e s was p r e s e n t i n Wade's Bog b u t d i d n o t f l o w e r t h e r e ) . P o l l i n a t i o n e n t o m o p h i l o u s , by s y r p h i d and m u s c i d f l i e s . 3\/3 i n d i v i d u a l s s e l f - i n c o m p a t i b l e ( a g r e e s w i t h f i n d i n g s o f Ka.wano et al. 1 9 6 8 ) . P r o p a g u l e a s a r c o c h o r e ( r e d b e r r y ) ; d i s p e r s a l e n d o z o o c h o r o u s ( R i d l e y 1 9 3 0 ) . E c o t y p i c v a r i a t i o n p r o b a b l e ( s e e Kawano et al. 1 9 7 1 ) . A p p a r e n t l y no h y b r i d i z a t i o n . D i s t r i b u t i o n : \"Shaded o r m o i s t s t r e a m b a n k s , and open t o dense woods where a t a l l m o i s t , f r o m n e a r s e a l e v e l up t o n e a r l y 3500 f t e l e v . i n o u r a r e a ; A l a s . s. f r o m a l o n g t h e c o a s t t o t h e Ca s c a d e s t o M a r i n Co., C a l i f . , e. t o c. B.C. and t o n. I d a . \" ( H i t c h c o c k et al. 1 9 6 9 ) . N o t e : see Kawano et al. ( 1 9 6 7 , 1968) and S u z u k i and Kawano (1967) f o r p e r t i n e n t d i s c u s s i o n s . 3 0 5 M y r i o a g a l e L. Deciduous shrub l a c k i n g vegetative reproduction. Flowers actinomorphic reduced, inconspicuous, odorless, n e c t a r l e s s , aggregated i n c a t k i n s ; p l a n t s dioecious flowering (1971) A p r i l 1-5 to 20-25 (Wade's Bog), A p r i l 1-7 to 24-30 (Ogg's Bog). P o l l i n a t i o n anemophilous. C o m p a t i b i l i t y i r r e l e v a n t (plants are d i o e c i o u s ) . Propagule a megasclerochore and pterochore ( f r u i t a drupe-like nut, ovoid, with l u s t r o u s resin-atoms, bearing two wing-like, adnate b r a c t s ) ; d i s p e r s a l hydrochorous, aa e&pchorous, epizoochorous (mud). Ec o t y p i c v a r i a t i o n very probable. Apparently no h y b r i d i z a t i o n . D i s t r i b u t i o n : \"Lowland to montane bogs and c o a s t a l swamps; Alas, to Newf. , s. to n.w. Oreg., B.C., Mich., W i s e , Va.-- and i n the Appalachians to Tenn. and N.C.; Eura s i a \" (Hitchcock et a l . 1964). Note: M. g a l e has ni t r o g e n -f i x i n g root nodules (Bond 1951, 1967; Rodriguez-Barrueco 1968). N e p h r o p h y l l i d i u m o r i s t a - g a l l i (Menzies) G i l g . Perennial herbs from t h i c k , f l e s h y , shallow rhizomes. Flowers stereomorphic, though f l o r a l tube only 2-4 mm long; p e r f e c t , moderately protandrous, copiously n e c t a r i f e r o u s , with a rank, sour smell suggestive of mildewed laundry; c o r o l l a tube t u r b i n a t e ; c o r o l l a white, r o t a t e , with erose-undulate membranes on the midnerves and margins, deliquescent w i t h i n a few days of anthesis; plants d i s t y l o u s , flowering (1971) May 26-June 1 to June 8-11 (Ogg's Bog). P o l l i n a t i o n entomophilous, by muscid and sarcophagid f l i e s . 3\/3 i n d i v i d u a l s at l e a s t p a r t i a l l y self-compatible. Propagule a megasclerochore; d i s p e r s a l hydrochorous and epizoochorous (mud). Ecotypic v a r i a t i o n u n l i k e l y . No h y b r i d i z a t i o n . D i s t r i b u t i o n : \"Bogs and swamps....; Olympic Peninsula, Wash., to A l a s . ; Japan\" (Hitchcock et a l . 1959). P l a n t a g o maorooarpa Cham. \u00a3 Schlecht. Scapose p e r e n n i a l from a deep, heavy r o o t , without vegetative reproduction. Flowers stereomorphic; p e r f e c t , s t r o n g l y proto-gynous, borne i n a dense, a c r o p e t a l l y - f l o w e r i n g spike, n e c t a r l e s s , o d o r l e s s ; gamopetalous c o r o l l a s reduced, s c a r i o u s ; anthers and elongate stigma well-exserted during anthesis; plants flowering (1971) A p r i l 20-25 to May 7-11 (Ogg's Bog). P o l l i n a t i o n anemophilous. 5\/5 i n d i v i d u a l s self-compatible. Propagule a mega-sclerochore ( l a r g e , indehiscent capsule) and desmochore (seeds mucilaginous when wetted); d i s p e r s a l hydrochorous and epizoochorous ( v i a mucilaginous seeds). Ecotypic v a r i a t i o n u n l i k e l y . Apparently no h y b r i d i z a t i o n ( c f . Hitchcock et a l . 1959; Bassett 1967; Calder and Taylor 1968). D i s t r i b u t i o n : \"Sphagnum bogs, lake shores, and other c o l d , wet places; subcoastal (but not maritime) from Grays Harbor, Wash., to the A l e u t i a n Islands\" (Hitchcock et a l . 1959). 306 Rhynchospora alba (I..) Vahl. Densely t u f t e d p e r e n n i a l , l a c k i n g vegetative repro-duction. Flowers amorphic; mostly p e r f e c t (terminal 1-2 s p i k e l e t s staminate or rudimentary), s t r o n g l y protogynous, reduced, inconspicuous, o d o r l e s s , n e c t a r l e s s ; blooming (1971) J u l y 10 -13 to 24-27 (Wade's Bog), June 29-July 6 to J u l y 21-26 (Ogg's Bog). P o l l i n a t i o n anemophilous. 5\/5 i n d i v i d u a l s s e l f - c o m p a t i b l e . Propagule a megasclerochore and desmochore (achene + 10-12 r e t r o r s e l y b a r b e l l a t e b r i s t l e s ) ; d i s p e r s a l hydrochorous and epizoochorous (mud and s t i c k t i g h t ) . Moderate e c o t y p i c v a r i a t i o n probable. No h y b r i d i z a t i o n . D i s t r i b u t i o n : \"Sphagnum bogs and other very wet places at moderate and low e l e v a t i o n s ; i n t e r r u p t e d l y circumboreal,...; i n Am. from Newf. to N.C., i n l a n d to the Great Lakes r e g i o n . . . ; Alas, panhandle to c. C a l i f \" (Hitchcock et al. 1969). Sanguisorba officinalis L. Perennial herb with mainly b a s a l leaves and a small amount of vegetative reproduction v i a s h o r t , t h i c k r o o t s t o c k s . Flowers pleomorphic p e r f e c t , homogamous, od o r l e s s , s t r o n g l y n e c t a r i f e r o u s , apetalous; sepals maroon to deep maroon-purple. Nectar secreted by an annulus at the base of the s t y l e . The orange, s t i c k y p o l l e n i s o f t e n shed i n bud, although at anthesis the anthers and stigmas are remote. Plants flowering (1971) J u l y 14-19 to Aug. 8-13 (Wade's Bog), J u l y 14-24 to Aug. 6-11 (Ogg's Bog). P o l l i n a t i o n entomophilous, by muscid and syrphid f l i e s ; some s e l f - p o l l i n a t i o n i n e v i t a b l e . 4\/4 i n d i v i d u a l s s e l f - c o m p a t i b l e . Propagule a megasclerochore and pterochore (achene + p e r s i s t e n t , enclosing> hardened, '4-wing-angled, pubescent c a l y x ) ; d i s p e r s a l anemochorous, hydrochorous, and epizoochorous (mud). Ec o t y p i c v a r i a t i o n very probable. H y b r i d i z a t i o n with S. canadensis (Calder and Taylor 1968). D i s t r i b u t i o n : \"Muskeg, swamps, and bogs, almost circumboreal; Iceland to much of n. Europe and A s i a , i n N. Am. from Alas, and Yukon...to n.w. C a l i f . . . . \" (Hitchcock et al. 1962). Scirpus cespitosus L. Densely t u f t e d p e r e n n i a l with short, f r e e l y r o o t i n g rhizomes Flowers amorphic; p e r f e c t , s t r o n g l y protogynous, reduced, inconspicuous, o d o r l e s s , n e c t a r l e s s , united i n a several-flowered, s o l i t a r y , t erminal s p i k e l e t ; blooming (1971) May 3-6 to June 6-10 (Wade's Bog), A p r i l 30-May 10 to May 27-June 5 (Ogg's Bog). P o l l i n a t i o n anemophilous. 6\/6 i n d i v i d u a l s s e l f - c o m p a t i b l e . Propagule a megasclerochore; d i s p e r s a l hydrochorous and epizoochorous (mud, perhaps aided by the 6 very f r a g i l e perianth b r i s t l e s that i n v e s t the achene). E c o t y p i c v a r i a t i o n almost c e r t a i n . Apparently no h y b r i d i z a t i o n . D i s t r i b u t i o n : \"Sphagnum bogs, marshes and other very wet places; circumboreal, extending s. i n Am. to N.C. , Term. , s.w. Mont., n.c. Utah, c. Ida.,...and Oreg....\" (Hitchcock ct al. 19C9). 307 Tofieldia glutinosa ( M i c h x . ) P e r s . P e r e n n i a l , s h o r t - r h i z o m a t o u s h e r b s . F l o w e r s p l e o m o r p h i c ; p e r f e c t , w e a k l y p r o t a n d r o u s , o d o r l e s s , m o d e r a t e l y n e c t a r i f e r o u s . N e c t a r s e c r e t e d a t t h e b a s e s o f t h e t h r e e f u r r o w s between t h e c a r p e l s and t h e a n g l e s between t h e o v a r y and t h e s i x stamens. P e r i a n t h creamy w h i t e t o g r e e n i s h - w h i t e , sometimes t i n g e d w i t h p u r p l e . As t h e f l o w e r w i t h e r s , t h e d e h i s c e d stamens bend b a c k i n f r o m t h e i r d e f l e x e d p o s i t i o n , f r e q u e n t l y e v e n t o u c h i n g t h e s t i g m a s , so t h a t s e l f -p o l l i n a t i o n i s p o s s i b l e i f t h e a n t h e r s r e t a i n any v i a b l e p o l l e n . P l a n t s f l o w e r i n g (1971) June 21-26 t o J u l y 19-22 (Wade's B o g ) , June 11-19 t o J u l y 5-11 (Ogg's B o g ) . P o l l i n a t i o n e n t o m o p h i l o u s , by s y r p h i d and m u s c i d f l i e s . 7\/7 i n d i v i d u a l s s e l f -c o m p a t i b l e . P r o p a g u l e a p t e r o c h o r e ( s e e d + s p o n g y , appendaged, +\/- i n f l a t e d t e s t a ) ; d i s p e r s a l anemochorous, h y d r o c h o r o u s , e p i z o o c h o r o u s (mud). E c o t y p i c v a r i a t i o n a l m o s t c e r t a i n ( c o n s i d e r t h e r a n g e and v a r i a b i l i t y o f t h e s p e c i e s and s e e t a x o n o m i c d i s c u s s i o n s i n C a l d e r and T a y l o r (1968) and H i t c h c o c k et al. ( 1 9 6 9 ) ) . A p p a r e n t l y no h y b r i d i z a t i o n . D i s t r i b u t i o n : \"Meadows, b o g s , and s t r e a m b a n k s t o a l p i n e r i d g e s ; A l a s . s. t o C a l i f . , e. t o t h e A t l . c o a s t and s. t o Wyo. and I d a . i n w. N. Am., and . . . t o N.C.\" ( H i t c h c o c k et al. 19 6 9 ) . Trientalis aretica F i s c h . H e r b a c e o u s p e r e n n i a l s f r o m s h o r t , t h i c k e n e d t u b e r s ; v e g e t a t i v e r e p r o d u c t i o n f a i r l y e x t e n s i v e v i a t u b e r f o r m a t i o n a t t h e ends o f u n d e r g r o u n d r h i z o m e s ( c f . A n d e r s o n and L o u c k s 1 9 7 3 ) . F l o w e r s a c t i n o m o r p h i c ( c o r o l l a , t h o u g h g a m o p e t a l o u s , d i v i d e d n e a r l y t o t h e b a s e ) ; p e r f e c t , homogamous, o d o r l e s s , n e c t a r l e s s ( t h i c k , f l e s h y , s t a m e n - b e a r i n g r i n g s u r r o u n d s o v a r y a n d . a t t r a c t s f l i e s ) . The w h i t e p e t a l s s p r e a d o u t i n t o a s h a l l o w l y b o w l - s h a p e d s t a r 12-16 mm b r o a d . P l a n t s f l o w e r i n g (1971) May 30-June 4 t o J u l y 4-9 (Wade's B o g ) , May 20-27 t o June 10-19 (Ogg's Bog). P o l l i n a t i o n e n t o m o p h i l o u s , by m u s c i d f l i e s and s m a l l , s h o r t - t o n g u e d b e e s . 7\/7 i n d i v i d u a l s s e l f - i n c o m p a t i b l e ( a g r e e s w i t h Crowe (1964) re t h e genus Trientalis). P r o p a g u l e a m i c r o s c l e r o c h o r e ; d i s p e r s a l anemochorous. E c o t y p i c v a r i a t i o n u n l i k e l y . H y b r i d i z a t i o n p r o b a b l e , i f t h i s s p e c i e s i s c o n s i d e r e d d i s t i n c t f r o m T. europea ( c f . H i t c h c o c k et al. 1959; C a l d e r and T a y l o r 1968; H u l t e n 1 9 6 8 ) . D i s t r i b u t i o n : \" A l a s . s. t o A l t a . , B.C. and i n n. I d a . , i n t h e Cascades t o n.Oreg. and a l o n g t h e c o a s t t o e x t r e m e s. O r e g . , i n bogs and swamps\" ( H i t c h c o c k et al. 1 9 5 9 ) . N o t e : see A n d e r s o n and L o u c k s (1973) f o r p e r t i n e n t a u t e c o l o g i c a l i n f o r m a t i o n . Vaccinium ovatum P u r s h . T a l l , e r e c t , e v e r g r e e n s h r u b . F l o w e r s s t e r e o m o r p h i c , b e l l -s h a p e d ; p e r f e c t , f e e b l y p r o t a n d r o u s , m i l d l y f r a g r a n t , c o p i o u s l y n e c t a r i f e r o u s . C o r o l l a g a m o p e t a l o u s , n a r r o w l y c a m p a n u l a t e , p i n k i s h - w h i t e . The s l i g h t l y e x s e r t e d 308 s t y l e i s c l o s e l y e n c i r c l e d by 10 s h o r t e r , i n c l u d e d stamens. N e c t a r i s s e c r e t e d a t t h e base o f t h e s t y l e and t o r e a c h i t an i n s e c t must push i t s t o n g u e b e t w e e n t h e c i r c l e o f st a m e n s , and i n so d o i n g d i s l o d g e s p o l l e n f r o m t h e t e r m i n a l l y p o r a t e a n t h e r s o f t h e g e n e r a l l y p e n d e n t f l o w e r s ( K n u t h 1906-1909; Karmo 1957; F r e e 1 9 7 0 a ) . p l a n t s f l o w e r i n g (1971) May 15-21 t o June 4-14 (Ogg's B o g ) . P o l l i n a t i o n entomo-p h i l o u s , by Bombus spp. and s y r p h i d f l i e s ; a l s o , t h e r e i s some s e l f - p o l l i n a t i o n ( s e e Hagerup 1954; R i t c h i e 1955, 1 9 5 6 ) . 3\/3 i n d i v i d u a l s s e l f - c o m p a t i b l e ( o t h e r w o r k e r s have f o u n d t h a t s e l f e d b l u e b e r r y f l o w e r s a r e f e r t i l e , b u t f r u i t s e t i s h i g h e r i n o u t c r o s s e d f l o w e r s ( A a l d e r s and H a l l 1961; B r e w e r et al. 1969; H a l l and B e i l 1 9 7 0 ) ) . P r o p a g u l e a s a r c o c h o r e ( b l a c k b e r r y ) ; d i s p e r s a l e n d o z o o c h o r o u s . E c o t y p i c v a r i a t i o n u n l i k e l y . A p p a r e n t l y no h y b r i d i z a t i o n . D i s t r i b u t i o n : \"From t h e w. s i d e o f t h e C a s c a d e s t o t h e c o a s t , B.C. t o t h e redwood a r e a o f C a l i f . , s p o r a d i c s o u t h w a r d t o s. C a l i f . \" ( H i t c h c o c k et al. 1 9 5 9 ) . Vaccinium oxycoccus L. E v e r g r e e n , c r e e p i n g , s l e n d e r h a l f - s h r u b ; . v e g e t a t i v e r e p r o d u c t i o n \" e x t e n s i v e by t h e c r e e p i n g stems ( s t o l o n s ) . F l o w e r s p l e o m o r p h i c ; p e r f e c t , s l i g h t l y p r o t a n d r o u s , o d o r l e s s , m o d e r a t e l y n e c t a r i f e r o u s . P e t a l s d i s t i n c t , deep p i n k i s h , r e c u r v e d . A n t h e r s f o r m a c y l i n d e r e n v e l o p i n g - t h e s h o r t - e x s e r t e d s t y l e . To g e t a t t h e n e c t a r , bumble bees must c l i m b on t o t h e f l o w e r s f r o m b e l o w and i n s e r t t h e i r p r o b o s c e s i n t o t h e a n t h e r c y l i n d e r w h i l e h a n g i n g u p s i d e down. U n p o l l i n a t e d f l o w e r s a r e v e r y l o n g - l i v e d . P l a n t s f l o w e r i n g (1971) June 1-14 t o J u l y 14-24 (Wade's B o g ) , May 22-30 t o June 19-26 (Ogg's B o g ) . P o l l i n a t i o n entomo-p h i l o u s , by s p e c i e s o f Bombus; when t h e f l o w e r s a r e d e n s e , I have s e e n t h e bumble bees w a l k , r a t h e r t h a n f l y , f r o m f l o w e r t o f l o w e r ( s e e C r o s s 1 966; M a r u c c i 1967; F r e e 1 9 7 0 a ) ; s e l f - p o l l i n a t i o n i s a l s o p o s s i b l e i n t h e s e p e n d e n t f l o w e r s (Hagerup 1 9 5 4 ) . 5\/5 i n d i v i d u a l s s e l f - c o m p a t i b l e ; C r o s s (1966) s t a t e s t h a t t h o u g h t h e r e l a t e d V. macrocavpus i s s e l f - c o m p a t i b l e , l a r g e r f r u i t i s p r o d u c e d by o u t c r o s s e d f l o w e r s . P r o p a g u l e a s a r c o c h o r e (deep r e d b e r r y ) and p e r h a p s an auxo h o r e ; d i s p e r s a l e n d o z o o c h o r o u s . E c o t y p i c v a r i a t i o n p r o b a b l e . H y b r i d i z a t i o n l i k e l y , a t l e a s t w i t h some o f t h e o t h e r p r o p o s e d s e g r e g a t e s i n t h e V. oxycoccus complex ( c f . Camp 1944; C a l d e r and T a y l o r 1 9 6 8 ) . D i s t r i b u t i o n : \" A c r o s s t h e c o n t i n e n t i n Can., s . , u s u a l l y i n sphagnum b o g s , t o Oreg. and I d a . i n t h e w e s t ; E u r a s i a \" ( H i t c h c o c k et al. 1 9 5 9 ) . Vaccinium uliginosum L. Low, much-branched, s p r e a d i n g , d e c i d u o u s s h r u b , l a c k i n g v e g e t a t i v e r e p r o d u c t i o n e x c e p t p e r h a p s by l a y e r i n g . F l o w e r s s t e r e o m o r p h i c ; p e r f e c t , f e e b l y p r o t a n d r o u s , v e r y f r a g r a n t , a b u n d a n t l y n e c t a r i f e r o u s . C o r o l l a g a m o p e t a l o u s 309 o v o i d - u r c e o l a t e , l i g h t p i n k . F l o r a l mechanism s i m i l a r t o t h a t o f V. ovatum. P l a n t s f l o w e r i n g . ( 1 9 7 1 ) June 1-5 t o J u l y 4-9 (Wade's B o g ) , May 23-30 t o Jun e 9-14 (Ogg's Bo g ) . P o l l i n a t i o n e n t o m o p h i l o u s , by Bombus s p p . ; a l s o by s e l f i n g ( s e e Hagerup 1954; R i t c h i e 1955, 19S&). 3\/3 i n d i v i d u a l s p a r t i a l l y s e l f - c o m p a t i b l e ( s e e r e m a r k s u n d e r V. ovatum). P r o p a g u l e a s a r c o c h o r e ( b l u e b e r r y ) ; d i s p e r s a l e n d o z o o c h o r o u s E c o t y p i c v a r i a t i o n a l m o s t c e r t a i n . H y b r i d i z a t i o n w i t h V. oooidentalis p o s s i b l e ( s e e Camp 1945; H i t c h c o c k et al. 1959; C a l d e r and T a y l o r 1 9 6 8 ) . D i s t r i b u t i o n : \"Bogs a l o n g t h e c o a s t , A l a s , t o n. C a l i f . , e. i n Can. t o t h e A t l a n t i c C o a s t ; E u r a s i a \" ( H i t c h c o c k et al. 1 9 5 9 ) . Vaooinium vitis-idaea L. s s p . minus (Lodd.) H u l t . Low, e v e r g r e e n , t a p r o o t e d ( s e e H a l l and B e i l 1 9 7 0 ) , m a t - f o r m i n g h a l f - s h r u b ; v e g e t a t i v e r e p r o d u c t i o n e x t e n s i v e v i a c r e e p i n g stems. F l o w e r s s t e r e o m o r p h i c ; p e r f e c t , f e e b l y p r o t a n d r o u s , w i t h s t r o n g v a n i l l a - l i k e o d o r , c o p i o u s l y n e c t a r i f e r o u s . C o r o l l a g a m o p e t a l o u s , open-c a m p a n u l a t e , p i n k i s h . F l o r a l mechanism s i m i l a r t o t h a t o f V. ovatum. P l a n t s f l o w e r i n g (1971) June 9-12 t o June 2 9 - J u l y 4 (Wade's B o g ) , May 26-31 t o June 8-15 (Ogg's B o g ) . P o l l i n a t i o n e n t o m o p h i l o u s , by Bombus s p p . ; a l s o by s e l f i n g ( s e e Hagerup 1954; R i t c h i e 1955, 1 9 5 6 ) . 3\/3 i n d i v i d u a l s p a r t i a l l y s e l f - c o m p a t i b l e ( s e e comments u n d e r V. ovatum). P r o p a g u l e a s a r c o c h o r e ( b r i g h t r e d b e r r y ) ; d i s p e r s a l e n d o z o o c h o r o u s . E c o t y p i c v a r i a t i o n p r o b a b l e . A p p a r e n t l y no h y b r i d i z a t i o n . D i s t r i b u t i o n : \" A l a s , t o B.C. and a c r o s s t h e c o n t i n e n t t o G r e e n l a n d ; E u r a s i a \" ( S z c z a w i n s k i 1962). N o t e : see R i t c h i e (1955) f o r an e c o l o g i c a l l i f e h i s t o r y o f V. vitis-idaea. C. S u b a l p i n e Meadow Achillea millefolium L. M o d e r a t e l y r h i z o m a t o u s , p e r e n n i a l h e r b . I n f l o r e s c e n c e compound; r a y f l o w e r s z y g o m o r p h i c , d i s c f l o w e r s s t e r e o m o r p h i c , c a p i t u l u m p l e o m o r p h i c ; r a y s p i s t i l l a t e ; d i s c f l o w e r s p e r f e c t , m a r k e d l y p r o t a n d r o u s , s t r o n g l y n e c t a r i f e r o u s , m o d e r a t e l y o d o r i f e r o u s . Ray and d i s c f l o w e r s \u2022 w h i t e . P o l l i n a t i o n e n t o m o p h i l o u s , by s y r p h i d and m u s c i d f l i e s , s h o r t - t o n g u e d b e e s , and b u t t e r f l i e s and s k i p p e r s . P l a n t s f l o w e r i n g (1972) Aug. 12-20 t o Aug. 27- S e p t . 5. 4\/4 i n d i v i d u a l s s e l f -i n c o m p a t i b l e ; C l a u s e n (1953) and C l a u s e n , Keck and H i e s e y (1940) f o u n d A. mille-folium t o be on t h e whole s e l f - i n c o m p a t i b l e , w i t h a few s e l f - c o m p a t i b l e r a c e s . P r o p a g u l e a m i c r o s c l e r o c h o r e (epappose a c h e n e ) ; d i s p e r s a l anemochorous. E c o t y p i c v a r i a t i o n s t r o n g and w e l l - d o c u m e n t e d ( C l a u s e n , Keck and H i e s e y 1940, 1948; H i e s e y and Nobs 1970; M u l l i g a n and B a s s e t t 1959; T y r l 1 9 6 9 ) . H y b r i d i z a t i o n w i t h o t h e r 3 1 0 c l o s e l y r e l a t e d species probable. D i s t r i b u t i o n : \"A h i g h l y v a r i a b l e circumboreal species... occurring rather commonly i n various h a b i t a t s throughout our range\" (Hitchcock et al. 1955). Agoseris aurantiaca (Hook.) Greene. Taprooted p e r e n n i a l herb. Flowers a l l zygo-morphic; capitulum c a p i t a t e ; flowers a l l l i g u l a t e and p e r f e c t , markedly protandrous, with a mild \"daisy\" odor, strongly n e c t a r i f e r o u s , burnt orange i n c o l o r ; blooming (1972) Aug. 4-7 to Aug. 21-28. P o l l i n a t i o n entomophilous, p r i m a r i l y by b u t t e r f l i e s and skippers, o c c a s i o n a l l y by bumble bees ( c f . Knuth's (1906-1909) comments on the p o l l i n a t i o n of Hieracium aurantiacum, which has a very s i m i l a r flower-type). SC*\u00bb^- iwcompatible; f r u i t i n g heads of 1\/4 s e l f e d i n d i v i d u a l s had ' a few r i p e achenes. Propagule a pogonochore (pappose achene); d i s p e r s a l anemochorous. Ecotypic v a r i a t i o n probable; Cronquist i n Hitchcock et al. (1955) d e l i m i t s a southern Rocky Mt. v a r i e t y that i s probably e c o t y p i c a l l y d i s t i n c t from the t y p i c a l , northern v a r i e t y . Apparently no h y b r i d i z a t i o n . D i s t r i b u t i o n : \"Meadows and woodlands at moderate to high e l e v a t i o n s i n the mts.; A l t a . and B.C. to C a l i f , and N.M. ; Que.\" (Hitchcock et al 1955). Anemone occidentalis Wats. Perennial herb from a stout, o f t e n branched caudex surmounting a taproot. Flowers actinomorphic, bowl-shaped; p e r f e c t , weakly proto-gynous, odorless, n e c t a r l e s s , apetalous; sepals creamy white. Knuth (19 0 6-19 09) and Wells \u00a3 Barling(1971)' mention that some other species of t h i s \"Pulsatilla\" group have staminodia modified in t o club-shaped n e c t a r i e s , but t h i s i s not the case with A. occidentalis , which appears to be s t r i c t l y a p o l l e n flower. E a r l y i n anthesis the stamens are bent outward, but l a t e r some bend back up near the stigmas, so that s e l f - p o l l i n a t i o n i s possible (although the stamens are shorter than the s t y l e s ) . Flowers blooming (1972) June 2 5-30 to J u l y 7-15. P o l l i n a t i o n entomophilous, by muscid and syrphid f l i e s , small, short-tongued bees, and o c c a s i o n a l l y bumble bees. 3\/4 i n d i v i d u a l s (one died) self-compatible. Propagule a pogonochore ( v i l l o u s achene tipped with a long, sinuous, silky-plumose s t y l e ) ; d i s p e r s a l anemochorous (toward which end, the flower stem elongates considerably a f t e r p o l l i n a t i o n ) . Ecotypic v a r i a t i o n p o s s i b l e ; Hitchcock et al. (1964) mention that \"plants from Calif....may be v a r i e t a l l y separable\". Apparently no h y b r i d i z a t i o n . D i s t r i b u t i o n : \"Mountain slopes and meadows at middle to higher e l e v a t i o n s ; n. B.C. southward, through the Cascades and i n the Olympic Mts., to the S i e r r a Nevada, e. to A l t a . , ...Oreg., n. Ida., and Mont.\" (Hitchcock et al. 1964). Note: see Wells and B a r l i n g (1971) f o r a d i s c u s s i o n of the autecology of the r e l a t e d Anemone Pulsatilla. 311 Antennaria lanata- (Hook.) Greene. White-wooly, non-stoloniferous p e r e n n i a l herb from a compactly branched caudex. Flowers stereomorphic; capitulum d i s c o i d , r a y l e s s plants dioecious; flowers moderately n e c t a r i f e r o u s , with unpleasant, cheesy odor, comparable to that of d i r t y socks; c o r o l l a s d i r t y greenish-white. Plants f l o w e r i n g (1972) J u l y 7-20 to Aug. 1-9. P o l l i n a t i o n entomophilous, p r i m a r i l y by muscid and syrphid f l i e s . C o m p a t i b i l i t y i r r e l e v a n t ; no agamospermy, as i s common i n other species of Antennaria (Gustaffson 1946-1947; Grant 1971); 3\/3 female i n d i v i d u a l s set no seed i n i n s e c t exclosure cages (Hitchcock et al. 1955: \"Commonly se x u a l \" ) . Propagule a pogonochore (pappose achene); d i s p e r s a l anemochorous. E c o t y p i c v a r i a t i o n u n l i k e l y . H y b r i d i z a t i o n probable; Hitchcock et al. (1955) and Beals (1961) point out that h y b r i d i z a t i o n accompanied by p o l y p l o i d y and apomixis i s common i n the genus. D i s t r i b u t i o n : \"Alpine and subalpine s t a t i o n s ( r a r e l y lower) i n the mts. from B.C. and A l t a . to Oreg., Ida., and Wyo.\" (Hitchcock et al. 1955). Note: Beals (1961) found evidence of a l l e l o p a t h y i n .species of Antennaria i n southern Wisconsin. Arenaria capillaris P o i r . Cespitose p e r e n n i a l herb from a branched crown, forming loose mats. Flowers pleomorphic; p e r f e c t , moderately protandrous (the outer whorl of stamens matures a day or two before the inner whorl, by which time the stigmas are u s u a l l y r e c e p t i v e ) , f a i n t l y f r a g r a n t , abundantly n e c t a r i f e r o u s . Stamens s l i g h t l y longer than s t y l e s . Nectar secreted by glandular perigynous d i s c with rather prominent g l a n d l i k e protuberances f l a n k i n g the f i l a m e n t s . P e t a l s white, forming a background against which the red anthers, appressed i n e a r l y a n t h e s i s , show up as nectar guides. Plants flowering (1972) J u l y 28-Aug. 2 to Aug. 19-72. P o l l i n a t i o n entomophilous, by small, short-tongued bees, syrphid f l i e s , and inf r e q u e n t l y by bumble bees. 5\/5 i n d i v i d u a l s s e l f - i n c o m p a t i b l e , although East (194 knew of no s e l f - i n c o m p a t i b i l i t y i n Caryophyllaceae. Propagule a microsclerochore and pterochore (small, f l a t t e n e d , wing-margined seed); d i s p e r s a l anemochorous. Ecotypic v a r i a t i o n probably strong (note s u b s p e c i f i c d i f f e r e n t i a t i o n by Maguire (1947)). t l y b r i d i z a t i o n highly probable; Hickman (1971) regards A. aapillaris as a highly v a r i a b l e species i n t e r g r a d i e n t with s i x other species. D i s t r i b u t i o n : \"Sagebrush p l a i n s to subalpine rocky slopes; Alas, southward,...to n. Oreg., e. to A l t a . , Mont., and n. Nev. ; E u r a s i a \" (Hitchcock et al. 1964). Arnica l a t i f o l i a Bong. Fibrous-rooted, f a i r l y e x t e n s i v e l y rhizomatous p e r e n n i a l herb. Ray flowers zygomorphic; d i s c flowers stereomorphic; capitulum actinomorphic ray f l s . p i s t i l l a t e ; d i s c f I s . p e r f e c t , markedly protandrous, with a mild \"daisy\" 312 odor, abundantly n e c t a r i f e r o u s ; ray and d i s c flowers both yellow; blooming (1972) Aug. 10-14 to Aug. 22-Sept. 3. P o l l i n a t i o n entomophilous, by Bombus spp., syrphid and bombylid f l i e s , b u t t e r f l i e s and skippers, and small, short-tongued bees. 3\/3 i n d i v i d u a l s s e l f - i n c o m p a t i b l e . Propagule a pogonochore and desmochore (achene + barbellate.pappus); d i s p e r s a l anemochorous and epizoochorous ( s t i c k t i g h t ) . E cotypic v a r i a t i o n probable, considering d i s t r i b u t i o n and v a r i a b i l i t y , of species (c f . taxonomic discussions i n Maguire 1943; Hitchcock et al. 1955; Calder and Taylor 1968). H y b r i d i z a t i o n probable, as t h i s i s a widespread, polymorphic species. D i s t r i b u t i o n : \"Rocky places... moist woods, meadows and moist open p l a c e s . . . i n the mountains; Alas, to Colo, and C a l i f . \" (Hitchcock et al. 1955). Arnica mollis Hook. Perennial herb from f r e e l y r o o t i n g rhizomes that may be shortened i n t o a l o o s e l y branched caudex. Ray flowers zygomorphic; d i s c f I s . stereomorphic; capitulum actinomorphic; f l o r a l biology as i.n'-A. lati folia, except that some c a p i t u l a of A. mollis are erad i a t e . P o l l i n a t i o n entomophilous, by Bombus spp., syrphid and bombylid f l i e s , b u t t e r f l i e s and skippers, and small, short-tongued bees. Plants flowering (1972) Aug. 12-19 to 23-28. 3\/3 i n d i v i d u a l s s e l f - i n c o m p a t i b l e . Propagule a pogonochore (achenes + subplumose pappus); d i s p e r s a l anemochorous. L a t i t u d i n a l ecotypic v a r i a t i o n p o s s i b l e . H y b r i d i z a t i o n probable, as A. mollis i s a h i g h l y polymorphic species (Maguire 1943; Hitchcock et al. 1955). D i s t r i b u t i o n : \"Moist places at moderate to high elevations i n the mts.; A l t a . and B.C. to Colo., Utah, and C a l i f . \" (Hitchcock et al. 1955); Hulten (1968) has l o c a t i o n s i n c e n t r a l Alaska and the N.W.T., als o . Carex rossii Boott. Cespitose per e n n i a l sedge, l a c k i n g vegetative reproduction. Flowers amorphic; plants monoecious, strong l y protogynous; f l s . u n i s e x u a l , reduced, inconspicuous, n e c t a r l e s s , odorless; blooming (1972) J u l y 18-23 to 27-31. P o l l i n -a t i o n anemophilous. 3\/3 i n d i v i d u a l s self-compatible. Propagule a megasclerochore and desmochore (achene + f i n e l y puberulent, beaked perigynium); d i s p e r s a l anemo-chorous, epizoochorous ( s t i c k t i g h t ) . Ecotypic v a r i a t i o n probable, i n view of d i s t r i b u t i o n . H y b r i d i z a t i o n with C. brevipes p o s s i b l e . D i s t r i b u t i o n : \"In dry to moderately moist s i t e s i n areas which w i l l support coniferous f o r e s t , and i n adjacent meadows and p r a i r i e s , from near sea l e v e l i n w. Oreg. and Wash, to near timb e r l i n e i n the mts.; widespread i n the western c o r d i l l e r a n region from Yukon to C a l i f . , A r i z . , and Colo., and i r r e g u l a r l y e. to n. Mich.\" (Hitchcock et al. 1969). 313 Carex speatabilis Dewey. Tufted, p e r e n n i a l sedge from a system of sho r t , branching rhizomes. Flowers amorphic; plants .monoecious, s t r o n g l y protogynous; f l s . unisexual reduced, inconspicuous, odorless, n e c t a r l e s s ; blooming (1972)'Aug. 3-8 to 15-21. P o l l i n a t i o n anemophilous. 2\/2 i n d i v i d u a l s self-compatible. Propagule a megasclero-chore; d i s p e r s a l anemochorous, hydrochorous.-Ecotypic v a r i a t i o n p o s s i b l e . Hybrid-i z a t i o n with C. kelloggii reported by Moore and Calder (1964) and i s probable, with other species i n the s e c t i o n Atratae. D i s t r i b u t i o n : \"Meadows and other moist or wet-places at moderate to high e l e v a t i o n s i n the mts., sometimes above timber l i n e ; c o r d i l l e r a n region, from Alas, and Yuk., s. through A l t a . and B.C. to C a l i f . , n. Ida., and w. Mont., and also i n e. A s i a \" (Hitchcock et al. 1969). Castilleja miniata Dougl. P e r e n n i a l , h e m i p a r a s i t i c herb, without v e g e t a t i v e repro-duction. Flowers zygomorphic; p e r f e c t , homogamous, od o r l e s s , abundantly nectar-i f e r o u s ; i n f l o r e s c e n c e b r i g h t red or s c a r l e t ; plants flowering (197 2) J u l y 29-Aug. 4 to Aug. 22-30. P o l l i n a t i o n o r n i t h o p h i l o u s , by hummingbirds (Selasophorus rufous); C. miniata l i s t e d by Grant and Grant (1968) as a d e f i n i t e hummingbird flower. 3\/3 i n d i v i d u a l s s e l f - i n c o m p a t i b l e ; t h i s agrees with Heckard's (1968) observation that Castilleja appears to be l a r g e l y s e l f - s t e r i l e . Propagule a pterochore (saccate-appendaged seed); d i s p e r s a l anemochorous. Ec o t y p i c v a r i a t i o n almost c e r t a i n (see taxonomic d i s c u s s i o n i n Hitchcock et al. 1959 and Calder and Taylor 1968). H y b r i d i z a t i o n almost c e r t a i n (see Heckard 1964, 1968). D i s t r i b u t i o n : \"Meadows and slopes, mostly at medium and lower e l e v a t i o n s i n the mountains of w. N. Am., occurring i n every state and province from Alas, to C a l i f . , A r i z . , and N.M. (Hitchcock et al. 1959). Note: A t s a t t ' s (1970) conclusions regarding Orthoaqrpus are p e r t i n e n t to Castilleja, another h e m i p a r a s i t i c genus: \"...genetic v a r i a b i l i t y i s maximized through o b l i g a t e outbreeding... and i s apparently maintained and r e i n f o r c e d by a 'permissive' host-environment that allows the s u r v i v a l of a wide range of. recombinant phenotypes i n n a t u r a l populations.\" Castilleja parviflora Bong. var. albida (Penn.) Ownbey. P e r e n n i a l , h e m i p a r a s i t i c herbs, l a c k i n g vegetative reproduction. Flowers zygomorphic; p e r f e c t , homogamous, odorless, abundantly n e c t a r i f e r o u s ; i n f l o r e s c e n c e showy, white or p i n k i s h ; plants flowering (1972) Aug. 2-9 to 24-28. P o l l i n a t i o n entomophilous, by Bombus spp. 3\/3 i n d i v i d u a l s self-incompatible (agrees with Heckard's (1968) observation that the genus appears to be l a r g e l y s e l f - s t e r i l e ) . Propagule a microsclerochore and pterochore ( s l i g h t l y saccate-appendaged seed); d i s p e r s a l anemochorous. Ec o t y p i c v a r i a t i o n probable, as t h i s i s a very v a r i a b l e species (see taxonomic treatment 314 i n Hitchcock et al. 1959). H y b r i d i z a t i o n probable (see Heckard 1964, 1968). D i s t r i b u t i o n : \"Subalpine meadows at 4000-7500 f t e l e v a t i o n i n the Cascade and Olympic mts., n. to s. Alas, (at 2000-3000 f t ) and the Rocky Mts. of B.C. and A l t a . \" (Hitchcock et al. 1959). Note: see comments under C. miniata. Claytonia lanaeolata Pursh. Perennial herb from a f a i r l y deep-seated corm. Flowers pleomorphic; p e r f e c t , moderately protogynous, with a moderate, sweet odor, abundantly n e c t a r i f e r o u s . Petals white with yellow splotches ( n e c t a r i e s ) at t h e i r bases. Stigmas borne 1-2 mm above the anthers. Plants flowering (1972) June 28-J u l y 5 to J u l y 18-22. P o l l i n a t i o n entomophilous, by numerous muscid and s y r p h i d f l i e s , and also small, short-tongued bees. The majority of the flowers of 4\/4 i n d i v i d u a l s were self-compatible, but the flowers are not a u t o m a t i c a l l y s e l f -p o l l i n a t i n g , due to the dichogamy and herkogamy of the sexual organs. Propagule a b a l l o c h o r e ; d i s p e r s a l autochorous ( b a l l i s t i c ) ( R i d l e y 1930). E c o t y p i c v a r i a t i o n h i g h l y probable; Douglas and Taylor (1972) presented evidence of e c o t y p i c v a r i a t i o n i n western Washington alone; Hitchcock et al. recognized 4 v a r i e t i e s i n the P a c i f i c Northwest, while Davis (1966) d e l i m i t e d 6 v a r i e t i e s i n a l l . Apparently no h y b r i d i z a t i o n . D i s t r i b u t i o n : \"Sagebrush f o o t h i l l s to alpine s l opes, u s u a l l y where moist at l e a s t e a r l y i n s p r i n g , often p a r t i c u l a r l y abundant near snowbanks; B.C. southward, on both sides of the Cascades, to s. C a l i f . , e. to A l t a . and N.M.\" (Hitchcock et al. 1964). Notes: Meiosis occurs i n the f a l l or e a r l y winter, and f l o r a l s t r u c t u r e s develop during the winter, under the snow (personal observations and Kimball et al. 1973). Lewis (1967) discusses the extensive aneuploidy i n Claytonia spp., i n c l u d i n g C. lanaeolata. Delphinium nuttallianum P r i t z . P e r e n nial herb, without vegetative reproduction. Flowers zygomorphic; p e r f e c t , moderately protandrous, odorless but abundantly n e c t a r i f e r o u s , b r i g h t blue. Nectar secreted at the ends of the two upper p e t a l s , which are spurred and themselves ensheathed i n the spur of the upper s e p a l . The immature stamens are d i r e c t e d downward, but upon dehiscence become e r e c t , thus ba r r i n g the entrance to the nectar spurs. As the stamens wither, they bend down again and t h e i r place i s taken by the s t y l e s and now-receptive stigmas. Plants flowering (1972) J u l y 31-Aug. 4 to Aug. 16-26. P o l l i n a t i o n entomophilous, by Bombus spp. 3\/4 i n d i v i d u a l s s e l f - c o m p a t i b l e ; t h i s f i n d i n g agrees with those of E p l i n g and Lewis (1952) who stated that the genus i s self-compatible. Propagule a megasclerochore and pterochore (seed wing-angled and with a shiny, l o o s e , and somewhat puckered t e s t a ) ; d i s p e r s a l anemochorous ( c f . E p l i n g and Lewis 1952). 315 Ecotypic v a r i a t i o n probable, i n view of the d i s t r i b u t i o n and great v a r i a b i l i t y of t h i s species (see Hitchcock et al. 1964). H y b r i d i z a t i o n very probable; Ewan (1945) maintained that there were'signs of \"extensive h y b r i d i z a t i o n i n nature\" f o r D. nuttallianum, and Legro (1961) e s t a b l i s h e d that many Delphinium species h y b r i d i z e f r e e l y . D i s t r i b u t i o n : \"Sagebrush desert to (more commonly) mountain v a l l e y s and slopes, most .frequent i n the ponderosa pine b e l t , u s u a l l y i n w e l l drained, g r a v e l l y s o i l ; s.w. B.C. southward, i n the f o o t h i l l s of the - Cascades, to n; C a l i f . , e. to A l t a . , Mont., Wyo., and Neb., s. to Colo, and A r i z . \" (Hitchcock et al. 1964). Elymus glaucus Buckl. Coarse, cespitose p e r e n n i a l grass, l a c k i n g vegetative repro-duction. Flowers amorphic; p e r f e c t , +\/- homogamous, reduced, inconspicuous, odor-l e s s , n e c t a r l e s s ; blooming (1972) Aug. 10-14 to 21-24. P o l l i n a t i o n by s e l f i n g and anemophily. 3\/3 i n d i v i d u a l s self-compatible (agrees with Snyder 1950, 1951; Stebbins 1957b). Propagule a megasclerochore and desmochore (caryopsis + awned, scabrid-pubescent lemmas); d i s p e r s a l epizoochorous ( s t i c k t i g h t ) . E c o t y p i c v a r i a t i o n almost c e r t a i n . H y b r i d i z a t i o n common i n the genus (Brown and P r a t t 1960), known between E. glaucus and E. mollis (Bowden 1964) and E. glaucus a n d E. hirsutus (Hitchcock et al. 1969). The taxonomic species E. glaucus i s composed of many i n t e r s t e r i l e s i b l i n g species but can h y b r i d i z e f r e e l y with other Elymus species and with c e r t a i n species of Agvopyron, Hordeum, and Sitanion (Snyder 1950, 1951; Stebbins 1957b; Grant 1971). D i s t r i b u t i o n : \" P r a i r i e s , open woods, and dry to moist h i l l s i d e s , from the lowlands to midmontane; s. A l a s , to s. C a l i f . , e. to Ont., Mich., Ind., l a . , Colo., and N.M.\" (Hitchcock et al. 1969). Epilobium alpinum L. Low, u s u a l l y matted, p e r e n n i a l herb; vegetative reproduction f a i r l y extensive, v i a rhizomes and stolons. Flowers pleomorphic; p e r f e c t , homogamous, small, odorless, minutely n e c t a r i f e r o u s . In the study population the p e t a l s are white,but the species sensu lato embraces white to deep pink or l i l a c - r o s e c o l o r forms. The stigma ( e n t i r e ) i s borne at about the same l e v e l as the stamens.-Plants flowering (1972) J u l y 23-28 to Aug. 6-13. 3\/3 i n d i v i d u a l s f u l l y s e l f - c o m p a t i b l e ; Trelease (1891) noted that the small-flowered species of Epilobium were homogamous and self-compatible, and Raven (1972) estimated that 90% of Epilobium sect. Epilobium i s predominantly autogamous. Propagule a pogonochore (comose seed); d i s p e r s a l anemochorous. Ecotypic v a r i a t i o n i n t h i s extremely v a r i a b l e , wide-ranging species i s h i g h l y probable. H y b r i d i z a t i o n probable, at l e a s t w i t h i n the small-flowered, North American Epilobium complex ( c f . taxonomic treatments i n Hitchcock et al. 1962 and Calder and Taylor 1968); Trelease (1891') considered h y b r i d i z a t i o n 316 frequent i n the genus. D i s t r i b u t i o n : \"Moist banks and rocks, t a l u s slopes, and mountain meadows, often above timber l i n e ; throughout the mts. of w. N. Am., south-ward as f a r as s. C a l i f , and Colo., e. to the n. A t l a n t i c coast; E u r a s i a \" (Hitchcock et al. 1962). Erigeron peregrinus (Pursh) Greene. Fibrous-rooted p e r e n n i a l herb from a short rhizome or short, stout caudex. Capitulum actinomorphic; ray flowers zygomorphic, p i s t i l l a t e ; d i s c flowers stereomorphic, u s u a l l y p e r f e c t , s t r o n g l y protandrous, moderately odoriferous, copiously n e c t a r i f e r o u s ; rays l i g h t pink to lavender, d i s c flowers yellow-orange. Plants flowering (1972) Aug. 1-4 to 22-29. P o l l i n a t i o n entomophilous, by a wide v a r i e t y of i n s e c t s , i n c l u d i n g bumble bees, s m a l l , short-tongued bees, b u t t e r f l i e s and skippers, muscid, s y r p h i d , and bombyliid f l i e s . 5\/5 i n d i v i d u a l s s e l f - i n c o m p a t i b l e . Propagule a pogonochore (pappose achene); d i s p e r s a l anemochorous (see Ridley 1930). Ecotypic v a r i a t i o n apparently strong ( c f . taxonomic treatments i n Cronquist 1947, Hitchcock et al. 1955, and Calder and Taylor 19 68). H y b r i d i z a t i o n probable, as E. peregrinus i s a p i v o t a l , p r i m i t i v e species and h y b r i d i z a t i o n i s common i n the genus (Cronquist .1947). D i s t r i b u t i o n : \"Moist meadows, streamsides, or bogs,, at moderate to high e l e v a t i o n s i n the mts.; mts. of C a l i f . , Utah, and n. N.M., n. to about 56\u00b0 i n the Canadian Rockies,... to the A l e u t i a n Islands to the Commander Islands\" (Hitchcock al. 1955). Er.y.thronium grandif'lorum Pursh. Perennial herb from a deep-seated corm. Flowers pleomorphic + stereomorphic; p e r f e c t , l a r g e , showy, homogamous, f a i n t l y r e d o l ent of lemon, abundantly n e c t a r i f e r o u s . Tepals b r i g h t yellow, f r e e , r e f l e x e d above, but forming a short, basal tube sealed at the top by a protruding c o l l a r of swellings on the three inner t e p a l s . The n e c t a r i e s are s i t u a t e d at the base of the t e p a l s . Narrow grooves i n the middle of the three inner t e p a l s furrow through t h i s c o l l a r and are covered by filaments. The grooves provide access f o r p o l l i n a t o r s ' probosces. Plants flowering (1972) June 25-July 2 to J u l y 15-20. P o l l i n a t i o n entomophilous, mainly by bumble bees, o c c a s i o n a l l y by sm a l l , short-tongued bees (see Michener and Rettenmeyer 19 56) and b u t t e r f l i e s and skippers. 3\/4 i n d i v i d u a l s self-compatible; the s t y l e u s u a l l y p r o j e c t s 1-2 mm beyond the dehiscent anthers, but the stigma lobes i n l a t e anthesis tend to c u r l back and sometimes contact the anthers of the longest stamens (there are u s u a l l y two s l i g h t l y unequal sets of three stamens). Propagule a megasclerochore; d i s p e r s a l anemochorous, myrmechory p o s s i b l e . Ecotypic v a r i a t i o n probable (see d i s c u s s i o n of geographic v a r i a b i l i t y i n Hitchcock et al. 1969). H y b r i d i z a t i o n p o s s i b l e ; Applegate 3 1 7 (19 35) maintained that there were many i n d i c a t i o n s of h y b r i d i z a t i o n w i t h i n the e n t i r e genus. D i s t r i b u t i o n : \"Sagebrush slopes to montane f o r e s t , sometimes to near t r e e l i n e ; general i n much of s. B.C., the Olympic and Cascade Mts., Wash-., and n. Oreg., e. to Mont., Wyo., and Colo.\" (Hitchcock et al. 1969). Note: meiosis occurs during the f a l l or e a r l y winter, and the f l o r a l s t r u c t u r e s develop during the winter, under the snow (personal observations and Kimball et al. 1973). Festuca viridula Vasey. Cespitose p e r e n n i a l grass, forming small clumps; no vege-t a t i v e reproduction. Flowers amorphic; p e r f e c t , moderately protandrous, reduced, inconspicuous, odorless, n e c t a r l e s s ; blooming (1972) J u l y 28-Aug. 1 to Aug. 9-14. P o l l i n a t i o n anemophilous. 5\/5 i n d i v i d u a l s p a r t i a l l y s e l f-compatible. Propagule a megasclerochore; d i s p e r s a l anemochorous. Ecotypic v a r i a t i o n d o u b t f u l . H y b r i d i z a t i o n with F. altaiaa or F. scabrella probable, although Piper (1906) makes no mention of h y b r i d i z a t i o n w i t h i n t h i s complex. D i s t r i b u t i o n : \"Subalpine or a l p i n e slopes and rock s l i d e s and meadows to w e l l above timber l i n e ; B.C. to C a l i f . , e. to w. A l t a . , Mont.', Ida., and n.e. Oreg., i n Wash, not w. of the Cascade Mts.\" (Hitchcock et al. 1969). Eieracium gvacile Hook. P e r e n n i a l , moderately rhizomatous herbs. Capitulum c a p i t a t e ; flowers a l l zygomorphic, l i g u l a t e and p e r f e c t , markedly protandrous, o d o r l e s s , moderately n e c t a r i f e r o u s , b r i g h t yellow; blooming (1972) Aug. 11-15 to 22-26. P o l l i n a t i o n entomophilous, by syrphid f l i e s ' and small, short-tongued bees. 3\/3 i n d i v i d u a l s s e l f-incompatible (no i n d i c a t i o n of agamospermy; p o l l e n mostly good, meiosis r e g u l a r ) . Propagule a pogonochore (pappose achene); d i s p e r s a l anemochorous. Ecotypic v a r i a t i o n probably strong (see taxonomic dis c u s s i o n s i n Hitchcock et al. 1955 and Calder and Taylor 1968). H y b r i d i z a t i o n with U. albiflorum reported by Kruckeberg (1967) but apparently uncommon. D i s t r i b u t i o n : \"Meadows and other open places i n the mts., often at high e l e v a t i o n s ; Alas, and Mack, to C a l i f , and n. N.M. ; a l s o i n the Andes of s. S. Am.,...\" (Hitchcock et al. 1955). Hydrophyllum fendleri (Gray) H e l l e r . P e r e n n i a l , moderately rhizomatous herbs. Flowers stereomorphic; p e r f e c t , weakly protandrous, m i l d l y f r a g r a n t , c o p i o u s l y . n e c t a r i f e r o u s . C o r o l l a white, campanulate; s t y l e exserted, 2 - c l e f t ; stamens exserted, each filament flanked by a p a i r of c i l i a t e , l i n e a r c o r o l l a appendages. Nectar concealed, secreted by the base of the ovary, stored i n p e t a l c a v i t i e s bordered by the c o r o l l a appendages. Flowers blooming (1972) J u l y 19-29 to Aug. 10-15. 318 P o l l i n a t i o n entomophilous, by Bombus spp. 2\/3 i n d i v i d u a l s s e l f - c o m p a t i b l e . Propagule a megasclerochore and desmochore ( i f s t i f f - c i l i a t e calyx p e r s i s t s ) ; d i s p e r s a l anemochorous, epizoochorous ( s t i c k t i g h t ) . Ecotypic v a r i a t i o n u n l i k e l y . Apparently no h y b r i d i z a t i o n . D i s t r i b u t i o n : \"Thickets and moist, open p l a c e s , from the v a l l e y s and f o o t h i l l s to w e l l up i n the mountains; ...s. B.C.,...Wash, and Oreg. to n. C a l i f . , . . . c . Ida.; s. Wyo. to N.M. and s.e. Utah\" (Hitchcock et al. 1959). Junous drummondii E. Meyer. Strongly t u f t e d , p e r e n n i a l rush, forming small mats. Flowers actinomorphic; p e r f e c t , strongly protogynous, reduced, inconspicuous, odorless, n e c t a r l e s s ; blooming (1972) Aug. 7-10 to 17-20. P o l l i n a t i o n anemophilous. 3\/3 i n d i v i d u a l s self-compatible.-Propagule a sporochore and pogonochore (seed \u2022 -with membranous appendages at l e a s t as long as the body at each end). L a t i t u d i n a l ecotypic v a r i a t i o n p o s s i b l e ; species i s quite v a r i a b l e (Hitchcock et al. 1969). Apparently no h y b r i d i z a t i o n . D i s t r i b u t i o n : \"Alpine to subalpine meadows, stream-banks, talus\" slopes, and r i d g e s ; Alas, to C a l i f . , e. to the Rocky Mts., from A l t a . to N.M.\" (Hitchcock et al. 1969). Lupinus latifolius Agardh. Perennial herb from a branched caudex, l a c k i n g vegetative reproduction. Flowers zygomorphic; p e r f e c t , e s s e n t i a l l y homogamous, n e c t a r l e s s but with a marked, sweet fragrance, b r i g h t blue, papilionaceous. The two lower, inner-most petals are connate along t h e i r adjacent margins and together form the k e e l that envelops the 10 stamens. The anthers of the 5 outer stamens dehisce before anthesis and t h e i r p o l l e n i s stored i n the hollow cone cone c o n s t i t u t e d by the t i p the k e e l . Under the weight of a bumble bee p o l l i n a t o r , the 5 inner stamens act as p i s t o n s , extruding a s t r i n g of p o l l e n from the keel apex and d e p o s i t i n g i t on the v i s i t o r ' s underside. The stigma also protrudes at a l a t e r stage, so that c r o s s i n g can be e f f e c t e d ( a f t e r Knuth 1906-1909). Plants flowering (1972) J u l y 22-29 to Aug. 14-Sept. 3. P o l l i n a t i o n entomophilous, by Bombus spp. 5\/5 i n d i v i d u a l s s e l f -incompatible. Propagule a b a l l o c h o r e ; d i s p e r s a l autochorous ( b a l l i s t i c ) ( R i d l e y 19 30) anemochorous. Ecotypic v a r i a t i o n probably strong; broad d i s t r i b u t i o n of the species complex strongly suggests ecotypic status f o r the subspecies recognized by Dunn and G i l l e t t (1966). H y b r i d i z a t i o n occurs f r e e l y with L. nootkatensis, L. argenteus, and \" p o s s i b l y others\" (Dunn and G i l l e t t 1966). D i s t r i b u t i o n : \"Cascade Mts. from B.C. to C a l i f . , w. to the c o a s t a l mountains, on open subalpine ridges to wooded slopes, o c c a s i o n a l l y on the 'lowland p r a i r i e s ' \" (Hitchcock et al-. 1962). \"Montane 319 n.. Oreg., Wash., and B.C. t o A l a s k a \" (Dunn and G i l l e t t 1966). Note: For p e r t i n e n t d i s c u s s i o n s o f breeding systems and p o p u l a t i o n g e n e t i c s o f Lupinus see Dunn (1956), Harding (19 70), and Harding and Mankinen (1967, 1971). Luzula hitohaoakii Hamet-Ahti. S t r o n g l y r h i z o m a t o u s . p e r e n n i a l . Flowers actinomorphi p e r f e c t , s t r o n g l y protogynous, s m a l l , i n c o n s p i c u o u s , o d o r l e s s , n e c t a r l e s s ; blooming (1972) J u l y 4-9 t o J u l y 20-Aug. 1. P o l l i n a t i o n anemophilous. 3\/3 i n d i v i d u a l s predominantly s e l f - i n c o m p a t i b l e . Propagule a m i c r o s c l e r o c h o r e ; d i s p e r s a anemochorous. E c o t y p i c v a r i a t i o n u n l i k e l y (Hamet-Ahti 1971: \"...a very d i s t i n c t and i n v a r i a b l e taxon...\"). Apparently no h y b r i d i z a t i o n . D i s t r i b u t i o n : \"...upper b o r e a l t o o r o h e m i a r c t i c zones from southern B r i t i s h Columbia t o Oregon and Montana\" (Hamet-Ahti 1971). Luzula spicata (L.) DC. Caespitose p e r e n n i a l l a c k i n g v e g e t a t i v e r e p r o d u c t i o n . Flowers actinomorphic; p e r f e c t , moderately protogynous, reduced, i n c o n s p i c u o u s , o d o r l e s s , n e c t a r l e s s ; blooming (1972) J u l y 18-22 t o 27-29. P o l l i n a t i o n anemophilous 2\/3 i n d i v i d u a l s s e l f - c o m p a t i b l e . Propagule a m i c r o s c l e r o c h o r e ; d i s p e r s a l anemo-chorous. E c o t y p i c v a r i a t i o n probable. Apparently no h y b r i d i z a t i o n (see Nordenskiold 1956, 19,57). D i s t r i b u t i o n : \"Open s l o p e s , moraines, and r i v e r b a r s , s u b a l p i n e - t o a l p i n e i n our area; A l a s , and B.C. to the Olympic and Cascade Mts., Wash., n.e. and Cascadian (?) Oreg., C a l i f . , and s. i n the Rocky Mts. t o c. I d a . , and t o Mont., Wyo. , and Colo.; E u r a s i a \" (Hitchcock et al. 1969). Microsteris gracilis (Hook.) Greene. Annual herb. Flowers stereomorphic; p e r f e c t , homogamous, o d o r l e s s , weakly n e c t a r i f e r o u s . C o r o l l a w i t h white o r y e l l o w i s h tube and pink t o lavender limb. Stamens and s t y l e s i n c l u d e d , adjacent. P l a n t s f l o w e r i n g (1972) J u l y 15-20 t o J u l y 28-Aug. 2. P o l l i n a t i o n by s e l f i n g and o c c a s i o n a l entomo-p h i l y by short-tongued bees ( c f . Grant and Grant 1965). 5\/5 i n d i v i d u a l s s e l f -compatible. Propagule a mic r o s c l e r o c h o r e and desmochore (seed mucilaginous when wetted); d i s p e r s a l anemochorous and epizoochorous (adhesion). E c o t y p i c v a r i a t i o n probable. No h y b r i d i z a t i o n . D i s t r i b u t i o n : \"Dry t o moderately moist, open p l a c e s , mostly i n the f o o t h i l l s and l o w l a n d s s o m e t i m e s i n meadows or along streams; s. B.C. t o Mont., southward t o Baja C a l i f , and N.M.; B o l i v i a , A r g e n t i n a , and C h i l e \" (Hitchcock et al. 1959). \u2022320 Pedioularis bracteosa Benth. P e r e n n i a l , h e m i p a r a s i t i c herbs without vegetative reproduction. Flowers zygomorphic; p e r f e c t , homogamous, odorless, abundantly nectariferous.' C o r o l l a creamy-yellow or yellow green and sometimes p a r t l y purple. Galea beakless. Nectar secreted by a u n i l a t e r a l s w e l l i n g on the lower side of the ovary. Flowers blooming (1972) J u l y 31-Aug. 4 to Aug. 9-18. P o l l i n a t i o n p r i m a r i l y entomophilous by Bombus spp. , o c c a s i o n a l l y o r n i t h o p h i l o u s by hummingbirds ( c f . Sprague 1962; Faegri and van der P i j l 1971; Macior 1973). 3\/3 i n d i v i d u a l s s e l f -incompatible (agrees with Macior 1973). Propagule a microsclerochore and ptero -chore ( s l i g h t l y winged seed); d i s p e r s a l anemochorous. Ecotypic v a r i a t i o n h i g h l y probable; some of the v a r i e t i e s i n Hitchcock et a\u00a3.(1959) are l i k e l y ecotypes. H y b r i d i z a t i o n unknown (Sprague 1962; Macior 1973). D i s t r i b u t i o n : \"Woods, meadows, and moist open slopes i n the mountains; B.C. and A l t a . to n. C a l i f , and Colo.\" (Hitchcock et al. 1959). Note: the comments by A t s a t t (1970) regarding the breeding system of Orthocarpus are a l s o apropos Pedioularis (see under Castilleja miniata). \u2022 Penstemon procerus Dougl. More or l e s s t u f t e d perennials from a s u r f i c i a l , woody rhizome-caudex. Flowers zygomorphic; p e r f e c t , weakly protandrous, m i l d l y f r a g r a n t abundantly n e c t a r i f e r o u s , deep blue-purple. Nectaries are located on the outer side of the bases of the two upper filaments. The p o s t e r i o r stamen i s transformed i n t o a long staminode, sharply bent at i t s base, and t r a v e r s i n g the c o r o l l a tube o b l i q u e l y u n t i l i t s t i p l i e s on the c e n t r a l lobe of the lower l i p (thus imposing a b a r r i e r to nectar thievery by small i n s e c t s (se'e Straw 1956)). Plants flowering (1972) J u l y 31-Aug. 4 to Aug. 21-26. P o l l i n a t i o n entomophilous, p r i m a r i l y by small, short-tongued bees ( e s p e c i a l l y osmian) , a l s o by syrphid f l i e s . 3\/3 i n d i v i d u a l s predominantly self-compatible. Propagule a microsclerochore and pterochore (small, angled seed with a r e t i c u l a t e t e s t a ) ; d i s p e r s a l anemochorous. L a t i t u d i n a l ecotypic v a r i a t i o n p o s s i b l e . H y b r i d i z a t i o n probable; Keck and Cronqui (19 57) state that P. procerus i s quite v a r i a b l e and h y b r i d i z a t i o n i s p o s s i b l e ; Viehmeyer (1958) found that i n t e r s p e c i f i c hybrids i n the genus are o f t e n h i g h l y f e r t i l e . D i s t r i b u t i o n : \"Dry meadows and open or timbered slopes from the f o o t h i l l to above timber l i n e ; Alas, and Yukon to C a l i f . , Colo., and Sask.\" (Hitchcock et al. 19 59). Note: see Straw (19 63) and Crosswhite and Crosswhite (19 66) f o r observations on p o l l i n a t i o n ecology. Phleum alpinum L. Strongly t u f t e d p e r e n n i a l grass; the culms often decumbent and 321 semi-rhizomatous. Flowers amorphic; p e r f e c t , e s s e n t i a l l y homogamous, s m a l l , inconspicuous, odorless, n e c t a r l e s s ; blooming (1972) Aug. 2-8 to 22-27. P o l l i n a t i o n anemophilous, and by s e l f i n g . 3\/3 i n d i v i d u a l s f u l l y s e l f-compatible. Propagule a desmochore (caryopsis + p e r s i s t e n t , awned, h i s p i d - c i l i a t e glumes); d i s p e r s a l anemochorous and.epizoochorous ( s t i c k t i g h t ) . E c o t y p i c v a r i a t i o n c e r t a i n ; the species extends from near sea l e v e l at the coast to a l p i n e meadows i n l a n d , and the work of Callaghan and Lewis (1971a \u00a3 b) i n d i c a t e s eoctypic d i f f e r e n t i a t i o n at even a l o c a l l e v e l . Apparently no h y b r i d i z a t i o n . D i s t r i b u t i o n : \"Alas, to Newf., s. to most of montane-subalpine w. U.S.; Mex., s. Am., Europe\" (Hitchcock et al. 1969), Phlox diffusa Benth. Cespitose, mat-forming p e r e n n i a l , l a c k i n g vegetative repro-duction. Flowers stereomorphic; p e r f e c t , homogamous, very f r a g r a n t , abundantly n e c t a r i f e r o u s . C o r o l l a salverform, lavender to mauve; both stigma and anthers w e l l included i n the c o r o l l a tube, thus f a v o r i n g tongue-tip p o l l i n a t i o n (Grant and Grant 1965). Plants flowering (1972) J u l y 7-13 to 25-31. P o l l i n a t i o n entomo-p h i l o u s , by b u t t e r f l i e s and skippers, bumble bees, and bee hawk moths. 3\/3 i n d i v i d u a l s s e l f - i n c o m p a t i b l e . Propagule a megasclerochore; d i s p e r s a l anemochorous. Ecotypic v a r i a t i o n p o s s i b l e (northern and southern v a r i e t i e s recognized by H i t c h -cock et al. 1959). Apparently no h y b r i d i z a t i o n . D i s t r i b u t i o n : \"Forests and open rocky slopes at moderate to high e l e v a t i o n s i n the mountains, wholly w. of the c o n t i n e n t a l d i v i d e ; mountains of s. Van. I . and n.w. W a s h . C a s c a d e regi o n from s. B.C. through Oreg., and i n to the S i e r r a Nevada...to n. Ida. and n.w. Mont.\" (Hitchcock et al. 1959). Poa cusickii Vasey var. epilis (Scribn.) C.L. Hitchc. Tufted p e r e n n i a l g r a s s , r a r e l y with short, t h i c k rhizomes. Flowers amorphic; plants nearly completely d i o e c i o u s ; f l s . unisexual, reduced, inconspicuous, o d o r l e s s , n e c t a r l e s s ; blooming (1972) J u l y 15-20 to Aug. 1-8. P o l l i n a t i o n anemophilous. Plants agamospermous; 5\/5 p i s t i l l a t e plants set abundant f r u i t when i s o l a t e d i n the greenhouse. Propagule a megasclerochore and desmochore; d i s p e r s a l anemochorous and epizoo-chorous ( s t i c k t i g h t ) . A l t i t u d i n a l ecotypic v a r i a t i o n p o s s i b l e . H y b r i d i z a t i o n probable, as t h i s species i s both agamospermous and p o l y p l o i d ( c f . Clausen 1954, 1961). D i s t r i b u t i o n : -\"Sagebrush p l a i n s to a l p i n e meadows and r i d g e s ; B.C. to A l t a . and Sask., southward ( i n both the Olympic and Cascade mts. of Wash.) to C a l i f . , Nev., Utah, and Colo.\" (Hitchcock et al. 1969). 322 P o t e n t i l l a d i v e r s i f o l i a Lehm. Perennial herb, without vegetative reproduction. Flowers pleomorphic; bowl-shaped, p e r f e c t , weakly protogynous, f a i n t l y f r a g r a n t , copiously n e c t a r i f e r o u s . Petals' deep yellow. Nectar secreted at the base of the saucer-shaped, gla n d - l i n e d hypanthium. Plants flowering (19 72) J u l y 27-Aug. 3 to Aug. 11-26. P o l l i n a t i o n entomophilous, by muscid and syrphid f l i e s , and small, short-tongued bees. 3\/3 i n d i v i d u a l s incompletely s e l f - i n c o m p a t i b l e ( s e l f e d f l s . o f t e n produced a few r i p e achenes). Propagule a microsclerochore (weakly r e t i c u l a t e achene); d i s p e r s a l anemochorous. Ecotypic v a r i a t i o n probable (see taxonomic treatment i n Hitchcock et a l . 1962). H y b r i d i z a t i o n probable, with members of the P. p l a t t e n s i s - pinnatiseota - wyomingensis and the P. concinna complexes (H i t c h -cock et a l . 1962). D i s t r i b u t i o n : \"From a l p i n e , where most common i n meadows and on ledges and rocky slopes, to subalpine or montane, where c h i e f l y along stream banks; Yukon and B.C. southward, i n the higher mts. of Wash, and Oreg., t o C a l i f , and Nev. , e. to Sask., and s. i n the Rocky Mts. to Utah and N.M.\" (Hitchcock et a l . 1962). P o t e n t i l l a f l a b e l l i f o l i a Hook. Perennial with branched crown and well-developed r o o t s t o c k s , l a c k i n g vegetative reproduction. Flowers pleomorphic; p e r f e c t , s l i g h t l y protogynous, f a i n t l y f r a g r a n t , abundantly n e c t a r i f e r o u s ; nectar secreted as a t h i n , s h i n i n g f i l m on a bla c k i s h - p u r p l e d i s c j u s t i n s i d e the filament bases. Pe t a l s deep yellow, forming a bowl. P o l l i n a t i o n entomophilous, u s u a l l y by short-tongued i n s e c t s that l i c k the n e c t a r i f e r o u s d i s c ; eg., numerous muscid and syrphid f l i e s , s m a l l , short-tongued bees, and very i n f r e q u e n t l y , bumble bees. Flowers blooming (1972) J u l y 8-18 to Aug; 1-9. 5\/5 i n d i v i d u a l s incompletely self-incom-p a t i b l e ( s e l f e d flowers often produced a few r i p e achenes). Propagule a micro-sclerochore (smooth achene); d i s p e r s a l anemochorous. Ecotypic v a r i a t i o n u n l i k e l y . H y b r i d i z a t i o n p o s s i b l e (many species of t h i s genus h y b r i d i z e f r e e l y ) , but none reported. D i s t r i b u t i o n : \"Wet meadows and stream banks to alpine or subalpine ridges and t a l u s slopes; B.C. southward i n the Cascades (and i n the Olympic Mts.) to the S i e r r a Nevada of C a l i f . , e. i n B.C. to the Rocky Mts. of s.e. A l t a . , and through most of montane Ida. and Mont., a l s o in...n.e. Oreg.\" (Hitchcock et a l . 1962). Ranunculus e s c h s c h o l t z i i Schlecht. P e r e n n i a l herb without vegetative reproduction. Flowers actinomorphic; p e r f e c t , weakly protogynous, moderately n e c t a r i f e r o u s , odorless. Petals yellow, bearing flap-covered nectar p i t s at t h e i r bases and forming shallow bowls. On dehiscence of the anthers the stamens d e c l i n e toward 323 the p e t a l s , so that p o l l e n does not r e a d i l y f a l l upon the stigmas. Plants flower-ing (1972) J u l y 16-20 to J u l y 25-Aug. 4. P o l l i n a t i o n entomophilous, p r i m a r i l y by muscid f l i e s , a l s o frequently by syrphid f l i e s and s mall, short-tongued bees. 3\/3 i n d i v i d u a l s self-compatible. Propagule a megasclerochore and desmochore (achene + s t y l a r beak); d i s p e r s a l anemochorous, hydrochorous, epizoochorous ( s t i c k t i g h t ) . Ecotypic v a r i a t i o n h i g h l y probable, as t h i s i s a \"widespread polymorphic species\" (Hitchcock et al. 1964). H y b r i d i z a t i o n with R. nivalis and R. pygmaeus p o s s i b l e (see Calder and Taylor 1968). D i s t r i b u t i o n : \"Mountain meadows and t a l u s slopes; A l a s . s. to C a l i f . , e. to A l t a . , the Rocky Mt. s t a t e s , and N.M. and A r i z . \" (Hitchcock et al. 1964). Note: see L i n s l e y and MacSwain (1959) f o r pertinent information about p o l l i n a t i o n ecology of Ranunculus. Sedum lanceolatum Torr. P e r e n n i a l , succulent herb; moderate vegetative reproduction by decumbent, s t e r i l e shoots. Flowers actinomorphic; p e r f e c t , homogamous, od o r l e s s , moderately n e c t a r i f e r o u s , b r i g h t yellow. Nectar secreted b a s a l l y between the petals and stamens. Plants flowering (1972) Aug. 12-18 to 25-29. P o l l i n a t i o n entomophilous, by bumble bees, small, short-tongued bees, and muscid and syrphid f l i e s . 3\/3 i n d i v i d u a l s s e l f - i n c o m p a t i b l e . Propagule a microsclerochore; d i s p e r s a l anemochorous. Ecotypic v a r i a t i o n h i g h l y probable, i n view of range. H y b r i d i z a t i o n probable (see taxonomic d i s c u s s i o n i n Hitchcock et al. 1964). D i s t r i b u t i o n : \"Open exposed p l a c e s , u s u a l l y on rocks or on g r a v e l l y or rocky s o i l , from near sea l e v e l to subalpine areas; s. Alas, and Yukon s. to Calif.', e.'to A l t a . , S.D., Neb., Colo., and N.M.\" (Hitchcock et al. 1964). Senecio integerrimus Nutt. Perennial herb from a very short, e r e c t crown, l a c k i n g vegetative reproduction. Capitulum pleomorphic; ray flowers l i g u l a t e , zygomorphic, p i s t i l l a t e ; d i s c flowers stereomorphic, p e r f e c t , markedly protandrous, moderately f r a g r a n t , copiously n e c t a r i f e r o u s . Ray and d i s c flowers both yellow. Plants flowering (1972) J u l y 18-25 to Aug. 3-10. P o l l i n a t i o n entomophilous, by bumble bees, small, short-tongued bees, bombyliid, syrphid, and muscid f l i e s . 3\/3 i n d i v i d u a l s s e l f - i n c o m p a t i b l e . Propagule a pogonochore (pappose achene); d i s p e r s a l anemochorous. Ecotypic v a r i a t i o n probable (see taxonomic d i s c u s s i o n s i n Hitchcock et al. 1955 and Barkley 1960). H y b r i d i z a t i o n p o s s i b l e . D i s t r i b u t i o n : \"Moderately dry to rather moist open places and open woods, from the v a l l e y s to near timber l i n e ; s. B.C. to C a l i f . , e. to Sask. and Minn.\" (Hitchcock et al. 1955). 324 Senecio triangularis Hook. T a l l , p e r e n n i a l herb, l a c k i n g vegetative reproduction. Capitulum pleomorphic; ray flowers zygomorphic, p i s t i l l a t e ; d i s c flowers stereo-morphic, p e r f e c t , markedly protandrous, mildy f r a g r a n t , s t r o n g l y n e c t a r i f e r o u s . Both ray and d i s c flowers yellow. Plants flowering (1972) Aug. 31-18 to Aug. 29-Sept. 4. P o l l i n a t i o n entomophilous, by bumble bees, small, short-tongued bees, b u t t e r f l i e s and skippers, muscid, syrphid, and bombyliid f l i e s . 3\/3 i n d i v i d u a l s s e l f - i n c o m p a t i b l e . Propagule a pogonochore (pappose achene); d i s p e r s a l anemo-chorous. Ecotypic v a r i a t i o n very probable; t h i s i s an extremely v a r i a b l e complex (see Hitchcock et al. 1955 and Calder and T a y l o r 1968). Apparently no h y b r i d i z a t i o n . D i s t r i b u t i o n : \"Streambanks and other moist p l a c e s , mostly at moderate and high e l e v a t i o n s i n the mts.; Alas, and Yukon to Sask., N.M., and C a l i f . \" (Hitchcock et al., 1955) . Sibbaldia procumbens L. Mat-forming, moderately rhizomatous p e r e n n i a l . Flowers pleomorphic; p e r f e c t , homogamous, odor l e s s , minutely n e c t a r i f e r o u s . P e t a l s small, about h a l f the length of the sepals, greenish-yellow. Nectar secreted by the shallow, g l a n d - l i n e d hypanthium. Plants flowering (1972) J u l y 9-18 to 26-29. P o l l i n a t i o n by s e l f i n g and o c c a s i o n a l l y entomophilous, by muscid f l i e s ( c f . Clapham, T u t i n and Warburg 1962); Coker (1966) says that i n the B r i t i s h I s l e s the species i s p o l l i n a t e d as w e l l by ants and bumble bees. 3\/3 i n d i v i d u a l s s e l f -compatible. Propagule a microsclerochore; d i s p e r s a l anemochorous. E c o t y p i c v a r i a t i o n probable, considering the d i s t r i b u t i o n . Apparently no h y b r i d i z a t i o n . D i s t r i b u t i o n : \"Circumpolar, extending southward i n N. Am., on open, dry to moist alpine slopes or meadows, to s. C a l i f . , Utah, Colo., Que., and N.H.\" (Hitchcock et al. 1962). Silene parryi (Wats.) Hitchc. S Maguire. Perennial herb from a simple to branched caudex surmounting a taproot. Flowers stereomorphic (by v i r t u e of a synsepalous c a l y x ) ; p e r f e c t , moderately protandrous, h e a v i l y perfumed, abundantly n e c t a r i f e r o u s . Anthesis n o c t u r n a l ; the 10 stamens are exserted and dehisce i n two successive sets of f i v e , and the stigmas are r e c e p t i v e the f o l l o w i n g night. P e t a l s creamy white, clawed; blade bilobed to n e a r l y e q u a l l y 4-lobed,' appendaged. Plants flower-ing (1972) Aug. 13-18 to 27-31. P o l l i n a t i o n entomophilous, by sphingid hawkmoths. 3\/3 i n d i v i d u a l s self-compatible; however, f l o r a l biology m i l i t a t e s against s e l f -p o l l i n a t i o n . Propagule a megasclerochore; d i s p e r s a l anemochorous. Ec o t y p i c v a r i a -t i o n u n l i k e l y . Kruckeberg (1955, 1961) has found most i n t e r s p e c i f i c hybrids among western North American Silenes to be s t e r i l e . D i s t r i b u t i o n : \"Montane, from B.C. 325 s. to the Olympic and Cascade Mts. of Wash., e. i n B.C. to the Rocky Mts., and s. to c. Ida. and n.w. Wyo.\" (Hitchcock et al. 1964). Thalictrum oacidentale Gray. Moderately rhizomatous, p e r e n n i a l herb. Flowers actinomorphic; plants incompletely dioecious\"; flowers reduced, r e l a t i v e l y inconspicuous, n e c t a r l e s s , odorless; blooming (1972) J u l y 18-25 to Aug. 3-7. P o l l i n a t i o n anemophilous (see Kaplan and Mulcahy 1971). C o m p a t i b i l i t y i r r e l e v a n t . Propagule a megasclerochore'and desmochore ( s h o r t - s t i p i t a t e achene with s h o r t , p e r s i s t e n t s t y l e ) ; d i s p e r s a l anemochorous, epizoochorous ( s t i c k t i g h t ) . E c o t y p i c v a r i a t i o n probable; \"This i s the most v a r i a b l e of our species...\" (Hitchcock et al. 1964). H y b r i d i z a t i o n with T. venulosum p o s s i b l e . D i s t r i b u t i o n : \"Common from B.C. southward, on both sides of the Cascades, to n. C a l i f . , e. to A l t a . , Mont., Wyo., Colo., and Utah.\" (Hitchcock et al. 1964). Trisetum spicatum (L.) Richter. Cespitose, perennial, grass; no vegetative repro-duction i n the study area although Hulten (1968) states that v i v i p a r o u s p l a n t s occur o c c a s i o n a l l y . Flowers amorphic; p e r f e c t , e s s e n t i a l l y homogamous, reduced, inconspicuous, o d o r l e s s , n e c t a r l e s s ; blooming (1972) Aug. 9-14 to 25-29. P o l l i n a t i o n anemophilous. 4\/4 i n d i v i d u a l s s e l f - i n c o m p a t i b l e . Propagule a desmochore + micro-sclerochore (caryopsis + d i v a r i c a t e l y awned, scabridulous lemmas); d i s p e r s a l anemochorous, epizoochorous ( s t i c k t i g h t ) . Ecotypic v a r i a t i o n strong (Tieszen and Bonde 1967; Hulten 1959 recognized 14 subspecies i n the complex). H y b r i d i z a t i o n probable, at l e a s t w i t h i n the whole species complex ( c f . Morrison 1959). D i s t r i -bution: \"Montane f o r e s t to a l p i n e slopes i n both dry and rocky, and moist h a b i t a t s , often somewhat weedy on waste areas; Alas, across Can. to Greenl., s. to most of the mountains of w. U.S., the Great Lakes area, and the Appalachian Mts., through Mex. to a n t a r c t i c S. Am.; a r c t i c - a l p i n e E u r a s i a \" (Hitchcock et al. 1969). Vaccinium deliciosutn Piper. Low, often matted, deciduous shrub. Flowers stereo-morphic; p e r f e c t , +\/- homogamous, f a i n t l y f r a g r a n t , copiously n e c t a r i f e r o u s . C o r o l l a gamopetalous, pink. F l o r a l mechanism s i m i l a r to that of V. ovatum. Flowers blooming (1972) J u l y 15-23 to Aug. 9-19. P o l l i n a t i o n entomophilous, by Bombus spp., and some s e l f i n g (see Hagerup 1954). 2\/2 i n d i v i d u a l s p a r t i a l l y self-compatible. Propagule a sarcochore (glaucous blue b e r r y ) ; d i s p e r s a l endo-zoochorous. Ecotypic v a r i a t i o n improbable. Apparently no contemporary. h y b r i d i z a t i o n . D i s t r i b u t i o n : \"Montane i n the Olympic Mts. and i n the Cascades from s. B.C. to n. Oreg.\" (Hitchcock et al. 1959). Note: Camp (1942) hypothesized that Vaccinium 326 deliciosum was the a l l o t e t r a p l o i d product of h y b r i d i z a t i o n between the d i p l o i d s V. saoparium and V. caespitosum. I have found n = 24 i n V. deliciosum and n - 12 i n V. saoparium; Camp's hypothesis may have been f u l l y accurate. Vaooinium saoparium Leiberg. More or l e s s matted, broomy, deciduous shrub. Flowers stereomorphic; p e r f e c t , +\/- homogamous, f a i n t l y f r a g r a n t , copiously n e c t a r i f e r o u s . C o r o l l a gamopetalous, pale pink. F l o r a l mechanism very s i m i l a r to that of V. ovatum. Plants flowering (197 2) J u l y 5-2 0 to Aug. 1-9. P o l l i n a t i o n entomophilous, almost e x c l u s i v e l y by Bombus spp.. The bumble bees v i s i t the flowers during almost a l l d a y l i g h t hours (8 A.M. to 8 P.M.), even i n c h i l l y weather. The flowers are -frequently borne very c l o s e to the ground, and the bumble bees f l y e n e r g e t i c a l l y from flower to flower, skimming j u s t above the surface l i k e h o v e r c r a f t . 3\/3 i n d i v i d u a l s p a r t i a l l y self-compatible (the majority .of s e l f e d flowers set f r u i t , but the development of the f r u i t was sometimes retarded or aborted). Propagule a sarcochore ( b r i g h t red b e r r y ) ; d i s p e r s a l endozoochorous. Ecotypic v a r i a t i o n u n l i k e l y . Apparently no h y b r i d i z a t i o n . D i s t r i b u t i o n : \"B.C. to n. C a l i f . , e. through Ida. to A l t a . and S.D., s. i n the Rocky Mts. to Colo., u s u a l l y r a t h e r w e l l up i n the mountains\" (Hitchcock et al. 1959). Valeriana sitchensis Bong. Perennial herb from a stout, branched rhizome or caudex. Flowers e s s e n t i a l l y stereomorphic ( c o r o l l a tube somewhat gibbous); p e r f e c t , moderately protandrous, white, h e a v i l y sweet-scented, c o p i o u s l y . n e c t a r i f e r o u s (nectar secreted and stored i n the c o r o l l a tube g i b b o s i t y ) . Three-lobed stigma borne 1-2 mm beyond the exserted anthers. P o l l i n a t i o n entomophilous, by a wide v a r i e t y of i n s e c t s , i n c l u d i n g bumble bees, small, short-tongued bees, muscid, syrphid, and bombyliid f l i e s , and b u t t e r f l i e s and skippers. Flowers blooming (1972) J u l y 27-Aug. 7 to Aug. 19-27. 4\/4 i n d i v i d u a l s self-compatible (agrees with East 1940). Propagule a pogonochore\" (nerved achene + p e r s i s t e n t calyx of 12-20 pappus-like segments); d i s p e r s a l anemochorous (Ridley 1930). E c o t y p i c v a r i a t i o n p o s s i b l e ; Meyer (19 51) mentioned subalpine and lowland v a r i a n t s . H y b r i d i z a t i o n with V. sopuleri p o s s i b l e . D i s t r i b u t i o n : \"Moist, open or wooded places at middle and upper a l t i t u d e s i n the mountains, often i n wet meadows; s. Yukon and s. Alas, to w. Mont., c. Ida., and n. C a l i f . . . . \" (Hitchcock et al. 1959). 327 Veratrum viride A i t . Robust, s l i g h t l y rhizomatous, per e n n i a l herb. Flowers p l e o -morphic ; plants polygamous, upper f I s , p e r f e c t and the lower staminate, weakly protandrous, moderately n e c t a r i f e r o u s , smelling f a i n t l y of musk. Tepals yellow-green to deep green, with nectar glands at t h e i r bases. Plants flowering (19 72) Aug. 3-7 to 18-22. P o l l i n a t i o n entomophilous, p r i m a r i l y by small, muscid f l i e s . Self-compatible ( ? ) . Propagule a pterochore ( f l a t seed broadly winged by the loose t e s t a ) ; d i s p e r s a l anemochorous, hydrochorous. Ecotypic v a r i a t i o n almost c e r t a i n . Apparently no h y b r i d i z a t i o n . D i s t r i b u t i o n : \"Wet t h i c k e t s , swamps, and lowland to montane meadows; Alas, s., through the Olympic and Cascade mts. of Wash., to the n. Cascades of Oreg., e. to the Rocky Mts., from B.C. and A l t a . to Mont, and Ida., and i n Can. to Ont. and Que. and s. to N.C.\" (Hitchcock et al. 1969). Veronica cusickii Gray. Perennial herb from a loose or compact system of shallow rhizomes. Flowers e s s e n t i a l l y stereomorphic ( s h o r t - t u b u l a r , s l i g h t l y i r r e g u l a r ) ; p e r f e c t , weakly protogynous, odorless, moderately n e c t a r i f e r o u s . C o r o l l a deep blue-purple; short c o r o l l a tube stores and conceals nectar secreted by a d i s c below the ovary. The conspicuously exserted s t y l e i s d i r e c t e d o b l i q u e l y downward and the two stamens diverge l a t e r a l l y . Flowers blooming (1972)- Aug.2-7 to 12-19. P o l l i n a t i o n entomophilous, almost e x c l u s i v e l y by syrphid f l i e s (see tex t f o r d e t a i l s ) . 4\/4 i n d i v i d u a l s self-compatible. Propagule a microsclerochore and pterochore (small, winged seed); d i s p e r s a l anemochorous. Ecotypic v a r i a t i o n p o s s i b l e (I have i n mind the C a l i f o r n i a v a r i e t y mentioned below). Apparently no h y b r i d i z a t i o n . D i s t r i b u t i o n : \"Moist, open rocky slopes, or sometimes along streams or i n meadows, at r a t h e r high e l e v a t i o n s i n the mountains, sometimes above timber l i n e ; Olympic and Cascade mts. of Wash., e. t o . . . c . Ida. and w. Mont., ...and n.e. Oreg.; a smaller-leaved, perhaps v a r i e t a l l y separable p l a n t occurs i n Yosemite Nat. Park, C a l i f . \" (Hitchcock et al. 1959). 328 L i t e r a t u r e c i t e d i n Appendix 3. Only those references not c i t e d i n the t e x t are l i s t e d below. A a l d e r s , L . E . , and I . 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