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Classification of epilithic bryophyte communities in south-western British Columbia Velzen, Johannes Petrus van 1981

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CLASSIFICATION OF EPILITHIC BRYOPHYTE COMMUNITIES IN SOUTH-WESTERN BRITISH COLUMBIA. by  K a n d i d a a t s ' degree S t a t e U n i v e r s i t y U t r e c h t / H o l l a n d , 1978  A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE  THE FACULTY OF GRADUATE STUDIES Department o f Botany  We a c c e p t t h i s t h e s i s as conforming to t h e r e q u i r e d standard  THE UNIVERSITY OF BRITISH COLUMBIA May 1981  ©  Johannes P e t r u s van V e l z e n , 1981  In p r e s e n t i n g t h i s  thesis i n p a r t i a l  f u l f i l m e n t o f the  requirements f o r an advanced degree a t the o f B r i t i s h Columbia, I agree t h a t it  freely  University  the L i b r a r y  a v a i l a b l e f o r r e f e r e n c e and study.  s h a l l make I further  agree t h a t p e r m i s s i o n f o r e x t e n s i v e copying o f t h i s f o r s c h o l a r l y purposes may  be granted by the head o f my  department o r by h i s o r her r e p r e s e n t a t i v e s . understood t h a t for  financial  copying or p u b l i c a t i o n  gain  of  J$>crt(*nij  The U n i v e r s i t y o f B r i t i s h 2075 Wesbrook P l a c e Vancouver, Canada V6T 1W5 Date  (2/79)  of t h i s  It i s thesis  s h a l l not be a l l o w e d without my  permission.  Department  thesis  Columbia  written  through a succession of sinuous rocky banks and of c l i f f s and moss-covered rocks I g l i d e d along  Liu Tsung-yuan  (773-819).  iii  ABSTRACT  The present study attempts to c l a s s i f y the e p i l i t h i c (=growing on rock) bryophyte vegetation of south-western  B r i t i s h Columbia,  Canada. During the summer f i e l d season of 1980 around 350 releves (=samples) were taken o f bryophyte assemblages from rocky habitats that ranged i n e l e v a t i o n from sea l e v e l to subalpine and alpine areas. Small scale v a r i a t i o n was expressed i n the segregation o f various types of micro-environments on rocks, a l l characterized by t h e i r own bryophyte vegetation. On a l a r g e r s c a l e , the releves of these e p i l i t h i c bryophyte assemblages were divided into four major groups: (1) exposed, dry rocks (2) shaded, humid rocks (3) s i l i c e o u s rocks along streams and (4) calcareous rocks. Within these major groups c l u s t e r analyses and tabular s o r t i n g , combined, were u t i l i z e d t o detect patterns i n the data set. The subsequent recognition o f groups of s i m i l a r releves formed the basis f o r the segregation of separate vegetation units (syntaxa). These syntaxa were defined by diagnostic species (character and d i f f e r e n t i a l  species)  and f u r t h e r characterized by m i c r o - s i t e s p e c i f i c i t y of species and/or communities and t h e i r geographic d i s t r i b u t i o n throughout the study area. Comparison to and i n t e g r a t i o n i n already e x i s t i n g c l a s s i f i c a t i o n schemes was attempted. In p a r t i c u l a r , f l o r i s t i c and s o c i o l o g i c a l a f f i n i t i e s assigned to the e p i l i t h i c bryophyte a s s o c i a t i o n s i n south  iv  western B r i t i s h Columbia were r e l a t e d t o those i n Europe. Where n e c e s s a r y and a p p r o p r i a t e ,  new s y n t a x a were d i s t i n g u i s h e d  F i n a l l y , the a p p l i c a t i o n o f ordination  and d e f i n e d .  techniques, although  l i m i t e d i n t h e p r e s e n t s t u d y , was u t i l i z e d as an a d d i t i o n a l means t o d i s p l a y and i n t e r p r e t p a t t e r n s i n t h e d a t a s e t . The p r i o r p a r a l l e l e d with  f i e l d d a t a on e c o l o g i c a l  classification,  s i t e c h a r a c t e r i s t i c s , already  showed some r e l a t i o n s h i p between r e l e v e s . T h i s p r o v i d e d a b a s i s f o r i n t e r p r e t i n g t h e o r d i n a t i o n s , w h i c h , i n t u r n , f u r t h e r e x p r e s s e d and c l a r i f i e d r e l a t i o n s h i p s o f releves ment.  t o one a n o t h e r and t o t h e e n v i r o n -  V  TABLE OF  CONTENTS  1. INTRODUCTION  .1  2. LOCATION OF THE STUDY AREA  :  '  5  3. PHYSIOGRAPHY AND GEOLOGY OF THE STUDY AREA....  8  3.1. P h y s i o g r a p h y  8  3.1.1. L a n d f o r m s 3.1.2. G l a c i a t i o n  8 i n British  Columbia  12  3.1.3. V o l c a n i s m  14  3.2. G e o l o g y  16  3.2.1. B e d r o c k t y p e s Intrusive  16 igneous  Sedimentary  rock  17  rock  18  Volcanic rock  19  Metamorphic rock..  19  4. CLIMATE OF THE STUDY AREA  20  5. VEGETATION OF THE STUDY AREA  25  5.1. The C o a s t a l D o u g l a s - F i r Z o n e . . .  •  27  5.2. T h e C o a s t a l W e s t e r n H e m l o c k Zone  28  5.3. T h e S u b a l p i n e M o u n t a i n H e m l o c k Zone  30  5.4. T h e S u b a l p i n e 5.5. The I n t e r i o r  Engelmann S p r u c e - S u b a l p i n e D o u g l a s - F i r Zone.  5.6. T h e A l p i n e T u n d r a Zone  F i r Zone  31 32 33  vi  6. METHODS  ,  36  6.1. D e l i m i t a t i o n o f e p i l i t h i c b r y o p h y t e communities  36  6.2. Sampling methods  37  6.2.1. Reconnaissance  37  6.2.2. Choice o f r e l e v e - s i t e and q u a d r a t - s i z e  38  6.2.3. E s t i m a t i o n o f s p e c i e s q u a n t i t y and s o c i a b i l i t y  40  6.2.4. E c o l o g i c a l c h a r a c t e r i z a t i o n o f t h e r e l e v e - s i t e  43  6.3. Data a n a l y s i s  45  6.3.1. Bryophyte i d e n t i f i c a t i o n  45  6.3.2. T a b u l a r comparison  46  6.3.3. Numerical t e c h n i q u e s  49  6.3.3.1. C l u s t e r a n a l y s i s  52  6.3.3.2. O r d i n a t i o n  54  7. SYNSYSTEMATICS  56  7.1. Terminology and nomenclature  56  8. ANALYSIS AND DESCRIPTION OF THE EPILITHIC BRYOPHYTE VEGETATION 8.1. Bryophyte a s s o c i a t i o n s from exposed, d r y r o c k s  60 62  8.1.1. S y n s y s t e m a t i c s  65  8.1.2. P h y t o g e o g r a p h i c a l s e t t i n g . . .  67  8.1.3.1. Order RACOMITRIETALIA HETEROSTICHI  67  8.1.3.2. Order POLYTRICHETALIA PILIFERI  81  8.2. Bryophyte a s s o c i a t i o n s from  shaded, humid r o c k s  88  8.2.1. S y n s y s t e m a t i c s  90  8.2.2. P h y t o g e o g r a p h i c a l s e t t i n g  91  8.2.3.1. Order LEPIDOZIETALIA REPTANTIS  93  vii  8.3. Bryophyte  a s s o c i a t i o n s from s i l i c e o u s r o c k s a l o n g streams  104  8.3.1. S y n s y s t e m a t i c s 8.3.2. P h y t o g e o g r a p h i c a l  107 setting  8.3.3.1. Order BRACHYTHECIETALIA PLUMOSI  110 I l l  8.3.3.2. Order PLATYHYPNIDIETALIA RUSCIFORMIS 8.4. Bryophyte  a s s o c i a t i o n s from c a l c a r e o u s r o c k s  134 139  8.4.1. S y n s y s t e m a t i c s  141  8.4.2. P h y t o g e o g r a p h i c a l s e t t i n g  143  8.4.3. A l l i a n c e E n c a l y p t i o n p r o c e r a e  144  8.4.4. E v a l u a t i o n o f t h e o r d i n a t i o n t e c h n i q u e s  153  9. SUMMARY  161  REFERENCES.  167  APPENDIX I . Abundance s c a l e  180  Sociability scale  181  APPENDIX I I . F i d e l i t y degrees APPENDIX I I I . Bryophyte  species l i s t :  182 Class Hepaticae.  184  C l a s s Musci  189  APPENDIX IV. O r d i n a t i o n data on t h e b r y o p h y t e v e g e t a t i o n from  APPENDIX V.  calcareous rocks  197  D i s t r i b u t i o n maps o f t h e b r y o p h y t e s y n t a x a  202  APPENDIX V I . T a b u l a r s o r t i n g t a b l e s and c l u s t e r a n a l y s e s f o r t h e f o u r major groups o f b r y o p h y t e v e g e t a t i o n r e l e v e s . . .  225  viii  LIST OF TABLES  TABLE I  S i m p l i f i e d o u t l i n e o f g e o l o g i c a l h i s t o r y i n south-western B r i t i s h Columbia  15  II  C l i m a t e data from weather s t a t i o n i n t h e study a r e a  24  III  Bryophyte  63  IV  R a c o m i t r i o - Andreaeetum p e t r o p h i l a e  V  Campy!opus a t r o v i r e n s - Paraleucobryum  VI  "Racomitrietum b r e v i p i s "  74  VII  Gymnomitrietum c o n c i n n a t i  77  VIII  Racomitrietum s u d e t i c i  80  IX  Racomitrio - Polytrichetum pi 1 i f e r i  83  X  Scapanietum americanae  86  XI  Bryophyte  89  XII  Diplophylletum albicans  94  XIII  M a r s u p e l l o - Scapanietum americanae  96  XIV  Claopodietum  XV  Bryophyte  XVI  Racomitrio - Seoulerietum aquaticae  a s s o c i a t i o n s from exposed, dry r o c k s . .  69 enerve  -facies  a s s o c i a t i o n s from shaded, humid r o c k s  bolanderi  a s s o c i a t i o n s from s i l i c e o u s r o c k s a l o n g streams.  Dichodontietum  72  101 105 113  pellucidi  XVII  B r a c h y t h e c i o - Conocephaletosum c o n i c a e  117  XVIII  Riccardio - Hygrobielletosum l a x i f o l i a e  121  XIX  B l i n d i o - Scapanietosum paludosae  123  XX  P h i l o n o t i s fontana - f a c i e s  125  XXI  B l i n d i a acuta - f a c i e s  126  ix  TABLE XXII  M a r s u p e l l o - Nardietum s c a l a r i s  129  XXIII  Hypnetum d i e c k i i  132  XXIV  Hygrohypnetum s m i t h i i  136  XXV  Bryophyte  a s s o c i a t i o n s from c a l c a r e o u s rocks  140  XXVI  Lophozio  - Bryoerythrophylletum r e c u r v i r o s t r i  145  XXVII  Metaneckeretum m e n z i e s i i  XXVIII B a r b u l a v i n e a l i s - P l a t y d i c t y a jungermannioides mannia a t r o v i r e n s - community  147 - Junger150  X  LIST OF FIGURES  F i g u r e 1.  Reference  map f o r t h e study area  7  F i g u r e 2.  P h y s i o g r a p h i c s u b d i v i s i o n s and g e n e r a l i z e d geology o f the s t u d y area  F i g u r e 3.  10  Annual p r e c i p i t a t i o n i n south-western  British  Columbia, given i n centimeters F i g u r e 4.  V e g e t a t i o n zones i n south-western  F i g u r e 5.  Nested  22 B r i t i s h Columbia...  26  p l o t t e c h n i q u e , used f o r t h e d e t e r m i n a t i o n o f  a minimum r e l e v e ( p l o t ) - s i z e  41  F i g u r e 6.  S p e c i e s - a r e a curve  42  F i g u r e 7.  Categories o f rock s u r f a c e h a b i t a t s  44  F i g u r e 8.  P r i n c i p a l Components A n a l y s i s o f t h e 27 l i m e s t o n e releves  F i g u r e 9.  154  S c a t t e r - d i a g r a m o f t h e 27 l i m e s t o n e r e l e v e s from t h e P.C.A.-cent  155  F i g u r e 10.  P o l a r o r d i n a t i o n o f t h e 27 l i m e s t o n e r e l e v e s  157  F i g u r e 11.  C l u s t e r a n a l y s i s o f t h e 27 l i m e s t o n e r e l e v e s  158  F i g u r e 12.  C l u s t e r a n a l y s i s o f P.C.A.-scores.(7 axes) o f t h e 27 limestone releves  160  xi  ACKNOWLEDGEMENTS  I am very g r a t e f u l t o my graduate  s u p e r v i s o r , Dr. W. B.  S c h o f i e l d , f o r h i s support o f my b r y o - s o c i o l o g i c a l s t u d y , i n a r r a n g i n g the f i n a n c e f o r my f i e l d w o r k and r e s e a r c h and i n enhancing and u n d e r s t a n d i n g  o f n o r t h e r n hemisphere b r y o p h y t e s .  my knowledge  I offer sincere  a p p r e c i a t i o n f o r h i s good a d v i c e d u r i n g t h e term o f my study and i n t h e w r i t i n g o f t h i s t h e s i s . I am a l s o i n d e b t e d t o him t h e use o f many u n p u b l i s h e d B r i t i s h Columbia d i s t r i b u t i o n maps o f l i v e r w o r t s and mosses. I thank Dr. G. E. B r a d f i e l d f o r t h e use o f h i s t a b u l a r s o r t i n g program VEGTAB and f o r h i s v a l u a b l e h e l p i n t h e computer a n a l y s e s o f the v e g e t a t i o n d a t a . A l s o , I thank Rob Scagel f o r h i s time and e f f o r t to h e l p me c a r r y i n g o u t these a n a l y s e s and d i s c u s s i n g t h e i r  results.  To my summer f i e l d a s s i s t a n t , A l l e n Leong, I o f f e r my apprec i a t i o n f o r h i s v a l u a b l e h e l p and p l e a s a n t companionship i n t h e f i e l d . I am i n d e b t e d t o my p a r e n t s , Mr. and Mrs. J . van Velzen-van Meer, f o r t h e i r encouragement throughout  the y e a r s o f my academic  education. In p a r t i c u l a r , g r a t e f u l l y r e c o g n i z e d i s t h e time I spent a t t h e I n s t i t u t e f o r S y s t e m a t i c Botany o f t h e S t a t e U n i v e r s i t y o f U t r e c h t , H o l l a n d where my i n t e r e s t i n b r y o l o g y was born and grew under t h e e l e c t r i f y i n g i n f l u e n c e o f Dr. S. R. G r a d s t e i n . I thank t h e f o l l o w i n g persons who h e l p e d i n t h e t y p i n g o f t h e manuscript  and t h e f i n a l  t h e s i s : E t a T h i e s s e n , Dr. R. T u r k i n g t o n .  xi i  I thank Deanne Brethower, O l i v i a Lee and John Pinder-Moss o f the Herbarium o f the U n i v e r s i t y o f B r i t i s h C o l u m b i a , f o r t h e i r h e l p i n t y p i n g t h e l a b e l s f o r the h e r b a r i u m specimens. To many p e o p l e I would l i k e t o e x p r e s s my s i n c e r e thanks and appreciation  f o r h e l p i n g me, m a t e r i a l l y o r m o r a l l y :  Vijko Lukkien,  Paul van Westendorp, A l l e n Banner, S h e i l a Humphrey, Leni  Gelten,  Nechemjah Cohen, B e n i t o Tan, Ross H i l l , T e r r y M c i n t o s h , John Wehr, Debby Donaldson, T r e v o r Goward, W i l f N i c h o l l s , Tim and Suzanne Thompson, M i c h a e l Dunn.  1  1.  INTRODUCTION.  The o b j e c t i v e s o f t h i s study a r e t o sample, a n a l y z e and c l a s s i f y t h e e p i l i t h i c b r y o p h y t e communities as t h e y o c c u r i n s o u t h western B r i t i s h Columbia.  D e s c r i p t i o n s o f the separate v e g e t a t i o n  u n i t s ( s y n t a x a ) a r e c o n c e n t r a t e d on t h e r e c o g n i t i o n o f d i a g n o s t i c s p e c i e s ( c h a r a c t e r s p e c i e s and d i f f e r e n t i a l  species), micro-site  s p e c i f i c i t y o f s p e c i e s and communities and t h e i r g e o g r a p h i c b u t i o n throughout  distri-  t h e study a r e a .  S e c o n d l y , comparison  t o and i n t e g r a t i o n i n a l r e a d y e x i s t i n g  c l a s s i f i c a t i o n systems i s attempted.  E p i l i t h i c bryophyte a s s o c i a t i o n s  have been d e s c r i b e d i n Europe ( a . o . H e r t e l 1974, Hebrard  1978 and  Neumayr 1971). T h e i r c l a s s i f i c a t i o n , however, becomes more meaningful w i t h t h e e x p a n s i o n o f knowledge o f e p i l i t h i c b r y o p h y t e a s s o c i a t i o n s o f North America America  and temperate A s i a . I t i s e v i d e n t t h a t i n North  ( i n c l u d i n g t h e west c o a s t ) s i m i l a r a l l i a n c e s and a s s o c i a t i o n s  a r e found (Barkman 1969, Hubschmann  1978).  F i n a l l y , m u l t i v a r i a t e s t a t i s t i c a l analyses are a p p l i e d to determine whether o b j e c t i v e t e c h n i q u e s o f data a n a l y s i s s u b s t a n t i a t e s t a b u l a t i o n methods, so e s s e n t i a l  i n the c l a s s i f i c a t i o n o f v e g e t a t i o n .  In t h e study a r e a and t h e a d j a c e n t r e g i o n s o f t h e p r e s e n t s t u d y , some e c o l o g i c a l r e s e a r c h has been done on t h e bryophyte  (and l i c h e n )  v e g e t a t i o n o f Vancouver I s l a n d ( S z c z a w i n s k i 1953, Hubschmann 1978) and t h e Olympic P e n i n s u l a i n Washington, U.S.A. (Hoffman and K a z m i e r s k i S z c z a w i n s k i ( 1 9 5 3 ) , i n an e x t e n s i v e study on t h e c o r t i c o l o u s  1969).  2  and 1 i g n i c o l o u s p l a n t communities i n the f o r e s t a s s o c i a t i o n s o f Douglas F i r f o r e s t on Vancouver I s l a n d , d e s c r i b e d  the  r e l a t i o n s between  o c c u r r e n c e , abundance, dominance, c o n s t a n c y a n d . v i g o u r o f b r y o p h y t e s and  l i c h e n s and  1ignicolous  he d i s t i n g u i s h e d t e n s o c i a t i o n s o f c o r t i c o l o u s  and  ( = e p i x y l i c ) cryptogams.  Hubschmann ( 1 9 7 8 ) , who  s t u d i e d moss communities on Vancouver  I s l a n d , a p p l i e d t h e Z i i r i c h - M o n t p e l 1 i e r School method o f  vegetation  c l a s s i f i c a t i o n . Beside aquatic, t e r r e s t r i a l  and  communities, he d i s t i n g u i s h e d and  seven e p i l i t h i c b r y o p h y t e  a s s o c i a t i o n s and  provided  described  e p i p h y t i c bryophyte  some i n f o r m a t i o n on physiognomy and  t r i b u t i o n of these vegetation  dis-  units.  Knowledge on the e c o l o g y o f b r y o p h y t e s i s s c a n t i n comparison with that of higher p l a n t s . Studies were m a i n l y l i s t s o f s p e c i e s  o f b r y o p h y t e s up t o about  i n c e r t a i n areas and  w i t h comments on the environmental r e l a t i o n s o r  1930  h a b i t a t s , sometimes micro-habitats.  L a t e r works c o n t a i n data on abundance, c o v e r and  frequency  of s p e c i e s . E s p e c i a l l y i n Europe, and t o a l e s s e r degree i n J a p a n , the a p p l i c a t i o n o f the BraurfBlariquet r e l e v e t e c h n i q u e l e d t o a  classifi-  c a t o r y system o f b r y o p h y t e communities on a v a r i e t y o f s u b s t r a t a as t r e e s (Barkman 1969,  Iwatsuki  1964), d e c a y i n g wood ( P h i l i p p i  1960), rocks  1965)  and  soil  ( H e r t e l 1974,  such  Sjogren  (Hubschmann 1975,  Wald-  heim 1947). North American b r y o - e c o l o g i c a l has  concentrated  m a i n l y on s t a t i s t i c a l  community s t r u c t u r e and c o m p o s i t i o n ,  research, e s p e c i a l l y recently, and m u l t i v a r i a t e a n a l y s i s  d i s t r i b u t i o n and  ecological  of rela-  3  t i o n s h i p s between b r y o p h y t e s , t h e i r s u b s t r a t e and environment Redfearn 1957, 1960, Foote 1963, 1966, S l a c k 1971, C u l b e r s o n  (a.o. 1955,  H a l e 1955). Earlier bryological  s t u d i e s , p a r t i c u l a r l y i n the United S t a t e s ,  have r e s u l t e d i n t h e r e c o g n i t i o n o f more o r l e s s d i s c r e t e v e g e t a t i o n u n i t s , e s p e c i a l l y on t r e e s . C a i n and Sharp ( 1 9 3 8 ) , u t i l i z i n g  coverage  and presence o f b r y o p h y t e s i n c e r t a i n f o r e s t t y p e s o f the Great Smokey M o u n t a i n s , were a b l e t o c l a s s i f y c o r t i c o l o u s ( = b a r k - i n h a b i t i n g ) communities. In a n o t h e r s t u d y from t h e same g e o g r a p h i c a r e a , B i l l i n g s and Drew (1938) i n v e s t i g a t e d bark f a c t o r s a f f e c t i n g the d i s t r i b u t i o n o f c o r t i c o l o u s b r y o p h y t e communities. These workers d e s c r i b e d " u n i o n s " o f bryophyt e s , dominated by one o r two s p e c i e s . At Mountain L a k e , V i r g i n i a , P a t t e r s o n (1940) r e c o g n i z e d w e l l developed c o r t i c o l o u s " s o c i e t i e s " based upon t h e dominant s p e c i e s i n q u a d r a t s on random mature t r e e s . The o c c u r r e n c e and d i s t r i b u t i o n o f t h e s e s o c i e t i e s was  found t o depend upon d i f f e r e n t l e v e l s o f e v a p o r a t i o n a l  s t r e s s and d i f f e r e n t m o i s t u r e h o l d i n g c a p a c i t y o f bark s u b s t r a t a . F i n a l l y , P h i l l i p s (1948, 1951)  r e c o g n i z e d a s s o c i a t i o n s o f bark-  i n h a b i t i n g b r y o p h y t e s i n t h r e e r e g i o n s o f t h e S t a t e o f M i c h i g a n . The a s s o c i a t i o n s were based on and named a f t e r t h e dominant s p e c i e s ( t h o s e w i t h h i g h e s t c o v e r a g e , c o n s t a n c y and  vitality).  A major Canadian s t u d y employing t h e same types o f methods as the p r e v i o u s works i s the i n v e s t i g a t i o n by L e B l a n c (1962) on the e c o l o g y and p h y t o s o c i o l o g y o f c r y p t o g a m i c e p i p h y t e s i n s o u t h e r n Quebec.  4  Coverage and dominance w i t h i n quadrats l e d t o t h e d i s t i n c t i o n and recognition  of epiphytic  s o c i e t i e s o f b r y o p h y t e s and l i c h e n s .  In s h o r t , p h y t o s o c i o l o g i c a l  work i n t h i s study attempts t o  r e c o g n i z e and d e s c r i b e e p i l i t h i c b r y o p h y t e assemblages. Major a t t e n t i o n i s p a i d t o t h e i n v e s t i g a t i o n o f t h e s i g n i f i c a n c e o f u s i n g the t a b u l a t i o n technique, essential  i n t h e Braun-Blanquet method o f v e g e t a t i o n  c a t i o n , p a r a l l e l e d with  the a p p l i c a t i o n of s t a t i s t i c a l  classifi-  multivariate  techniques. In t h e hope t h a t some c o o r d i n a t i o n  can be a c h i e v e d between  t h e s e approaches i n t h e i r a p p l i c a t i o n t o b r y o - e c o l o g y , t h e p r e s e n t study has been undertaken.  5  2. LOCATION OF THE STUDY AREA.  The  area o f t h e p r e s e n t  study i s l o c a t e d i n s o u t h - w e s t e r n  B r i t i s h Columbia, Canada's westernmost p r o v i n c e . I t s l o c a t i o n i s shown i n F i g u r e 1 . B o u n d a r i e s o f t h e area a r e e s s e n t i a l l y t h e m a i n l a n d c o a s t l i n e on t h e w e s t ; t h e Canada-U.S.A. b o r d e r on t h e s o u t h ; t h e mount a i n range south o f t h e C h i l l i w a c k R i v e r and C h i l l i w a c k Lake t o the s o u t h - e a s t ;  and, f i n a l l y , Pemberton t o t h e n o r t h .  To p r o v i d e f a m i l i a r i t y w i t h t h i s r e g i o n and a b a s i s f o r understanding  the v a r i a t i o n i n e p i l i t h i c vegetation, the physio-  graphy, g e o l o g y , c l i m a t e and v e g e t a t i o n o f t h e s t u d y a r e a a r e d e s c r i b e d i n t h e f o l l o w i n g c h a p t e r s . Regional  climate  (macroclimate),  p h y s i o g r a p h y , and geology i n f l u e n c e t h e ( v a s c u l a r ) v e g e t a t i o n which develops i n a r e g i o n . A l l f o u r elements determine t h e a v a i l a b i l i t y of habitats f o r bryophytes,  i n g e n e r a l , and t h e development o f  e p i l i t h i c b r y o p h y t e assemblages, i n p a r t i c u l a r . The  p r o x i m i t y t o t h e P a c i f i c Ocean and t h e v a r i e d topography  o f t h e study a r e a , c r e a t i n g a g r e a t d i v e r s i t y o f h a b i t a t s , make south-western B r i t i s h Columbia an i d e a l l y s u i t a b l e r e g i o n f o r a study o f e p i l i t h i c b r y o p h y t e communities.  7  F i g u r e 1. R e f e r e n c e map f o r t h e study a r e a .  1.  B l a c k Tusk Meadows  2.  Brandywine  3.  Bridal  4.  Brittania  5.  C a l l a g h a n Creek  6.  Cheakamus R i v e r V a l l e y  7.  C h i l l i w a c k Lake  8.  Chipmunk Creek  9.  Diamond Head  Falls  Veil  Falls  Beach  10. Ford Mountain 11. G a r i b a l d i  Lake  12. Cypress Bowl 13. L i g h t h o u s e Park 14. L o n e t r e e Creek 15. Lynn Canyon 16. Mount Seymour 17. M u r r i n P r o v i n c i a l 18. N a i r n  Park  Falls  19. Panorama Ridge 20. P o r t e a u Camp 21. Shannon F a l l s 22. S l e s s e Creek 23. Stoney Creek 24. Sunset Creek 25. W h i s t l e r Mountain 26. White C l i f f P a r k , Horseshoe Bay 27. Sumas Mountain 28. A g a s s i z Mountain  8  3. PHYSIOGRAPHY AND GEOLOGY OF THE STUDY AREA.  3.1. P h y s i o g r a p h y .  3.1.1. Landforms.  The a r e a o f t h i s s t u d y l i e s w i t h i n t h e C o r d i l l e r a n R e g i o n , one o f t h e f i v e major p h y s i o g r a p h i c s u b d i v i s i o n s r e c o g n i z e d f o r Canada ( H o l l a n d 1976). An extreme v a r i e t y i n topography i s shown i n t h e presence o f an i m p r e s s i v e f i o r d - i n c i s e d c o a s t l i n e , v a l l e y s , mountains and canyons, r i v e r s and l a k e s . Of t h e s e v e r a l B r i t i s h Columbian p h y s i o g r a p h i c s u b d i v i s i o n s ( H o l l a n d 1976), two a r e r e p r e s e n t e d i n t h e s t u d y a r e a : t h e C o a s t a l Trough and t h e Coast Mountain Area which encompasses t h e Coast Mountains and t h e N o r t h e r n Cascade M o u n t a i n s . To the e a s t t h e s t u d y a r e a i s bounded by t h e I n t e r i o r P l a t e a u ( F i g u r e 2 ) . In t h e study a r e a t h e C o a s t a l Trough (1) i s r e p r e s e n t e d by a minute s e c t i o n o f t h e G e o r g i a Lowland (a narrow s t r i p on t h e m a i n l a n d southeastward from Horseshoe Bay) and by t h e F r a s e r Lowland. The G e o r g i a Lowland i s u n d e r l a i n by g r a n i t e r o c k s , J u r a s s i c and o l d e r , a f f e c t e d by e r o s i o n . The F r a s e r Lowland i s a p a r t o f t h e G e o r g i a Lowland, from which i t d i f f e r s i n b e i n g a l o w - l y i n g a r e a o f d e p o s i t i o n a l r a t h e r than e r o s i o n a l o r i g i n . I t s t r i a n g u l a r shape  includes the Fraser River d e l t a , extending  eastward from P o i n t Grey f o r 110 km t o L a i d l o w and s o u t h e a s t w a r d t o t h e c o a s t a t B e l l i n g h a m , Washington, U.S.A. The F r a s e r Lowland c o n s i s t s o f  10  F i g u r e 2. P h y s i o g r a p h i c s u b d i v i s i o n s o f the study a r e a .  and g e n e r a l i z e d geology  LEGEND. Physiographic 1. Coastal 2. I n t e r i o r  subdivisions:  Trough Plateau  3. Coast Mountain Area A. Coast  Mountains  B. Cascade  Height o f Mountains:  Generalized  Mountains  A  below 800 m  over 800 m  geology:  F o l i a t e d metamorphic r o c k .  ^  V  A .  A  Intrusive  igneous  r o c k , L a t e Mesozoic o r T e r t i a r y  Folded and f a u l t e d v o l c a n i c and sedimentary rock c h i e f l y Mesozoic  F l a t o r g e n t l y d i p p i n g sedimentary r o c k . Cretaceous and younger  11  e x t e n s i v e low h i l l s r a n g i n g i n e l e v a t i o n from 15 t o 300 meters s e p a r a ted  by w i d e , f l a t - b o t t o m e d v a l l e y s (Armstrong 1954). The d e l t a o f t h e  F r a s e r R i v e r has been the s i t e o f s e d i m e n t a r y d e p o s i t i o n s i n c e the l a t e C r e t a c e o u s . Deep d r i l l i n g has shown t h a t the g r a n i t e basement i s o v e r l a i n by C r e t a c e o u s , T e r t i a r y and Quaternary s e d i m e n t a r y r o c k s . Recent d e p o s i t s , s t i l l  i n the p r o c e s s o f f o r m a t i o n , c o n s i s t o f  d e l t a i c and f l o o d - p l a i n d e p o s i t s o f the F r a s e r R i v e r as i t b u i l d s i t s d e l t a seaward at t h e r a t e o f about 8.5 m  a y e a r ( H o l l a n d 1976).  The major p h y s i o g r a p h i c s u b d i v i s i o n ( H o l l a n d 1976) r e p r e s e n t e d i n the study a r e a i s t h e Coast Mountain Area (3) which encompasses two r e g i o n s , t h e Coast Mountains (3A) and t h e Cascade Mountains ( 3 B ) . As an unbroken mountain c h a i n t h e Coast Mountains e x t e n d from t h e i r s o u t h end at  t h e F r a s e r R i v e r northwestward a l o n g t h e m a i n l a n d c o a s t l i n e o f  B r i t i s h Columbia i n t o t h e Yukon. Of t h e s e Coast Mountains t h e P a c i f i c Ranges, making up t h e major p a r t o f t h e a r e a o f t h e p r e s e n t s t u d y , comprise t h e e s s e n t i a l l y g r a n i t i c mountains e x t e n d i n g s o u t h e a s t w a r d from the  B e l l a C o o l a R i v e r f o r about 480 km t o the F r a s e r R i v e r mouth. The  P a c i f i c Ranges c o n t a i n the h i g h e s t peaks i n the Coast Mountains: Mount Haddington ( e l e v a t i o n 4017 m) l i e s n o r t h o f t h e study a r e a . High  mountains  c l o s e , o r w i t h i n , t h e s t u d y area i n c l u d e Mount G i l b e r t (3110 m) and Mount Garibaldi  (2679). The Coast Mountains comprise s e d i m a n t a r y and v o l c a n i c  r o c k s ( m i d d l e J u r a s s i c and o l d e r age) w i t h Coast i n t r u s i o n s o f r o c k s t h a t a r e e s s e n t i a l l y g r a n o d i o r i t e and q u a r t z d i o r i t e . The Cascade Mountains extend northward from the n o r t h e r n b o r d e r of  C a l i f o r n i a , through Oregon and Washington  i n t o south-western B r i t i s h  12  Columbia. The  Cascades are composed o f P a l e o z o i c and Mesozoic sedimen-  t a r y and v o l c a n i c r o c k s , s t r o n g l y f o l d e d , metamorfosed and g r a n i t e b a t h o l i t h s . The approximately  intruded  summits o f t h e peaks and r i d g e s a t t a i n  by  an  u n i f o r m e l e v a t i o n , w i t h the e x c e p t i o n o f some o f  the  h i g h e s t peaks i n Oregon and Washington such as Mount Baker (3286 m) G l a c i e r Peak (3180  m),  Mount S t . Helens (3225 m) and  Mount Hood (3745  A l l are v o l c a n i c cones b u i l t up by t h e a c c u m u l a t i o n o f the and r e c e n t l a v a and ash  and m).  pleistocene  upon t h e T e r t i a r y e r o s i o n s u r f a c e . R i c h i n  mountain f e a t u r e s such as hanging v a l l e y s , t a r n s and g l a c i a t e d rock s u r f a c e s , t h e Cascades' rugged peaks, r i d g e s and  cirques give evidence  of extensive alpine g l a c i a t i o n . To the e a s t the study area s u b d i v i s i o n ( H o l l a n d 1976)  i s bounded by a n o t h e r  physiographic  termed here t h e I n t e r i o r P l a t e a u ( 2 ) . Most  o f t h i s area i s d r a i n e d by t h e F r a s e r R i v e r and  i t s tributaries. Its  f l a t o r g e n t l y r o l l i n g s u r f a c e , which l i e s f o r most p a r t below 900 i s covered with g l a c i a l  d r i f t and has  3.1.2. G l a c i a t i o n i n B r i t i s h  few exposures o f bedrock.  Columbia.  A b a s i c knowledge o f g l a c i a t i o n i s important  i n the  o f landforms i n B r i t i s h Columbia, because the P r o v i n c e was g l a c i a t e d d u r i n g the P l e i s t o c e n e . The effects of g l a c i a l consinan  i s considered  consideration intensely  topography everywhere shows t h e  e r o s i o n and d e p o s i t i o n , almost e n t i r e l y o f t h e  stage. Holland  m,  (1976) s t a t e s t h a t g l a c i a t i o n i n mountain  to f o l l o w a c y c l e . I t begins w i t h the formation  Wisregions  of small  13  c i r q u e g l a c i e r s w h i c h , by c o n t i n u e d growth, expand i n t o mountain  and  v a l l e y g l a c i e r s , then f o r m a t i o n o f a mountain i c e - c a p whose movement i s c o n t r o l l e d by u n d e r l y i n g topography, and  u l t i m a t e l y , a t the maximum  s t a g e , by a r e g i o n a l i c e - s h e e t c o n t r o l l e d f o r t h e most p a r t by c l i m a t e . As the i c e c o v e r d i m i n i s h e s t h e r e i s r e g r e s s i o n t o the mountain i c e cap w h i l e l o b e s o f the main i c e - s h e e t p e r s i s t l o n g e r i n c o l d e r , p r o t e c t e d v a l l e y s (Mathews 1951). The  l a s t i c e t o remain i s i n s m a l l c i r q u e  g l a c i e r s as i n t h e b e g i n n i n g o f t h e The  cycle.  C o r d i l l e r a n Region i n B r i t i s h Columbia was  by t h e C o r d i l l e r a n i c e - s h e e t . The  covered  entirely  P l e i s t o c e n e began w i t h t h e accumu-  l a t i o n o f i c e a t s e v e r a l major c e n t r e s and numerous s u b o r d i n a t e ones ( H o l l a n d 1976). The  d i s t r i b u t i o n o f p r e s e n t day g l a c i e r s , t o some-extent  i n d i c a t e major, centres.;;of: f o r m a t i o n o f P l e i s t o c e n e i c e . With r e s p e c t t o t h e p r e s e n t s t u d y area t h e s e were i n the P a c i f i c Ranges o f the Coast Mountain area between B e l l a Cool a and  Garibaldi.  I c e moved a c r o s s B r i t i s h Columbia from the Coast Mountains i n a g e n e r a l e a s t e r l y d i r e c t i o n a c r o s s t h e F r a s e r P l a t e a u . Elsewhere i n t h e v i c i n i t y o f t h e p r e s e n t study area t h e i c e has r e c o r d e d t h e d i r e c t i o n o f i t s movement by t h e landforms  i t c r e a t e d . The  i c e moved westward from  the mainland  a c r o s s t h e n o r t h e r n end o f Vancouver I s l a n d , southward and  southeastward  down t h e S t r a i t o f Georgia and southwestward a c r o s s t h e  s o u t h e r n end o f Vancouver I s l a n d and, f i n a l l y , southward down Pudget Sound i n t o Washington. Evidence has accumulated t o i n d i c a t e t h a t t h e r e were a t l e a s t two major advances o f t h e C o r d i l l e r a n i c e - s h e e t , s e p a r a t e d by an glacial  s t a g e . L a t e P l e i s t o c e n e events  i n south-western  British  interColumbia  14  a r e summarized i n T a b l e I . The  P l e i s t o c e n e i c e , w i t h a t h i c k n e s s o f as  much as 2400 m i n some a r e a s , depressed e a r l i e r l e v e l . Since disappearance  the l a n d w i t h r e s p e c t t o i t s  o f t h e i c e - s h e e t , the l a n d s u r f a c e  has r i s e n , and r e g a i n e d i t s mountainous topography from b e f o r e t h e  onset  of the P l e i s t o c e n e .  3.1.3. V o l c a n i s m .  In B r i t i s h Columbia v o l c a n i c a c t i v i t y has o c c u r r e d s i n c e midT e r t i a r y t i m e . With r e s p e c t t o the p r e s e n t study area s e v e r a l c e n t r e s o f P l e i s t o c e n e and r e c e n t v o l c a n i c a c t i v i t y extend  in a line  northward  from Mount G a r i b a l d i . There i s evidence o f some a c t i v i t y w i t h i n the few hundred y e a r s , much o f i t h a v i n g taken p l a c e s u f f i c i e n t l y  last  recent  t h a t the c h a r a c t e r i s t i c v o l c a n i c l a n d f o r m s , such as cones and l a v a p l a i n s have been l i t t l e m o d i f i e d by e r o s i o n and g l a c i a t i o n (McKee 1972). Within h i s t o r i c  times t h e r e have been a c t i v e volcanoes  i n the  Cascade Mountains o f Washington and Oregon. Of these p l e i s t o c e n e s t r a t o v o l c a n o e s , Mount Baker e m i t t e d steam and t o x i c fumes d u r i n g 1975 melted  glacial  which  i c e . More r e c e n t l y Mount S t . Helens has been more a c t i v e  w i t h a major e r u p t i o n i n May  1980  and a subsequent decrease  in size.  15  Table I . S i m p l i f i e d o u t l i n e o f g e o l o g i c a l h i s t o r y i n south-western B r i t i s h Columbia ( a f t e r Godfrey 1977).  AGE ERA  (10  6  yrs)  GEOLOGICAL EVENTS  PERIOD (Epoch) (Recent)  c a . 2.5  QUATERNARY (PIeistocene) cu c OJ  27  cm  >o  o cu  65  o cu  to  Q_  CRETACEOUS (Upper)  o I—I  o  OO  Glacial erosion; deposits of d r i f t , outwash; i c e - m a r g i n a l d e p o s i t s . L o w - r e l i e f surface u p l i f t e d again along I s l a n d and Coast M o u n t a i n s ; i n c r e a s e d steam e r o s i o n ; f o r m a t i o n o f mountainous topography. D e p o s i t i o n s o f sediments w i t h i n c o a s t a l t r o u g h and on w e s t e r n s i d e o f I s l a n d Mtns r e d u c t i o n o f l a n d s u r f a c e t o one o f low r e l i e f ; v o l c a n i s m o f s o u t h e r n Vancouver Island.  cu cu  O  u p l i f t o f land.  Di  o  M  Post-glacial  110  Ancient erosion land surface u p l i f t e d , 2 axes o f g r e a t e s t u p l i f t were a l o n g I s l a n d and Coast Mtns. ( s e p a r a t e d by an ancestral trough). Land s u r f a c e o f low r e l i e f ; e r o s i o n o f mountains l e d t o t h e f o r m a t i o n o f s e d i ments which were d e p o s i t e d i n marine and b r a c k i s h water b a s i n s .  16  3.2.  Geology.  The Coast Range orogeny, which l a s t e d from e a r l i e s t  Jurassic  time through post Lower C r e t a c e o u s , t o g e t h e r w i t h e x t e n s i v e stream and g l a c i a l  e r o s i o n have dominated the g e o l o g i c a l h i s t o r y o f the study  a r e a . Much o f t h i s g e o l o g i c a l h i s t o r y i s r e c o r d e d by landforms many o f which a r e s c u l p t u r e d on bedrock by t h e f o r c e s o f e r o s i o n ( H o l l a n d  1976).  Since t h i s study deals w i t h e p i l i t h i c bryophyte vegetations i t i s i m p o r t a n t t o t r e a t t h e d i f f e r e n t bedrock t y p e s on t h e b a s i s o f the p h y s i c a l and chemical c h a r a c t e r i s t i c s t h a t i n f l u e n c e t h e o v e r - a l l  form  o f t h e topography. S u b s e q u e n t l y , t h e s e l a n d f o r m t y p e s r e f l e c t t h e a v a i l a b i l i t y o f r o c k s u b s t r a t a and h a b i t a t s and determine t h e of  development  v a s c u l a r p l a n t v e g e t a t i o n , on w h i c h , i n t u r n , some e p i l i t h i c  b r y o p h y t e assemblages  are e c o l o g i c a l l y  dependent.  3.2.1. Bedrock t y p e s .  The landforms w i t h i n t h e p h y s i o g r a p h i c r e g i o n s d i s c u s s e d b e f o r e are  commonly u n d e r l a i n by r e l a t e d bedrock t y p e s and have undergone t h e  same geomorphic  h i s t o r y . The c h a r a c t e r o f t h e s e landforms i s i n f l u e n c e d  by t h e r e a c t i o n o f bedrock t o w e a t e r i n g and e r o s i o n . The r a t e o f p h y s i c a l w e a t h e r i n g o r mechanical d e s i n t e g r a t i o n o f a r o c k i s i n f l u e n c e d by t h e presence o f Tines o f weakness and pore spaces t h a t a l l o w w a t e r , . a i r and temperature changes t o p e n e t r a t e beneath t h e s u r f a c e . Thus, d e n s e l y j o i n t e d r o c k s such as b a s a l t , o r v e r y porous m a t e r i a l such as  17  sandstone, a r e more r e a d i l y g r a n i t e s (Ryder  f r a c t u r e d than m a s s i v e , s p a r s e l y j o i n t e d  1978).  The m i n e r a l o g i c a l c o m p o s i t i o n o f a r o c k determines  i t s suscep-  t i b i l i t y t o chemical w e a t h e r i n g , s i n c e m i n e r a l s d i f f e r i n t h e i r  resis-  tance t o chemical a t t a c k .  I n t r u s i v e igneous rock ( s e e F i g u r e 2)  These a r e t y p i c a l l y c o a r s e c r y s t a l l i n e r o c k s t h a t i n B r i t i s h Columbia range from s y e n i t e , through g r a n i t i c They a r e r e l a t i v e l y r e s i s t a n t largely  types, to d i o r i t e .  t o w e a t h e r i n g s i n c e they a r e composed  o f d u r a b l e m i n e r a l s ( q u a r t z , h o r n b l e n d e , f e l d s p a r ) a r r a n g e d as  a cohesive f a b r i c o f i n t e r l o c k i n g  crystals.  As a r e s u l t , s l o p e s on  t h e s e r o c k s a r e g e n e r a l l y s t e e p and topography  i s rugged.  However, d e s p i t e t h e i r r e l a t i v e r e s i s t a n c e t o w e a t h e r i n g , some disintegration  and d e c o m p o s i t i o n o f t h i s t y p e o f bedrock  inevitably  o c c u r s . Mechanical breakdown through p r o c e s s e s such as f r o s t  shattering  tends t o produce r e l a t i v e l y l a r g e fragments. Extremely c o a r s e r u b b l e on c o l l u v i a l s l o p e s and l a r g e b o u l d e r s i n t i l l  characterize this  t y p e . A c o m b i n a t i o n o f p h y s i c a l and chemical w e a t h e r i n g - t y p i c a l l y granular disintegration  terrain causes  (Ryder 1978). Components l i k e c a l c i c f e l d s p a r a r e  most s u s c e p t i b l e t o a t t a c k by chemical w e a t h e r i n g ; c r y s t a l s  o f quartz  and potash and s o d i c f e l d s p a r a r e r e l e a s e d t o form a sandy o r g r i t t y r e s i d u e . T i l l s o f t h i s r e s i d u e and much sandy outwash a r e thus a s s o c i a t e d w i t h areas o f c o a r s e c r y s t a l l i n e r o c k s .  18  Sedimentary r o c k s (see F i g u r e 2)  The r e s i s t a n c e o f s e d i m e n t a r y r o c k s t o w e a t h e r i n g and e r o s i o n depends upon f a c t o r s such as m i n e r a l o g i c a l c o m p o s i t i o n , degree o f i n d u r a t i o n , i . e . t h e h a r d e n i n g through c e m e n t a t i o n , p r e s s u r e and h e a t . P o o r l y i n d u r a t e d r o c k s , such as c l a y , mudstone and s h a l e , d i s i n t e g r a t e r e a d i l y and a r e e a s i l y eroded. Well i n d u r a t e d , massive sandstone ( e.g. d e p o s i t i o n s on t h e I s l a n d s i n t h e S t r a i t o f Juan de Fuca and G e o r g i a 2 S t r a i t as w e l l as t h e Sumas E s c a r p m e n t ) , l i m e s t o n e s  (occurring within  t h e study a r e a i n t h e C h i l l i w a c k R i v e r v a l l e y ) and conglomerates a r e r e l a t i v e l y r e s i s t a n t to erosion. P h y s i c a l d i s i n t e g r a t i o n o f s e d i m e n t a r y r o c k s produces  fragments  o f a s i z e r e l a t e d t o the s p a c i n g o f l i n e s o f weakness i n t h e bedrock. Thus s h a l e s , which a r e t h i n l y bedded, produce s m a l l , p l a t y p a r t i c l e s t h a t r e a d i l y d i s i n t e g r a t e f u r t h e r . At the o t h e r extreme conglomerates shed l a r g e b l o c k s which s u r v i v e i n t a c t d u r i n g g l a c i a l  transport. Besides t h i s  physical d i s i n t e g r a t i o n , the weathering of sedimentary rocks a l s o r e s u l t s from chemical removal o f t h e cementing agent by s o l u t i o n o r o t h e r means. The t e r r a i n developed upon f a u l t e d , f o l d e d and s t e e p l y d i p p i n g s e d i m e n t a r y r o c k s shows a s t r o n g r e l a t i o n s h i p between r o c k t y p e , s t r u c t u r e and topography. I n d i v i d u a l  r i d g e s o r mountains a r e commonly asymmetric  w i t h s t e e p e r s c a r p and g e n t l e r d i p s l o p e s ; d i p s o f o v e r 45^ produce e x t r e m e l y j a g g e d r i d g e s , w h i l e prominent c l i f f s commonly c o i n c i d e w i t h f a u l t p l a n e s o r s t e e p bedding p l a n e s (Ryder 1978).  19  V o l c a n i c r o c k s ( s e e F i g u r e 2)  Topography upon t h e s e r o c k types tends t o be s i m i l a r t o t h a t developed  on sedimentary  rocks o f s i m i l a r s t r u c t u r e (Ryder  1978).  F l a t l y i n g l a v a f l o w s form p l a t e a u s bounded by escarpments and stepped h i l l s i d e s . T e r r a i n o f t h i s type i s widespread  i n South-central  British  Columbia.  Metamorphic r o c k s ( s e e F i g u r e 2 ) .  No p a r t i c u l a r t y p e o f t e r r a i n can be a s c r i b e d t o metamorphic rocks i n general. T h e i r r e s i s t a n c e t o weathering upon t h e i r i n d i v i d u a l  and e r o s i o n depends  characteristics.  Many metamorphic rocks such as g n e i s s and s c h i s t a r e f o l i a t e d , that i s , t h e minerals are segregated morphic q u a r t z i t e s a r e extremely In g e n e r a l , w e a t h e r i n g t e x t u r e d igneous  i n t o bands w i t h i n t h e r o c k . Meta-  r e s i s t a n t to weathering.  p r o d u c t s a r e s i m i l a r t o those o f c o a r s e  r o c k s o f t h e same c o m p o s i t i o n .  1. T h i s i s t h e c a l c u l a t e d r a t e o f advance a t a depth o f 90 m. 2. The l i m e s t o n e s o c c u r r i n g i n t h e study area a r e o f C a r b o n i f e r o u s age and a r e a s s o c i a t e d w i t h q u a r t z i t e and a r g i l l i t e i n s t e e p l y d i p p i n g d e p o s i t s . The l i m e s t o n e as sampled and a n a l y z e d by Goudge (1944) c o n t a i n e d over 3% o f s i l i c a , some had over 10%. The magnesium carbonate c o n t e n t , however, i s low as i t ranged from 1.3 t o 2.5%.  20  4. CLIMATE OF THE  STUDY AREA.  On a m a c r o - s c a l e , c l i m a t e a s s o c i a t e d w i t h t h e major  landscape  r e g i o n i s a product o f g e o g r a p h i c a l l o c a t i o n and l a r g e s c a l e topography. On t h e m i c r o - s c a l e , c o n d i t i o n s a t t h e e a r t h ' s s u r f a c e ( i . e . rock s u r f a c e ) a r e m o d i f i e d by l o c a l  p h y s i o g r a p h i c f a c t o r s such as s l o p e , a s p e c t ,  e l e v a t i o n and s u r r o u n d i n g v e g e t a t i o n ( S c h a e f e r 1978).  Two  f a c t o r s p l a y a dominant r o l e i n d e t e r m i n i n g the c l i m a t e o f t h e  study a r e a : (1) the P a c i f i c Ocean and  (2) the Coast Mountains (Kendrew  and K e r r 1955). Onshore w e s t e r l y o c e a n i c a i r masses impinge upon s u c c e s s i v e l a r g e - s c a l e mountain b a r r i e r s a l i g n e d i n a northwest  to southeast  direc-  t i o n . S i n c e these are r o u g h l y p e r p e n d i c u l a r t o the mean winds a l o f t , they determine  t o a major e x t e n t the o v e r a l l d i s t r i b u t i o n o f p r e c i p i -  t a t i o n . Weather systems c a r r i e d by the p r e v a i l i n g w e s t e r l y winds a l o f t drop c o n s i d e r a b l e m o i s t u r e over e x t e n s i v e a r e a s , p a r t i c u l a r l y w e s t e r l y exposures and a t h i g h e r e l e v a t i o n s . - East f a c i n g s l o p e s and r e c e i v e s u b s t a n t i a l l y l e s s . S n o w f a l l accounts annual  p r e c i p i t a t i o n near sea  lowlands  f o r a s m a l l f r a c t i o n o f the  level.  In s h o r t , the c l i m a t e o f the study area i s c h a r a c t e r i z e d by moderate year-round  temperatures,  l a c k o f temperature  extremes, high  amount o f p r e c i p i t a t i o n and f r e q u e n t c l o u d - c o v e r and f o g . C l i m a t e data (from Environment Canada 1970)  as o b t a i n e d by weather s t a t i o n s c l o s e  to c e r t a i n s i t e s i n the study area are g i v e n i n t a b l e I I .  71  22  F i g u r e 3. Annual p r e c i p i t a t i o n i n 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 , g i v e n i n c e n t i m e t e r s . Measurements i n c e n t i m e t e r s c o n v e r t e d from measurements i n i n c h e s g i v e n by Chapman and T u r n e r (1956).  LEGEND.  over 381  254-381  A  7v 152-254  101-152  76-101  23  Annual p r e c i p i t a t i o n i s shown i n F i g u r e 3. The map was sketched f o l l o w i n g University  a map c o m p i l e d by t h e D i v i s i o n o f Geography,  Of B r i t i s h Columbia (Chapman and Turner 1956).  c  Latitude  Longitude  UJ  r—  "(0 <1  Annuc mean snowl  Station  ta >  •c ta  Annual mean rainfal  o  •r™•  •  cu E  Q-  • ta  o  C 4-> CU ,— ,—, c O  «t +->  >-  Q.  1—' C r— 03 QJ ("0  LO  4-  -c -a +-> ta  =t*= s  7.2  128  8.9  84  5.1  164  1068  9.8  57  1034  1805  10.9  36  782  1503  9.4  76  846  1648  10.2  998  1396  PEMBERTON  50 27 N  122 56 W  223  742  2824  1024  GARIBALDI  49 59 N  123 08 W  366  1301  4229  1724  SQUAMISH  49 42 N  123 09 W  1.5  1916  1455  2061  WOODFIBRE  49 40 N  123 16 W  6  2856  1466  3003  HOLLYBURN RIDGE  49 22 N  123 12 W  951  2128  8108  2939  HOLLYBURN  49 20 N  123 09 W  46  1852  658  1917  SEYMOUR  49 26 N  122 57 W  823  2364  4862  2850  SEYMOUR FALLS  49 26 N  122 58 W  201  3505  2258  3733  VANCOUVER INTERNATIONAL AIRPORT  49 11 N  123 10 W  5  1018  523  SUMAS CANAL  49 07 N  122 07 W  6  1702  ABOTTSFORD  49 01 N  122 22 W  60  1425  BRIDAL VEIL FALLS  49 12 N  121 45 W  61  CHILLIWACK RIVER (Mt. T h u r s t o n )  49 05 N  121 46 W  198  1276  "OSt  T a b l e I I . C l i m a t e data from weather s t a t i o n s i n t h e study a r e a (from Environment Canada 1970).  25  5. VEGETATION OF THE  STUDY AREA.  R e l a t e d t o q u i t e d i s t i n c t s e a s o n s , c l i m a t e s and a wide v a r i e t y o f t e r r a i n and s o i l t a t i o n . The  c o n d i t i o n s , B r i t i s h Columbia has a very d i v e r s e vege-  d e s c r i p t i o n o f the v e g e t a t i o n o f the p r e s e n t  study  area  f o l l o w s K r a j i n a ' s (1965) c l a s s i f i c a t i o n which d i v i d e s t h e p r o v i n c e 4 b i o g e o c l i m a t i c formations comprising  into  7 b i o g e o c l i m a t i c r e g i o n s which  i n c l u d e 12 b i o g e o c l i m a t i c zones ^. A l l are based on b i o t a ( v e g e t a t i o n  and  fauna) and s o i l s , b e i n g the p r o d u c t s o f g e o l o g i c a l p a r e n t m a t e r i a l s , s o i l animal a c t i v i t y , t o p o g r a p h y , anf the e f f e c t s o f c l i m a t e and t i m e upon each o f  these. G e n e r a l l y each b i o g e o c l i m a t i c zone (=geographic area  c h a r a c t e r i s t i c v e g e t a t i o n w i t h a s s o c i a t e d a n i m a l s , s o i l s and  having climate)  be broken i n t o subzones which are d i s t i n g u i s h e d by the c l i m a x  can  plant  a s s o c i a t i o n (=grouping o f s i m i l a r p l a n t communities) on a mesic s i t e . T h i s s i t e i s the m i d - p o i n t o f the n u t r i e n t and m o i s t u r e range o c c u r r i n g w i t h i n t h e subzone; i t i s t h e r e f o r e not s u b j e c t t o e x c e s s i v e d r a i n a g e t o seepage w a t e r s and  nor  hence the v e g e t a t i o n b e s t r e f l e c t s the m a c r o - c l i m a t e  (Jones and Annas 1978). Each subzone possesses a s e r i e s o f communities i n h a b i t a t s w h i c h , o t h e r than by m a c r o - c l i m a t e , are i n f l u e n c e d by f a c t o r s such as s o i l s , drainage, The  m o i s t u r e r e g i m e , s l o p e , a s p e c t and  micro-climate.  b i o g e o c l i m a t i c zones o c c u r r i n g i n the p r e s e n t  are shown on the map  different  study  i n F i g u r e 4 which i s d e r i v e d from K r a j i n a ' s  " B i o g e o c l i m a t i c Zones o f B r i t i s h C o l u m b i a " , p u b l i s h e d by t h e  area, map,  British  26 F i g u r e 4. V e g e t a t i o n zones i n s o u t h - w e s t e r n  B r i t i s h Columbia.  VEGETATION  ZONES  SW BRITISH  COLUMBIA.  ]  IN  C o a s t a l Douglas F i r C o a s t a l Western  Hemlock  S u b a l p i n e Mountain Hemlock Subalpine  Engelmann SpruceSubalpine F i r  I n t e r i o r Douglas F i r Alpine  Tundra  Glaciers  27  Columbia E c o l o g i c a l Reserves Committee. These v e g e t a t i o n zones and characteristic  v a s c u l a r p l a n t s a r e d i s c u s s e d below.  S i n c e t h i s study f o c u s e s on b r y o p h y t e v e g e t a t i o n w i t h i n some major v e g e t a t i o n z o n e s , l i v e r w o r t s and mosses a r e mentioned t h a t a r e characteristic  5.1.  t a x a o c c u r r i n g i n t h e s e zones.  The C o a s t a l D o u g l a s - F i r Zone.  T h i s zone [termed B o r e o m e r i d i o n a l  zone by Hamet-Ahti ( 1 9 6 5 ) ] i s  s i t u a t e d a l o n g t h e south c o a s t a l r e g i o n i n t h e rainshadow o f t h e Vancouver I s l a n d and t h e Olympic Mountains o f Washington, U.S.A. I t ranges i n e l e v a t i o n from sea l e v e l t o 450 m on t h e e a s t c o a s t Of Vancouver I s l a n d and from sea l e v e l t o 150 m on t h e m a i n l a n d and G u l f I s l a n d s . .. 2 In t h e t r e e s t r a t u m , t h e dominant t r e e , Pseudotsuga m e n z i e s n , o c c u r s t o g e t h e r w i t h s p e c i e s such as Tsuga h e t e r o p h y l l a , A b i e s g r a n d i s and Cornus n u t t a l l i i . C h a r a c t e r i s t i c u n d e r s t o r y v e g e t a t i o n i n c l u d e s t a x a such as G a u l t h e r i a s h a l l o n , B e r b e r i s n e r v o s a , Ribes sanguineum,  Vaccinium  p a r v i f o l i u m , P o l y s t i c h u m munitum, S t o k e s i e l l a oregana and Rhizomnium glabrescens. K r a j i n a (1956) d i v i d e s t h e C o a s t a l D o u g l a s - F i r Zone i n t o two subzones. The w e t t e r subzone i s c h a r a c t e r i z e d by Arb ut us me nz i es i i , Pinus c o n t o r t a and Thuja p l i c a t a . The v e g e t a t i o n o f t h e d r i e r  subzone  i n c l u d e s s p e c i e s such as Quercus g a r r y a n a , Camassia quamash, P l e c t r i t i s congesta  and Zigadenus venenosus.  Bryophytes:  A n t i t r i c h i a c a l i f o r n i c a , A. c u r t i p e n d u l a , B r a c h y t h e c i u m  28  asperrimum, Buxbaumia pi p e r i , C a l y p o g e i a m u l l e r i a n a , C e p h a l o z i a b i c u s p i d a t a , Claopodium b o l a n d e r i , C. c r i s p i f o l i u m , Conocephalum conicum, Dendroal s i a a b i e t i n a , D i c r a n o w e i s i a c i r r a t a , Dicranum f u s c e s c e n s , JJ. scoparium, D o u i n i a o v a t a , Drepanocladus  aduncus, Homalothecium  Hypnum subimponens, I s o p t e r y g i u m e l e g a n s , Lophocolea  nutallii,  c u s p i d a t a , L.  h e t e r o p h y l l a , M e t z g e r i a c o n j u g a t a , Metaneckera m e n z i e s i i , Neckera dougl a s i i , O r t h o t r i c h u m c o n s i m i l e , 0. l y e l l i i ,  Philonotis fontana, Plagiochila  a s p l e n i o i d e s , P I a g i o t h e c i u m d e n t i c u l a t u m , P o l y t r i c h u m a l p i n u m , P_. j u n i perinum, P_. p i l i f e r u m , P o r e l l a cordaeana,  P_. n a v i c u l a r i s , P_. r o e l l i i ,  P o r o t r i c h u m b i g e l o v i i , P t i 1 i d i u m c a l i f o r n i c u m , P_. pulcherrimum, b o l a n d e r i , Racomitrium  canescens,  Radula  R. h e t e r o s t i c h u m , R h y t i d i a d e l p h u s  s q u a r r o s u s , R. t r i q u e t r u s , Scapania americana, S_. b o l a n d e r i , S c i e r o p o d i u m c a e s p i t a n s , Sphagnum c a p i l l a c e u m , S^. fuscum, S_. p a l u s t r e , S_. p a p i l l o s u m , S_. recurvum, S_. t e n e l 1 um,  w a r n s t o r f i i , T e t r a p h i s pel l u c i d a .  5.2. The C o a s t a l Western Hemlock Zone.  W i t h i n t h e study a r e a t h i s zone [ e n c l o s e d i n t h e Hemiboreal  zone by  Hamet-Ahti (1965)] encompasses p r o d u c t i v e temperate m a r i t i m e , c o n i f e rous r a i n f o r e s t and i s l o c a t e d above t h e d r i e r C o a s t a l D o u g l a s - F i r Zone, but below t h e S u b a l p i n e Mountain Hemlock Zone. The most c h a r a c t e r i s t i c p l a n t a s s o c i a t i o n s a r e hemlock-moss communities ( K r a j i n a 1965)  r e c o g n i z e d by a c o m b i n a t i o n o f s p e c i e s ,  i n c l u d i n g Tsuga h e t e r o p h y l l a , Thuja p i i c a t a , Blechnum s p i c a n t , Hylocomium s p l e n d e n s , I s o t h e c i u m s t o l o n i f e r u m , P I a g i o t h e c i u m undulatum and  29  R h y t i d i a d e l p h u s l o r e u s . In the d r i e r subzone, Pseudotsuga m e n z i e s i i i s a common s u c c e s s i o n a l t r e e s p e c i e s t o the c l i m a x Tsuga h e t e r o p h y l l a . In the w e t t e r subzone, t h i s l a t t e r s p e c i e s i s o f t e n accompanied by Abies a m a b i l i s . P l a n t communities w i t h i n the C o a s t a l Western Hemlock Zone vary w i t h l o c a l topography and s l o p e p o s i t i o n . Rock o u t c r o p s  and areas w i t h  w e l l d r a i n e d s o i l s , p a r t i c u l a r l y t h o s e w i t h s o u t h e r l y a s p e c t s , commonly support  Pseudotsuga m e n z i e s i i , G a u l t h e r i a s h a l l o n , and a c a r p e t o f  l i c h e n s and b r y o p h y t e s .  Depressional  areas and lower s l o p e s , where t h e r e  i s permanent seepage w a t e r , m a i n t a i n a mixed f o r e s t o f Tsuga h e t e r o p h y l l a and Thuja p i i c a t a w i t h an e x t e n s i v e u n d e r s t o r y o f f e r n s . Bryophytes:  Andreaea r u p e s t r i s , Apometzgeria pubescens, B a z z a n i a  denudata,  B_. t r i c r e n a t a , Blepharostoma t r i c h o p h y l l u m , C a l y p o g e i a n e e s i a n a ,  Cephalozia  b i c u s p i d a t a , Claopodium b o l a n d e r i , Dichodontium pel 1ucidum, Dicranum f u s c e s c e n s , D_. s c o p a r i u m , Heterocladium  F r u l l a n i a t a m a r i s e r ssp.  m a c o u n i i , H_. p r o c u r r e n s , H o o k e r i a  nisquallensis,  1 ucens, Hylocomi um s p l e n -  dens, Hypnum subimponens, L e p i d o z i a r e p t a n s , Lophocolea h e t e r o p h y l l a , Lophozia  i n c i s a , Metaneckera m e n z i e s i i , M e t z g e r i a c o n j u g a t a , M y l i a t a y l o r i ,  01igotrichum  p a r a l l e l u m , Orthotrichum  l y e l l i i , P e l l i a neesiana,  Philonotis  f o n t a n a , P l a g i o c h i l a a s p l e n i o i d e s , PIagiomnium i n s i g n e , P l a g i o t h e c i u m d e n t i c u l a t u m , P_. l a e t u m , P o l y t r i c h u m a l p i n u m , P_. formosum, P o r e l l a n a v i c u l a r i s , Radula b o l a n d e r i , R. c o m p l a n a t a , Racomitrium  heterostichum,  R. lanuginosum, Rhizomnium g l a b r e s c e n s , I*, punctatum, R. venustum, R h y t i d i a d e l p h u s t r i q u e t r u s , Sphagnum f i m b r i a turn, S_. f us cum, S_. mageTlanicum, S^. russowi  , Tetraphis pellucida  S_.  imbricatum,  , U l o t a megalospora.  30  5.3. The S u b a l p i n e Mountain Hemlock  Zone.  O c c u r r i n g above t h e C o a s t a l Western Hemlock Zone and below t h e A l p i n e Zone, t h i s zone i n c l u d e s t e r r a i n a l o n g t h e P a c i f i c Coast o f B r i t i s h Columbia a t a l t i t u d e s from 915 t o 1680 m. The S u b a l p i n e Mountain Hemlock Zone [ i n c l u d e d i n Hamet-Ahti's (1965) Upper O r o b o r e a l z o n e ] i s c h a r a c t e r i z e d by rugged t o p o g r a p h y , w i t h rock o u t c r o p s , r i d g e s , t a l u s s l o p e s , narrow s t e e p - s i d e d v a l l e y s , f a s t f l o w i n g mountain streams and heavy snow a c c u m u l a t i o n d u r i n g w i n t e r t i m e . W i t h i n t h e S u b a l p i n e Mountain Hemlock Zone, two subzones a r e d i s t i n g u i s h e d : t h e F o r e s t Subzone and t h e P a r k l a n d  Subzone.  F o r e s t Subzone. In t h i s lower subzone ( a t e l e v a t i o n s o f a p p r o x i m a t e l y 900 - 1100 m) t h e major t r e e s p e c i e s a r e Tsuga m e r t e n s i a n a , A b i e s a m a b i l i s and Chamaecyparis n o o t k a t e n s i s , w i t h t h e l a t t e r  generally  o c c u p y i n g h a b i t a t s w i t h abundant m o i s t u r e . Dominant shrubs on mesic s i t e s a r e V a c c i n i u m a l a s k a e n s e , V_. membranaceum and M e n z i e s i a f e r r u g i n e a . The herb l a y e r i s u s u a l l y not w e l l developed w i t h o n l y Rubus pedatus and C l i n t o n i a u n i f l o r a common. Dicranum p a l l i d i s e t u m and R h y t i d i o p s i s r o b u s t a dominate u s u a l l y e x t e n s i v e moss l a y e r o f mesic s i t e s . On l o w e r and m i d d l e s l o p e p o s i t i o n s , seepage w a t e r i s p l e n t i f u l  e x p r e s s e d by an u n d e r s t o r y o f S t r e p t o p u s  r o s e u s , Blechnum s p i c a n t , Veratrum e s c h s c h o l t z i i and T i a r e l l a  unifoliata.  P a r k l a n d Subzone. In t h i s subzone, o c c u r r i n g a t e l e v a t i o n s o f a p p r o x i mately 1070 - 1220 m, v a r i a t i o n s i n topography and i t s subsequent e f f e c t  31  on t h e d u r a t i o n o f t h e s n o w - l i e r e s u l t i n a v e g e t a t i o n which i s broken i n t o a mosaic o f f o r e s t p a t c h e s , shrubby areas and meadows. These meadows, formed on s i t e s w i t h l o n g e r snow d u r a t i o n and i r r i g a t i o n from snowmelt c r e a t i n g h y g r i c and h y d r i c c o n d i t i o n s , can be d i v i d e d i n t o  drier  f l o w e r ( f o r b ) meadows and m o i s t h e a t h e r (heath) meadows (Brooke e t a l . 1970, B r i n k 1959, A r c h e r 1963). Frequent s p e c i e s i n t h e s e r i c h f l o w e r meadows i n c l u d e C a l t h a l e p t o s e p a l a , A r n i c a  floristically  1atifolia,  C a s t i l l e j a m i n i a t a , E r i g e r o n p e r e g r i n u s , L e p t a r r h e n a pyrol'i f o l i a , V a l e r i a n a s i t c h e n s i s and Veratrum v i r i d e . The v e g e t a t i o n o f m o i s t h e a t h e r meadows i s c h a r a c t e r i z e d by P h y l l o d o c e e m p e t r i f o r m i s , Vacciniurn d e l i ciosum, C a s s i o p e m e r t e n s i a n a and Luetkea  pectinata.  B r y o p h y t e s : Andreaea n i v a l i s , A. r o t h i i , B a r b i l o p h o z i a  floerkei,  B a z z a n i a denudata, Dicranum f u s c e s c e n s , D_. p a l l i d i s e t u m , H o o k e r i a 1 ucens, Hylocomium s p l e n d e n s , Hypnum c i r c i n a l e , Mniurn s p i n u l o s u m , O l i g o t r i c h u m a l i g e r u m , 0_. h e r c y n i c u m , P h i l o n o t i s f o n t a n a , P l a g i o t h e c i u m d e n t i c u l a t u m , P_. undulatum,  P l e u r o z i u m s c h r e b e r i , Pseudoleskea b a i l e y i ,  Ptilidium  pulcherrimum, Racomitrium s u d e t i c u m , R h y t i d i a d e l p h u s l o r e u s , R h y t i d i o p s i s r o b u s t a , S c a p a n i a b o l a n d e r i , S_. g i r g e n s o h n i i , S_. m a g e l l a n i c u m , S.. s q u a r rosum, S_. t e r e s .  5.4. The S u b a l p i n e Engelmann S p r u c e - S u b a l p i n e F i r Zone.  As i s shown i n F i g u r e 4, t h i s zone o c c u r s toward t h e n o r t h e r n p a r t o f t h e study a r e a ( G a r i b a l d i P a r k ) . The S u b a l p i n e Engelmann SpruceS u b a l p i n e F i r Zone [ p a r t o f t h e M i d d l e Oroboreal zone (Hamet-Ahti  1965)]  32  encompasses v e g e t a t i o n o f a more c o n t i n e n t a l c h a r a c t e r : s u b a l p i n e c o n i f e r o u s f o r e s t , r e p l a c i n g the Mountain Hemlock f o r e s t o f c o a s t a l subalpine e l e v a t i o n s . Main d i f f e r e n c e s between i n t e r i o r and c o a s t a l s u b a l p i n e are i n d i c a t e d by lower annual and annual  temperature,  l e s s t o t a l annual  precipitation  s n o w f a l l , and h i g h e r p r o b a b i l i t y o f summer drought and  ground i n w i n t e r ( K r a j i n a 1965,  frozen  F r a s e r 1970).  Throughout the e n t i r e zone the dominant t r e e s p e c i e s a r e P i c e a e n g e l m a n n i i , A b i e s l a s i o c a r p a , Pinus c o n t o r t a v a r . ( a f t e r f i r e s ! ) and Pinus  latifolia  albicaulis.  P r o d u c t i v e seepage s i t e s o f t e n show presence o f p l a n t s p e c i e s 1 i k e Alnus  i n c a n a , Gymnocarpium d r y o p t e r i s and V a l e r i a n a s i t c h e n s i s .  In the d r i e r , l e s s p r o d u c t i v e p a r t s o f t h i s zone many s p e c i e s c h a r a c t e r i s t i c o f dry c o n d i t i o n s o c c u r f r e q u e n t l y e.g. uva-ursi, Calamagrostis Bryophytes:  rubescens and A s t e r  Arctostaphylos  conspicuus.  B a r b i l o p h o z i a f l o e r k e i , B_. l y c o p o d i o i d e s , Dicranum  JJ. scoparium,  Mnium s p i n u l o s u m ,  Pogonatum u r n i g e r u m , P o l y t r i c h u m  P_. j u n i p e r i n u m , Racomitrium s u d e t i c u m , c a p i l l a c e u m , S_. f i m b r i a t u m ,  fuscescens, alpinum,  R h y t i d i o p s i s r o b u s t a , Sphagnum  fuscum, S_. recurvum, S_. r u b e l 1 um,  S_.  squarrosum, Timmia a u s t r i a c a .  5.5.  The  I n t e r i o r D o u g l a s - F i r Zone.  Although  not encountered i n t h e p r e s e n t s t u d y , t h i s zone [termed  Hemiboreal zone by Hamet-Ahti (1965)] marks the n o r t h e a s t e r n boundary o f the study a r e a . Located  i n the shadow o f the Coast M o u n t a i n s , t h e  Interior  33  D o u g l a s - F i r Zone o c c u p i e s e l e v a t i o n s o f 610 - 1340 m i n south e a s t e r n B r i t i s h Columbia C h a r a c t e r i s t i c s p e c i e s o f open f o r e s t s a t lower e l e v a t i o n s are Pinus ponderosa,  Pseudotsuga m e n z i e s i i and Agropyron  s p i c a t u m . In t h e s e  areas g r a z i n g a c t i v i t y and f i r e h i s t o r y t e n d t o determine and p a t t e r n o f p r e s e n t day v e g e t a t i o n (Jones and Annas  the composition  1978).  In m o i s t e r and more c l o s e d f o r e s t s , a t h i g h e r e l e v a t i o n s o f t h e zone, t h e two dominant t r e e s p e c i e s , Pinus ponderosa and menziesi i ,  Pseudotsuga  are accompanied by L a r i x o c c i d e n t a l i s , Thuja p l i c a t a , P i c e a  engelmannii  and Pinus c o n t o r t a . C l o s e d f o r e s t s p e c i e s P a x i s t i m a m y r s i n i t e s  and C h i m a p h i l a u m b e l l a t a a r e more f r e q u e n t i n w e t t e r p a r t s o f the zone. Bryophytes:  B a r b i l o p h o z i a h a t c h e r i , B_. l y c o p o d i o i d e s , B r a c h y t h e c i u m  a l b i c a n s , Bryum c a n a r i e n s e , Ceratodon Drepanocladus  p u r p u r e u s , Picranurn f u s c e s c e n s ,  u n c i n a t e s , Hylocomium s p l e n d e n s , Mnium s p i n u l o s u m ,  nutans, P t i l i d i u m pulcherrimum,  RhytidiadeTphus  Pohlia  t r i q u e t r u s , Timmia  austriaca, Tortula rural i s .  5.6.  The A l p i n e Tundra Zone.  T h i s zone [ i n c l u d e d by Hamet-Ahti (1965) i n t h e O r o h e m i a r c t i c z o n e ] c o n t a i n s t e r r a i n a t h i g h e r e l e v a t i o n s , l y i n g above c o n t i n u o u s f o r e s t , p a r k l a n d and meadows, and below g l a c i e r s o r permanent s n o w f i e l d s . As encountered  i n the study a r e a , some l a n d s c a p e f e a t u r e s i n the A l p i n e  Zone a r e d e p r e s s i o n s and b a s i n s , such as l a t e snow s i t e s , r u b b l e , b o u l d e r f i e l d s , bare r o c k o u t c r o p s , rock w a l l s , exposed summits and r i d g e s w i t h  34  t h i n s o i l s . Of major i n f l u e n c e i n s o i l  f o r m a t i o n , type o f v e g e t a t i o n ,  and p l a n t s u c c e s s i o n i s snow i . e . p r e s e n c e , d e p t h , and movement o f snow and snow p e r s i s t e n c e . Vegetation  i s composed o f h e r b s , b r y o p h y t e s and l i c h e n s and  o n l y very low shrubs.  No t r e e s t h r i v e i n t h i s zone, but some c o n i f e r s  and a few deciduous woody p l a n t s develop a dwarf o r s t u n t e d  growth  form a t the l o w e s t e l e v a t i o n s i n the zone. Among them are Tsuga mertens i a n a , Abies  1 a s i o c a r p a and  Pinus a l b i c a u l i s . The  v e g e t a t i v e season i s  very s h o r t . Bryophytes:  Amphidium l a p p o n i c u m , A n a s t r o p h y l l u r n minutum, Andreaea  b l y t t i i , A. n i v a l i s , A. r o t h i i , A n t h e l i a j u r a t z k a n a , A r c t o a Bartramia  fulvella,  i t h y p h y l l a , BI i n d i a a c u t a , Bryum a l p i n u m , B_. w e i g e l i i , Desmatodon  l a t i f o l i u s , Dichodontium Olympicurn, Dicranodontiurn denudatum, D i c r a n o w e i s i a c r i s p u l a , D i p l o p h y l 1 urn t a x i f o l i u m , Grimmia a l p e s t r i s , (3. d o n n i a n a , Gymnomitrion coneinnaturn, H y g r o b i e l l a l a x i f o l i a , Hygrohypnum b e s t i i , H_. s m i t h i i , Jungermannia e x s e r t i f o l i a , K i a e r i a b l y t t i i , Lophozia  v e n t r i c o s a , Marsupella  K_. s t a r k e i ,  e m a r g i n a t a , M. s p h a c e l a t a ,  Oligotrichum  h e r c y n i c u m , P a r a l eucobryum e n e r v e , Pohl i a c a r d o t i i , P_. l u d w i g i i , P o l y trichum sexangulare,  Pseudoleskea a t r i c h a , P. i n c u r v a t a , P_. r a d i c o s a ,  Racomitrium s u d e t i c u m , S c a p a n i a u l i g i n o s a , Sphagnum l i n d b e r g i i , T o r t u l a norvegica, Tritomaria  quinquedentata.  1. Hamet-Ahti (1965) has d e s c r i b e d the v e g e t a t i o n zones o f Western Canada by a p p l y i n g the f o r e s t s i t e type system, commonly used i n S c a n d i n a v i a . T h i s one i s not p u r e l y f l o r i s t i c , but l a r g e l y emphasizes the c l i m a t e i n i t s e f f e c t on the v e g e t a t i o n . T h i s a u t h o r d i s t i n g u i s h e s ' : s i x 1 a t i -  35  tud.ional zones, which a r e a l s o r e c o g n i z e d v a l l e y , thus both h o r i z o n t a l and v e r t i c a l under t h e same u n i t s .  i n mountains from peak t o z o n a t i o n s are combined  2. Nomenclature f o r v a s c u l a r p l a n t s i s t h a t o f H i t c h c o c k and Cronquist 1973; t h a t f o r b r y o p h y t e s f o l l o w s I r e l a n d , B i r d , B r a s s a r d , S c h o f i e l d and V i t t 1980.  36  6. METHODS.  6.1.  Delimitation of e p i l i t h i c  Bryophytes seldom cracks, depressions  b r y o p h y t e communities.  grow d i r e c t l y on rock s u r f a c e s . They occupy  and o c c u r a l o n g  ' m i c r o - r i d g e s ' . Moreover,  do not c o l o n i z e r o c k s not i n f l u e n c e d by w e a t h e r i n g ,  bryophytes  but can invade  the  p h y s i c a l l y and c h e m i c a l l y d i s i n t e g r a t e d rock s u r f a c e . Because t r a n s i t i o n s between rock s u r f a c e and m i n e r a l gradual, i t i s d i f f i c u l t  ' s o i l ' (product of weathering) are o f t e n  t o determine whether b r y o p h y t e communities  are e p i g e i c ( = t e r r e s t r i a l ) on m i n e r a l  soil  or e p i l i t h i c  on the weathered  rock s u r f a c e . With t i m e , humus m a t e r i a l w i l l  accumulate beneath t h e b r y o p h y t e  c o v e r . E s p e c i a l l y under the e x t e n s i v e b r y o p h y t e cushions  and c a r p e t s  p r o g r e s s i v e s u c c e s s i o n a l stages humus l a y e r s o f s e v e r a l c e n t i m e t e r s to develop.  of tend  In t h i s case t h e d e l i m i t a t i o n from e p i g e i c b r y o p h y t e  communities becomes even more d i f f i c u l t .  With t h i c k e n i n g humus l a y e r s ,  s i t e s l become invaded by f o r e s t f l o o r mosses such as R h y t i d i a d e l p h u s P l e u r o z i u m s c h r e b e r i , R h y t i d i o p s i s r o b u s t a , S t o k e s i e l 1 a oregana  spp.,  and  P l a g i o t h e c i u m d e n t i c u l a t u m . These e p i g e i c b r y o p h y t e s p e c i e s can comp l e t e l y cover s m a l l e r stones  (up t o 20 c e n t i m e t e r i n d i a m e t e r ) . Where  they o c c u r on more e x t e n s i v e r o c k s u r f a c e s , however, the c o n t r i b u t i o n of predominantly  epilithic  bryophytes  i s high ( S j o g r e n 1964).  Several species tend to p r e f e r s p e c i f i c s u b s t r a t a . Sjogren d i v i d e s t h e mosses o f South Sweden i n t o c a t e g o r i e s r e l a t e d t o t h e i r  (1964)  37  s u b s t r a t e p r e f e r e n c e . With r e s p e c t t o e p i l i t h i c bryophytes  he  distin-  guishes o b l i g a t o r y , p r e f e r e n t i a l , f a c u l t a t i v e and a m p h i s u b s t r a t i c s p e c i e s w i t h t r a n s i t i o n s between t h e groups. i s very common, t h e substratum  In r e g i o n s where t h e s p e c i e s  p r e f e r e n c e i s u s u a l l y weaker than where  i t i s r a r e . Hence, d i a s p o r e i n p u t i s e s s e n t i a l substrate preference (Hertel  i n t h e judgement o f  1974).  T h i s change i n s u b s t r a t e , f o l l o w i n g weakened s u b s t r a t e p r e f e r e n c e , becomes apparent  i n comparison o f s e v e r a l f l o r i s t i c a r e a s . S j o g r e n  p r e s e n t s o b s e r v a t i o n s comparing t h e bryophyte  (1964)  f l o r a s o f South Sweden and  West Europe. Gams (1932) i n d i c a t e s t h e change from e p i p h y t i c t o e p i l i t h i c s u b s t r a t e p r e f e r e n c e i n s u b a l p i n e and a l p i n e a r e a s . Herzog (1926) desc r i b e s t h e t r a n s i t i o n from e p i l i t h i c  into epigeic conditions i n a r c t i c  regions. In t h e p r e s e n t study s a m p l i n g was c o n f i n e d t o rock s u r f a c e s which were covered by a l a y e r o f m i n e r a l m a t e r i a l t h a t o r i g i n a t e d from d i s i n t e g r a t e d r o c k , o r a m i x t u r e o f d e p o s i t e d o r g a n i c and i n i t i a l  mineral  ' s o i l ' up t o a t h i c k n e s s o f 2 cm.  6.2. Sampling methods.  6.2.1.  Reconnaissance.  An e s s e n t i a l a c t i v i t y  b e f o r e t a k i n g any r e l e v e s i s t h e s e a r c h  f o r and f a m i l i a r i z a t i o n w i t h p o t e n t i a l r e l e v e - s i t e s . The (meaning " p r e l i m i n a r y e x a m i n a t i o n  reconnaissance  o f a given area") i n the present  study  38  i n c l u d e d an assessment o f : a. a v a i l a b i l i t y o f r o c k s u b s t r a t e t y p e s ( b o u l d e r s , o u t c r o p s and rockwalls) b. i n f l u e n c e o f t o p o g r a p h y ,  v e g e t a t i o n and m a c r o - c l i m a t e on t h e rock  s u b s t r a t e s p r e s e n t ( e x p o s u r e , m o i s t u r e , accumulated c. t h e b r y o p h y t e f l o r a ; i n i t i a l  soil)  c o l l e c t i o n s were made and specimens  identified d. minimum area measurement f o r r e l e v e s e. geographic d i s t r i b u t i o n o f t a x a t o i d e n t i f y p o s s i b l e r e g i o n a l variation  6.2.2. Choice o f r e l e v e - s i t e and q u a d r a t - s i z e .  The d e c i s i v e c r i t e r i o n i n t h e c h o i c e o f t h e r e l e v e s i t e s i s t h e homogeneity o f t h e q u a d r a t . Only rock s u r f a c e s w i t h u n i f o r m c o n d i t i o n s and homogeneous v e g e t a t i o n were sampled. The homogeneity o f a s i t e i s high when, f o r i n s t a n c e , a s i n g l e f a c t o r i s o f d e c i s i v e i n f l u e n c e . S i n c e several  ( e n v i r o n m e n t a l ) f a c t o r s c o i n c i d e on t h e s i t e , c o n d i t i o n s on r o c k  s u r f a c e s a r e f r e q u e n t l y v a r i a b l e . S u r f a c e s w i t h changing i n c l i n a t i o n and exposure were e x c l u d e d from s a m p l i n g , as were r o c k s w i t h i n s i d e - p l o t changes i n rock s u r f a c e roughness, amount o f accumulated  s o i l o r changing  m o i s t u r e c o n d i t i o n s (seepage w a t e r ) . In g e n e r a l , q u a d r a t s were t a k e n i n homogeneously developed e p i l i t h i c b r y o p h y t e assemblages which c o v e r a rock s u r f a c e o f a p p a r e n t l y u n i f o r m e c o l o g i c a l c o n d i t i o n s o f a t l e a s t 1/2 m  (Homogeneous r e f e r s t o  38a  LEAP 39 OMITTED IN NUMBERING  40  v e g e t a t i o n l a r g e enough t o reach i t s f u l l  floristic  and s t r u c t u r a l  compo-  s i t i o n , as i s the case when more space i s a v a i l a b l e and the v e g e t a t i o n has s u f f i c i e n t time t o become w e l l d e v e l o p e d ) .  Types o f s i t e s chosen  i n c l u d e b o u l d e r s , detached p i e c e s o f stone o f a t l e a s t 1 meter i n d i a m e t e r , o u t c r o p s , p a r t s o f bedrock p r o j e c t i n g from s o i l and  rock-walls  which encompass steep r o c k s u r f a c e s . For t a b u l a r comparison and s y n t h e s i s o f a s s o c i a t i o n s i t i s important t o t a k e o n l y one r e l e v e a t each s i t e . T h i s representation of a c c i d e n t a l l y l o c a l  t a x a , fragments \  ( e . g . dominance/abundance, s p e c i e s c o m b i n a t i o n s ) requirement  or characters  i n the t a b l e s . T h i s  c o u l d not always be f u l f i l l e d and sometimes i n s i t e s , where  a community was Hertel  avoids"over-  w e l l developed,  more r e l e v e s were taken  (see W i r t h  1972,  1974). Within t h i s b r y o s o c i o l o g i c a l research a constant p l o t - s i z e of  625  cm  (25 x 25 cm) was  combination  found a p p r o p r i a t e t o encompass the  characteristic  o f s p e c i e s . T h i s i s i n d i c a t e d i n the t a b l e s o f the  separate  bryophyte syntaxa. As a c o n t r o l f o r the q u a d r a t - s i z e the Nested P l o t Technique was  a p p l i e d ( F i g u r e 5 ) . The  s p e c i e s number p l o t t e d over the s i z e o f  sample area r e s u l t s i n a s p e c i e s / a r e a curve  ( F i g u r e 6 ) , a f t e r which the  d e c i s i o n i s made on t h e q u a d r a t s i z e .  6.2.3. E s t i m a t i o n o f s p e c i e s q u a n t i t y and  sociability.  In o r d e r t o e s t i m a t e t h e cover o f b r y o p h y t e s p e c i e s w i t h i n the  41  cn o  10 CM  ro o O  40 CM  F i g u r e 5. Nested p l o t t e c h n i q u e , used f o r the o f a minimum r e l e v e ( p 1 o t ) - s i z e .  determination  s a p a d s j.o uaqwriN  43  quadrat a s c a l e i s used, as developed by Frey (1933) and a p p l i e d by many o t h e r c r y p t o g a m - ( l i c h e n - and b r y o p h y t e - )  s o c i o l o g i s t s , In t h i s  scale,  c o v e r v a l u e s below 50%, e s p e c i a l l y , a r e more a c c u r a t e l y measurable (Appendix I ) . In a l l r e l e v e s , attempts a r e made t o e s t i m a t e t h e degree o f s o c i a b i l i t y . A l t h o u g h s o c i a b i l i t y i s s p e c i e s - s p e c i f i c i t may r e v e a l i n f o r m a t i o n on t h e c o n t r i b u t i o n o f t h e s p e c i e s t o t h e o v e r a l l s t r u c t u r e (physiognomy).  community  For t h e e s t i m a t i o n o f t h e degree o f s o c i a b i l i t y  a s c a l e was used, developed by Klement (1955) (Appendix I ) .  6.2.4. E c o l o g i c a l c h a r a c t e r i z a t i o n o f t h e r e l e v e - s i t e .  The l o c a t i o n on t h e r o c k s u r f a c e was r e c o r d e d f o l l o w i n g F r e y ' s (1922) t y p e s o f r o c k s u r f a c e h a b i t a t s ( F i g u r e 7 ) . To o b t a i n i n f o r m a t i o n on environmental  f a c t o r s d e t e r m i n i n g and  c o n t r o l l i n g t h e b r y o p h y t e assemblages on r o c k s , d a t a were c o l l e c t e d on the f o l l o w i n g  parameters:  l o n g i t u d e / l a t i t u d e : taken from g e o g r a p h i c a l maps a l t i t u d e : measured w i t h an a l t i m e t e r d i r e c t i o n o f slope (exposure): measured w i t h B e z a r d Compass i n c l i n a t i o n of slope: soil  accumulation:  d i v i d e d i n two c l a s s e s 0-1 cm  roughness  1-2 cm  o f rock s u r f a c e : s u b j e c t i v e l y  assessed  44  1  F i g u r e 7. C a t e g o r i e s o f rock s u r f a c e h a b i t a t s . 1. t o p s u r f a c e o r n e a r l y l e v e l 2. g e n t l e s l o p e (20-70°) 3. steep s l o p e (70-90°) 4. overhanging 5. g r o t t o  slope  (0-20°)  45  exposure:  l i g h t was  e s t i m a t e d as e i t h e r b r i g h t s u n l i g h t o r  shade and a s i t e was  c o n s i d e r e d shaded o n l y i f i t was  shaded by t r e e s ( o r o t h e r r o c k s ) f o r o n e - h a l f o r more o f each d a y l i g h t p e r i o d m o i s t u r e : e x p r e s s e d as e i t h e r wet o r d r y . The wet  sites include  o n l y those which a r e v i s i b l y m o i s t from r u n - o f f w a t e r , t e m p o r a r i l y submerged o r e f f e c t e d by s p r a y  6.3.  Data  analysis.  6.3.1. Bryophyte  identification.  In t h e p r e l i m i n a r y i d e n t i f i c a t i o n o f t h e b r y o p h y t e from the r e l e v e s , a s e r i e s o f f l o r a s and s p e c i a l  specimens  g e n e r i c and s p e c i e s -  complex t r e a t m e n t s were used. For t h e H e p a t i c a e t h e s e were: Flowers ( 1 9 6 1 ) , Frye and C l a r k (1937-1947),  Godfrey  S c h u s t e r (1966, 1969,  ( 1 9 7 7 ) , M a c v i c a r ( 1 9 2 6 ) , M u l l e r (1957) and 1974).  For t h e Musci t h e f o l l o w i n g f l o r a s were used: Crum ( 1 9 7 6 ) , Flowers ( 1 9 7 3 ) , Grout (1928-1940),  Lawton ( 1 9 7 1 ) , Nyholm (1954-1969)  and Smith (1978). I d e n t i f i c a t i o n o f t h e genus P l a g i o t h e c i u r n f o l l o w e d I r e l a n d ( 1 9 6 9 ) , t h a t o f Hygrohypnum f o l l o w e d Jamieson  (1976), that of  Dicranum f o l l o w e d P e t e r s o n (1979) and t h a t o f S e l i g e r i a f o l l o w e d V i t t (1976). F i n a l l y , Koponen (1974) was  used i n t h e i d e n t i f i c a t i o n o f the  f a m i l y Mniaceae and H o i s i n g t o n (1979) was  c o n s u l t e d i n the r e c o g n i t i o n  46  o f t a x a out o f the B r a c h y t h e c i u m asperrimum  - frigidum  complex.  B a s i c a l l y , t h e nomenclature f o l l o w s Crum, S t e e r e , and (1973) and the s p e c i e s l i s t  Anderson  (Appendix I I I ) i s based on S c h o f i e l d (1981).  A t o t a l o f 2200 b r y o p h y t e specimens were examined, the i d e n t i f i c a t i o n o f which formed the b a s i s f o r the bryophyte a s s o c i a t i o n  tables.  T h i s p l a n t m a t e r i a l has been d e p o s i t e d i n the Herbarium o f the U n i v e r s i t y of B r i t i s h  Columbia.  I n f o r m a t i o n on the p h y t o g e o g r a p h i c a l s e t t i n g o f t h e e p i l i t h i c b r y o p h y t e v e g e t a t i o n s o f s o u t h - w e s t e r n B r i t i s h Columbia was d e r i v e d from Godfrey ( 1 9 7 7 ) , S c h o f i e l d (1968a, 1968b, 1969, 1976, 1980) and Worley (1972).  6.3.2. T a b u l a r comparison.  The raw d a t a s e t was s u b d i v i d e d i n t o f o u r major groups which c o n s t i t u t e c o l l e c t i o n s o f r e l e v e s from the f o l l o w i n g h a b i t a t t y p e s : 1. exposed, dry r o c k s (129 r e l e v e s , 152 s p e c i e s ) sandstone and i n t r u s i v e ( s i l i c e o u s ) r o c k s 2. shaded, humid r o c k s (83 r e l e v e s , 128 s p e c i e s ) 3. s i l i c e o u s r o c k s on and a l o n g streambanks (102 r e l e v e s , 117 s p e c i e s ) 4. l i m e s t o n e , both exposed  and.shaded  (27 r e l e v e s , 61 s p e c i e s ) A crucial  procedure i n the a n a l y s i s o f p h y t o s o c i o l o g i c a l  data i s  47  the development o f t a b l e s . Braun-Blanquet (1951a,b) wrote from W e s t h o f f and Maarel  (transl.  1973):  " A p p r o p r i a t e l y e l a b o r a t e d a s s o c i a t i o n t a b l e s a r e comparable w i t h a thorough s p e c i e s d i a g n o s i s . By means o f the t a b l e s t h e c o n s i d e r a b l y d e t a i l e d work from the minute f l o r i s t i c a n a l y s i s o f the lower v e g e t a t i o n , u n i t s become a c c e s s i b l e and e v a l u a b l e . From the t a b l e i t a l s o shows whether one has worked s e r i o u s l y and r e l i a b l y ; t h e t a b l e s a r e the p r o p e r t o u c h - s t o n e o f the concerned p l a n t s o c i o l o g i s t " . E l l e n b e r g ( 1 9 5 6 ) , Knapp ( 1 9 7 1 ) , Shimwell (1971) and Dombois and E l l e n b e r g  Mueller-  (1974) p r e s e n t e d d e t a i l e d a c c o u n t s o f the t a b l e  t e c h n i q u e which was performed on the r e l e v e s o f each major group i n t h i s s t u d y . I t s procedure can be summarized  as f o l l o w s :  a. E n t r a n c e o f the r e l e v e data i n t o a raw t a b l e . S p e c i e s v a l u e s (cover and s o c i a b i l i t y ) are r e c o r d e d i n rows, r e l e v e s i n columns, w h i l e t h e o r d e r o f e n t r y i s i n c o n s e q u e n t i a l .  b. Count o f presence o f s p e c i e s  and c a l c u l a t i n g t h e i r c o n s t a n c y  v a l u e , ranked i n c l a s s e s from I t o V. In t h i s presence t a b l e i t becomes apparent whether r e l e v e s v a r y i r r e g u l a r l y , o r whether c e r t a i n c o m b i n a t i o n o f s p e c i e s a r e found r e c u r r e n t l y . In the l a t t e r cases such groups a r e more o r l e s s m u t u a l l y e x c l u s i v e and s e r v e as groups o f d i f f e r e n t i a t i n g Those a r e not l i k e l y and I ; t h e y w i l l III  species.  t o be found i n the presence c l a s s e s V  m o s t l y b e l o n g t o the i n t e r m e d i a t e  classes I I ,  and IV. These d i f f e r e n t i a t i n g s p e c i e s , which a r e p r o v i -  s i o n a l s e t s of d i a g n o s t i c s p e c i e s , a r e used t o r e a r r a n g e the table.  48  c. E x t r a c t i o n o f these d i f f e r e n t i a t i n g s p e c i e s and u s i n g them to  rearrange  rows i n t h e i r c o r r e s p o n d i n g  columns i n t h e i r c o r r e s p o n d i n g  groups, and t h e  r e l e v e s . T h i s rearrangement  seeks t o show a d i a g o n a l o r d e r o f s p e c i e s groups from t h e l e f t to t h e r i g h t o f t h i s , now c a l l e d , d i f f e r e n t i a t e d  d. Replacement o f p a r t i a l  table.  t a b l e by a column i n w h i c h , f o r each  p a r t i c i p a t i n g s p e c i e s , t h e presence degree i s i n d i c a t e d . A f t e r comparison o f t h i s s y n o p t i c t a b l e (Westhoff  and Maarel  '1973) w i t h those from o t h e r types o f v e g e t a t i o n from the same r e g i o n , an i d e a i s formed about t h e l o c a l d i a g n o s t i c s p e c i e s groups i n t h e t a b l e under s t u d y . With these s p e c i e s as w e l l as the c o n s t a n t t a x a , a phytocoenon [ = v e g e t a t i o n and Maarel  unit  (Westhoff  (1973)] i s then i d e n t i f i e d and d e s c r i b e d .  e. Comparison o f t h e s y n o p t i c t a b l e w i t h those a v a i l a b l e i n t h e l i t e r a t u r e and w i t h t a b l e s from s i m i l a r v e g e t a t i o n types  from  other r e g i o n s , r e s u l t s i n the d e s i g n a t i o n o f character t a x a , and a syntaxanomic i n t e r p r e t a t i o n f o l l o w s .  f. Comparison o f t a b l e s c o v e r i n g t h e v e g e t a t i o n o f a g i v e n  area  makes i t p o s s i b l e t o d i s c e r n c h a r a c t e r s p e c i e s on c r i t e r i a o f d i s t r i b u t i o n o f t h e s p e c i e s i n d i f f e r e n t phytocoena. T h i s e n t a i l s t h e d e t e r m i n a t i o n o f f i d e l i t y o f the t a x a ( f i d e l i t y i s the degree t o which a taxon i s c o n c e n t r a t e d  i n one phytocoenon  49  v s . d i s p e r s e d w i t h more even o c c u r r e n c e i n s e v e r a l  phytocoena).  F i v e degrees have been d i s t i n g u i s h e d (Braun-Blanquet 1964) i n t h e r e l a t i o n o f a s p e c i e s t o a g i v e n phytocoenon  (Appendix  II).  The  ' c l a s s i c a l ' arrangement t e c h n i q u e by manual o r d e r i n g o f  t a x a and r e l e v e s i s a time consuming procedure and a p o t e n t i a l  source  o f e r r o r . A number o f c o m p u t e r i z e d t e c h n i q u e s f o r t a b l e arrangement have been developed o f which t h e more r e c e n t ones have been r e p o r t e d t o be remarkably s u c c e s s f u l and e f f i c i e n t (Moore 1972, Janssen 1972, M a a r e l , Janssen and Louppen 1978, Meulen, M o r r i s and W e s t f a l l 1978, Louppen and Maarel 1979). In o r d e r t o s i m p l i f y and mechanize t h e procedure o f t h e t a b u l a t i o n t e c h n i q u e a computer program was used, which was d e s i g n e d t o s o r t t h e rows and columns o f a p h y t o s o c i o l o g i c a l t a b l e a u t o m a t i c a l l y . T h i s program, 'VEGTAB'. developed by Dr. G. B r a d f i e l d , Department o f Botany o f t h e U n i v e r s i t y o f B r i t i s h Columbia, t a k e s c a r e o f s t e p s a , b and c o f t h e t a b u l a t i o n p r o c e d u r e : p r o d u c i n g a raw t a b l e , t r a n s f o r m i n g t h i s i n t o a presence t a b l e , a n d , f i n a l l y , r e a r r a n g i n g t h i s l a t t e r one into a differentiated  table.  6.3.3. Numerical t e c h n i q u e s .  A d d i t i o n a l t o t h e t r a d i t i o n a l methods o f t h e Z u ' r i c h - M o n t p e l l i e r s c h o o l , q u a n t i t a t i v e t e c h n i q u e s have been developed and a p p l i e d t o study b r y o p h y t e v e g e t a t i o n on a v a r i e t y o f s u b s t r a t a .  50  In most s t u d i e s c o n c e r n i n g the q u a n t i t a t i v e e c o l o g y o f b r y o phytes (and l i c h e n s ) , workers have used random s a m p l i n g t e c h n i q u e s . Their s t a t i s t i c a l ecological  a n a l y s e s o f t h e v e g e t a t i o n o f q u a d r a t s and  the  f a c t o r s w i t h i n them, m o s t l y showed v e g e t a t i o n a l c o n t i n u a  r e l a t e d t o some environmental and Maycock 1968,  g r a d i e n t (a.o. J e s b e r g e r 1973,  C u l b e r s o n 1955, Hale  Lambert  1955).  With r e s p e c t to q u a n t i t a t i v e e c o l o g i c a l work on  epilithic  bryophyte v e g e t a t i o n , i n p a r t i c u l a r , some i n v e s t i g a t o r s have emphasized the continuum  concept and have attempted t o show r e l a t i o n s h i p s w i t h  e n v i r o n m e n t a l parameters  (Foote 1963, 1966, Y a r r a n t o n 1967a,b,c).  Other  workers have endeavoured t o i n t e r p r e t q u a n t i t a t i v e d a t a i n an  attempt  to r e c o g n i z e more o r l e s s d i s c r e t e v e g e t a t i o n u n i t s (Redfearn  1957,  N o r r i s 1964, West and S t o t l e r Foote (1963, 1966)  1977).  s t u d i e d sandstone and l i m e s t o n e o u t c r o p s i n  south-western W i s c o n s i n by means o f random s a m p l i n g t e c h n i q u e . O r d i n a t i o n s o f bryophyte  " s t a n d s " d i s p l a y e d a continuum  o f the h a b i t a t s from  xeric  to m o i s t s i t e s . The m o i s t u r e r e l a t i o n s o f the i n d i v i d u a l stands  appeared  to most i n f l u e n c e t h e frequency o f the s p e c i e s . Other e c o l o g i c a l  factors  such as exposure and d i r e c t i o n and i n c l i n a t i o n o f the rock s l o p e seemed to be secondary .  i n t h e i r i n f l u e n c e on the d i s t r i b u t i o n o f most o f t h e s e  2  species. . Y a r r a n t o n (1967a,b,c) produced  a quantitative analysis  (Principal  Components A n a l y s i s ) o f v a r i a t i o n i n the e p i l i t h i c b r y o p h y t e v e g e t a t i o n a t Steps B r i d g e , Devon ( E n g l a n d ) . R e s u l t s o f t h e a n a l y s i s i n d i c a t e d a c o r r e l a t i o n between t h i s v a r i a t i o n and e n v i r o n m e n t a l f a c t o r s such as  51  shade, c r e v i c e s and s o i l  depth.  Bunce (1967) s t u d i e d t h e e p i l i t h i c v e g e t a t i o n o f a Snowdonian c l i f f on t h e w e s t e r n seaboard o f B r i t a i n . The o r d i n a t i o n s , c o n s t r u c t e d on frequency d a t a from q u a d r a t - v e g e t a t i o n s , showed r e l a t i o n s h i p s between s p e c i e s d i s t r i b u t i o n s and a major environmental dry  g r a d i e n t from  o l i g o t r o p h i c t o wet e u t r o p h i c c o n d i t i o n s . Redfearn  t a t i o n on exposed  ( 1 9 5 7 ) , i n h i s quadrat s t u d i e s o f t h e b r y o p h y t e  vege-  l i m e s t o n e i n c e n t r a l and n o r t h e r n F l o r i d a , U.S.A.,  used h i g h f r e q u e n c y s p e c i e s i n d e l i m i t i n g d i f f e r e n t b r y o p h y t e assemb l a g e s . C a l c u l a t i o n s o f Cole's Index o f A s s o c i a t i o n between t h e most f r e q u e n t s p e c i e s made i t p o s s i b l e t o r e c o g n i z e f o u r t e e n b r y o p h y t e M o i s t u r e was t h e predominant  environmental  "unions".  f a c t o r c o n t r o l l i n g the d i s -  t r i b u t i o n o f t h e bryophytes and t h e i r u n i o n s . Norn's'  (1964) study o f e p i l i t h i c h a b i t a t s i n t h e A p p a l a c h i a n  Spruce-Zone r e s u l t e d i n an arrangement o f quadrats i n t o a number o f b r y o p h y t e communities  o r " u n i o n s " . Groups o f q u a d r a t s ( i n which  coverage  o f s p e c i e s was e s t i m a t e d i n increments o f t e n p e r c e n t ) were s o r t e d i n a way as t o m i n i m i z e t h e number o f 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 p a i r s . S i g n i f i c a n c e o f a s s o c i a t i o n i s judged by t h e C h i square method f o r each p o s s i b l e s p e c i e s p a i r . The a u t h o r used t h i s o b j e c t i v e  classification  o f t h e v e g e t a t i o n because " p r e l i m i n a r y i n v e s t i g a t i o n s i n d i c a t e d a c l o s e r c o r r e l a t i o n w i t h h a b i t a t s between p l o t s o f s i m i l a r s p e c i e s c o m p o s i t i o n than between p l o t s o f s i m i l a r s p e c i e s dominance" ( N o r r i s 1964, p. 8 ) . His  study r e s u l t e d i n t h e r e c o g n i t i o n o f 12 e p i l i t h i c b r y o p h y t e  unions,  8 o f which were c o n f i n e d t o t h i s t y p e o f s u b s t r a t e and 4 had a w i d e r  52  ecological  range ( s o i l , t r e e s ) . Major f a c t o r s i n t h e d i s t r i b u t i o n o f  bryophytes and t h e i r unions i n c l u d e d s o u r c e o f w a t e r , f r e q u e n c y and d u r a t i o n o f drought and l i g h t  intensity.  West and S t o t l e r (1977) a n a l y z e d t h e b r y o p h y t e - m a c r o - l i c h e n f l o r a o f sandstone canyons i n s o u t h e r n I l l i n o i s by u s i n g q u a d r a t s i n which r e l a t i v e c o v e r and f r e q u e n c y was determined f o r each s p e c i e s . I n t h e i r data a n a l y s e s t h e y computed t h e importance v a l u e s and community s i m i l a r i t y c o e f f i c i e n t s , once from t h e c o v e r and f r e q u e n c y v a l u e s and a g a i n from presence/absence d a t a . However, t h e data and t h e a u t h o r s ' difcussion o f t h e r e s u l t s f a i l  t o d e l i m i t a s s o c i a t i o n s and r e c o g n i z e  d i s t i n c t groupings. In t h e p r e s e n t s t u d y , c l u s t e r a n a l y s i s was used t o d e l i m i t , c h a r a c t e r i z e and c l a s s i f y t h e e p i l i t h i c b r y o p h y t e v e g e t a t i o n i n t h e study a r e a . F u r t h e r m o r e , attempts were made t o o r d i n a t e some r e l e v e s t o enhance t h i s c l a s s i f i c a t i o n by showing r e l a t i o n s h i p s o f a s s o c i a t i o n s and lower s y n t a x a t o one another and t o t h e environment.  6.3.3.1. C l u s t e r a n a l y s i s .  C l u s t e r a n a l y s i s was performed on each o f t h e f o u r major g r o u p s , u s i n g t h e MIDAS s t a t i s t i c a l  package  (Fox and G u i r e 1976). C l u s t e r  a n a l y s i s i s an a g g l o m e r a t i v e method i n which t h e c l u s t e r i n g "proceeds from t h e i n d i v i d u a l  samples  ( r e l e v e s ) , and combines them i n  p r o g r e s s i v e l y i n c r e a s i n g groups o f d e c r e a s i n g i n t e r n a l the whole s e t o f samples i s combined"  algorithm  ( G o o d a l l 1973).  similarity  until  53  The c l u s t e r a l g o r i t h m used here t o d e l i m i t the  bryophyte  v e g e t a t i o n at t h e v a r i o u s s i t e s i s t h e procedure d e s c r i b e d by Ward (1963). I t i s a h i e r a c h i a l method ( u s i n g E u c l i d e a n d i s t a n c e measure) g r o u p i n g , i n t h i s c a s e , r e l e v e s i n t o c l u s t e r s , based on c o v e r v a l u e s o f s p e c i e s i n t h e r e l e v e s . The t o t a l d e c r e a s e i n i n t e r n a l s i m i l a r i t y i s r e f l e c t e d i n t h e t o t a l sum o f squared d e v i a t i o n s o f every p o i n t from the mean o f the c l u s t e r t o which  i t b e l o n g s . At each s t e p i n the  a n a l y s i s , union o f every p o s s i b l e p a i r o f c l u s t e r s i s c o n s i d e r e d and the two c l u s t e r s whose f u n c t i o n r e s u l t s i n t h e minimum i n c r e a s e i n the e r r o r sum squares  o f squares a r e combined ( E v e r i t t 1974). The e r r o r sum  of  (E.S.S.) i s g i v e n by  E.S.S. =  N p -j S X, - -rj- (S i=l  y  x.)  N  The aim o f these c l u s t e r a n a l y s e s i s t o e x p l o r e and s u b d i v i d e the l a r g e s e t s o f r e l e v e s i n t o a number o f c l u s t e r s , each o f which i s c h a r a c t e r i z e d by one o r more c o n s t a n t s p e c i e s . These c l u s t e r s  are  then compared to the g r o u p i n g o f r e l e v e s a r r i v e d a t through t h e t a b u l a t i o n t e c h n i q u e o f the Braun-Blanquet was  method. In t h i s comparison  (which  o n l y c a r r i e d out on t h e group o f r e l e v e s from c a l c a r e o u s r o c k s ) connec-  are sought between the v a l u e o f s i m i l a r i t y index ( g i v e n as a  54  Euclidean  d i s t a n c e measure) and  a t i o n s and  6.3.3.2.  Ordination.  (Gaugh 1977)  t e c h n i q u e s are used t o s e a r c h f o r p a t t e r n s and  In v e g e t a t i o n s a n o t h e r and  i n a data  r e s u l t i n an arrangement o f e n t i t i e s  in a uni- or multidimensional  one  (alliances, associ-  subassociations).  Ordination matrix  syntaxonomic l e v e l  (releves)  order.  with associations generally discontinuous  with  t h e e n v i r o n m e n t , o r d i n a t i o n has been a p p l i e d t o under-  s t a n d the r e l a t i o n s o f a s s o c i a t i o n s and between t h e r e l e v e s  lower s y n t a x a t o one  ( M a t u s z k i e w i c z and T r a c z y k 1958,  another  Falinski  1960).  On a d i f f e r e n t l e v e l o r d i n a t i o n can be a p p l i e d t o the r e l a t i o n s o f a s s o c i a t i o n s and h i g h e r s y n t a x a as w h o l e s , t o one ronment (Westhoff and  a n o t h e r and  the  Maarel 1973).  In the p r e s e n t study a major f u n c t i o n o f o r d i n a t i o n i s t o p l a y groups o f s i m i l a r r e l e v e s  i n the c o l l e c t i o n o f l i m e s t o n e  niques t o the d a t a seeks t o enhance the c l a s s i f i c a t i o n by  and  and  another  a n a l y s e s were performed u s i n g the ORDIFLEX  computer program developed by Gaugh (1977) a t C o r n e l l U n i v e r s i t y , New  1.  tech-  further  c l a r i f y i n g r e l a t i o n s h i p s o f a s s o c i a t i o n s t o one  the environment. The  dis-  vegetation  samples (27 r e l e v e s , 61 s p e c i e s ) . T h i s a p p l i c a t i o n o f o r d i n a t i o n  expressing  envi-  Ithaca,.  York, U.S.A.  "A community can be c a l l e d fragmentary i f i t l a c k s s p e c i e s t h a t are u s u a l l y p r e s e n t i n the r e c u r r i n g p l a n t assemblages o f t h i s k i n d "  55  (Mueller-Dombois and E l l e n b e r g 1974). N e i t h e r species poor, nor fragmentary v e g e t a t i o n s , nor " a t y p i c a l " t r a n s i t i o n communities a r e o m i t t e d i n t h e p r e s e n t s t u d y , s i n c e they may be i m p o r t a n t i n t h e u n d e r s t a n d i n g o f s y n e c o l o g i c a l h a b i t a t d i f f e r e n c e s and s y n t a x o n o m i c a l p o s i t i o n o f s p e c i e s groups and v e g e t a t i o n types (Barkman 1969, Hale 1955, C u l b e r s o n 1955, H e r t e l 1974) 2. W i t h i n t h e frame work and s e t o f o b j e c t i v e s i n the p r e s e n t s t u d y , i t i s i n t e r e s t i n g t o note t h e f o l l o w i n g f i n d i n g s o f Foote (1963): " S e v e r a l stands on p o r t i o n s o f t h e continuum a r e d i s c u s s e d and t h e v e g e t a t i o n s o f t h e s e stands compared t o a s s o c i a t i o n s r e c o g n i z e d i n Europe. Many o f t h e European a s s o c i a t i o n s a r e s i m i l a r t o t h e v e g e t a t i o n o f c e r t a i n stands i n t h i s i n v e s t i g a t i o n . I t appears t h a t t h e bryophyte and l i c h e n v e g e t a t i o n o f m i d d l e Europe i s p r o b a b l y a continuum s i m i l a r t o t h a t found i n t h e p r e s e n t s t u d y . Because t h e European method o f samp l i n g includes only " t y p i c a l " stands, intermediate vegetations are i g n o r e d . Such s t u d i e s r e s u l t i n a c l a s s i f i c a t i o n o f s e p a r a t e , w e l l d e f i n e d a s s o c i a t i o n s r a t h e r than a continuum o f s p e c i e s " .  56  7. SYNSYSTEMATICS.  7.1. T e r m i n o l o g y and nomenclature.  C e n t e r i n g on t h e n a t u r e o f p l a n t communities i . e . e p i l i t h i c bryophyte communities, i t i s i m p o r t a n t on t h e one hand t o r e c o g n i z e the  h e t e r o g e n e i t y o f s p e c i e s d i s t r i b u t i o n , y e t t o emphasize on t h e o t h e r  hand t h e i n t e r a c t i o n s between p l a n t s i n t h e community. In t h i s the of  respect,  p l a n t community o r phytocoenose has a c e r t a i n i n d i v i d u a l i t y r e l a t i v e d i s c o n t i n u i t i e s between communities i n t h e f i e l d .  because  This  a p p l i e s , e s p e c i a l l y , t o rocky s u b s t r a t a which, i n g e n e r a l , are s t r o n g l y d i s c o n t i n u o u s i n space. Moreover, t h e i n d i v i d u a l  rock s i t e s  (boulders,  o u t c r o p s , r o c k w a l l s ) d i f f e r w i d e l y i n form and s i z e . Each s i t e , i n t u r n , may be v e r y heterogeneous i n e c o l o g i c a l f e a t u r e s ( a s p e c t , i n c l i n a t i o n , exposure, m o i s t u r e ) .  W i t h i n t h e c o n t e x t o f t h i s t h e s i s phytocoenose and phytocoenon are  d e f i n e d as f o l l o w s ( a f t e r Westhoff and Maarel 1973): phytocoenose: a p a r t o f a v e g e t a t i o n c o n s i s t i n g o f i n t e r a c t i n g p o p u l a t i o n s growing i n a u n i f o r m environment and showing a f l o r i s t i c  c o m p o s i t i o n and s t r u c t u r e t h a t  i s r e l a t i v e l y u n i f o r m and d i s t i n c t from t h e surrounding vegetation phytocoenon:  a t y p e o f phytocoenose d e r i v e d from t h e c h a r a c t e r i z a t i o n o f a group o f phytocoenoses  corres-  57  ponding w i t h each o t h e r i n a l l c h a r a c t e r s t h a t a r e considered t y p o l o g i c a l l y relevant In a p p l y i n g t h e Z i i r i c h - M o n t p e l l i e r s c h o o l method o f v e g e t a t i o n analysis  ,  t h e p r e s e n t study seeks t o c l a s s i f y e p i l i t h i c  assemblages, as p h y t o c o e n a , i n t o s y n s y s t e m a t i c A l l i a n c e , A s s o c i a t i o n , e t c . ) based on constancy  bryophyte  u n i t s ( C l a s s , Order, and f i d e l i t y o f t a x a  and d e f i n e d by c h a r a c t e r s p e c i e s , d i f f e r e n t i a t i n g s p e c i e s and companions. The  fundamental u n i t o f t h e . h i e r a r c h y i s t h e a s s o c i a t i o n , a u n i t  t h a t corresponds idiotaxonomy,  i n f u n c t i o n t o t h e s p e c i e s as t h e fundamental u n i t i n  o r t h e c l a s s i f i c a t i o n o f i n d i v i d u a l organisms. The  a s s o c i a t i o n can be d e f i n e d as "a phytocoenose i d e n t i f i e d by i t s c h a r a c teristic  species combination,  i n c l u d i n g one o r more c h a r a c t e r - t a x a o r  d i f f e r e n t i a t i n g taxa". Character species are species t h a t are r e l a t i v e l y r e s t r i c t e d t o samples o f a g i v e n phytocoenon, and t h e r e f o r e c h a r a c t e r i z e i t and i n d i c a t e i t s environment. One o r more c h a r a c t e r s p e c i e s a r e used f o r t h e c h a r a c t e r i z a t i o n o f any syntaxon  from t h e a s s o c i a t i o n t o t h e c l a s s . Degrees o f  r e s t r i c t i o n t o a g i v e n taxon a r e e x p r e s s e d  by t h e degree o f f i d e l i t y  (Appendix I I ) . Hence, c h a r a c t e r s p e c i e s c h a r a c t e r i z e s y n t a x a by t h e i r normal ( o r sometimes h i g h ) o c c u r r e n c e  i n phytocoenose o f t h a t  c o n t r a s t e d w i t h t h e i r absence o r l e s s f r e q u e n t o c c u r r e n c e of other  syntaxon,  i n phytocoenoses  syntaxa. I t i s p o s s i b l e a l s o t o d i s t i n g u i s h c l o s e l y r e l a t e d s y n t a x a by  the presence and absence o f c e r t a i n s p e c i e s w i t h o u t concern broader  about t h e  d i s t r i b u t i o n o f those s p e c i e s i n o t h e r phytocoenoses. As s u c h , two  58  s u b a s s o c i a t i o n s o f an a s s o c i a t i o n may  be d i s t i n g u i s h e d and c h a r a c t e r i z e d  by means o f d i f f e r e n t i a t i n g s p e c i e s . F i n a l l y , a t h i r d group o f d i a g n o s t i c s p e c i e s w i t h d i a g n o s t i c v a l u e are t h e companions which o c c u r i n most r e l e v e s o f a s y n t a x o n but are not d e s i g n a t e d  as c h a r a c t e r or d i f f e r e n t a t i n g s p e c i e s . Companions  are added t o the c h a r a c t e r t a x a t o form the " c h a r a c t e r i s t i c combination"  f o r a s s o c i a t i o n s and h i g h e r  species  syntaxa.  With r e s p e c t t o e p i l i t h i c b r y o p h y t e assemblages, v a r i o u s have d e s c r i b e d a l a r g e number o f b r y o p h y t e communities w i t h o u t  authors  proper  a s s o c i a t i o n r e c o g n i t i o n . These b r y o p h y t e communities a r e based on dominance r a t h e r than f i d e l i t y o f t a x a . As W e s t h o f f and Maarel s t a t e " t h e r e i s no n e c e s s a r y  r e l a t i o n between f i d e l i t y and  (1973)  dominance;  c h a r a c t e r t a x a can as w e l l be minor as major ones". In the p r e s e n t study attempts a r e made t o f o l l o w as as p o s s i b l e t h e p r i n c i p l e s o f the Z i i r i c h - M o n t p e l l i e r s c h o o l  stringently i n the  d e f i n i t i o n o f s y n t a x a by means o f c h a r a c t e r and d i f f e r e n t i a t i n g Where p o s s i b l e , the s y n s y s t e m a t i c  u n i t s as d i s t i n g u i s h e d i n t h i s  taxa. study  a r e i n c o r p o r a t e d i n e x i s t i n g s y n t a x a , many o f them d e s c r i b e d from e p i l i t h i c b r y o p h y t e assemblages i n Europe. In accordance t o nomenclature p r o p o s a l s as t h o s e o f Neuhausl (1968) and name-assigning r u l e s o f Bach, Knock and Moor (1962) new  u n i t s a r e d e s c r i b e d . For the s t a n d a r d i z a t i o n  o f syntaxon names, t h e f o l l o w i n g procedures are used. To t h e g e n e r i c p a r t o f the names o f one o r two  characteristic  t a x a o f a syntaxon a s u f f i x i s added. These s u f f i x e s a r e s p e c i f i c f o r the d i f f e r e n t syntaxonomic l e v e l s :  59  -etalia  f o r order  -ion  for alliance  -etum  for association  -etosum  f o r subassociation  When d e a l i n g w i t h an a s s o c i a t i o n o r a l l i a n c e t h a t i s s u f f i c i e n t l y c h a r a c t e r i z e d by one taxon  ( e . g . by a f a i t h f u l  t e r t a x o n ) , t h e genus name o f t h e taxon suffix  dominant  charac-  i s used w i t h t h e a p p r o p r i a t e  (-etum, - i o n ) , f o l l o w e d by t h e s p e c i e s name i n t h e g e n i t i v e . When a syntaxon was t o be named a f t e r  two c h a r a c t e r t a x a , t h e  name o f t h e second taxon was p r o v i d e d w i t h t h e a p p r o p r i a t e s u f f i x ,  while  the name o f t h e f i r s t taxon was j o i n e d w i t h t h e second one by t h e s u f f i x - e t o . (The f i r s t name may be a dominant s p e c i e s , g e n e r a l l y t h e second name r e p r e s e n t s t h e c h a r a c t e r taxon c o n s i d e r e d most i m p o r t a n t ) .  diagnostically  60  8. ANALYSIS AND DESCRIPTION OF THE EPILITHIC BRYOPHYTE VEGETATION.  The raw data s e t , containing a l l releves i n columns and a l l species i n rows, i s subdivided into the f o l l o w i n g major groups:  1. releves from exposed, dry rocks 2. releves from shaded, humid rocks 3. releves from s i l i c e o u s rocks along streams 4. releves from calcareous rocks, both exposed and shaded  A c l u s t e r a n a l y s i s i s performed on each o f these major groups, in order t o recognize groups of "similarr.eTe.ves". In analysing the phytos o c i o l o g i c a l data these c l u s t e r s function as the s t a r t i n g point i n the preparation o f the a s s o c i a t i o n t a b l e s . A f t e r the preparation o f the raw t a b l e and the presence t a b l e , VEGTAB i s given the command to rearrange the order of re-1 eves according to the sequence shown i n the c l u s t e r a n a l y s i s . This r e s u l t s i n the organization o f a " d i f f e r e n t i a t e d " t a b l e , i n which, subsequently, the c l u s t e r s are drawn i n by v e r t i c a l l y separating columns i n t o groups of releves (see Appendix Vi),. The f l o r i s t i c composition of the c l u s t e r s i s studied and r e l a t e d to f i e l d data with respect to e.g. geographic d i s t r i b u t i o n and habitat c h a r a c t e r i s t i c s . Then, w i t h i n these c l u s t e r s , attempts are made to recognize diagnostic species: recognition o f order, a l l i a n c e and a s s o c i a t i o n character species and, where possible and appropriate, d i s t i n c t i o n  61  o f s u b a s s o c i a t i o n s , v a r i a n t s and f a c i e s based on the presence and d e s i g n a t i o n o f d i f f e r e n t i a t i n g s p e c i e s . The r e s u l t s a r e shown i n T a b l e I I I , X I , XV, XXV and Appendix V ( w i t h i n t h i s Appendix a l i s t i s p r o v i d e d f o r each major group, which " t r a n s l a t e s " t h e releve-number i n t h e c l u s t e r a n a l y s e s i n t o t h e releve-number as taken i n the f i e l d given i n the separate a s s o c i a t i o n  tables).  F i n a l l y , t h e c l u s t e r a n a l y s e s may  s e r v e t o i n d i c a t e syntaxono-  mic r e l a t i o n s h i p s a t the v a r i o u s l e v e l s o f the c l a s s i f i c a t i o n and between  and  hierarchy  s y n t a x a a t t h e same l e v e l . To enhance and c l a r i f y t h e s e  r e l a t i o n s h i p s r e f e r e n c e s a r e made t o f l o r i s t i c c o m p o s i t i o n and  habitat  o f the b r y o p h y t e assemblages.  The c l u s t e r a n a l y s i s p r o c e d u r e , i t s r e s u l t i n each major group, a r e shown and d i s c u s s e d on the f o l l o w i n g pages. In t h e o f major group number 4, the a p p l i c a t i o n o f o r d i n a t i o n the d a t a w i l l  discussion  t e c h n i q u e s on  a l s o be e v a l u a t e d . These methods a r e used t o enhance  c l a s s i f i c a t i o n as a c h i e v e d by the t a b u l a t i o n l y s i s . They a l s o may  the  t e c h n i q u e and c l u s t e r ana-  c l a r i f y and f u r t h e r e x p r e s s r e l a t i o n s h i p s  groups o f r e l e v e s and t h e i r a f f i n i t i e s t o the environment.  between  62  8.1  BRYOPHYTE ASSOCIATIONS FROM EXPOSED,  DRY  ROCKS.  T a b l e I I I . B r y o p h y t e a s s o c i a t i o n s from exposed, dry r o c k s .  Order RACOMITRIETALIA HETEROSTICHI P h i l i p p i  1956  A l l i a n c e ANDREAION PETROPHILAE Hadac and K l i k a  C l u s t e r 1.  1944  A s s o c i a t i o n R a c o m i t r i o ( h e t e r o s t i c h i ) - Andreaeetum  p e t r o p h i l a e Frey 1922  A s s o c i a t i o n - typicum Gymnomitrion obtusum - v a r i a n t  Campy1 opus a t r o v i r e n s - Paraleucobryum enerve - f a c i e s  C l u s t e r 2.  Association  " R a c o m i t r i e t u m b r e v i p i s " a s s . nov.  C l u s t e r 5.  A s s o c i a t i o n Gymnomitrietum  concinnati  Herzog  1943  A s s o c i a t i o n - typicum T r i t o m a r i a quinquedentata - Andreaea b l y t t i i Dicranum f u s c e s c e n s - B a r b i l o p h o z i a Andreaea r o t h i i - v a r i a n t  - variant  floerkei - variant  Table I I I (continued).  C l u s t e r 6.  A s s o c i a t i o n R a c o m i t r i e t u m s u d e t i c i Herzog 1943  Order POLYTRICHETALIA PILIFERI v. Hubschmann 1967  A l l i a n c e POLYTRICHION PILIFERI Norr 1969  C l u s t e r 3.  Association Racomitrio - Polytrichetum  p i l i f e r i Herzog 1943  C l u s t e r 4.  A s s o c i a t i o n Scapanietum americanae a s s . nov.  65  8.1.  Bryophyte a s s o c i a t i o n s from exposed, d r y r o c k s (Table I I I ) .  8.1.1.  Synsystematics.  W i t h i n the a l l i a n c e A n d r e a e i o n p e t r o p h i l a e Hadac and 1944,  Philippi  (1956) d i s t i n g u i s h e d t h r e e a s s o c i a t i o n s : the R a c o m i t r i o -  Andreaeetum p e t r o p h i l a e Frey 1922,  the Andreaeetum r o t h i i  and the Gymnomitrietum c o n c i n n a t i Herzog 1943. was  Klika  o r i g i n a l l y d e s c r i b e d as two  (1943): a Racomitrium sudeticum-  Philippi  1956  This l a s t a s s o c i a t i o n  s e p a r a t e v e g e t a t i o n u n i t s by Herzog and a Gymnomitrion concinnatum -  Andreaea r u p e s t r i s - a s s o c i a t i o n . In t h e p r e s e n t s t u d y , t h e s e two  v e g e t a t i o n types a r e  recognized  s e p a r a t e l y a g a i n and a r e g i v e n the s t a t u s o f a s s o c i a t i o n , based on the dominance and f i d e l i t y o f t h e i r r e s p e c t i v e c h a r a c t e r s p e c i e s , Racomitrium sudeticum  and Gymnomitrion concinnatum, r e s p e c t i v e l y .  F i n a l l y , a new  a s s o c i a t i o n i s proposed, the  "Racomitrietum  b r e v i p i s " . o f which v a r i o u s a l l i a n c e c h a r a c t e r s p e c i e s (Andreaea r u p e s t r i s , K i a e r i a s t a r k e i , Lophozia s u d e t i c a ) show i t s a f f i l i a t i o n w i t h the o t h e r t h r e e a s s o c i a t i o n s , t h u s , i n d i c a t i n g i t s syntaxonomic p o s i t i o n w i t h i n the A n d r e a e i o n p e t r o p h i l a e . B o t h , the R a c o m i t r i o - Andreaeetum p e t r o p h i l a e and t h e "Racom i t r i e t u m b r e v i p i s " show a wide e l e v a t i o n a l range. The occurs predominantly  f i r s t association  a t lower e l e v a t i o n s , whereas the l a t t e r one  shows  i t s b e s t development i n s u b a l p i n e a r e a s . On exposed a l p i n e r o c k s  these  a s s o c i a t i o n s o c c u r o n l y r a r e l y : h e r e , t h e Gymnomitrietum c o n c i n n a t i and  66  the R a c o m i t r i e t u m s u d e t i c i a r e the most f r e q u e n t and L i k e Neumayr (1971) and  Hertel  ( 1 9 7 4 ) , t h i s s t u d y , synsystema-  t i c a l l y , p l a c e s the a l l i a n c e A n d r e a e i o n p e t r o p h i l a e mitrietalia heterostichi Philippi  dominant a s s o c i a t i o n s .  i n the o r d e r Raco-  1956.  From exposed r o c k s , a second group o f a s s o c i a t i o n s described,  The  are c o n f i n e d  elevations  t o low  two  syntaxa, distinguished and  s l i g h t l y s l o p i n g rock surfaces  s o i l and  i n the p r e s e n t study  c h a r a c t e r i s t i c a l l y grow on  Herzog 1943  o v e r l a i n by a m i x t u r e o f m i n e r a l  f i r s t a s s o c i a t i o n , the R a c o m i t r i o - P o l y t r i c h e t u m has been d e s c r i b e d  Norr 1969,  Neumayr 1971,  from s e v e r a l  been p l a c e d i n the  alTv  piliferi  l o c a t i o n s i n Europe (Herzog  Hubschmann 1975)  and  i s also  from Vancouver I s l a n d (Hubschmann 1978). S y n s y s t e m a t i c a l l y has  horizontal  humus. The  1943,  been  which are c l a s s i f i e d i n the o r d e r P o l y t r i c h e t a l i a p i l i f e r i  V. Hubschmann 1967.  and  has  Polytrichion p i l i f e r i  reported the  by Norr (1969).  In the p r e s e n t s t u d y , a newly proposed a s s o c i a t i o n , the nietum americanae, i s d e s c r i b e d conditions  from s i m i l a r h a b i t a t s and  as the p r e v i o u s a s s o c i a t i o n . I t s own  based on c h a r a c t e r  Scapa-  ecological  i d e n t i t y , however, i s  as w e l l as t y p i c a l accompanying  species.  Its  syntaxonomic a f f i n i t y t o the R a c o m i t r i o - P o l y t r i c h e t u m  piliferi  its  subsequent placement i n the same a l l i a n c e and  additional  species  association  companion  species.  order i s expressed  and by  67  8.1.2. P h y t o g e o g r a p h i c a l s e t t i n g .  Both o r d e r s R a c o m i t r i e t a l i a h e t e r o s t i c h i and P o l y t r i c h e t a l i a p i l i f e r i c o n t a i n many c i r c u m b o r e a l b r y o p h y t e s p e c i e s d e s i g n a t e d as a s s o c i a t i o n c h a r a c t e r s p e c i e s o r i m p o r t a n t companions.  Among t h e s e t h e most  f r e q u e n t and abundant a r e D i p l o p h y l l u m t a x i f o l i u m , Gymnomitrion  concin-  natum, Marsupel1 a e m a r g i n a t a , Racomitrium h e t e r o s t i c h u m , R. c a n e s c e n s , K i a e r i a s t a r k e i , P o h l i a nutans and Andreaea r u p e s t r i s. The r e p r e s e n t a t i o n o f s p e c i e s , endemic t o w e s t e r n North America i s o f minor importance i n most b r y o p h y t e a s s o c i a t i o n s from exposed, d r y r o c k s . An obvious e x c e p t i o n i s "Racomitrium b r e v i p e s " , which i s t r e a t e d as a s e p a r a t e t a x o n o m i c a l e n t i t y i n t h i s s t u d y . Because o f i t s c o n s t a n c y , t h i s w e s t e r n North American endemic i s d e s i g n a t e d as t h e c h a r a c t e r s p e c i e s f o r t h e newly proposed a s s o c i a t i o n " R a c o m i t r i e t u m b r e v i p i s " . In a d d i t i o n , o t h e r s i g n i f i c a n t l y accompanying  endemics  i n r e l e v e s from low e l e v a t i o n s  a r e S c a p a n i a americana and Hypnum c i r c i n a l e . F r e q u e n t l y o c c u r r i n g b i p o l a r d i s j u n c t b r y o p h y t e s a r e s p e c i e s such as B a r b i l o p h o z i a f l o e r k e i , B_. h a t c h e r i and B a r t r a m i a p o m i f o r m i s .  8.1.3.1. Order RACOMITRIETALIA HETEROSTICHI P h i l i p p i 1956  A l l i a n c e A n d r e a e i o n p e t r o p h i l a e Hadac and K l i k a 1944.  R a c o m i t r i o ( h e t e r o s t i c h i ) - Andreaeetum p e t r o p h i l a e Frey 1922 ( T a b l e I V ) .  In t h e s t u d y a r e a t h i s a s s o c i a t i o n i s found and d e s c r i b e d from  68  e l e v a t i o n s r a n g i n g from sea l e v e l  ( L i g h t h o u s e Park) t o a l p i n e a r e a s .  T h i s apparent e l e v a t i o n independence d i f f e r s from the s i t u a t i o n i n Europe, where Andreaea r u p e s t r i s reaches i t s l o w e s t e l e v a t i o n around 450 m, and t h i s a s s o c i a t i o n ranges from s u b a l p i n e t o a l p i n e a r e a s . Reports and documentation on t h e p r e s e n t syntaxon a r e g i v e n by Frey ( 1 9 2 2 ) , Herzog ( 1 9 4 3 ) , P h i l i p p i Hertel  (1974). From Vancouver  a.o.  (1956, 1965), Neumayr (1971)  and  I s l a n d Hubschmann (1978) d e s c r i b e d s m a l l  fragments o f t h i s a s s o c i a t i o n . H o r i k a w a , Ando and Kawai (1961)  describe  Andreaea r u p e s t r i s v a r . f a u r i e r i - and Racomitrium h e t e r o s t i c h u m communities from the a l p i n e zone o f Mount Hakusan, Japan. A l t h o u g h t h e i r sample methods d i f f e r e d s i g n i f i c a n t l y , t h e data i n d i c a t e s u b s t a n t i a l a f f i n i t i e s with s i m i l a r vegetations i n south-western B r i t i s h and  Columbia  Europe. The R a c o m i t r i o - Andreaeetum i n t h e p r e s e n t s t u d y shows a wide  g e o g r a p h i c d i s t r i b u t i o n w i t h i n t h e Coast Mountains area o f B r i t i s h Columbia, from l o c a l i t i e s d i r e c t l y n o r t h o f Vancouver t o N a i r n  Falls  ( F i g u r e 1 ) . S i n c e both c h a r a c t e r s p e c i e s , Racomitrium h e t e r o s t i c h u m and Andreaea r u p e s t r i s , o c c u r o v e r a much l a r g e r a r e a t h r o u g h o u t the p r o v i n c e ( S c h o f i e l d 1976, Tan 1980), t h e a s s o c i a t i o n i s l i k e l y t o be more widespread than encountered i n the p r e s e n t s t u d y . The a s s o c i a t i o n p r e d o m i n a n t l y o c c u r s on dry and exposed  rocks,  where i t forms l u s h b r y o p h y t e assemblages. A t lower e l e v a t i o n s t h e s e rocks a r e m o s t l y i n t r u s i v e i n n a t u r e ; i n a l p i n e a r e a s t h e a s s o c i a t i o n i s a l s o d e s c r i b e d from metamorphic r o c k s . Frequent a s s o c i a t e s o f t h e c h a r a c t e r s p e c i e s a r e b r y o p h y t e t a x a  Table IV.  Number of releves Releve-number Elevation(m) Direction of exposure Inclination ) Total cover(%) Q  Racomitrium heterostichum var. heterostichum Andreaea rupestris Gymnomitrion obtusum Diplophyll urn taxi folium Lophozia ventricosa var. ventricosa Pohlla nutans Kiaeria starkei Marsupella emarginata Lophozia sudetica Scapania americana Racomitrium sudeticum Polytrichum alpinum var. macounii Polytrichum piliferum Grimmia torquata Cephaloziella byssacea var. asperifolia Anastrophyllum minutum Dicranella heteromalla Oryptodon patens Homalothecium nevadense Cephaloziella byssacea var. byssacea Racomitrium heterostichum var. affine Scapania scandica Pohlia cruda Bartramia pomiformis Dicranum scoparium Cynodontium jenneri Anthelia juratzkana Marsupella sparsifolla Hypnum circinale Isopterygium elegans Marsupella condensata Pseudobraunia californica Plagiochila asplenioldes Bryum caespiticium Herbertus aduncus Cephaloziella turnerl Chandonanthus setiformis Ptilidium c i l i a r e  1 2 2 9 11 36 61 E-SE E-SE 78 30 80 75  3 15 61 S-SW 46 80  4 5 179 119 1433 610 E-SE N 68 32 68 85 100  6 181 1433 E-SE 84~ 84 100  5.5 5 .5  4 4.4 .4 1.2  4 4.5 .5  3 .4 3.4  4.4 4 .4  1 1.4 .4  _  1.2 +.1 + .1 1.2  1.4 1 .4 2.4  1.2 2.2 3.4 3 .4  3 .4 3.4 1.2 +.1 + .1 2.2 1.4 1.2  2.2  Racomitrio (heterostichi) - Andreaeetum 7 8 9 10 11 12 7 11 8 13 26 243 245 244 331 288 18 1492 1492 1492 610 610 E NM N NW E-NE NW E ~66~ 32 46 24 24" 32 66 32 24 76 75 80 90 80 90 85 75 80 75 1.2 2.2  3 .4 3.4 3.2 3.2 2 .4 2.4 2.4 2.4  4 .4 4.4 1.1  1 .4 1.4  1.2  2 .4 2.4  3 .4 3.4 2 .4 2.4 1-4 1.4 +.4.4  2 .4 2.4 1 .4 1.4 2 .4 2.4 1 .4 1.4  3 .4 3.4 1 .4 1.4 +.4.4 1 .4 1.4  2.4 2.4 1.4 +.4  2.2  +-4 .4 1.1 2.4 2-*  22.4 .4 22.4 .4 +.1 + .1 1.2 1-2 1.2 1-2 1.2  1.4 1.4  1.4 1 .4  1 1.4 .4  1.4 +.1 1-2 .4 1.4  +  +  3.3  1 .4 1.4  14 289 610 E 102 90  15 290 610 E 98 90  16 17 18 65 17 332 61 61 610 NW N NW NW 66 64 74 90 60 100  3.4 1.2 2.4  4.4 2.4 2.4  3.4 2.4 2.2 + .4 4.4 3.4  1.4  1.4  1.4  1.2  + .4 + .4  2.4  1.2  +.1  2.4 3.4 1.4 + .1  19 20 178 134 1433 427 E SE N 72 50 90 100  21 135 427 NW 62 100  3.4 2.4 3.4  3.4 2.4 3.4 2.4  2.4 3.4 1.4 1.4  2.4 2.4 1.4 2.4  1-1  + +  +.4  1.4 1.2 3.4  petrophilae Frey 1922  1.4 2.2  .l -2  2.4  1.2  1.2  1.2  2.4  2.4  3.4  ]-2 1-4  1.2 1-4  1.4 1.2 2.2  1 .2  +.4  +.4 2.2  1.4  +.1 +.1  2.4 2.2  4.4 2.4  1.4  1.4  4.5  1.4  4.5  3.4  1 .4 1.4  + .4  1.1  + .1  1.1  1.2  2.3  3.4  4.4  3.3  • 1.2  1.4  1.4  +.4  1 - 2  3.4 1.4  2.4  2.4  1.2  + .1  +.1 1  +,4 1.4  2.4 3.4 3.4 3.4  25 21 18 N 84 90  4.4 2.2  + .1 + .1 + .1  '.2  1.4  2.4 2.4 2.4 1.4  24 223 610 W 68 80  1.4  +.4  1.2  3.4 + .1 1.2  22 23 182 222 1433 610 E SE W 88 70 100 70  +.1  4 +,4 '  4  +.4 +.1  1.2  1.1 1.4  +.4  1.4 '  1.4 '  +1 11 ' i 4  3  2  > « 4  -2  *°  T a b l e IV ( c o n t i n u e d ) .  Found o n l y once:  Gymnomitrion concinnatum, Racomitrium canescens, B a r b i l o p h o z i a f l o e r k e i , Andreaea n i v a l i s , D i c r a n o w e i s i a c i r r a t a , Hypnum subimponens, Claopodium b o l a n d e r i , Ceratodon purpureus, Dicranum f u s c e s c e n s , P h i l o n o t i s f o n t a n a , Barbilophozia h a t c h e r i , P t i l i d i u m c a l i f o r n i c u m , PIagiothecium p i l i f e r u m , P o l y t r i c h u m j u n i p e r i n u m , Chandonanthus f i 1 i f o r m i s , D i p l o p h y l l u m a l b i c a n s , Racomitrium lanuqinosum, A n a c o l i a m e n z i e s i i , Scleropodium o b t u s i f o l i u m , Campy!opus paradoxus, Amphidium c a l i f o r n i c u m , L o p h o z i a w e n z e l i i , Timmia a u s t r i a c a , Racomi t r i um fas cn c u l a r e , B a r b i 1 o p h o z i a l y c o p o d i o i des, B a z z a n i a t r i c r e n a t a , H e t e r o c l a d i u m macouni i .  R e l e v e - l o c a t i o n s : 9,11,13 15,17 178,179,181 ,182 119 21,25,26 243,244,245 331,332,333 288,289,290, 65 134,135 222,223 237,238,239  Horseshoe Bay M u r r i n P r o v i n c i a l Park Mount Seymour Cypress Bowl L i g h t h o u s e Park G a r i b a l d i Lake C a l l a g h a n Creek Nairn F a l l s Shannon F a l l s Brandywine F a l l s Cheakamus R i v e r V a l l e y B l a c k Tusk Meadows  71  such as Gymnomitrion obtusum, D i p l o p h y l 1 u m t a x i f o l i u m and L o p h o z i a ventricosa var. ventricosa. Rock i n c l i n a t i o n up t o about 70^ p e r m i t s some s o i l  and humus  a c c u m u l a t i o n underneath t h e b r y o p h y t e v e g e t a t i o n . In such s i t u a t i o n s f r e q u e n t accompanying  s p e c i e s a r e P o h l i a nutans and B a r b i l o p h o z i a  f l o e r k e i . On h o r i z o n t a l r o c k s u r f a c e s , where o f t e n a c o n s i d e r a b l e depos i t i o n o f humus can be f o u n d , t h e two c h a r a c t e r s p e c i e s o f t h e p r e s e n t association w i l l  e v e n t u a l l y be outcompeted  by Racomitrium c a n e s c e n s ,  P o l y t r i c h u m p i l i f e r u m and Dicranum f u s c e s c e n s , t h e f i r s t two t a x a b e i n g the  character species o f the Racomitrio - Polytrichetum p i l i f e r i .  exposed c o n d i t i o n s w i t h o n l y a s l i g h t humus l a y e r o r no s o i l  Under  accumu-  l a t i o n w h a t s o e v e r , Gymnomitrion obtusum, Marsupel1 a emarginata and C e p h a l o z i e l l a byssacea a r e i m p o r t a n t accompanying  species. At higher  e l e v a t i o n s e v e r a l s p e c i e s , c h a r a c t e r t a x a o f o t h e r a s s o c i a t i o n s from the  same a l l i a n c e become abundant w i t h i n the relieves. Examples  are  Racomitrium s u d e t i c u m , Gymnomitrion  and P s e u d o l e s k e a i n c u r v a t a . Accompanying  o f these  concinnatum, K i a e r i a b l y t t i i taxa with rare occurrences  ( a c c i d e n t a l s p e c i e s ) i n t h e s e reTeves from s u b a l p i n e - a l p i n e s i t e a r e M a r s u p e l l a c o n d e n s a t a , Andreaea n i v a l i s and A n t h e l i a j u r a t z k a n a .  Finally, a local  " f a c i e s " i s r e c o g n i z e d from s t e e p , wet r o c k s  at Shannon F a l l s ^. A t p l a c e s where some s o i l  had accumulated - i n and  around c r a c k s and toward t h e base o f t h e s e r o c k s - major s p e c i e s from the  R a c o m i t r i o - Andreaeetum p e t r o p h i l a e had e s t a b l i s h e d  r u p e s t r i s , A. r o t h i i ,  (Andreaea  Marsupel1 a e m a r g i n a t a , R a c o m i t r i um h e t e r o s t i c h u m  T a b l e V. R a c o m i t r i o ( h e t e r o s t i c h i ) - Andreaeetutn p e t r o p h i l a e Frey 1922. Campy!opus a t r o v i r e n s - Paraleucobryum enerve - f a c i e s Number o f r e l e v e s Releve-number Elevation(m) D i r e c t i o n ofgexposure Inclination( ) Total cover(%)  1 46 61 NW 60 75  2 347 61 N-NW 72 80  3 345 61 NW 58 75  4 348 61 NW 43 85  Campylopus a t r o v i r e n s Andreaea r u p e s t r i s M a r s u p e l l a emarginata Racomitrium h e t e r o s t i c h u m v a r . h e t e r o s t i c h u m Gymnomitrioni obtusum C e p h a l o z i e l l a byssacea v a r . byssacea Paraleucobryum enerve Chandonanthus f i l i f o r m e Diplophyllum albicans Andreaea r o t h i i Anastrophyllum a s s i m i l e Marsupella s p a r s i f o l i a  3.4 4.4 2.4 1.2 1.4 + .4 2.2 + .4  2.4 3.4 2.4 1.2 1.4 T.4 2.2  4.4 2.4 2.4 1.2 1.4 + .4 2.2 1.4 1.4 + .2 1.2  4.4 2.4 2.4 1.2 2.4 + .4 2.2 1.4 1.4 + .2 1.2  L o c a t i o n o f a l l r e l e v e s : .Shannon F a l l s .  1.4 + .2 + .2 + .4  5 346 61 N-NW 62 80 3.4 2.4 3.4 2.3 2.4 + .4 1.4 + .4 + .2  73  Gymnomitrion obtusion). However, under t h e s e c o n d i t i o n s , a c o m b i n a t i o n o f species, not o c c u r r i n g i n the present a s s o c i a t i o n , c o n s t i t u t e a f a c i e s c h a r a c t e r i z e d by t h e f o l l o w i n g t a x a : Campy!opus a t r o v i r e n s , P a r a l e u c o bryum e n e r v e , A n a s t r o p h y l 1 urn a s s i m i l e , D i p l o p h y l l u r n a l b i c a n s , Chandonanthus f i 1 i f o r m e . and C e p h a l o z i e l l a byssacea  var. byssacea-(Table V ) .  S i n c e most s p e c i e s i n t h i s f a c i e s have a d i s t r i b u t i o n  extending  c o n s i d e r a b l y beyond t h e p r e s e n t study a r e a , f u r t h e r r e s e a r c h i s needed in order t o designate t h i s  ' a c c i d e n t a l ' c o m b i n a t i o n o f s p e c i e s as a  s u b a s s o c i a t i o n o f t h e R a c o m i t r i o - Andreaeetum p e t r o p h i l a e , o r even r a i s e i t to a s s o c i a t i o n  rank.  "Racomitrietum b r e v i p i s " a s s . nov. ( T a b l e V I ) .  Problems have a r i s e n here around t h e taxon d e s i g n a t e d as c h a r a c t e r s p e c i e s f o r t h e a s s o c i a t i o n on hand. "Racomitrium b r e v i p e s " i s c l o s e l y r e l a t e d t o R. h e t e r o s t i c h u m , b u t d i f f e r s from i t by t h e m i x t u r e o f l o n g and s h o r t a p i c a l l e a f c e l l s and t h e r i d g e d ( r e s e m b l i n g p a p i l l a e ) c r o s s w a l l s i n l e a f c r o s s s e c t i o n (Banu 1969). As Ramsay and S c h o f i e l d (1981) d i s c u s s , t h e name "Racomitrium b r e v i p e s " Kindb. appears t o be synonymous w i t h R. sudeticum (Funck)~B.S.G. based on t h e apparent t y p e . A c c o r d i n g t o Lawton (1971) t h i s type c o l l e c t i o n o f R. b r e v i p e s Kindb. l a c k s t r u e p a p i l l a e and has b i s t r a t o s e l e a f margins. However, f o l l o w i n g Persson's (1947) s u g g e s t i o n s and Banu's (1969) c o n c e p t s , "j?. b r e v i p e s " has been t r e a t e d i n t h i s study as a s e p a r a t e t a x o n o m i c a l e n t i t y . As s t a t e d by Ramsay and S c h o f i e l d (1981) t h e name "R_. b r e v i p e s " Kindb. i s used f o r c o n v e n i e n c e , s i n c e t h e w e s t e r n North American m a t e r i a l i s c l e a r l y n o t i d e n t i c a l t o the European concept o f JR. h e t e r o s t i c h u m .  The  geographic d i s t r i b u t i o n o f "Racomitrium  b r e v i p e s " extends  c o n s i d e r a b l y beyond t h e p r e s e n t study a r e a : from t h e west c o a s t o f Vancouver I s l a n d t o t h e Queen C h a r l o t t e I s l a n d s , a l l a l o n g t h e n o r t h c o a s t and eastward  i n t o t h e P u r c e l l Mountains i n t h e Kootenays (Tan 1980).  Table VI. "Racomitrietum brevipis" ass. nov. Number of releves Releve-number Elevation(m) Direction of exposure Inclinat1on( ) Total cover(%)  1 2 252 236 1487. 1487 S-SE NE 68 38 100 100  3 258 1492 NE 50 90  "Racomitrium brevipes" Andreaea rupestris Kiaeria starkei Lophozia sudetica Dicranoweisia crispula Pseudoleskea radicosa Lophozia ventricosa var. ventricosa Barbilophozia floerkei Racomitrium sudeticum Diplophyllum taxifolium Scapania americana Polytrichum piliferum Pohlia nutans Racomitrium canescens Gymnomitrion concinnatum Ptilidium californicum Andreaea nivalis Marsupella stableri Polytrichum alpinum var. macounii Hypnum circinale Kiaeria b l y t t i i  2.4 1.2  1.2  2.4 2.4  n  Found only once:  4.5 3.4  1.2  4.4  4.4 3.4 1.4  1.4 1.2 3.4 1.1 1.4  4 5 265 264 1487 1487 S-SW S-SW 52 68 90 100  6 131 457 SW 38 100  7 133 457 NW 52 100  8 132 457 E 30 80  3.4 2.2 1.2  3.4 2.2 2.4 2.4  3.4 1.2 1.4 1.4 2.4  1.4 1.2 1.2 2.6 1.4 1.4  4.4 3.4 2.4  1.4  1.4  + .2 2.4 1.2  3.4  1.2 + .1 2.2  1.2 + .1 2.2  1.4  1.4  1.2 3.4  9 10 11 235 263 248 1487 1487 1487 NW W E-SE 44 62 26 90 100 90 4.4  1.4 1.4 + .2 1.2  1.4 1.2 2.2  1.2  4.4 1.2 1.4 1.4 1.4 2.4  1.2  1.4  + .1  + .1  16 15 257 262 1492 1487 NW NW 76 78 90 90  3.4  3.4  4.4  3.4 2.4  2.4 2.4  3.4 1.4  1.4  2.4  1.4  2.2 1.2 3.4 1.4 1.4  1.4  1.4 1.4 1.4  3.4  5.5 1.4  1.2  1.2 1.4  + .4  2.2  4.4  5.5 1.2  14 12 13 249 254 255 1487 1492 1492 E-SE NE NE 28 80 64 100 100 80  1.1 1.4  1.4  1.4 1.2  1.4  2.2 1.4  -p. 3.4  1.2  + .1  1.1  2.2  2.2  Cephaloziella byssacea var. byssacea, Scleropodium obtusifolium, Marsupella sparsifolia, Brachythecium albicans, Scapania scandica, Ceratodon purpureus, Anastrophyllum minutum, Bartramia ithyphylla, Polytrichum juniperinum, Dichodontium pellucidum, Hygrohypnum ochraceum, Pseudoleskea patens, Kiaeria falcata, Chiloscyphus polyanthos, Porotrichum bigelovii, Campylopus paradoxus, Cephalozia bicuspidata, Claopodium bolanderi, Barbilophozia hatcheri, Racomitrium heterostichum var. affine.  Releve-locations: 235,236.248,249,252,262,263,264,265  Black Tusk Meadows  254,255,257,258  Panorama Ridge  131,132.133  Brandywine Falls  ^1  75  Hence, t h e a s s o c i a t i o n i s l i k e l y t o o c c u r more w i d e s p r e a d than encount e r e d i n t h e p r e s e n t s t u d y . As such i t i s , a p a r t from low e l e v a t i o n l o c a l i t i e s such as M u r r i n c r i b e d from s u b a l p i n e  P r o v i n c i a l Park and Brandywine F a l l s , des-  s i t e d w i t h i n G a r i b a l d i P r o v i n c i a l Park.  Except f o r r a r e o c c u r r e n c e s i n o t h e r a s s o c i a t i o n s brevipes"  i s h i g h l y c o n s t a n t i n , hence, a good c h a r a c t e r  "Racomitrium species f o r  the "Racomitrietum b r e v i p i s " .  The a s s o c i a t i o n ' s r e l a t i o n s h i p t o t h e t h r e e . o t h e r the A n d r e a e i o n p e t r o p h i l a e character  syntaxa o f  i s i n d i c a t e d by t h e f r e q u e n c y o f t h e a l l i a n c e  s p e c i e s Andreaea r u p e s t r i s , K i a e r i a s t a r k e i and L o p h o z i a  s u d e t i c a . The " R a c o m i t r i e t u m b r e v i p i s " forms l u s h v e g e t a t i o n s  on g e n t l y  sloping surfaces  (sub)alpine  character  o f exposed i n t r u s i v e r o c k s . The predominant  o f t h i s a s s o c i a t i o n i s r e f l e c t e d i n t h e presence o f accompa-  nying species  such as Racomitrium s u d e t i c u m , Gymnomitrion concinnatum,  Bartramia i t h y p h y l l a , K i a e r i a b l y t t i i  and M a r s u p e l l a  stableri  Gymnomitrietum c o n c i n n a t i Herzog 1943 ( T a b l e V I I ) .  T h i s a s s o c i a t i o n has been r e p o r t e d and  and d e s c r i b e d  from  subalpine  a l p i n e areas i n t h e s o u t h e r n p o r t i o n o f West Germany (Herzog 1943,  Philippi  1956 and H e r t e l  1974)  . T h e i r s as w e l l as t h e p r e s e n t d a t a  show t h a t t h e Gymnomitrietum c o n c i n n a t i  replaces  Andreaeetum p e t r o p h i l a e a t t h e s e h i g h e r  elevations. Likewise i t s  character  species  the Racomitrio -  - Gymnomitrion concinnatum -, t h i s a s s o c i a t i o n i s  76 strictly  confined to subalpine and a l p i n e s i t e s . I t forms a low but  extensive bryophyte cover over steeply s l o p i n g exposed rocks surfaces. The a s s o c i a t i o n ' s most frquent and abundant accompanying species are Diplophyllum taxi f o l i u m, Marsupella emargi nata and Lophozia sudetica. Important a l l i a n c e character species are Lophozia ventricosa var. ventricosa and Andreaea r u p e s t r i s . This s t r i c t  subalpine-alpine charac-  ter of the a s s o c i a t i o n i s also r e f l e c t e d by accompanying species such as Andreaea n i v a l i s , A. r o t h i i ,  A. b l y t t i i , Pseudoleskea  baileyi,• Ahthelia  juratzkana, Marsupella condensata, K i a e r i a b l y t t i i and Bartramia i t h y 3  p h y l l a . A d d i t i o n a l associates c o n s t i t u t e three l o c a l 'variants' The Gymnomitrietum concinnati - typicum (1 i n Table VII) predominantly  occurs i n Cypress Bowl P r o v i n c i a l Park and Mount Seymour  (1433 m). Most frequent and abundant accompanying species i n these areas are Andreaea n i v a l i s and K i a e r i a s t a r k e i . Along G a r i b a l d i Lake (1492 m), Tritomaria quinquedentata, A n t h e l i a juratzkana, Nardia japonica, Andreaea b l y t t i i and Marsupel!a condesata contribute considerably to the Gymnomitrietum concinnati.This-combination of species c o n s t i t u t e s the Tritomaria quinquedentata-Andreaea b l y t t i i variant (2). In the Diamond Head area (1524 m), i n the southwest corner of Garibaldi P r o v i n c i a l Park, Barbilophozia f l o e r k e i , Dicranum fuscescens, Dryptodon patens and Racomitriurn heterostichum var. a f f i n e were frequent accompanying species. This group of species c o n s t i t u t e s the Dicranum fuscescens-Barbilophozia f l o e r k e i - variant ( 3 ) , which, c h a r a c t e r i s t i c a l l y , occurs on gently s l o p i n g rock surfaces with considerable s o i l accumu-  Table VII. Gymnomitrietumconcinnati Herzog 1943 Number of releves Releve-number Elevation(m) Direction of exposure Inclination( ) Total coverU)  1 116 610 N-NW 106 100  Gymnomitrium concinnatum Diplophyllum taxifollum Marsupella emarginata Lophozia sudetica Racomitrium sudeticum Lophozia ventricosa var. ventricosa Andreaea rupestris Pohlia nutans "Racomitrium brevipes" Polytrichum alpinum var. alpinum Pohlia cruda Bartramia ithyphylla Cephalozia bicuspidata Andreaea nivalis Kiaeria starkei Pseudoleskea baileyi Anastrophyl1 urn minutum Dicranella heteromalla BIindia acuta Marsupella sparsifolia Oligotrichum hercynicum Tritomaria quinquedentata Anthelia juratzkana Nardia japonica Andreaea b l y t t i i Marsupella condensata Cephaloziella byssacea var. byssacea Scapania scandica Barbilophozia floerkei Dicranum fuscescens Dryptodon patens Racomitrium heterostichum  4.4 1.2 4.5  n  u  var. affine Cynodontium jenneri Pt111di urn californicum Andreaea rothii Amphidium californicum Campylopus fragilis Arctoa fulvella Kiaeria b l y t t i i  2 3 4 170 117 169 610 1433 1433 NW N-NW NW 76 100 72 95 100 90 3.4 1.2 2.4 2.4  2.4  4.5  2.4 1.4  1.4 2.4  5 122 610 N 74 80  6 123 610 N 64 100  7 174 1433 NW 80 80  8 175 1433 N-NW 62 100  2.4 1 .2 2.4 2.4 1.2  + .4 2.4 2.4 + .4  2.4  3.4 2.4  1.4 3.4  2.4 1.2  4.4 2.2  4.4 2.2  2.3 +.2  4.5 1.4  4.5 2.4  2.4 3.4  3.4 1.4  1.4 2.2  1.4 1.2  1.4  2.4  3.4  1.4 1.2  1.4 1.2  1.2  2.4  1.4  1.2  4.4 2.2  + .1 + .1 1.2  1.2 2.2  3.4  3.4 1.2 1.4  +.4 1.4  4.5 2.4  1.2 2.4  1.2  11 247 1487 N 60 100  3.4 2.4 1.4  +.2 3.3  1.1 4.4  4.4  +.4  1.4  2.2  3.3  2.4 1.2  1.2 4.4 1.4 1.4  3.4  3.4  1.4  1.4  1.4 1.4  1.4 1.4  12 240 1492 SW 58 100  14 13 242 241 1492 1492 SW SW 48 60 90 100  9 10 173 246 1433 1487 NE N 68 54 100 100  15 176 1433 E 76 80  16 177 1433 E 84 100  3.4  2.4 1.2  1.4 1.2  + .1  + .1 1.2  1.1 1.2  2.4 1.4 1.4 1.2 +.4  3.4 2.4 1.4 1.2 1.4  1.4 2.4 2.4 1.2 +.4  2.2  17 18 334 335 1524 1524 NW N NE 32 52 95 80  19 20 338 336 1524 1524 NW S 58 58 100 80  1.2 1.2 1.4 1.4  3.4 + .2 1.4  3.4 + .2 1.4  2.4 + .2 + .2  2.4 + .2 + .2  2.4 + .2 2.3  21 337 1524 SE 34 90  3.4  22 23 311 312 1219 1219 N NW NE 62 40 100 90  24 315 121 N NE 74 100  1.4  4.5 1.4 4.4 1.4 1.2  4.5 2 1.2 3.4 3 + .2 1 1.2 +  + .2 2.2  + .2 2.2  1.2  + .1 + .1  1.1  1.2 1.2 1.2 1.2 1.2  3.4 +.2 1.2 +.2 +.2  25 313 1219 E NE 78 100  26 314 1219 C NE 66 100  4  2.4  2.4  4 4 2  2.4 2.4 2.4  3.4 1.4 2.4  f  1.2 1.2  1 -2  3.4  4.4  + 1  1.4  2  2.4  1.4  3  3.4 3.3 1.2  2.4 2.2 1.2  2.2 3.4  3.3 2.4  +.4 1.2 3.3  +.4 2.3 1.2  +.4 1.2 1.2  2.4 2.2  1.4 1.2  1.2  +.2  1.2  1.2  4.4  4  2.4 1.2  Table VII (continued).  Found o n l y once:  Hypnum c i r c i n a l e , P o h l i a l u d w i g i i , Racomitrium lanuginosum, Herbertus aduncus, Hyqrohypnum ochraceum, Racomitrium a c i c u l a r e , M a r s u p e l l a s p h a c e l a t a , Pel 1 i a n e e s i a n a , Brachytheciurn s t a r k e i , Pseudoleskea i n c u r v a t a , Jungermannia confertissima.  R e l e v e - l o c a t i o n s : 116,117,122,123 169,170,173,174,175,176,177  oJS'oi?  240,241,242 311,312,313,314,315 334,335,336,337,338  B  Cypress Bowl Mount Seymour Meadows G a r i b a l d i Lake W h i s t l e r Mountain Diamond Head 1  a  c  k  T  u  s  k  79  lation. On Mount W h i s l e r (2286 m) Andreaea r o t h i i becomes an  important  a s s o c i a t e o f t h e a s s o c i a t i o n , and forms a t h i r d v a r i a n t ( 4 ) . Whether t h e s e l o c a l companions can be d e s i g n a t e d as  differen-  t i a t i n g s p e c i e s t o d i s t i n g u i s h s u b a s s o c i a t i o n s needs f u r t h e r  investi-  g a t i o n . E s p e c i a l l y , because a l l t a x a have a d i s t r i b u t i o n which  extends  beyond the l i m i t s o f the p r e s e n t study a r e a , t h i s cannot be done w i t h out more r e s e a r c h .  R a c o m i t r i e t u m s u d e t i c i Herzog 1943  (Table V I I I ) .  Herzog (1943) r e p o r t s and d e s c r i b e s a Racomitrium  sudeticum-  a s s o c i a t i o n from the B l a c k F o r e s t r e g i o n i n s o u t h e r n West Germany. In the study a r e a t h e R a c o m i t r i e t u m  s u d e t i c i o c c u r s on exposed r o c k s a t  s u b a l p i n e and a l p i n e e l e v a t i o n s o f Cypress Bowl P r o v i n c i a l Mount Seymour and areas i n G a r i b a l d i P r o v i n c i a l  Park,  Park. In c o n t r a s t t o  the Gymnomitrietum c o n c i n n a t i , which o c c u r s on s t e e p ( o r even o v e r hanging) rock s l o p e s , t h e R a c o m i t r i e t u m s u d e t i c i p r e d o m i n a n t l y  grows  on g e n t l y s l o p i n g r o c k s u r f a c e s . G e n e r a l l y , more humus and m i n e r a l  soil  has been formed underneath  the  t h e l u s h b r y o p h y t e assemblages i n which  a s s o c i a t i o n c h a r a c t e r s p e c i e s , Racomitrium  s u d e t i c u m , i s t h e dominant  taxon. Most f r e q u e n t accompanying s p e c i e s a r e Gymnomitrion Andreaea n i v a l i s , K i a e r i a s t a r k e i and D i c r a n o w e i s i a c r i s p u l a .  concinnatum, Although  they were found o n l y a few t i m e s , t h r e e t a x a out o f t h e genus Grimmia  Table VIII. Racomitrietum sudetici Herzog 1943 Number of releves Releve-number Elevation(m) Direction of exposure Inclination( ) Total cover(!fc)  1 118 610 W 58 60  2 124 610 S 26 .100  3 309 1219 N 112 75  Racomitrium sudeticum Gymnomitrion concinnatum Andreaea rupestris Andreaea nival 1s Kiaeria starkei Dicranoweisia crispula Diplophyllum taxifollum Racomitrium heterostichum var. heterostichum Pseudoleskea radicosa Kiaeria b l y t t i i Grimmia alpestris var. holzingerl Lophozia sudetica Marsupella sparsifolia Amphidium californicum Grlnrnia donniana Pseudoleskea atricha Pohlia cruda Grimmia alpestris var. alpestris Racomitrium canescens Lophozia ventricosa var. ventricosa Lophozia i n d s a Marsupel la emarginata. Grimmia montana Pohlia nutans  3.4  4.4  3.4 2.4 3.3  2.4  4.4 2.4 1.2  n  u  Found only once:  1.2  1.4  4 5 330 310 1219 1219 E-SE E-NE 60 108 85 85 4.5 2.4 2.3 1.4  1.4 1.2 1.2  6 125 610 SE 78 100 4.4 + .2 3.4  1.2  7 266 1492 W-NW 62 85 4.5  9 8 253 302 1492 1219 W NW 14 24 100 90 4.4  1.2  2.4 2.3  1.4  1.4  2.4  11 12 304 120 1219 610 W S-SW 52 36 80 90  1.4 1.2 2.4  3.4 1.2 1.2  1.2 1.4  1.2 1.4  2.4 3.4 1.2  10 303 1219 N 48 80  1.2 + .4 2.4 2.2  2.4 2.2 1.2 + .2 1.2 1.4 1.4  1.4 2.2 1.2  2.2  1.2  1.2 3.2  16 15 14 256 171 172 1492 1433 1433 NW W-NW NW 58 42 84 100 40 90  2.4  1.4 1.2  3.4 1.2  1.4 2.2  3.4 2.4  3.4 3.4 2.4 2.2  1.4 4.4  1.4  4.4  3.4 1.4  3.4  3.4  1.4  2.4 1.4  1.4 3.4  1.1  1.2  1.4 1.4 +.2  1.4 1.2  3.4 1.2 1.2 3.4 3.2  13 121 610 S 70 100  1.4  3.4  1.4 1-2  1.4 1.2  1.2  Andreaea b l y t t i i , Kiaeria falcata, Marsupella brevissima, M. stablerl, Tritomaria quinquedentata, Hypnum subimponens, "Racomitrium brevipes". Scapania' americana, Barbilophozia floerkei, Polytrichum piliferum, Grimmia torquata, Ceratodon purpureus, Bartramia ithyphylTa, Nardia japonica, Schistidium strictum, Pohlia ludwigii, Cephalozia bicuspidata, Pleuroclada albescens, Hygrohypnumochraceum, Anoectangium aestivum, Racomitrium aciculare, Lophozia wenzelii, Brachythecium albicans, Racomitrium varium, Pseudoleskea stenophylTa", Polytrichum sexangulare, Pohlia cardotii, Barbula vinealis, Cephaloziella byssacea var. byssacea ,~C~. byssacea var. asperifolia. Pseudoleskea patens, Aneura pinguis.  Releve-locatlons: 118,120,121,124,125  Cypress Bowl  302,303,304,309,310,330  Whistler Mountain  253,256,266  Panorama Ridge  171,172  Mount Seymour  81  were c o n f i n e d t o t h i s a s s o c i a t i o n : Grimmia a l p e s t r i s v a r . a l p e s t r i s and v a r . h o l z i n g e r i and Grimmia donniana. Herzog ( 1 9 4 3 ) , i n h i s o r i g i n a l d e s c r i p t i o n o f t h i s syntaxon from s o u t h e r n West Germany, mentions Grimmia commutata and G_. e l o n g a t a as t y p i c a l  f o r h i s Racomitrium  sudeticum - a s s o c i a t i o n . These s p e c i e s , t o g e t h e r w i t h a s s o c i a t e s such as Pseudoleskea r a d i c o s a , P_. a t r i c h a , K i a e r i a b l y t t i i  and Marsupel!a  sparsifolia  r e f l e c t t h e s u b a l p i n e - a l p i n e c h a r a c t e r o f the R a c o m i t r i e t u m s u d e t i c i .  8.1.3.2. Order POLYTRICHETALIA PILIFERI v. Hubschmann  Alliance Polytrichion p i l i f e r i  Racomitrio  Norr  - Polytrichetum p i l i f e r i  The R a c o m i t r i o  1967  1969.  Herzog 1943  (Table I X ) .  - P o l y t r i c h e t u m p i l i f e r i was  first  reported  by  Herzog (1943) from the B l a c k F o r e s t r e g i o n i n s o u t h e r n West Germany. The a u t h o r d e s c r i b e d the a s s o c i a t i o n from sandy s o i l ,  as d i d a.o.  Koppe ( 1 9 5 5 ) , Norr ( 1 9 6 9 ) , Neumayr(1971) and Hubschmann (1967, 1975). The l a s t a u t h o r a l s o r e p o r t e d the syntaxon from Vancouver a g a i n from sandy s o i l trichetum p i l i f e r i for  Island,  (Hubschmann 1978). Separate a s s o c i a t i o n s ( P o l y -  and R a c o m i t r i e t u m c a n e s c e n t i s ) have been d e s c r i b e d  both c h a r a c t e r s p e c i e s , P o l y t r i c h u m p i l i f e r u m and  Racomitrium  c a n e s c e n s , r e s p e c t i v e l y ( K r u s e n s t j e r n a 1945, F r e g r i 1933, Smarda  1947,  82  Giacomini 1951, Dunk 1971). In t h e p r e s e n t s t u d y t h i s p r e d o m i n a n t l y t e r r e s t r i a l  bryophyte  a s s o c i a t i o n i s found on h o r i z o n t a l tops and s l i g h t l y s l o p i n g  exposed  rock s u r f a c e s a t low e l e v a t i o n s o n l y . In both m i c r o - s i t e t y p e s t h e amount of  accumulated humus and/or m i n e r a l s o i l  c o n s t a n t l y ranged from 1 - 2  cm. The c h a r a c t e r s p e c i e s , P o l y t r i c h u m p i l i f e r u m and Racomitrium c a n e s c e n s , a r e commonly accompanied  by o t h e r ' t e r r e s t r i a l '  species  such as Ceratodon purpureus and P o l y t r i c h u m j u n i p e r i num. Other f r e q u e n t a s s o c i a t e s , which o f t e n o c c u r on r o c k s w i t h a thinner soil  l a y e r a r e s p e c i e s such as C e p h a l o z i e l l a byssacea v a r .  b y s s a c e a , Hedwigia c i l i a t a and S c h i s t i d i u m apocarpum v a r . s t r i c t u m . In t h e d i f f e r e n t i a t e d t a b l e o f t h e f i r s t major group, a c o l l e c t i o n o f f o u r r e l e v e s (305 t o 308) i s s p l i t o f f from the r e l e v e s  consti-  t u t i n g t h e p r e s e n t a s s o c i a t i o n . These f o u r r e l e v e s t a k e n a t Mount W h i s t l e r , a r e d i s c u s s e d f u r t h e r i n t h e d i s c u s s i o n o f major group number 3 (8.3.3.1 A s s . D i c h o d o n t i e t u m p e l l u c i d i ; s u b a s s . B l i n d i o - S c a p a n i e tosum p a l u d o s a e ) .  Scapanietum americanae a s s . nov. ( T a b l e X ) .  The d i s t r i b u t i o n o f t h i s a s s o c i a t i o n shows b a s i c a l l y t h e same p a t t e r n as t h a t o f t h e R a c o m i t r i o - P o l y t r i c h e t u m p i l i f e r i . With r e s p e c t to  i t s h a b i t a t i t a l s o o c c u r s on h o r i z o n t a l o r s l i g h t l y s l o p i n g rock  s u r f a c e s o v e r l a i n by s o i l  and/or humus. The c h a r a c t e r s p e c i e s o f t h e  a s s o c i a t i o n i s Scapania americana. a w e s t e r n North American endemic. I t  Table IX. Racomitrio - Polytrichetum p i l i f e r i Herzoa 1943 Number of releves Releve-number Elevation(m) Direction of exposure Inclination( ) Total cover(%) Q  Racomitrium canescens Polytrichum piliferum Ceratodon purpureus Hypnum -subimponens Schistidium strictum Cephaloziella byssacea var. byssacea Racomitrium heterostichum var. heterostichum Pohlia nutans Hedwigia c i l i a t a Grimmia torquata Lophozia ventricosa var. ventricosa Polytrichum juniperinum Scapania americana Homalothecium nevadense Racomitrium sudeticum Aulacomnium androgynum Barbilophozia floerkei Amphidium lapponicum Tortula princeps Dryptodon patens Homalothecium n u t a l l i l Claopodium bolanderi Pseudoleskea radicosa "Racomitrium brevipes" Porella navicularis Douinia ovata Anacolia menziesii Reboulla hemisphaerica Cephaloziella byssacea var. asperifolia Lophocolea cuspidata Radula bolanderi Encalypta c i l i a t a Found only once:  1 12 36 E-SE 54 60  2 22 - 18 S-SW 38 95  3 27 18 SE 52 80  3.4 3.3  5.4 2.2 2.2  3.4 3.3 1.2  5 4 68 31 18 61 U-NU S-SW 38 46 85 90 4.4 1.2 2.3  4.4 2.2 2.2 3.4  2.4  2.4  7 157 266 SW 56 90  8 158 266 S 54 90  9 286 610 E-NE 48' 60  10 287 610 E 36 60  11 294 610 E 42 80  12 296 610 E-NE 68 100  13 295 610 E-NE 30 100  3.4  1.2 1.2 1.2 3.4 3.4  1.2  1.2 1.2  2.4 + .2  1.2  3.4 1.2 1.2 2.4 1.2  1.4  1.2  1.2 2.2 1.2 1.4 3.4  1.4  1.4  2.3  1.4  1.2 + .2 3.4  1.2 + .2 4.4  1.2 1.2 + .2 + .2  1.4 1.1  + .4  1.4  +.4  6 32 61 N-NE 54 100  + .4  1.4  2.3 1.4  2.4  1.4 1.2 1.2  1.2 + .4 + .1  + .2 + .1 + .4 2.2 1.4  3.4 2.3 1.4  1.2 1.4  3.4  1.4  1.4  1.4  1.2  +.2  1.4  2.3 2.3  1.2  1.2  1.2 1.1  1.2 1.2  1.2 1.2 1.2  3.4 1.2 1.2  3.3  3.3  1.2 1.2  1.2  1.2 1.4  1.4 2.3  3.4 2.4  2.2  1.2 2.4  14 357 354 E 52 85 3.4  1.2  15 358 354 E-SE 44 100  16 359 354 E 62 90  4.4 1 .2 1.2 2.4 1.2  3.4 1.2 1.2 2.4  1.4  5.4  1.2 1.2 1.4  2.4 2.3  1.4 2.2 1.4  1.1 1.4  1.4  1.2 2.2  1.2  1.4  1.4 2.2  2.4 2.4  1.4  1 .2 ^_  1-4  2.4  1.2  3.2  3.4 2.3  1-4  2.4  2.4 1.4  1.4  Gymnomitrion obtusum, Bartramia pomiformis, Racomitrium varium, Dicranoweisia cirrata, Schistostega pennata, Distichium capillaceum, Atrichum undulatum, Heterocladium dimorphum, Pseudobraunia californica, Diplophyllum taxifolium, Racomitrium heterostichum var. affine, Cynodontium jenneri, Preisia quadrata.  Table IX ( c o n t i n u e d ) .  Rel e v e - l o c a t i o n s : 12  Horseshoe Bay  22,27  L i g h t h o u s e Park  68  B r i t t a n i a Beach H i s t o r i c  31,32  Bridal  157,158  Chilliwack  286,287,294,295,296  Nairn  357,358,359  Agassiz  Veil  Falls River Valley  Falls Mountain  Site  85  i s o f t e n a s s o c i a t e d w i t h P o l y t r i c h u m alpinum nutans and B a r t r a m i a pomiformis. Lophozia  Other f r e q u e n t companions a r e  v e n t r i c o s a v a r . v e n t r i c o s a , Scapania  scoparium.  var. macounii, P o h l i a  s c a n d i c a and Dicranum  The s y n t a x o n o m i c a l l y c l o s e r e l a t i o n s h i p o f t h e Scapanietum  americanae t o t h e p r e v i o u s l y d e s c r i b e d syntaxon  i s expressed  by s p e c i e s  such as Racomitrium c a n e s c e n s , Hypnum subimponens and Ceratodon purpur e u s . However, i n t h i s a s s o c i a t i o n , P o l y t r i c h u m p i l i f e r u m , P_. j u n i perinum and s p e c i e s l i k e C e p h a l o z i e l l a byssacea v a r . byssacea and Hedwigia c i l i a t a o c c u r as r a r e a s s o c i a t e s . In c o n t r a s t t o t h e R a c o m i t r i o - P o l y t r i c h e t u m p i l i f e r i , w h i c h , w i t h no e x c e p t i o n , occurs  i n exposed h a b i t a t s , t h e c u r r e n t a s s o c i a t i o n  i s sometimes found on rock s u r f a c e s shaded o r somewhat p r o t e c t e d by surrounding  low v e g e t a t i o n o r o t h e r b o u l d e r s . In these  situations,  the c h a r a c t e r s p e c i e s as w e l l as the o t h e r w i s e f r e q u e n t companions, can be a s s o c i a t e d w i t h s p e c i e s such as Claopodium b o l a n d e r i , P l a g i o t h e c i u m p i l i f e r u m and P t e r i g y n a n d r u m f i l i f o r m e . These t a x a p l a y a major r o l e i n b r y o p h y t e assemblages d e s c r i b e d from shaded and p r o t e c t e d r o c k s .  1. A ' f a c i e s ' i s a more o r l e s s a c c i d e n t a l combination an a s s o c i a t i o n o r s u b a s s o c i a t i o n .  of species w i t h i n  2. Hebrard (1978) r e p o r t s on an a s s o c i a t i o n w i t h Andreaea r u p e s t r i s and Gymnomitrion concinnatum from a l p i n e areas i n south west France. The newl y proposed a s s o c i a t i o n Andreaeetum r u p e s t r i s shows a f f i n i t i e s , i n s p e c i e s c o m p o s i t i o n , t o t h e p r e s e n t data as w e l l as t o p h y t o s o c i o l o g i c a l t a b l e s i n P h i l i p p i (1956) and H e r t e l (1974). The low constancy o f Andreaea r u p e s t r i s and t h e presence o f s p e c i e s l i k e Racomi t r i um s u d e t i c u m , Gymnomitrion concinnatum, D i c r a n o w e i s i a c r i s p u l a , B a r t -  Table X. Scapanietum americanae ass, nov. Number of releves Rel eve-number Elevation(m) Direction of exposure Incl ination( ) Total cover(X)  1 16 61 N-NW 32 100  2 38 61 E 48 100  3 37 61 S-SW 38 90  Scapania amerlcana Polytrichum alpinum var. macounil Pohlia nutans Lophozia ventricosa var. ventricosa Marsupella emarginata Ceratodon purpureus Racomitrium canescens Dlcranum scoparium Scapania scandica Hypnum subimponens Lophozia incisa "Racomitrium brevipes" Barbilophozia hatcherl Pleurozium schreberi Polytrichum piliferum Dryptodon patens Polytrichum juniperlnum Racomitrium heterostichum var. afflne Gymnomitrion obtusum Pseudoleskea radicosa Cephaloziella byssacea var. byssacea Racomitrium lanuginosum Oligotrichum aligerum Ditrichum heteromallum Lophozia excisa Hedwigia c i l i a t a Plagiothecium piliferum Pseudobraunia californica Pterigynandrum filiforme Lophocolea cuspidata Claopodium bolanderi  1.4  2.4  5.4  3.2 1.2  3.2 1.2  1.2 1.2  Q  Found only once:  2.3 1.2  1.2 1.2 + .2 1.4 2.4 2.4  2.4 1.2 2.3 1.4  4 44 61 NE 48 100  5 43 61 NE 26 100  6 291 610 E 50 90  2.4  1.4  3.4  2.2 + .1  2.2  3.4  1.4  1.2 1.2  1.4  2.3  7 293 610 E 60 80 3.4  1.4  1.4  8 292 610 E 66 90  9 316 266 NW 44 100  10 317 266 N 56 100  11 318 266 NW 32 100  12 13 319 33 266 61 N-NW E-NE 64 60 100 100  14 15 35 154 61 266 N-NW N 56 50 90 90  2.4  1.4  2.4  3.4  3.4  1.4  3.4  2.4  2.4  1.2  2.2  + .1 1.2  + .1  4.3  + .1 1.2  1.2 2.2  2.2 1.2  1.4 3.4  1.4 2.4  3.4 1.4  1.4 1.4 + .2  1.4  2.4  4.3 1.2 1.2 2.4  3.3 1.2  1.4  1.4-  + .4 4.4 + .4  2.4 3.2  1.4 2.2  1.4 1.2  1.4  + .4 + .2  1.4  + .4 + .2  + .4 1.4  2.4  2.4  2.4  1.4  4.3 3.3  1.4  3.2  2.2  4.3 2.2 1.2  + .1 1.1  1.2 1.4  1.2  2.2 1.4 1.4  + .4  1.2 1.4  2.4  2.2  1.1  1.4 1.4  1.4 1.2  +.4  + .4  1.4 2.4 + .2 2.4  1.4  2.2 1.2  1.4  Murrin Provincial Park Bridal Veil Falls Nairn Falls Chilliwack River Valley SI esse Creek  2.2 2.2  1.4 + .4  Lophozia sudetica, Pohlia cruda, Cynodontium jenneri, Anastrophyl 1um minutum, Encalypta procera, Anoectangium aestivum, Aulacomnium androqynum, Rhytidiadelphus triquetrus, Preissia quadrata, Bazzania denudata, Plagiopus oederi.  Releve-locations: 16 33,35,37,38,43,44 291 ,292,293 154,155 316,317,318,319  2.4 1.4  1.1  +.2 1.4 1.4  3.4  16 155 266 NE 40 90  1.4  CO  87  rami a i t h y p h y l l a and Pseudoleskea i n c u r v a t a i n d i c a t e t h i s newly desc r i b e d a s s o c i a t i o n ' s n a t u r e as fragments o f high e l e v a t i o n s s y n t a x a as the Gymnomitrietum c o n c i n n a t i Herzog 1943 and R a c o m i t r i e t u m s u d e t i c i Herzog 1943. 3. A ' v a r i a n t ' i s a u n i t w i t h i n an a s s o c i a t i o n w i t h one p a r t i c u l a r taxon b e i n g dominant, which i s absent o r o n l y o c c a s i o n a l l y (and w i t h low coverage) o c c u r r i n g i n the a s s o c i a t i o n - " t y p i c u m " .  88  8.2  BRYOPHYTE ASSOCIATIONS FROM SHADED, HUMID ROCKS.  T a b l e X I . Bryophyte a s s o c i a t i o n s from shaded, humid r o c k s .  Order LEPIDOZIETALIA REPTANTIS ( P h i l i p p i  1956)  A l l i a n c e DIPLOPHYLLION ALBICANTIS ( P h i l i p p i  C l u s t e r 1.  Hertel  1956)  Association Diplophylletum  1974  Hertel  1974  a l b i c a n t i s Schade  1923  A l l i a n c e SCAPANION AMERICANAE a l l . nov.  C l u s t e r 2.  A s s o c i a t i o n M a r s u p e l l o - Scapanietum americanae a s s .  C l u s t e r 3.  A s s o c i a t i o n Claopodietum  nov.  bolanderi  A s s o c i a t i o n - typicum Subassociation  I s o p t e r y g i e t o s u m e l e g a n t i s subass.  Subassociation  Porelletosum  roellii  subass.  nov.  nov.  90  8.2.  Bryophyte a s s o c i a t i o n from shaded, humid r o c k s ( T a b l e X I ) .  8.2.1.  Synsystematics.  Philippi  (1956) proposed the o r d e r D i p l o p h y l l e t a l i a  albicantis  to c o n t a i n b r y o p h y t e a s s o c i a t i o n s from humid and shaded s i l i c e o u s  rocks.  W i t h i n t h i s o r d e r he a l s o proposed t h e a l l i a n c e D i p l o p h y l l i o n a l b i c a n t i s , i n c l u d i n g a.o. s t j e r n a 1945.  the a s s o c i a t i o n D i p l o p h y l l e t u m a l b i c a n t i s  T h i s l a t t e r syntaxon  had been r e p o r t e d and  Krusen-  described  e a r l i e r by Schade ( 1 9 2 3 ) , hence, as s t a t e d by H e r t e l (1974). i t s h o u l d be termed D i p l o p h y l l e t u m a l b i c a n t i s Schade 1923. D i p l o p h y l l e t a l i a a l b i c a n t i s prov. P h i l i p p i 1956 Diplophylletalia albicantis Philippi Although Hertel  1963,  However, the o r d e r c o n f l i c t s w i t h the  d e s c r i b e d from m i n e r a l  P h i l i p p i s t r e s s e s t h e d i s s i m i l a r i t i e s between these two  order soil.  syntaxa,  (1974) r i g h t l y s t a t e d t h a t a s p e c i e s can not be a c h a r a c t e r s p e c i e s  f o r two o r d e r s . M i n e r a l s o i l species combination  communities w i t h D i p l o p h y l l u m a l b i c a n s show a  s i m i l a r t o samples from rocky h a b i t a t s . Hence, they  s h o u l d , on f l o r i s t i c  criteria,  be grouped i n one c a t e g o r y . T h i s  syntaxon,  the a l l i a n c e D i p l o p h y l l i o n a l b i c a n t i s thus c o n t a i n s a s s o c i a t i o n s from humid, shaded r o c k s as w e l l as m i n e r a l s o i l . Together w i t h the T e t r a p h i d i o n p e l l u c i d a e K r u s e n s t j e r n a 1945  ( a l s o c o n t a i n i n g a s s o c i a t i o n s from  humid and shaded rocks and s o i l ) the D i p l o p h y l l i o n a l b i c a n t i s 1956) em.,  em.  (Philippi  i s p l a c e d i n the o r d e r L e p i d o z i e t a l i a r e p t a n t i s ( P h i l i p p i  following Hertel  1963)  (1974).  In the p r e s e n t s t u d y , a new  a l l i a n c e i s p r o p o s e d , the Scapanion  91  americanae, c o n t a i n i n g two newly d e s c r i b e d a s s o c i a t i o n s M a r s u p e l l o Scapanietum americanae and Claopodietum  b o l a n d e r i . Based on t h e f o l l o -  wing c r i t e r i a t h e s e s y n t a x a a r e c l a s s i f i e d i n t h e o r d e r  Lepidozietalia  r e p t a n t i s ( P h i l i p p i 1963) H e r t e l 1974: (1) a l l t h r e e a s s o c i a t i o n s share a c o n s i d e r a b l e number o f s p e c i e s ; (2) they o c c u r i n t h e same t y p e s o f h a b i t a t s and (3) many s p e c i e s ( c h a r a c t e r s p e c i e s as w e l l as companions) w i t h i n t h e r e l e v e s a r e known from m i n e r a l s o i l  and humus-covered-rocks. As Neumayr  (1971) and H e r t e l (1974) s t a t e , c l o s e r e l a t i o n s h i p s between communities from m i n e r a l s o i l  and a t h i n humus l a y e r over r o c k s , do not j u s t i f y a  separation at higher synsystematic  8.2.2. P h y t o g e o g r a p h i c a l  levels.  setting.  In both a l l i a n c e s , D i p l o p h y l l i o n a l b i c a n t i s and Scapanion canae, t h e presence  o f endemic and c i r c u m b o r e a l  ameri-  t a x a i s predominant.  Although o n l y r e p r e s e n t e d by l e s s f r e q u e n t companions, two o t h e r phytogeographical (Blepharostoma  elements a r e shared by these s y n t a x a : b i p o l a r d i s j u n c t s  t r i c h o p h y l l u m , Polytrichum alpinum var. macounii, B a r t -  ramia pomiformis)  and c i r c u m b o r e a l - m i s s i n g - i n - J a p a n  var. v e n t r i c o s a , Isopterygium  (Lophozia v e n t r i c o s a  elegans).  In t h e D i p l o p h y l l e t u m a l b i c a n t i s t h e c i r c u m b o r e a l  character  s p e c i e s , D i p l o p h y l l u m a l b i c a n s , i s f r e q u e n t l y accompanied by a w e s t e r n North American endemic, Scapania americana (which r e p l a c e s  Scapania  nemorosa front European samples o f t h i s a s s o c i a t i o n . Other endemic t a x a  92  are Rhizomnium g l a b r e s c e n s  and t h e more i n f r e q u e n t D i p l o p h y l l u m  f o l i u m . The most i m p o r t a n t c i r c u m b o r e a l  companions o f the D i p l o p h y l l e t u m  a l b i c a n t i s i n t h e p r e s e n t study a r e M a r s u p e l l a e m a r g i n a t a , b i c u s p i d a t a , Racomitrium h e t r o s t i c h u m v a r . h e t e r o s t i c h u m alpinum  taxi-  Cephalozia  and  Polytrichum  ( i n f a c t the v a r . m a c o u n i i , an endemic t a x o n ) . The  a l l i a n c e , Scapanion a m e r i c a n a e , r e f l e c t s a n o t a b l e  increase  i n c o n t r i b u t i o n by w e s t e r n North American endemics. For i n s t a n c e , Scapania  americana and Claopodium b o l a n d e r i a r e t h e r e s p e c t i v e c h a r a c t e r  s p e c i e s f o r the two a s s o c i a t i o n s d i s t i n g u i s h e d i n t h i s a l l i a n c e . t a n t companions a r e I s o t h e c i u m  Impor-  s t o l o n i f e r u m , Rhizomnium g l a b r e s c e n s ,  P o r e l l a r o e ! 1 i i and F r u l l a n i a t a m a r i s c i s s p . n i s q u a l l e n s i s . However, there s t i l l  i s a l a r g e c o n t r i b u t i o n by c i r c u m b o r e a l  Marsupella emarginata,  P l a g i o c h i l a a s p l e n i o i d e s , Diplophyllum albicans  and t o a l e s s e r e x t e n t by Racomitrium h e t e r o s t i c h u m and P o l y t r i c h u m alpinum  species i n c l u d i n g  var.  var.  heterostichum  macounii.  Among c h a r a c t e r i s t i c a s s o c i a t e s o f both a s s o c i a t i o n s o f the Scapanion americanae a r e r e p r e s e n t a t i v e s from t h r e e a d d i t i o n a l g r a p h i c a l elements:  phytogeo-  w e s t e r n North America - w e s t e r n Europe - d i s j u n c t s  (Scapania s c a n d i c a , P l a g i o t h e c i u m p i l i f e r u m ) , a w e s t e r n North America Japan - d i s j u n c t (Hypnum subimponens) and B i p o l a r d i s j u n c t s ( P o l y t r i c h u m alpinum, Bartramia pomiformis, Plagiothecium  denticulatum  S t o k e s i e l l a praelonga  var. praelonga  and  93  8.2.3. Order LEPIDOZIETALIA REPTANTIS ( P h i l i p p i  Alliance Diplophyllion albicantis (Philippi  D i p l o p h y l l e t u m a l b i c a n t i s Schade 1923  Bryophyte communities on m i n e r a l s o i l  1956) H e r t e l  1956) H e r t e l  1974  1974.  (Table X I I ) .  containing Diplophyllum a l b i c a n s occur  as w e l l as on r o c k . T h i s o c e a n i c a s s o c i a t i o n (Gams  i s w i d e l y d i s t r i b u t e d over c e n t r a l and n o r t h e r n Europe.  Frequent r e p o r t s  from shaded e p i l i t h i c h a b i t a t s a r e g i v e n by Schade (1923', 1934), (1943), P h i l i p p i  1952)  Herzog  (1956, 1965), Norr ( 1 9 6 9 ) , Neumayr ( 1 9 7 1 ) , Dunk ( 1 9 7 1 ) ,  M a r s t a l l e r (1973) and H e r t e l  (1974).  In the s t u d y area t h e D i p l o p h y l l e t u m a l b i c a n t i s occurs" widespread a t p r e d o m i n a n t l y low but sometimes a t s u b a l p i n e e l e v a t i o n s . I t s h a b i t a t i s c o n f i n e d t o shaded r o c k s , a d j a c e n t t o o r i n the d i r e c t neighbourhood  o f w a t e r , on humid r o c k w a l l s i n canyons or a t the base  o f b i g b o u l d e r s i n s i d e f o r e s t s . The a s s o c i a t i o n forms l u s h assemblages c h a r a c t e r i z e d by m o s t l y low-growing  b r y o p h y t e s i n f r e q u e n t l y mixed w i t h  l u x u r i o u s growth o f some mosses and  liverworts.  The c h a r a c t e r s p e c i e s i s D i p l o p h y l l u m a l b i c a n s , which i s most f r e q u e n t l y accompanied by Scapania a m e r i c a n a , a w e s t e r n North  American  endemic which r e p l a c e s Scapania nemorosa from t h e D i p l o p h y l l e t u m a l b i c a n t i s i n Europe ^. Another s p e c i e s used as c h a r a c t e r s p e c i e s f o r the l a t t e r syntaxon i s H e t e r o c l a d i u m heteropterum  (Philippi  1956, Norr  1969).  P r e s e n t data show H e t e r o c l a d i u m macounii w i t h o n l y o c c a s i o n a l o c c u r r e n c e s  Table  XII.  Diplophylletum albicantis Schade 1923 Number of releves Releve-number Elevation(m) Direction of exposure Inclination ) Total cover(%)  1 2 1 2 1150 1150 NE NE 85 40 90 85  3 4 30 SW 64 80  4 75 18 NE 76 90  5 80 61 E 52 90  6 180 1433 E-SE 86 100  7 59 61 N 70 100  8 95 30 SW 56 100  9 100 610 SW 64 100  Diplophyllum albicans Scapania americana Racomitrium heterostichum var. heterostichum Bazzania denudata Marsupella emarginata Rhizomnlum glabrescens Isopteryglum elegans Chandonanthus filiforme Polytrichum alplnum var. macounii Cephalozia bicuspidata Bazzania tricrenata Mylia taylori Dicranella heteromalla Lepidozia reptans Calypogeia muelleriana Bartramia pomiformls Plagiochila asplenioides Andreaea rupestris Diplophyllum taxlfolium Douinia ovata Nardia scalaris Herbertus aduncus Gyrothyra underwoodiana Pogonatum contortum Marsupella sparslfolia Blepharostoma trichophyllum Lophozia ventricosa var. ventricosa Heterocladium macounii PI agiothecium undulatum  5.5  • 4.5  2.4 1.4  4.4  4.5  3.4  5.4 1.4  2.4 4.4  3.4 2.4  3.3  3.3  2.2  + .2  + .1 3.4 + .4 + .4 + .1 + .1 1.4  Q  + .4 + .1  2.4  1.4 1.2 1.4 1.1  + .2 + .4  2.4 2.2  1.4 +.1 +.4  + .4 + .1 + .4  11 50 61 N 74 95  12 13 54 60 30 61 N-NW N-NW 56 76 100 70  1.4 55 61 N 78 80  15 61 61 N-NE 86 60  2.4 3.4  3.4 2.4  + .4 2.4 4.4  3.4  IA  2.2 4.4  + .1  4,5  3,1  3.3 1-4  1.4  3.4  4.4  4.5  2.2 +.4  1.4  2.2  + .1  + .4  10 45 61 NW 78 100  2.2 1.2 1.4  + .1  2.2  +.4  +.4  2.4  1.1  + .4  Found only once: Claopodium bolanderi, Frullania tamarisci ssp. nisquallensis, Pohlia cruda, Hypnum circinale, Anastrophyllum minutum, Plagiothecium piliferum, Cephaloziella byssacea var. asperifolia, Lophozia incisa, Racomitrium canescens, Scapania undulata, Racomitrium brevipes, Rhytidiadelphus loreus, Dryptodon patens, Gymnomitrion concinnatum, G. obtusum, Calypogeia trichomams. Releve-locations:!,2,180 4,60,95 75 61,80 45,50,54,55,59 100  Mount Seymour Lynn Canyon Lighthouse Park Bridal Veil Falls Shannon Falls Cheakamus River Valley  95  and i s not t r e a t e d as a d i a g n o s t i c s p e c i e s f o r t h e a s s o c i a t i o n . A d d i t i o n a l a s s o c i a t e s i n d i c a t i n g a f f i n i t i e s t o the European r e p r e s e n t a t i o n o f the D i p l o p h y l l e t u m a l b i c a n t i s from e p i l i t h i c h a b i t a t s a r e : Marsupella emarginata, B a r t r a m i a pomiformis  Isopterygium elegans, Cephalozia b i c u s p i d a t a ,  and L o p h o z i a v e n t r i c o s a v a r . v e n t r i c o s a .  However, D i c r a n e l l a h e t e r o m a l l a , a f r e q u e n t and  important  c h a r a c t e r s p e c i e s f o r t h e a l l i a n c e and o r d e r i n Europe o c c u r s i n the study a r e a o n l y o c c a s i o n a l l y . W i t h i n the D i p l o p h y l l e t u m a l b i c a n t i s the p r e s e n t data  distin-  guish a B a z z a n i a denudata - v a r i a n t , w h i c h , i n a d d i t i o n , can be  charac-  t e r i z e d by s p e c i e s such as Chandonanthus f i l i f o r m i s and t o a l e s s e r e x t e n t by H e r b e r t u s aduncus. I t s d i s t r i b u t i o n i s c o n f i n e d t o very shaded and p r o t e c t e d s t e e p b o u l d e r s w i t h i n f o r e s t v e g e t a t i o n around Shannon F a l l s and B r i d a l V e i l  Falls.  Releves 59, 95 and 100 i n d i c a t e a B a z z a n i a t r i c r e n a t a - C a l y p o g e i a m u e l l e r i a n a - f a c i e s through t h e a c c i d e n t a l s p e c i e s c o m b i n a t i o n o f t h e s e two bryophyte  taxa.  A l l i a n c e Scapanion  americanae a l l .  nov.  M a r s u p e l l o - Scapanietum americanae a s s . nov.  (Table X I I I ) .  The a s s o c i a t i o n a l s o o c c u r s on r o c k s and b o u l d e r s , not exposed but shaded and s h e l t e r e d by overhanging  and/or s u r r o u n d i n g v e g e t a t i o n .  I t forms l u s h assemblages on g e n t l y s l o p i n g rock s u r f a c e s w i t h o n l y a  Table XIII. Marsupello - Scapanietum americanae ass. nov. Number of releves Releve-number Elevation(m) Direction of exposure Inclination( ) Total cover(%)  1 2 3 63 1150 61 NE W 74 74 80 100  Marsupella emarginata Scapania amerlcana Racomitrium heterostichum var. heterostichum Plagiochila asplenioides Isopterygium elegans Diplophyllum albicans Isothecium stoloniferum Rhizomnium glabrescens Heterocladium macounii Andreaea rupestrls Diplophyllum taxi folium Cynodontium jenneri Dicranum scoparium Hypnum subimponens Lophozia ventricosa var. ventricosa Plagiothecium denticulatum Lophocolea cuspidata Claopodium bolanderi Polytrichum alpinum var. macounii Frullania tamarisci ssp. nisquallensls Douinia ovata Lophozia incisa Cephaloziella byssacea var. asperifolla Nardia scalaris Plagiothecium laetum Pogonatum urnigerum Herbertus aduncus Plagiothecium undulatum  4.5 3.4  5.5 3.4  2.4  3.4 + .1  3.4  2.4  0  Found only once:  1.4 + .2  4 3 98 101 30 • 30 NW SE 72 52 100 100  5 13 36 SE 36 95  6 23 18 S-SE 82 95  7 29 61 S-SW 72 80  3.4 3.4  3.4 + .4  2.4 1.4  3.4 3.4  3.4 3.4  2.4 4.4  1.2 2.4 1.4 4.5  2.4 +.1  2.4  2.4 1.4  + .1  1.2 1.4 1.4  2.2  3.4  2.4  2.2 + .4  2.4 + .4 5.3  1.4  1.4  + .1  16 17 67 62 30 61 E-SF NW 78 70 90 100  18 103 30 SE 24 80  3.4 3.4  3.4 5.5  2.4 2.4  2.4 3.4  1.4 1.4  2.4 4.4 1.4 + .4 1.4 + .4 1.4 + .1  2.4 .2.4  2.4  4.4  3.4  4.4  3.4  1.4 2.2 + .4  1.2 1.4  1.2 1.4  + .4  + .4  11 74 18 NW 58 80  12 58 61 N 70 85  2.4 3.4  4.5 1.4  4.4 2.4  1.4 4.5  1.2 1.4 1.4  +.2 +.1 +.4  + .4  + .1  +.4  + .1  15 69 18 S-SI' 52 100  10 24 18 E-NE 62 90  9 273 266 W 52 90  1.4  + .2 1.2  1.4  8 272 266 W 54 90  1.4 1.2  + .4 1.2  1.4  1.4  13 140 30 S-SE 64 100  14 139 30 S-SE 78 100  2.4 3.4  1.4  1.4 1.4  1.4  1.4  + .4  2.2  2.2 1.4 + .4 + .4  2.4 1.4  + .1  2.4 3.4 1.4  1.4  1.4 + .4  1.4 + .1  1.2  +.2 1.4 + .2 2.2  + .1  + .1  1.2  + .1  Bazzania denudata, Pohlia cruda, Bartramia pomiformis, Hypnum circinale, Plagiopus oederi, Stokesiella praelonga var. praelonga, Anastrophyl1 urn minutum, Pohlia nutans, Scapania scandica, Claopodium crispifolium, Dicranella heteromalla, Dicranum fuscescens, Porella navTcularis, Marsupella sparsifolia, Polytrichum formosum, Diplophyllum obtusum, Racomitrium heterostichum var. affine, Brachythecium plumosum, Jamesoniella autumnalis, Pohlia proligera, Cephaloziella phyllacantha, Stokesiella praelonga var. stokesii, Lophozia gillmanii  CD  Table XIII  Releve-locations: 3  (continued).  Mount Seymour  58,63  Shannon F a l l s  98,101,103  Lynn Canyon  13  Horseshoe Bay  23,24  L i g h t h o u s e Park  29,62  Bridal  272,273  Chilliwack River Valley  74  Sunset Creek  67,139,140  M u r r i n P r o v i n c i a l Park  69  Porteau Camp  Veil  Falls  ^  98  t h i n l a y e r o f humus o r s o i l  accumulation.  I t s d i s t r i b u t i o n , as r e c o g n i z e d  by t h e p r e s e n t d a t a , i s c o n f i n e d t o t h e s o u t h e r n p o r t i o n o f t h e s t u d y area and ranges from L i g h t h o u s e  Park i n t o t h e C h i l l i w a c k R i v e r V a l l e y .  I t s a f f i l i a t i o n t o t h e p r e v i o u s a s s o c i a t i o n i s , a s i d e from h a b i t a t s i m i l a r i t i e s , r e f l e c t e d i n t h e presence and abundance o f major s p e c i e s w i t h i n t h e r e l e v e s . The c h a r a c t e r s p e c i e s o f t h e p r e s e n t c i a t i o n a r e Scapania  americana and, c o n s t a n t l y a s s o c i a t e d w i t h i t ,  M a r s u p e l l a emarginata.  Other f r e q u e n t companions a r e such s p e c i e s as  Racomitrium h e t e r o s t i c h u m and I s o p t e r y g i u m  asso-  var. heterostichum, P l a g i o c h i l a a s p l e n i o i d e s  e l e g a n s . Less f r e q u e n t accompanying s p e c i e s t h a t  still  i n d i c a t e t h i s a s s o c i a t i o n ' s r e l a t i o n s h i p to the Diplophylletum a l b i c a n t i s are D i p l o p h y l l u r n a l b i c a n s and Rhizomnium g l a b r e s c e n s . A l l i a n c e c h a r a c t e r s p e c i e s , which a r e c o n f i n e d t o t h e Scapanion americanae and, t h u s , d i f f e r e n t i a t e i t from t h e D i p l o p h y l l i o n a l b i c a n t i s are: Isothecium  s t o l o n i f e r u m , Hypnum subimponens, Lophocolea c u s p i d a t a  and P l a g i o t h e c i u m d e n t i c u l a t u m . S p e c i e s which were, d e s p i t e t h e i r few occurrences,  found o n l y i n t h i s a s s o c i a t i o n were Dicranum s c o p a r i u m and  Cynodontium j e n n e r i . However, they cannot be d e s i g n a t e d as c h a r a c t e r s p e c i e s f o r t h e M a r s u p e l l o - Scapanietum americanae, s i n c e they  occur  f r e q u e n t l y as major a s s o c i a t e s i n v e g e t a t i o n t y p e s from open and exposed s i l i c e o u s rocks, contained i n the order R a c o m i t r i e t a l i a h e t e r o s t i c h i P h i l i p p i 1956.  99  Claopodietum  b o l a n d e r i a s s . nov.  (Table XIV).  This widely d i s t r i b u t e d a s s o c i a t i o n occurs  characteristically  on s t e e p rock s u r f a c e s o f rock w a l l s i n s h e l t e r e d c r e e k canyons, and  on  s l i g h t l y s l o p i n g s u r f a c e s o f shaded b o u l d e r s which are p r o t e c t e d by surrounding vegetation. C h a r a c t e r s p e c i e s f o r t h e a s s o c i a t i o n i s Claopodium b o l a n d e r i , a western North American endemic. A l l i a n c e c h a r a c t e r s p e c i e s f r e q u e n t l y a s s o c i a t e d w i t h i t are P l a g i o c h i l a a s p l e n i o i d e s , S c a p a n i a americana Isopterygium  and  elegans.  Hubschmann (1978) r e p o r t s on a newly d e s c r i b e d a s s o c i a t i o n , Claopodio - Porotrichetum b i g e l o v i i  from shaded, humid b o u l d e r s  some Vancouver I s l a n d f o r e s t v e g e t a t i o n s . B e s i d e s P o r o t r i c h u m  inside  bigelovii  and Claopodium c r i s p i f o l i u m as t h e most abundant and c o n s t a n t c h a r a c t e r s p e c i e s , a l s o Claopodium b o l a n d e r i i s mentioned as an a d d i t i o n a l  charac-  t e r i z i n g s p e c i e s . In a d d i t i o n t o the s i m i l a r i t i e s i n h a b i t a t t y p e s , t h e r e are f l o r i s t i c a f f i n i t i e s between t h i s C l a o p o d i o - P o r o t r i c h e t u m b i g e l o v i i and t h e Claopodietum  b o l a n d e r i , as a newly d e s c r i b e d a s s o c i a -  t i o n i n t h i s s t u d y . These s i m i l a r i t i e s are e x p r e s s e d i n a s s o c i a t e d s p e c i e s such as I s o t h e c i u m s t o l o n i f e r u m , Rhizomnium g l a b r e s c e n s , S t o k e s i e l l a p r a e l o n g a v a r . p r a e l o n g a , Apometzgeria  pubescens and Metaneckera menzie-  sii . The p r e s e n t data d i s t i n g u i s h a Claopodietum cum  bolanderi - t y p i -  and f u r t h e r d i f f e r e n t i a t e two s u b a s s o c i a t i o n s w i t h i n t h i s  syntaxon.  The a s s o c i a t i o n - t y p i c u m (1 i n T a b l e XIV) i s c h a r a c t e r i z e d by s p e c i e s  100  such as I s o t h e c i u m  s t o l o n i f e r u m , P l a g i o t h e c i u m p i l i f e r u m , Pseudo-  l e s k e a i n c u r v a t a , Hypnum c i r c i n a l e and t o a l e s s e r degree by P l a g i o theciurn c a v i f o l i u m and Racomitrium sudeticum.  In t h i s C l a o p o d i e t u m  bolanderi - typicum the c o n t r i b u t i o n o f a l l i a n c e character species 1ike  Scapania  a m e r i c a n a , Marsupel!a emarginata and P l a g i o c h i l a a s p l e n i o -  i d e s i s minor. These b r y o p h y t e t a x a p l a y a more i m p o r t a n t r o l e i n t h e overall  f l o r i s t i c composition The  o f t h e two d i s t i n g u i s h e d s u b a s s o c i a t i o n s .  f i r s t subassociation, the Isopterygietosum  i s d e f i n e d by t h e f o l l o w i n g d i f f e r e n t i a t i n g s p e c i e s : e l e g a n s , Hypnum subimponens, Rhizomnium g l a b r e s c e n s macounii.  elegantis ( 2 ) , Isopterygium  and H e t e r o c l a d i u r n  Other s p e c i e s c o n t r i b u t i n g t o t h i s s u b - u n i t and i n d i c a t i n g  i t s s p e c i f i c m i c r o - h a b i t a t , a r e P o l y t r i c h u m alpinum  var.  macounii,  P l a g i o t h e c i u m undulatum and P_. 1 aetum. T h i s s u b a s s o c i a t i o n c h a r a c t e r i s t i c a l l y o c c u p i e s lower p o r t i o n s o f b o u l d e r s and r o c k w a l l s where  moisture  and r e l a t i v e h u m i d i t y a r e found t o be h i g h e r from s o i l i n f l u e n c e ( S j o g r e n 1964). The  second s u b a s s o c i a t i o n o f t h e C l a o p o d i e t u m b o l a n d e r i  on very s t e e p t o overhanging  rock s u r f a c e s w i t h o u t any s o i l  and s h e l t e r e d by dense v e g e t a t i o n . The major d i f f e r e n t i a t i n g for  t h i s syntaxon  Porelletosum r o e l l i i  occurs  accumulation species  (3) a r e P o r e ! l a r o e l l i i ,  t a m a r i s c i s s p . n i s q u a l l e n s i s and Amphidium c a l i f o r n i c u m . Other  Frullania bryophytes  o c c a s i o n a l i n t h i s type o f m i c r o - h a b i t a t a r e A n a c o l i a m e n z i e s i i , D o u i n i a o v a t a and P l a g i o p u s o e d e r i .  As i n d i c a t e d i n t h e c l u s t e r a n a l y s i s and t h e subsequent  "differ-  Table XIV. Claopodietum bolanderi ass, nov. Number of releves Rel eve-number Elevation(m) Direction of exposure Inclination( ) Total cover(iS) Q  Claopodium bolanderi Plagiochila asplenioldes Scapania americana Isothecium stoloniferum Marsupella emarginata Racomitrium heterostichum var. heterostichum Pohlia cruda Apometzgeria pubescens Polytrichum alpinum var. macounii Diplophyllum albicans Mnium spinulosum Bartramia ithyphylla Diplophyllum taxifolium Cephaloziella byssacea var. asperifolla Metzgeria conjugata Radula bolanderi Plagiothecium denticulatum Plagiothecium laetum Grimnia torquata Blepharostoma trlchophyllum Lophocolea heterophylla Plagiothecium piliferum Pseudoleskea incurvata Hypnum circinale Plagiothecium roeseanum Racomitrium sudeticum Isopterygium elegans Hypnum subimponens Rhizomnium glabrescens Heterocladium macounii Lophocolea cuspidata Plagiothecium undulatum Porella r o e l l i i Frullania tamarisci var. nisquallensls Amphidium californicum Anacolia menziesii. Plagiopus oederl Douinia ovata  1 5 30 SW 20 80  2 7 30 N 40 80  1.4  3.4 3.4 1.1 + .4  1.1  2.4  1.4  3 230 610 SW 66 100  4 18 18 N 84 90  2.4 2.4 3.4 2.4 1.4 1.1 1.1 + .4  6 5 19 156 18 266 N NE N 68 76 90 100  9 7 8 8 96 97 30 30 30 N NW W NW W NW 64 68 54 100 80 100  2.4 2.4 4.5  3.4 + .4 3.4  2.4 3.4  4.5 + .4 1.4  1.4  1.4  2.4  4.4  3.4  1.1  1.4 2.4 1.4  4.5 1.4 1.4 3.4  1.4 4.4 1.4  4.4  1.4 3.4 + .4 + .4 1.4 + .1  1.1  1.1  + .1  1.1  1.1 1.4  1.4 1.2  1.4  3.4  1.1  + .1 +.1 +.1  +.1  1.4 1.2  2.4  +.4  +.1  1.2 2.4  l-  2  1.4 v  1.4 +.4  + .4  2.2  +.4  1.4  2.4  2.4  1.4 +.4  +  +  5.5  1.4 3.4 1.4  4.5  4.4 ^ + .1 1.4 1.4 + .1  3.4 + .1 + .4 + .1  2.4 3.4 + .1 2.4 2.4  2.4  + .4  3.3  4.3 1.4  2.4  -4  -4  +.4 +.4  2 .4 3 .4 1,.1 + .1  2.2 2.4  + .1 + .4  2.4  2.4  1.4  3.4  -4  2.4 2.4  1.4  /  I-  1.4 1-4 1.4 1.4 1.4 +.1  2.4 3.4 2.4 1.1  21 138 30 E 94 80  -' 2.2  2  1.4  3.4 + .4 3.4  20 71 18 W NW 84 90  +  +  1.4  19 18 72 70 18 18 W Nil W NW 84 92 80 90  17 47 61 N 96 90  14 225 610 E 68 100  12 340 1524 SW 108 100  3.4 3.4 + .4 1.4 1.4 2.4 2.4 + .4  15 16 226 30 610 61 N NE S SW 68 100 100 80  13 102 30 NW 26 100  11 10 231 339 610 1524 SE SW 82 56 100 100  1  1.4 2.4 2.4 1.4  ,  2  1.4  2.4 2.4 1.4 2.4  4.5  +.4  +.4  2.1  2.4  3.4  1.4  2.4  3.4 3.3  3.4 3.3  4.5  4.5  2.4 1.2 2.4  2.3  +.1  1.4  1.4  T a b l e XIV ( c o n t i n u e d )  Found o n l y once:  B a r b i l o p h o z i a l y c o p o d i o i d e s, S c h i s t i d i u m s t r i c t u m , S t o k e s i e l l a praelonga v a r . p r a e l o n g a , L o p h o z i a v e n t r i c o s a v a r . v e n t r i c o s a , Andreaea r u p e s t r i s , B a z z a n i a denudata, C e p h a l o z i a b i c u s p i d a t a , Anastrophyl1um minutum, P o h l i a n u t a n s , S c a p a n i a s c a n d i c a , C e p h a l o z i e l l a byssacea v a r . b y s s a c e a , H e r b e r t u s aduncus, Metaneckera m e n z i e s i i , Dicranum f u s c s c e n s , P o r e l l a n a v i c u l a r i s , Pogonatum u r n i g e r u m , Drepanocladus u n c i n a t u s , Racomitrium canescens, Racomitrium h e t e r o s t i c h u m v a r . a f f i n e , R h y t i d i a d e l p h u s l o r e u s , B r a c h y t h e c i u m plumosum, Grimmia' anomala ,'. Radula complanata, Scapania mucronata, Barbilophozia f l o e r k e i , Tritomaria quinquedentata, Pseudoleskeella t e c t o r u m , Homalothecium nut-all i i , B a r b i l o p h o z i a h a t c h e r i , Racomitrium a c i c u l a r e , Encalypta c i l i a t a .  R e l e v e - l o c a t i o n s : 5,7,8,96,97,102 18,19 156 225,226,230,231 339,340 30,47 70,71,72 138  Lynn Canyon L i g h t h o u s e Park Chilliwack River Valley Cheakamus R i v e r V a l l e y Diamond Head Bridal Veil Falls Lonetree Creek M u r r i n Creek  103  e n t i a t e d " t a b l e (Appendix  V ) , t h e r e s t - g r o u p o f r e l e v e s from shaded,  humid rocks i s t o o fragmentary  and too heterogeneous t o a l l o w any  arrangement. T h i s r e s u l t s from e i t h e r l a c k o f r e l i a b l e d i a g n o s t i c s p e c i e s o r t o o few r e l e v e s w i t h i n c l u s t e r s t o d e s i g n a t e as s e p a r a t e s y n t a x a . Hence, a f f i n i t i e s w i t h a l r e a d y r e c o g n i z e d a s s o c i a t i o n s were c o n s i d e r e d t o o vague and obscure t h e s e p o s s i b l e f l o r i s t i c and phytosociological  relationships.  As s u c h , t h i s r e s t - g r o u p i s n o t f u r t h e r d i s c u s s e d here. tional  Addi-  r e s e a r c h may r e v e a l t h e i r a f f i n i t i e s t o a l r e a d y e s t a b l i s h e d  s y n t a x a , both i n B r i t i s h Columbia and i n Europe, o r t h e i r "own" n a t u r e , based on which they may become d e s i g n a t e d as s e p a r a t e  syntaxa.  1. Reports on communities, i n which both s p e c i e s a r e d e s i g n a t e d as t h e co-dominant c h a r a c t e r s p e c i e s a r e g i v e n by Dunk ( 1 9 7 1 ) , W i l c z y n s k a ( 1 9 7 4 ) , Hubschmann (1975) and Mars t a l l e r (1980).  104  8.3  BRYOPHYTE ASSOCIATIONS FROM SILICEOUS ROCKS ALONG STREAMS.  Table XV. Bryophyte  a s s o c i a t i o n s from s i l i c e o u s rocks a l o n g  streams.  Order BRACHYTHECIETALIA PLUMOSI ( P h i l i p p i 1956) prov.  A l l i a n c e RACOMITRION ACICULARIS K r u s e n s t j e r n a 1945  C l u s t e r 1 + 3.  A s s o c i a t i o n R a c o m i t r i o - S c o u l e r i e t u m a q u a t i c a e a s s . nov.  A s s o c i a t i o n Dichodontietum  p e l l u c i d i v. Hubschmann 1967  C l u s t e r 4.  S u b a s s o c i a t i o n B r a c h y t h e c i o - Conocephaletosum c o n i c a e subass. nov  C l u s t e r 7.  S u b a s s o c i a t i o n R i c c a r d i o - H y g r o b i e l l e t o s u m l a x i f o l i a e subass. nov  C l u s t e r 9.  S u b a s s o c i a t i o n B l i n d i o - Scapanietosum paludosae  C l u s t e r 5.  P h i l o n o t i s fontana - f a c i e s  C l u s t e r 8.  B l i n d i a acuta - f a c i e s  C l u s t e r 10.  A s s o c i a t i o n M a r s u p e l l o - Nardietum  s c a l a r i s a s s . nov.  subass. nov.  T a b l e XV  C l u s t e r 11.  (continued).  A s s o c i a t i o n Hypnetum d i e c k i i a s s . nov.  A s s o c i a t i o n - typicum S u b a s s o c i a t i o n Scapanio - Jungermannietosum obovatae subass. nov.  Order PLATYHYPNIDIETALIA RUSCIFORMIS P h i l i p p i 1956  A l l i a n c e PLATYHYPNIDION RUSCIFORMIS P h i l i p p i 1956  C l u s t e r 6.  A s s o c i a t i o n Hygrohypnetum s m i t h i i a s s . nov.  £  107  8.3. Bryophyte a s s o c i a t i o n s from s i l i c e o u s r o c k s a l o n g streams ( T a b l e XV).  8.3.1. S y n s y s t e m a t i c s .  Bryophyte assemblages growing on b o u l d e r s and r o c k s a l o n g s t r e a m s , e i t h e r i n open areas o r under shaded c o n d i t i o n s , have t r a d i t i o n a l l y been i n c o r p o r a t e d i n t h e " a q u a t i c b r y o p h y t e communities" s c h a f t e n " ; a.o. P h i l i p p i  ("Wassermoosgesell-  1956, 1965, Neumayr 1971 , H e r t e l  1974).  Hubschmann (1957) has proposed a s y n s y s t e m a t i c c l a s s i f i c a t i o n o f t h e a q u a t i c b r y o p h y t e communities. T h i s t r e a t m e n t i s u n s a t i s f y i n g because a s s o c i a t i o n s from d i f f e r e n t h a b i t a t s a r e grouped t o g e t h e r , and o t h e r v e g e t a t i o n u n i t s o c c u r r i n g under s i m i l a r e c o l o g i c a l  conditions  a r e c l a s s i f i e d i n t o n o n - a l l i e d s y n t a x a . The p r e s e n t s t u d y f o l l o w s t h e p r o p o s a l s by P h i l i p p i  ( 1 9 5 6 ) , who emphasizes more c o n s i s t e n t l y t h e r e l a t i o n  between h a b i t a t and t h e i r b r y o p h y t e assemblages, and c l a s s i f i e s t h e s e v e g e t a t i o n s a c c o r d i n g l y . The bryophyte v e g e t a t i o n s from stream a d j a c e n t r o c k s , because o f t h e i r unique s p e c i e s c o m p o s i t i o n and s p e c i e s combin a t i o n s , can h a r d l y be l i n k e d w i t h b r y o p h y t e assemblages from r o c k s not i n f l u e n c e d by s u r r o u n d i n g o r a d j a c e n t w a t e r b o d i e s . The y e a r l y p e r i o d i c i t y i n w a t e r i n - f l u x '(varying w a t e r l e v e l through snow m e l t and p r e c i p i t a t i o n ) has a d e c i s i v e i n f l u e n c e on t h e development o f stream-accompanying causes a z o n a t i o n p a t t e r n  b r y o p h y t e assemblages. T h i s phenomenon  which i n c l u d e s v e g e t a t i o n s i n u n d a t e d f o r most  o f t h e y e a r , v e g e t a t i o n which l a y d r y f o r s h o r t e r o r l o n g e r p e r i o d s o f t i m e and, f i n a l l y , bryophyte assemblages o n l y o c c a s i o n a l l y submerged a t  108  the t i m e o f the y e a r ' s h i g h e s t water l e v e l s . The many a s s o c i a t i o n s described species  by European b r y o - s o c i o l o g i s t s a r e based m o s t l y on dominant  (designated  as c h a r a c t e r  s p e c i e s ) w i t h i n these zones. Dominance  i n and f i d e l i t y t o t h e s e 'communities' f a i l e d t o u n i t e them i n t o a sociological  s y s t e m , and f o r each o f them h i g h e r  were e r e c t e d .  synsystematic units  Hence, e c o l o g i c a l c r i t e r i a were used i n t h e proposal  f o r a c l a s s i f i c a t i o n o f the " a q u a t i c b r y o p h y t e communities". A f t e r an extensive  d i s c u s s i o n o f c l a s s i f i c a t i o n schemes, proposed by  b r y o - s o c i o l o g i s t s , Hertel study) synsystematic  (1974) p r o v i d e s t h e most u s e f u l  various  (for this  hierarchy.  In t h e p r e s e n t s t u d y , p r e d o m i n a n t l y , t h o s e a q u a t i c  bryophyte  communities were sampled, which a r e emerged f o r most o f t h e y e a r and t h o s e which a r e o n l y a f f e c t e d ( o c c a s i o n a l l y submerged) a t t h e t i m e s o f the y e a r ' s h i g h e s t w a t e r l e v e l s . These assemblages, "amphibische g e s e l l schaften"(Hertel Krusenstjerna,  1974), u n i t e d i n the a l l i a n c e R a c o m i t r i o n a c i c u l a r i s  are contained  as proposed by K r u s e n s t j e r n a and H e r t e l  w i t h i n the o r d e r B r a c h y t h e c i e t a l i a p l u m o s i , (1945) and a c c e p t e d by P h i l i p p i  (1974). T h i s o r d e r i n c l u d e s a l s o the a l l i a n c e o f submerged  b r y o p h y t e a s s o c i a t i o n s , t h e Scapanion u n d u l a t a e P h i l i p p i not s t u d i e d  (1956)  1956, as such  here.  Another a s s o c i a t i o n has been r e c o g n i z e d  and proposed as a new  s y n t a x o n . I t i s t h e Hygrohypnetum s m i t h i i , an a s s o c i a t i o n  described  from s o i l - c o v e r e d r o c k s a l o n g a l p i n e streams. I t s syntaxonomic p o s i t i o n , w i t h i n the  a l l i a n c e Platyhypnidion  r u s c i f o r m i s P h i l i p p i 1956 and t h e  order P l a t y h y p n i d i e t a l i a rusciformis P h i l i p p i  1956, i s f u r t h e r  elaborated  109  on i n the d i s c u s s i o n o f t h i s newly proposed  syntaxon.  The two a l l i a n c e s , R a c o m i t r i o n a c i c u l a r i s and Scapanion c o n t a i n s p e c i e s f o r which  European b r y o - s o c i o l o g i s t s have e r e c t e d  s e p a r a t e a s s o c i a t i o n s ^. Although the p r e s e n t data show apparent ships ( f l o r i s t i c a l l y s p e c i e s (Dichodontium  undulatae,  relation-  and e c o l o g i c a l l y ) w i t h t h e s e s y n t a x a , most o f these p e l l u c i d u m , Brachythecium  plumosum,  Racomitrium  a c i c u l a r i s , Scapania u n d u l a t a ) have been d e s i g n a t e d as a l l i a n c e o r as o r d e r c h a r a c t e r s p e c i e s . Order c h a r a c t e r s p e c i e s f o r t h e B r a c h y t h e c i e t a l i a plumosi  ( P h i l i p p i 1956)  prov. are Brachythecium  plumosum, Scapania  undu-  l a t a and Pel 1 i a n e e s i a n a . A l l i a n c e c h a r a c t e r s p e c i e s f o r t h e . R a c o m i t r i o n a c i c u l a r i s K r u s e n s t j e r n a 1945  i s Racomitrium  acicularis.  The f o l l o w i n g a s s o c i a t i o n s and s u b a s s o c i a t i o n s have been r e c o g n i z e d w i t h i n the a l l i a n c e R a c o m i t r i o n  acicularis:  (1) A s s . R a c o m i t r i o - S c o u l e r i e t u m a q u a t i c a e a s s . (2) Ass. Dichodontietum  p e l l u c i d i v. Hubschmann  nov.  1967  Subass. B r a c h y t h e c i o - Conocephaletosum c o n i c a e Subass. R i c c a r d i o - H y g r o b i e l l e t o s u m Subass. B l i n d i o - Scapanietosum  laxifoliae  paludosae  (3) Ass. M a r s u p e l l o - Nardietum  s c a l a r i s ass.  (4) Ass. Hypnetum d i e c k i i a s s .  nov.  nov.  In a d d i t i o n , t w o . f a c i e s have been r e c o g n i z e d w i t h i n the  Dichodontietum  p e l l u c i d i : a P h i l o n o t i s f o n t a n a f a c i e s and a B l i n d i a a c u t a - f a c i e s .  no  8.3.2. P h y t o g e o g r a p h i c a l  setting.  The p h y t o g e o g r a p h i c a l s e t t i n g a l s o i n d i c a t e s a f f i n i t i e s between s y n t a x a o f t h e o r d e r B r a c h y t h e c i e t a l i a p l u m o s i , as descrbed i n Europe and d i s t i n g u i s h e d i n t h i s s t u d y f o r B r i t i s h Columbia. T h i s i s , predomin a n t l y , r e f l e c t e d i n the large c o n t r i b u t i o n o f circumboreal  bryophytes  to t h e o v e r a l l s p e c i e s c o m p o s i t i o n o f t h e s y n t a x a . Among t h e h e p a t i c s , as major r e p r e s e n t a t i v e s o f t h e c i r c u m b o r e a l element, a r e Scapania u n d u l a t a , Pel 1 i a n e e s i a n a , M a r s u p e l l a e m a r g i n a t a , Jungermannia obovata  and N a r d i a s c a l a r i s . T h e i r i m p o r t a n t  c o n t r i b u t i o n to the syntaxa o f t h i s order i s o b v i o u s l y r e f l e c t e d i n t h e i r d e s i g n a t i o n as c h a r a c t e r s p e c i e s . A l s o H y g r o b i e l l a l a x i f o l i a i s a f r e q u e n t h e p a t i c o f stream accompanying bryophyte  v e g e t a t i o n , m o s t l y as an a s s o c i a t e ,  but foremost as t h e d i f f e r e n t i a t i n g s p e c i e s o f t h e s u b a s s o c i a t i o n R i c c a r d i o H y g r o b i e l l e t o s u m l a x i f o l i a e . The s p e c i e s has a b r o k e n - c i r c u m b o r e a l t r i b u t i o n , o c c u r r i n g i n o c e a n i c areas i n Europe, Japan and North  disAmerica  and a l s o f r e q u e n t l y found i n ( s u b ) a l p i n e areas o f C e n t r a l Europe. Among major companions a r e l i v e r w o r t s p e c i e s such as C h i l o s c y p h u s p o l y a n t h o s , P l a g i o c h i l a a s p l e n i o i d e s , D i p l o p h y l l u r n a l b i c a n s and Cepjhalozia bicuspidata. Mosses, i n d i c a t i n g t h e p r e d o m i n a n t l y  circumboreal aspect o f the  s y n t a x a w i t h i n t h e B r a c h y t h e c i e t a l i a plumosi a r e Dichodontium B l i n d i a a c u t a , Racomitrium l e s s e r degree, Pogonatum  a c i c u l a r e , Brachythecium  pellucidum,  piumosum and, t o a  urnigerum.  As w e s t e r n North American endemics o n l y S c o u l e r i a a q u a t i c a and  in  Rhizomnium g l a b r e s c e n s p l a y a f l o r i s t i c a l l y b i g e l o v i i and S c l e r o p o d i u m  important r o l e .  Porotrichum  o b t u s i f o l i u m o c c u r o n l y as o c c a s i o n a l l y 2  accompanying s p e c i e s i n some a s s o c i a t i o n s , . Among the d i s j u n c t p h y t o g e o g r a p h i c a l element o n l y a few  bryo-  phyte s p e c i e s s t a n d o u t as s i g n i f i c a n t c o n t r i b u t o r s to a s s o c i a t i o n s w i t h i n the B r a c h y t h e c i e t a l i a p l u m o s i . B i p o l a r d i s j u n c t r e p r e s e n t a t i v e s are Brachythecium  plumosum, and, among h e p a t i c s , R i c c a r d i a  multifida  and Blepharostoma  t r i c h o p h y l l u m . A w e s t e r n North American - Japan  d i s j u n c t , the moss Hypnum d i e c k i i , i s an almost c o n s t a n t companion i n most a s s o c i a t i o n s and the c h a r a c t e r s p e c i e s f o r the Hypnetum d i e c k i i . In t h i s s y n t a x o n , and i n t h e M a r s u p e l l o - Nardietum  scalaris,  r e p r e s e n t a t i v e o f t h e l a t t e r p h y t o g e o g r a p h i c a l element, chaceous moss O T i g o t r i c h u m  the  another  Polytri-  p a r a l l e l u m o c c u r s as an a d d i t i o n a l  8.3.3.1. Order BRACHYTHECIETALIA PLUMOSI ( P h i l i p p i  1956)  companion.  prov.  A l l i a n c e Racomitrion a c i c u l a r i s Krusenstjerna  1945  R a c o m i t r i o - S c o u l e r i e t u m a q u a t i c a e a s s . hov.  (Table XVI).  T h i s a s s o c i a t i o n occurs over a wide g e o g r a p h i c a l range w i t h i n the study a r e a , from Lynn Canyon to Brandywine F a l l s . I t was most f r e q u e n t l y a t Tow  found  e l e v a t i o n s and o n l y a few r e l e v e s were t a k e n  from a l p i n e s i t e s w i t h i n the B l a c k Tusk Meadows a r e a . Both c h a r a c t e r species f o r t h i s a s s o c i a t i o n , S c o u l e r i a a q u a t i c a , a western  North  112  American endemic, and Racomitrium B r i t i s h Columbia,  a c i c u l a r e , are widespread  throughout  i n c l u d i n g Vancouver I s l a n d and t h e Queen C h a r l o t t e  I s l a n d s . Thus, the d i s t r i b u t i o n o f the a s s o c i a t i o n i s l i k e l y t o e x t e n d beyond the range o f the p r e s e n t study a r e a . Hubschmann (1978) has d e s c r i b e d a S c o u l e r i e t u m a q u a t i c a e Vancouver I s l a n d . A c o n s t a n t companion was  from  Scleropodium o b t u s i f o l i u m ,  with S c h i s t i d i u m rivuTare.and F o n t i n a l i s a n t i p y r e t i c a also frequently o c c u r r i n g . The s h o r t s i t e d e s c r i p t i o n s show t h a t the a u t h o r has  taken  h i s r e l e v e s from s t r e a m s i d e r o c k s which a r e f r e q u e n t l y i n u n d a t e d . Under such c o n d i t i o n s s o l i d stands o f S c o u l e r i a a q u a t i c a develop w i t h S c l e r o p o d i u m o b t u s i f o l i u m as a f r e q u e n t a s s o c i a t e . However, f o r t h i s study the r e l e v e s from bryophyte  vegetations  a s s o c i a t e d w i t h streams were taken on r o c k s emerged f o r most p a r t o f t h e y e a r o r o n l y a f f e c t e d by spray i n p e r i o d s o f high water  levels.  Under such c o n d i t i o n s , S c o u l e r i a a q u a t i c a , a l t h o u g h sometimes becoming the dominant b r y o p h y t e s p e c i e s , was nions such as Racomitrium  o f t e n a s s o c i a t e d w i t h major compa-  a c i c u l a r e , Scapania s u b a l p i n a , B r a c h y t h e c i u m 3  plumosum and.Hypnum d i e c k i i  . Other o r d e r c h a r a c t e r s p e c i e s , o c c u r r i n g  as a d d i t i o n a l a s s o c i a t e s , are Jungermannia o b o v a t a , Dichodontium cidum, C h i l o s c y p h u s p o l y a n t h o s and S c a p a n i a The s i m i l a r i t i e s  pellu-  undulata.  i n f l o r i s t i c c o m p o s i t i o n ( e x c e p t f o r the endemic  c h a r a c t e r s p e c i e s S c o u l e r i a a q u a t i c a ) and i n h a b i t a t , i n d i c a t e t h e a s s o c i a t i o n ' s s y n s y s t e m a t i c a f f i n i t i e s t o the a l l i a n c e  Racomitrion  a c i c u l a r i s . S u b s e q u e n t l y , as suggested by Hubschmann (1978) - who posed a 'probable' own  a l l i a n c e , Scleropodion o b t u s i f o l i a e - the  prosyntaxon  Table XVI. Racomitrio - Seoulerietum aquaticae ass. nov. Number of releves Releve-number Elevation(m) Direction ofgexposure Inclination! ) Total cover(%)  1 6 30 NE 48 95  2 259 1487 S 80 100  Scouleria aquatica Racomitrium aciculare Brachythecium plumosum Scapania subalpina Hypnum dieckii Jungermannia obovata Scapania undulata Dichodontium pellucidum Blepharostoma trichophyllum Chiloscyphus polyanthos Pellia neesiana Schistidium rivulare Plagiochila asplenioides Marsupella emarginata Jungermannia pumila Jungermannia atrovirens Scleropodium obtusifolium Drepanocladus unclnatus Racomitrium aquaticum Blindia acuta Hygrohypnum ochraceum Hygrohypnum luridum Brachythecium frigidum Brachythecium velutinum Bryum pseudotriquetrum Isothecium stoloniferum Jungermannia exsertifolia Porotrichum bigelovii Dryptodon patens Racomitrium lanuginosum Riccardia multifida Pogonatum urnigerum Bryum pallescens Schistidium strictum Philonotis fontana  4.5 1.2  3.4  2.4  1.4 1.4  1.4 1.4  1.4 4.3  + .4 1.2  + .1  + .4  1.4 1.4 2.2  1.4 + .4 2.2  + .4  4 3 261 104 30 1487 N-NE N 10 58 90 95  6 106 30 S 54 95  2:4  9 224 610 N 26 100  10 126 457 SU 30 100  11 127 457 SW 22 90  12 128 457 SW 56 100  4.5 1.2 3.4 + .4  3.4 2.3 2.4 1.4  3.4 3.3 2.4 3.4 2.4 1.4  2.4 2.3 3.4 2.4 2.4  4.5 2.2 2.4 2.4 3.4  1.2 + .4 1.4  1.2 + .4 1.4 1.4  3.4 + .2 3.4  3.4 4.5 + .2 + .2 3.4 1.4  2.4 2.4 2.4  1.4 + .4 2.4  1.4 2.4 + .4 2.4 2.4  13 210 457 E 24 100  2.4  2.2  2.2 1.4 3.4 2.4 2.4  + .4  1.4 2.2 1.4  15 227 610 SW 66 90  16 228 610 SW 64 90  17 212 457 E 12 100  18 213 457 S 48 100  2.4 2.3 1.4 4.5-  2.4 2.3  2.,4 1..2  4.5  2.4 1.4 1.4  1.4 1.4 1.4  4..5 + .4 , 3..4 1..4 2..3  + .2 1.2 1.4 3.4 1.4 4.5  2.4  2.4  + .4 1.4  + .2  2.4  1.4  1,.4 2,.4  1.4  2.3  1.4 2.4 1 .4  2.4  + .2  1.2  1.2  1,.2  2.4  3.4  2.4  14 211 457 NE 26 90  5.5 3.3 1.4 3.4 1.4 1.4  2.2  1.4  2.4  8 229 610 SE 62 90  2.4 1.4  3.4  7 107 30 NE 50 100  3.4 1.2 4.5  1.4  + .4  5 105 30 W-SW 22 100  1.4  2.4  2.4  2.4  2.3  2.3  1.2  2.2  + .1  2..4 2.4  2.2 1.4 1.1 1.2 1.2  1.4 \  2.4  + .2 + .1 + .1 + .1  2.4  T a b l e XVI ( c o n t i n u e d ) .  Found o n l y once: H y g r o b i e l l a l a x i f o l i a , S t o k e s i e l l a p r a e l o n g a v a r . p r a e l o n g a , Aneura pingu Conocephalum conicum, Ceratodon purpureus, Scapania s c a n d i c a , M e t z g e r i a c o n j u g a t a , H e t e r o c l a d i u m m a c o u n i i , I s o p t e r y g i u m p u l c h e l l u m , Anoectangium a e s t i v u m , Pseudoleskea patens, F i s s i d e n s b r y o i d e s , P o r e l l a r o e l l i i , Plagiomnium r o s t r a t u m , Grimmia t o r q u a t a .  Rel  eve-locations:6,104,105,106,107 259,261 224,227,228,229 126,127,128,210,211,212,213  Lynn Canyon B l a c k Tusk Meadows Cheakamus R i v e r V a l l e y Brandywine F a l l s  115  s h o u l d be c l a s s i f i e d i n t h e o r d e r B r a c h y t h e c i e t a l i a plumosi 1956.  Philippi  <  Dichodontietum p e l l u c i d i  v. Hubschmann 1967  T h i s a s s o c i a t i o n w i t h Dichodontium p e l l u c i d u m as i t s c h a r a c t e r s p e c i e s , i s d e s c r i b e d by Hubschmann (1967) from s t r e a m s i d e r o c k s , c o v e r e d w i t h s o i l b u t submerged f o r most o f t h e y e a r . From  similar  h a b i t a t s i t has been r e p o r t e d and documented by Norr ( 1 9 7 0 ) , Neumayr ( 1 9 7 1 ) , M a r s t a l l e r (1973) and as a s u b a s s o c i a t i o n o f t h e B r a c h y t h e c i e t u m plumosi K r u s e n s t j e r n a 1945 by M a r s t a l l e r (1980). Hubschmann (1978) a l s o d e s c r i b e d t h e a s s o c i a t i o n from Vancouver  Island.  Frequent a s s o c i a t e s i n t h e D i c h o d o n t i e t u m p e l l u c i d i are  Mnium punctatum,  Conocephalum com'cum, Eurhynchium  Platyhypnidium r i p a r i o i d e s .  i n Europe  schwartzii,  Further associated species, contributing  t o t h e p r e s e n t syntaxon both i n Europe and t h e Vancouver  Island vegetations  documented by Hubschmann ( 1 9 7 8 ) , a r e b r y o p h y t e s such as B r a c h y t h e c i u m rivulare,  C h i l o s c y p h u s p o l y a n t h o s , S t o k e s i e l l a p r a e l o n g a and Brachy-  thecium plumosum. The p r e s e n t data do n o t c o n t a i n any r e p r e s e n t a t i o n o f t h e Dichodontietum p e l l u c i d i  - "typicum". D e s p i t e i t s constancy t h e c h a r a c t e r  s p e c i e s u s u a l l y o c c u r r e d w i t h low c o v e r v a l u e s , w i t h t h e e x c e p t i o n o f r e l e v e s 90 and 92. In comparison  to t h e t y p e o f h a b i t a t , d e s c r i b e d i n  European r e p o r t s on t h i s syntaxon and t h e s i t e d e s c r i p t i o n s by Hubschmann (1978) from Vancouver  I s l a n d , the t a b l e d r e l e v e s o f the Dichodontietum  116  pellucidi  i n the p r e s e n t treatment were t a k e n from d i f f e r e n t m i c r o -  h a b i t a t s . These w i l l  be e l a b o r a t e d  on i n the d i s c u s s i o n s o f the  several  d i s t i n g u i s h e d u n i t s w i t h i n the a s s o c i a t i o n . W i t h i n the D i c h o d o n t i e t u m p e l l u c i d i t h i s study r e c o g n i z e s subassociations  Brachythecio  and two f a c i e s .  - Conocephaletosum c o n i c a e  T h i s s u b a s s o c i a t i o n was  subass. nov.  (Table  XVII).  found a t s e v e r a l s i t e s i n Lynn Canyon  and o c c u r s a l s o northward a t Brandywine F a l l s , a t the base o f Mountain and, Its  three  f i n a l l y , was  described  from the C h i l l i w a c k R i v e r V a l l e y .  h a b i t a t i s bases o f b o u l d e r s and  under shady and  Whisler  s t e e p r o c k w a l l s , where,  s h e l t e r e d c o n d i t i o n s , the b r y o p h y t e v e g e t a t i o n  never  becomes i n u n d a t e d but i s e x t e n s i v e l y a f f e c t e d by s p r a y , e i t h e r from a d j a c e n t streams o r permanent seepage. The soil  on the rock s u r f a c e was  a c c u m u l a t i o n o f sandy-loamy  never measured t h i c k e r than 1  cm.  In b r y o s o c i o l o g i c a l work c a r r i e d out i n Europe, most communities w i t h Conocephal urn com'cum as t h e dominant s p e c i e s , have been as a s s o c i a t i o n s from c a l c a r e o u s  rocks  1955,  J e z e k and Vondracek 1962,  H o f l e r 1959,  Smarda 1947,  described  (Herzog and H o f l e r 1944,  Koppe  Philippi  1965,  Hebrard 1975). As such t h e i r f l o r i s t i c c o m p o s i t i o n i s s u b s t a n t i a l l y d i f f e r e n t from t h a t e x h i b i t e d i n the p r e s e n t d a t a , which i n d i c a t e c l o s e a f f i n i t i e s t o the Conocephalurn conicum - community d e s c r i b e d  by  Poelt  ( 1 9 5 4 ) , the B r a c h y t h e c i u m r i v u l a r e - Conocephal urn conicum - community o f Hagel (1966) and  the F e g a t e l l e t u m  conicae  Schade 1934,  reported  on  Table X V I I . D i c h o d o n t i e t u m p e l l u c i d i v. Hubschmann 1967: Brachythecio-Conocephaletosum Number o f r e l e v e s Releve-number E l e v a t i o n (m) D i r e c t i o n o f ^exposure Inclination ( ) T o t a l c o v e r {%)  1 86 30 NE 74 90  2 90 30 E 72 100  3 88 30 E 84 80  Dichodontium p e l l u c i d u m Scapania u n d u l a t a Conocephalurn conicum Geocalyx g r a v e d ens Jungermannia pumila S t o k e s i e l l a praelonga var. praelonga Plagiochila asplenioides Porotrichum b i g e l o v i i Brachythecium f r i g i d u m Hypnum d i e c k i i B l i n d i a acuta Riccardia muVtifida. Scapania s u b a l p i n a Rhizomnium g l a b r e s c e n s Racomitrium a c i c u l a r e Metzgeria conjugata Pel 1 i a n e e s i a n a  3.3  4.3 1.4 5.5 + .4 1.4 1.4 + .4  1 .2  4.5 2.4 2.4 + .4 + .4 + .4  2.4  3.4 1.4 2.4 2.4 + .4  c o n i c a e subass.  nov.  4 87 30 SE 78 90  5 89 30 E-NE 76 90  6 276 266 W 56 100  7 329 1219 SW 48 90  8 206 457 NE 48 100  9 208 457 S-SW 56 100  1 .2  2.2 1.4 3.4  2.4  1.2 1.4  2.2 4.4  2.2 4.4  3.4 1 .4 2.4 4.5  1.4 1.4 2.4  1.4  + .1 + .1 1.4  + .4 4.5  3.4 1.4 1.2  3.4 1.4 + .1 1.4  1.1  1.2 + .4 + .1  2.4  2.2 2.3  2.4 + .1  Found o n l y once: Jungermannia o b o v a t a , C e p h a l o z i a b i c u s p i d a t a , Blepharostoma t r i c h o p h y l 1 urn, C h i l o s c y p h u s p o l y a n t h o s , P h i l o n o t i s f o n t a n a , Bryum p a l l e s c e n s , Aneura p i n g u i s , Jungermannia a t r o v i r e n s , S c h i s t i d i u m r i v u l a r e , S c a p a n i a americana, Scleropodium obtusum, Hygrohypnum ochraceum. Ceratodon p u r p u r e u s , T r i t o m a r i a q u i n q u e d e n t a t a , Brachythecium v e l u t i n u m , P o h l i a c r u d a , Amphidium c a l i f o r n i c u m , Cynodontium j e n n e r i , A t r i c h u m undulatum, M y l i a t a y l o r i , S t o k e s i e T l a p r a e l o n g a v a r . s t o k e s i i , B l a s i a p u s i l l a , Lophozia s u d e t i c a  Table XVII (continued). Releve - l o c a t i o n s : 86. 87. 88, 89, 90  Lynn Canyon  276  C h i l l i w a c k River Vail  329  Whistler Mountain  206, 208  Brandywine F a l l s  119  by Hiibschtnann ( 1 9 6 7 ) , Norr (1969) and M a r s t a l l e r (1973) . H  These  a f f i n i t i e s are e x p r e s s e d by t h e two major d i f f e r e n t i a t i n g s p e c i e s f o r t h i s s u b a s s o c i a t i o n : Conocephalum conicum and Brachythecium f r i g i d u m . F u r t h e r i n d i c a t i o n i s found i n t h e presence o f s p e c i e s such as S t o k e s i e l l a p r a e l o n g a v a r . p r a e l o n g a , P l a g i o c h i l a a s p l e n i o i d e s and R h i z o mnium g l a b r e s c e n s , a w e s t e r n North American  endemic.  However, t h e r e l e v e s c o m p r i s i n g the p r e s e n t s u b a s s o c i a t i o n express major d i f f e r e n c e s i n f l o r i s t i c European b r y o - s o c i o l o g i c a l may  c o m p o s i t i o n i n comparison  s t u d i e s mentioned  show t h i s t o be i n accordance  to  before. Further research  to M a r s t a l l e r ' s (1973) statement t h a t  "the F e g a t e l l e t u m c o n i c a e Schade 1934, due t o i t s wide h a b i t a t amplitude,^has to be s u b d i v i d e d i n numerous subassociations"  As encountered  i n the present study t h i s c o l l e c t i o n o f r e l e v e s  has been g i v e n s u b a s s o c i a t i o n rank based on two main r e a s o n s : (1) A l t h o u g h Conocephalum conicum and B r a c h y t h e c i u m  f r i g i d u m are domi-  nant t a x a , t h e r e l e v e s do not c o n t a i n ' o n e - s p e c i e s - v e g e t a t i o n s  1  but  w e l l developed c o m b i n a t i o n s o f s p e c i e s ; (2) The c o l l e c t i o n o f r e l e v e s i s not c o n f i n e d t o one s e c t i o n o f the study a r e a but i s w i d e s p r e a d . The B r a c h y t h e c i o - Conocephaletosum c o n i c a e J s ,  besides i t s  c h a r a c t e r s p e c i e s , f u r t h e r d e f i n e d by s p e c i e s such as Scapania  undulata,  Jungermannia p u m i l a , Geocalyx g r a v e o l e n s . Less f r e q u e n t b r y o p h y t e s i n c l u d e Hypnum d i e c k i i , P l a g i o c h i l a a s p l e n i o i d e s and P o r o t r i c h u m b i g e l o v i i .  120  Riccardio - Hygrobielletosum  l a x i f o l i a e subass.  nov.  (Table X V I I I ) .  T h i s s u b a s s o c i a t i o n i s d e s c r i b e d from s e v e r a l s i t e s i n Lynn Canyon where i t o c c u r r e d c h a r a c t e r i s t i c a l l y on s h e l t e r e d and r o c k s which become o c c a s i o n a l l y inundated from snow melt o r e x t r e m e l y  shaded  at high water l e v e l s  resulting  high p r e c i p i t a t i o n . I t s micro-habitat i s  h o r i z o n t a l and g e n t l y s l o p i n g rock s u r f a c e s covered w i t h o n l y a t h i n layer of deposited s i l t , containing p a r t i c l e s of d i s i n t e g r a t e d rock. Although  t h e s e r e l e v e s were t a k e n from one g e o g r a p h i c  the r e c u r r i n g combination communities", was o f t h e i r constancy  o f s p e c i e s , as opposed t o  area,  "one-species-  d e c i s i v e i n the r e c o g n i t i o n as s u b a s s o c i a t i o n . Because and near f i d e l i t y t o t h i s s e t o f r e l e v e s , Hygro-  b i e l l a l a x i f o l i a and R i c c a r d i a m u l t i f i d a have been d e s i g n a t e d as characterizing differential  s p e c i e s . F u r t h e r c o n s t a n t , and  sometimes  h i g h l y abundant, d i f f e r e n t i a t i n g t a x a a s s o c i a t e d w i t h t h e two s p e c i e s are Scapania  the  previous  u n d u l a t a , N a r d i a s c a l a r i s , Hypnum d i e c k i i and  to  a l e s s e r degree Jungermannia obovata and C e p h a l o z i a b i c u s p i d a t a . C o n s t a n t , but not d i f f e r e n t i a t i n g , companions a r e Dichodontium p e l l u c i d u m and B I i n d i a a c u t a .  B l i n d i o - Scapanietosum paludosae subass.  The  nov.  (Table X I X ) .  p r e s e n t s u b a s s o c i a t i o n i s d e s c r i b e d from s e v e r a l s i t e s  i n f r o n t o f the Brandywine F a l l s . Exposed and h e a v i l y a f f e c t e d by  spray,  the B l i n d i o - Scapanietosum paludosae o c c u r r e d on h o r i z o n t a l and g e n t l y s l o p i n g b o u l d e r s u r f a c e s w i t h o n l y a t h i n l a y e r o f accumulated  Table XVIII.  D i c h o d o n t i e t u m p e l l u c i d i v. Hubschmann 1967: R i c c a r d i o - H y g r o b i e l l e t o s u m l a x i f o l i a e subass. nov. Number o f r e l e v e s Releve-number Elevation(m) D i r e c t i o n ofpexposure Inclination( ) Total cover  1 195 30 SE 48 100  2 196 30 S-SW 46 100  3 200 30 SE 24 100  4 197 30 SE 58 100  5 198 30 SE 70 90  6 199 30 S 28 90  Scapania undulata Hygrobiella l a x i f o l i a Dichodontium p e l l u c i d u m Nardia s c a l a r i s B l i n d i a acuta Riccardia multifida Hypnum d i e c k i i Jungermannia obovata Cephalozia bicuspidata Jungermannia p u m i l a Oxystegus t e n u i r o s t r i s  4.5 3.4 3.3 2.4 2.3 2.4 1.4 2.4 2.4 2.4 2.2  4.4 3.4 1.2 3.4 2.3 2.4 1.4 2.4 1.4 1.4 1.2  4.5 2.4 3.3 2.4 1.2 2.4 1.4 2.4 2.4  3.4 4.4 2.2 2.4 2.2 1 .4 2.4  3.4 4.4 2.2 2.4 1 .2 1.4 1.4'  3.4 4.4 + .2 1 .4 4.3 1 .4 + .1  L o c a t i o n o f a l l r e l e v e s : Lynn Canyon.  122  ., 6 soil The c h a r a c t e r i z i n g d i f f e r e n t i a l  species f o r t h i s  subassociation  a r e S c a p a n i a p a l u d o s a and B l i n d i a a c u t a . Other b r y o p h y t e s , w h i c h , because o f t h e i r near confinement t o t h i s s u b a s s o c i a t i o n , a r e h i g h l y c h a r a c t e r i s t i c f o r i t are P r e i s s i a q u a d r a t a and F i s s i d e n s osmundoides. C o n s t a n t companions t o the B l i n d i o - Scapanietosum p a l u d o s a e a r e s p e c i e s such as Dichodontium p e l l u c i d u m , Blepharostoma t r i c h o p h y l T u r n , Bryum p a l l e s c e n s , Aneura p i n g u i s , Jungermannia a t r o v i r e n s and Hygrobiella  laxifolia. G e i s s l e r ( 1 9 7 6 ) , i n her study o f t h e h y g r o p h i l i c b r y o p h y t e  v e g e t a t i o n o f the e a s t e r n Swiss A l p s , proposes a new a l l i a n c e , t h e M a r s u p e l l o - S c a p a n i o n . A s s o c i a t i o n s w i t h i n t h i s s y n t a x o n o c c u r on s i t e s where streams o r i g i n a t e i n s u b a l p i n e and a l p i n e a r e a s . Predominant environmental f a c t o r s f o r t h e b r y o p h y t e v e g e t a t i o n a r e (1) the w a t e r volume and i t s low c u r r e n t v e l o c i t y and (2) the i m p e r m e a b i l i t y o f the ( m a i n l y g r a n i t i c ) s u b s t r a t e . Both f a c t o r s cause an environment w i t h h i g h m o i s t u r e s t a t u s and h i g h a i r h u m i d i t y . The p r e s e n t s u b a s s o c i a t i o n shows a p p a r e n t f l o r i s t i c  relationships  t o the a s s o c i a t i o n s , B l i n d i o - Scapanietum u n d u l a t a e and B l i n d i o Scapanietum u l i g i n o s u m , as d e s c r i b e d by G e i s s l e r (1976) i n t h e a l l i a n c e M a r s u p e l l o - S c a p a n i o n . F u r t h e r r e s e a r c h might s u b s t a n t i a t e A particularly allied assemblage was  this.  [ t o G e i s s l e r ' s (1976) d a t a ] b r y o p h y t e  found a t e c o l o g i c a l l y s i m i l a r s i t e s as d e s c r i b e d by t h i s  Swiss b r y o l o g i s t . Is i s r e p r e s e n t e d by t h e r e l e v e s 305 t o 308, t a k e n from  a l p i n e s i t e s on W h i s t l e r Mountain. These assemblages a r e predomi-  Tab'le XIX. Dichodontietum p e l l u c i d i v. Hubschmann 1967: B I i n d i o - S c a p a n i e t o s u m paludosae a s s , nov. Number o f r e l e v e s Releve-number Elevation(m) D i r e c t i o n ofgexposure Inclination( ) Total cover{%)  1 216 457 W 26 100  2 219 457 SW 40 100  3 218 457 W 32 100  4 220 457 S 68 100  5 217 457 E 40 100  6 221 457 S 40 100  B I i n d i a acuta S c a p a n i a paludosa Dichodontium p e l l u c i d u m P r e i s s t a quadrata F i s s i d e n s osmundoides BIepharostoma t r i c h o p h y l l u r n Hygrobiell a l a x i f o l i a Aneura p i n g u i s Jungermannia a t r o v i r e n s Bryum p a l l e s c e n s Pel 1 i a n e e s i a n a Jungermannia p u m i l a Rhizomnium g l a b r e s c e n s Jungermannia e x s e r t i f o l i a Scapania s u b a l p i n a  4.4 3.4 2.3 2.4 1.2 1.4 1.4 1.4 1.4  4.4 4.5 2.2 1.4 1.2 1.4 1.4 + .4 1.4 1.2 + .1 1.4  4.5 4.5 2.3 1.4 1 .2 + .4 1 .4 + .4  3.4 4.5 1.2 2.4 + .1 1.4 + .4 3.4 2.4 + .2 + .1  4.5 2.4 2.2 2.4 3.3 1.4  4.4 2.4 2.2 + .1 1.2 1 .4 4.5  Location of a l l  r e l e v e s : Brandywine  1.1  1 .2 3.4  2.4 1.4 2.3  1 .4 1 .2  2.4 1.1 2.4 1.4  Falls  124  n a n t l y c h a r a c t e r i z e d by A n t h e l i a j u r a t z k a n a , a f r e q u e n t companion s p e c i e s i n t h e a s s o c i a t i o n Nardietum compressae d e s c r i b e d by  Geissler  (1976) and c l a s s i f i e d i n the a l l i a n c e M a r s u p e l l o - S c a p a n i o n . F u r t h e r c h a r a c t e r s p e c i e s o f t h e b r y o p h y t e v e g e t a t i o n o f r e l e v e s 305 t o are  b r y o p h y t e s s u c h as P h i l o n o t i s f o n t a n a , Pseudoleskea  308  atricha,  Hypnum subimponens and S c a p a n i a u l i g i n o s a . A d d i t i o n a l a s s o c i a t e s a r e Racomitrium s u d e t i c u m , L o p h o z i a s u d e t i c a , K i a e r i a s t a r k e i , D i c r a n o w e i s i a c r i s p u l a , Bartramia i t h y p h y l l a , Pohlia l u d w i g i i , Pleuroclada albescens, Jungermannia e x s e r t i f o l i a and S c a p a n i a p a l u d o s a . The r e l e v e s 305 t o 308 a l s o c o n t a i n f l o r i s t i c a f f i n i t i e s w i t h the Pseudoleskea - B r a c h y t h e c i um g l a c i a l e - v a r i a n t and t h e R a c o m i t r i um sudeticum - v a r i a n t o f t h e a s s o c i a t i o n Dermatocarpetum r i v u l o r u m as newly proposed and d e s c r i b e d by G e i s s l e r  (1976).  F u r t h e r r e s e a r c h , e s p e c i a l l y i n s u b a l p i n e and a l p i n e areas o f w e s t e r n North A m e r i c a , may  e l a b o r a t e on the s t r u c t u r e and n a t u r e o f t h e s e  v e g e t a t i o n s and c l a r i f y p o s s i b l e r e l a t i o n s h i p s w i t h European v e g e t a t i o n s .  In  a d d i t i o n t o the t h r e e s u b a s s o c i a t i o n s w i t h i n t h e a s s o c i a t i o n  D i c h o d o n t i e t u m p e l l u c i d i , t h e p h y t o s o c i o l o g i c a l t a b l e r e v e a l s t h e presence of  two groups o f r e l e v e s , r e c o g n i z e d here as ' f a c i e s ' . These are b r y o p h y t e  assemblages which show f l o r i s t i c a f f i n i t i e s to t h e i r but are dominated  'parent' a s s o c i a t i o n  by an a c c i d e n t a l c o m b i n a t i o n o f s p e c i e s , s i g n i f i c a n t l y  d i f f e r e n t from the a s s o c i a t i o n w i t h i n which these ' f a c i e s ' are r e c o g n i z e d . Extremes a r e e x p r e s s e d by  "one-species-communities".  As s u c h , t h e p r e s e n t data d i s t i n g u i s h a P h i l o n o t i s f o n t a n a - f a c i e s  Table XX.  Dichodontietum  p e l l u c i d i v. Hubschmann 1967: P h i l o n o t i s f o n t a n a - f a c i e s .  Number o f r e l e v e s Releve-number Elevation(m) Direction of exposure Inclination^ ) Total cover(%)  1 269 305 SW 52 100  P h i l o n o t i s fontana Dichodontium p e l l u c i d u m RiccardiamuTtifida P o h l i a nutans Cephalozia bicuspidata Chiloscyphus polyanthos Scapania u n d u l a t a  4.3 4.3 3.3 2.2 3.3 4.3 2.4 2.4 3.4 1 . 2 1 . 2 1 . 2 +.4 +.4 +.4 +.1 3.4 +.1  n  Location of a l l  r e l e v e s : C h i l l i w a c k Lake  2 271 305 W 38 100  3 270 305 W 60 100  T a b l e XXI. D i c h o d o n t i e t u m p e l l u c i d i v. Hubschmann 1967: B l i n d i a a c u t a - f a c i e s . Number o f r e l e v e s Releve-number Elevation(m) D i r e c t i o n ofgexposure Inclination( ) Total cover(%)  1 207 457 E 104 60  2 267 305 SW 58 90  3 268 305 S 32 95  4 215 457 NW 26 100  5 275 305 NW 32 100  B l i n d i a acuta Dichodontium p e l l u c i d u m Bryum c a p i l l a r e Scapania paludosa Scapania undulata P h i l o n o t i s fontana Barbula r e f l e x a S c a p a n i a mucronata Radula b o l a n d e r i P r e i s s i a quadrata Jungermannia a t r o v i r e n s Bryum miniatum P e l l i a neesiana Rhizomnium g l a b r e s c e n s Chiloscyphus polyanthos Jungermannia e x s e r t i f o l i a Brachythecium f r i g i d u m Aneura p i n g u i s I s o p t e r y g i u m elegans Isopterygium pulchellum Plagiochila asplenioides Racomitrium a c i c u l a r e  4.5  4.5 1.2 1.2 1 .4  4.5  4.5 3.3  4.5 3.3 1.2  1 .2 + .1  1.4 2.4 1.2  1.4 2.2 1 .2 + .1  1.2 + .1  3.4 3.4 2.4 2.2 1.1 1.1 1.4 1.4 1.1 1.4 1 .4 1.2 + .1  R e l e v e - l o c a t i o n s : 207, 215 Brandywine F a l l s 267, 268, 275 C h i l l i w a c k Lake  + .1  127  and a B l i n d i a a c u t a - f a c i e s . The f i r s t f a c i e s i s d e s c r i b e d from s o i l - c o v e r e d r o c k s a t e x t e n s i v e , open seepage a r e a s a l o n g C h i l l i w a c k Lake. Under t h e s e c o n d i t i o n s P h i l o n o t i s f o n t a n a was c o n s t a n t l y  associated  w i t h R i c c a r d i a m u l t i f i d a , Dichodontium pel 1ucidum, P o h l i a nutans and Cephalozia bicuspidata. The second f a c i e s i s r e p o r t e d from the same a r e a as the P h i l o n o t i s f o n t a n a - f a c i e s but r e c u r r e d under t h e same c o n d i t i o n s a t Brandywine  F a l l s . Frequent companions i n t h i s B l i n d i a a c u t a - f a c i e s  were Dichodontiurn pel 1ucidum, S c a p a n i a p a l u d o s a and Bryum c a p i l l a r e .  M a r s u p e l l o - Nardietum s c a l a r i s a s s . nov. ( T a b l e X X I I ) .  T h i s a s s o c i a t i o n i s d e s c r i b e d from r o c k s a d j a c e n t t o streams which do not become i n u n d a t e d b u t a r e a f f e c t e d o n l y by s p r a y a t t i m e s o f h i g h e s t w a t e r l e v e l s . The M a r s u p e l l o - Nardietum s c a l a r i s  characteris-  t i c a l l y o c c u r s on h o r i z o n t a l top s u r f a c e s and g e n t l y s l o p i n g p a r t s o f r o c k s under shaded and s h e l t e r e d c o n d i t i o n s . Under such c o n d i t i o n s most b r y o p h y t e s p e c i e s were found growing on o n l y a t h i n l a y e r o f s o i l t h a t had accumulated on t h e s e r o c k s . Geographically, the present a s s o c i a t i o n i s widely  distributed  t h r o u g h o u t the s t u d y a r e a . I t i s documented from s t r e a m s i d e r o c k h a b i t a t s i n Lynn Canyon, M u r r i n Creek and Stoney Creek canyons a l o n g t h e Highway, northward from t h e Cheakamus R i v e r banks and Brandywine  Squamish Falls  and, f i n a l l y , eastward from s i t e s a l o n g the C h i l l i w a c k R i v e r . A l t h o u g h o c c u r r i n g i n an e c o l o g i c a l l y d i f f e r e n t e n v i r o n m e n t , t h e  128  M a r s u p e l l o - Nardietum scalaris Philippi  1956,  s c a l a r i s shows some a f f i n i t i e s t o the  an a s s o c i a t i o n r e p o r t e d and documented by  European b r y o - s o c i o l o g i s t s ( P h i l i p p i  1956,  1963, Hubschmann 1967,  1969, Neumayr 1971, Dunk 1971, M a r s t a l l e r 1973, all  d e s c r i b e t h e Nardietum  s c a l a r i s as a t e r r e s t r i a l  t r a i l s and uprooted t r e e s . T h i s syntaxon u r n i g e r i K r u s e n s t j e r n a 1945  D i c r a n e l l e t a l i a heteromallae P h i l i p p i  Norr *  1980). These a u t h o r s association  open, d i s t u r b e d s i t e s where i t o c c u r s on loamy-sandy s o i l  Pogonation  Nardietum  from  along roads,  i s c o n t a i n e d i n the a l l i a n c e  and c l a s s i f i e d i n the o r d e r 1956, an o r d e r from a c i d l y  reacting  s o i l . T h i s i s r e f l e c t e d i n the major companions, d e s i g n a t e d as o r d e r and a l l i a n c e c h a r a c t e r s p e c i e s : D i c r a n e l l a h e t e r o m a l l a , D i t r i c h u m heteromal1 urn,  A t r i c h u m undulatum, C e p h a l o z i a b i c u s p i d a t a , I s o p t e r y g i u r n  e l e g a n s , Pogonatum urnigerum.  In t h e p r e s e n t s t u d y , a l l t h e s e s p e c i e s  o c c u r e i t h e r o c c a s i o n a l l y ( e x c e p t f o r C e p h a l o z i a . b i c u s p i d a t a , which i s an almost c o n s t a n t companion i n t h i s a s s o c i a t i o n ) o r a r e not p r e s e n t at  a l l . P r e s e n t b r y o p h y t e t a x a , i n d i c a t i n g f l o r i s t i c a f f i n i t i e s between  t h e "European" syntaxon and t h e M a r s u p e l l o - Nardietum  scalaris  recog-  n i z e d i n t h i s s t u d y a r e : D i p l o p h y l l u m a l b i c a n s , O l i g o t r i c h u m p a r a l l e l urn, 0_. a l igerum and A t r i c h u m s e l w y n i i . D e s p i t e t h e s e a f f i n i t i e s i n f l o r i s t i c c o m p o s i t i o n , two major reasons f o r t h e r e c o g n i t i o n o f t h e M a r s u p e l l o - Nardietum as a s e p a r a t e syntaxon and i t s syntaxonomic m i t r i o n a c i c u l a r i s K r u s e n s t j e r n a 1945  p o s i t i o n w i t h i n the Raco-  are  (1) the h a b i t a t o f t h e p r e s e n t syntaxon ' t e r r e s t r i a l ' Nardietum  scalaris  i s d e c i s i v e l y d i f f e r e n t from  scalaris Philippi  1956  and  the  Table  XXII.  Marsupello - Nardietum scalaris ass. nov. Number of releves Releve-number Elevation(m) Direction ofgexposure Inclination! ) Total cover(%)  1 2 48 115 61 61 NW N-NW 12 82 90 80  3 81 30 NE 52 95  4 203 610 SE 62 100  Nardia scalaris Marsupella emarginata Hypnum dieckii Rhizomnium glabrescens Cephalozia bicuspidata Diplophyllum albicans Pellia neesiana Scapania undulata Jungermannia obovata Dichodontium pelluddum Blindia acuta Oligotrichum parallelum Hygrobiella l a x i f o l l a Racomitrium aciculare Pogonatum urnlgerum Atrichum selwynii Oligotrichum allgerum Anastrophyllum minutum Scapania subalplna Brachythecium frigidum Scapania americana Riccardia mul'tifida  4.4  3.4  1.4 1.2 1.4  1.2 2.4  3.4 2.4 2.4 + .1 + .4  2.4  +.1  3.4 1.4 2.4 1.2 2.4 1.4 1.4 2.4 2.4 1.2  Found only once:  2.3 1.2  1.4 2.3  + .1 2.4  2.4 2.4 3.4  1.4  + .4  2.4  1.4 3.4 2.4  9 194 457 S 68 100  10 202 610 SE 44 100  11 205 610 S-SE 82 100  12 76 266 NE 18 95  13 77 266 NE 26 90  14 111 30 NW 50 100  2.4 3.4 4.4 3.3 2.4 1.4 + .4 + .1 1.4 + .1 2.4 2.3  2.4 1.4 4.5 2.2 2.4 2.4 1.4  2.4 4.5 3.4 1.2  + .4 5.5 1.4 2.3 1.4 1.4  3.4 2.4 2.2 2.4 1.4  3.3  1.2  2.3  2.3  1.2 1.4  2.4 1.4 1.4 1.2 2.4 1.4 1.4 2.4 2.4 2.3 1.2 4.3  3.4 2.4 2.4 2.2 1.4 2.4 1.4 2.4  1.2 2.3  1.4 2.4  6 7 8 344 191 192 61 457 457 N-NW S-SE S-SW 32 52 48 80 90 100  5.5 1.4 3.4  + .1 + .1  3.4  5 83 30 N 34 100  1.4 2.3  3.4 2.4 4.5 2.3  1.4 1.4 2.4 + .1  Brachythecium plumosum, Plagiochila asplenioides, Philonotis fontana, Pohlia nutans, Diplophyl1um taxifolium. Isopterygium elegans , Oxystegus tenuirostris, Diplophyllum piicatum, Heterocladium macounli, Bartramiopsis l e s c u r i i , "Racomitrium brevipes", Ditrichum heteromallum, Pohlia longebracteata, Jamesoniella autumnal i s , Philonotis arnelli i  Releve-locations: 48,344 115 78,79,81,83,84,91.92,93,94 201,202.203,204,205 191 ,192,193,194 111 76,77  Shannon Falls Murrin Creek Lynn Canyon Cheakamus River Valley Brandywine Falls Stoney Creek Chilliwack River Valley  1.4  2.4 2.2  1.2  + .4 1.4  1.4 + .1  1.4  1.4 1.4  1.4 + .2  2.4 1.4 + .1  2.3 + .1  15 16 79 84 30 30 NW NE 28 64 100 .100  17 78 30 NE 36 100  18 92 30 W 62 95  19 91 30 NE 62 95  20 193 457 S 50 95  21 93 30 NE 30 100  22 94 30 SW 52 80  23 201 610 SE 82 100  24 204 610 E-SE 68 100  1.4 2.4 4.4 3.4 + .4 + .4  4.5 2.4 1.4 3.3  3.4 3.4 + .4 3.3  4.4 3.4 2.4 2.2  2.4 2.4  1.4  2.4 4.5 + .4 + .1 1.2 + .4 1.4 + .4 + .4  3.4 2.4  1.4  4.5 2.4 2.4 2.3 1.4 2.4 + .1  + .4  1.4  1.4 1.4 2.4 1.2  1.2  1.2  + .1  3.4 4.5  2.3  1.2  1.2 2.4 3.4  + .1  1.2  + .1  1.2 2.4  + .1  + .1  1.4 3.3  2.2  2.4 1.4  1.2  1.2  1.2  1.4  2.4 + .1  130  (2) t h e presence and frequency  of alliance  c h a r a c t e r s p e c i e s as major  companions. Because o f t h e i r constancy  and abundance, N a r d i a s c a l a r i s and  M a r s u p e l l a emarginata a r e d e s i g n a t e d as t h e c h a r a c t e r s p e c i e s o f t h e p r e s e n t a s s o c i a t i o n . Other s p e c i e s , c h a r a c t e r i s t i c a l l y , w i t h major occurrence  i n t h i s syntaxon  i n c l u d e C e p h a l o z i a b i c u s p i d a t a , Rhizomnium  g l a b r e s c e n s , D i p l o p h y l l u m a l b i c a n s and O l i g o t r i c h u m p a r a l l e l u m . Order and a l l i a n c e c h a r a c t e r s p e c i e s as s i g n i f i c a n t a s s o c i a t e s are Scapania  u n d u l a t a , P e l l i a n e e s i a n a , Dichdontium p e l l u c i d u m ,  whereas Racomitrium a c i c u l a r e , Jungermannia obovata and H y g r o b i e l l a l a x i f o l i a occur a l s o f r e q u e n t l y i n t h i s a s s o c i a t i o n .  Hypnetum d i e c k i i a s s . nov. ( T a b l e X X I I I ) .  The  a s s o c i a t i o n i s r e p o r t e d and documented from Lynn Canyon,  i n t h e southern  p o r t i o n o f t h e study a r e a , t o Brandywine F a l l s i n t h e  n o r t h , where t h e syntaxon  i s a l s o d e s c r i b e d from two s u b a l p i n e  sites  i n t h e B l a c k Tusk Meadows a r e a . I t s h a b i t a t i s h o r i z o n t a l t o s l i g h t l y s l o p i n g rock s u r f a c e s with mineral s o i l accumulation  up t o 1 cm i n t h i c k n e s s ( i n o n l y a few  i n s t a n c e s t h i s was measured t o be almost 2 cm t h i c k ! ) . These s u b s t r a t a f o r t h e a s s o c i a t i o n o c c u r r e d under shaded and s h e l t e r e d c o n d i t i o n s as w e l l as i n open and exposed s i t e s a d j a c e n t t o sreams. The  c h a r a c t e r s p e c i e s f o r t h e Hypnetum d i e c k i i i s Hypnum d i e c k i i ,  a w e s t e r n North American - Japan d i s j u n c t b r y o p h y t e t a x o n . Most commonly  131  i t was accompanied  by M a r s u p e l l a e m a r g i n a t a , a s p e c i e s which o c c u r s  c o n s i d e r a b l y l e s s f r e q u e n t l y i n the r e c o g n i z e d s u b a s s o c i a t i o n Scapanio Jungermannietosum  obovatae. Other s p e c i e s , which a r e s h a r e d by both  s y n t a x a and i n d i c a t e t h e i r syntaxonomic p o s i t i o n w i t h i n t h e  alliance  R a c o m i t r i o n a c i c u l a r i s a r e Dichodontium p e l l u c i d u m , Pel 1 i a n e e s i a n a , Rhizomniurn g l a b r e s c e n s and Brachytheciurn piumosum. However, b r y o p h y t e s p e c i e s which d i f f e r e n t i a t e t h e Hypnetum d i e c k i i - " t y p i c u m " (1 i n T a b l e X X I I I ) from the Scapanio -  Jungermannie-  tosum obovatae, and, t h u s , c h a r a c t e r i z e t h i s syntaxon a r e S c a p a n i a s u b a l p i n a , BIepharostoma t r i c h o p h y l l u m , Pogonatum u r n i g e r u m , S c l e r o podium o b t u s i f o l i u m , D i p l o p h y l l u m t a x i f o l i urn and O l i g o t r i c h u m p a r a l l e l urn.  Scapanio - Jungermannietosum  obovatae s u b a s s . nov.  O c c u r r i n g a t low e l e v a t i o n s o n l y , t h i s s u b a s s o c i a t i o n i s r e p o r t e d and documented from s t r e a m s i d e r o c k s i n Lynn Canyon, a t Shannon F a l l s and a t Brandywine  Falls.  Characteristically,  the Scapanio - Jungermannietosum  obovatae  (2 i n T a b l e X X I I I ) grows on g e n t l y s l o p i n g r o c k s u r f a c e s under  shaded  and s h e l t e r e d c o n d i t i o n s , o n l y a f f e c t e d by d i r e c t m o i s t u r e - i n p u t , through s p r a y , a t times o f h i g h e s t water l e v e l s . Only a t h i n l a y e r o f s o i l  or  sometimes no accumulated m a t e r i a l a t a l l was found on the r o c k s u r f a c e . The abundance and constancy o f Hypnum d i e c k i i i n d i c a t e t h e p r e s e n t syntaxon's a f f i n i t y t o the Hypnetum d i e c k i i - t y p i c u m . The d i f f e r e n t i a t i n g s p e c i e s , however, which c h a r a c t e r i z e t h i s  major  subassociation  Table XXIII. Hypnetum dieckii ass, nov. Number of releves Rel eve-number Elevation(m) Direction of exposure Inclination( ) Total cover(X)  1 52 61 N 22 100  Hypnum dieckii Dichodontium pelluddum Pellia neesiana Brachythecium plumosum Rhizomnium glabrescens Cephalozia bicuspidata Diplophyllum albicans Drepanocladus uncinatus Blindia acuta Philonotis fontana  4.5 1.4  Marsupella emarginata Blepharostoma trichophyllum Pogonatum urnigerum Claopodium bolanderi Diplophyllum taxifolium Scleropodium obtusifolium Scapania subalpina Oligotrichum parallelum Scapania americana Bazzania tricrenata Schistidium rivulare Nardia scalaris Mnium b l y t t i i Platydictya jungermmanioides Bartramia ithyphylla Tritomaria quinquedentata  3.4  Q  Jungermannia obovata Scapania undulata Riccardla multifida Stokesiella praelonga var. praelonga Hypnum subimponens Porotrichum blgelovii Pleuroclada albescens Hygroblella l a x i f o l l a Scouleria aquatica Geocalyx graveolens  2 53 61 N 24 100  3 56 61 E 30 90  4 130 457 SW 24 90  3.4  2.4 + .1  2.4  1.4  1.4  5 85 457 W 20 100 4.5  2.2  + .4  6 190 30 NW 78 100  7 250 1487 E 40 100  8 251 1487 E-SE 26 100  9 186 30 NW 76 90  10 189 30 W 82 80  11 12 108 184 30 30 W-NW NW 48 62 100 100  2.4  4.5 2.2  5.5 2.2 1.4 1.4  2.4 1.2 2.4  2.2 2.4  4.5 + .1 + .1 2.4  1.2  1.4  2.4 1.2  + .1  3.4 + .4 + .1  4.5  + .4 +.1  3.4 2.4  3.4  2.4  1.4  + .4  2.4 1.4 1.2 2.4 2.4 1.4  + .1  1.4  1.4  1.1  1.4  1.4 1.2 1.2  2.2  1.2  1.2 1.4 1.2 + .1  1.2 1.4  1.2 2.4  1.2 1.4 2.4  4.5 1.2 2.4 1.2 2.4  13 14 109 110 61 61 W-SW S-SW 32 36 95 100  15 209 457 E-SE 66 100  16 183 30 W 70 100  17 185 30 N-NW 34 100  2.3  3.4  3.4  2.4  1.4 3.5  2.4 1.4 1.2  2.4 2.3 2.4  2.2  1.2  1.4 2.4  1.4 2.2  2.4  2.4 1.4  2.4  2.4 2.4 4.5  2.4 2.4 2.4  2.2  1.1  2.4  2.4 2.4  1.2  1.4  1.4  +.4  18 187 457 W 58 80  19 188 457 NW 48 100  2.4 1.2  2.4 3.3 2.4  1.4  1.4 1.2  CO  ro  Table XXIII  Found o n l y once:  (continued).  Racomitrium a c i c u l a r e , P l a g i o c h i l a a s p l e n i o i d e s , Bryum p a l l e s c e n s , Jungermannia e x s e r t i f o l i a , Brachythecium f r i g i d u m , Aneura p i n g u i s , Conocephalurn conicum, A t r i c h u m s e l w y n i i , 01 i g o t r i c h u m a l i g e r u m , Racomitrium a q u a t i c u m , P o l y t r i c h u m a l p i n u m v a r . m a c o u n i i , Racomitrium f a s c i c u l a r e , Scapania s c a n d i c a , I s o p t e r y g i u m e l e g a n s , Lophocolea h e t e r o p h y l l a , D i p l o p h y l l u m p l i c a t u m , Racomitrium heterostichum var. a f f i n e , P o h l i a p r o l i g e r a , Bartramiopsis l e s c u r i i , P l a g i o t h e c i u m roeseanum, P o r e l l a cordaeana, D i c r a n e l l a h e t e r o m a l l a , B a r b i l o p h o z i a l y c o p o d i o i d e s , T o r t e l l a f r a g i l i s , P l a g i o t h e c i u m laetum, Rhytidiadelphus squarrosus.  R e l e v e - l o c a t i o n s : 52,53,56,109,110 85,130,187,188,209 108,183,184,185,186,189,190 250,251  Shannon F a l l s Brandywine F a l l s Lynn Canyon B l a c k Tusk Meadows  134  a r e Jungermannia obovata and S c a p a n i a u n d u l a t a . A d d i t o n a l  differentia-  t i n g species i n c l u d e R i c c a r d i a m u l t i f i d a , S t o k e s i e l l a praelonga praelonga,  Porotrichum  b i g e l o v i i , Pleuroclada albescens  var.  and Hypnum  subimponens.  8.3.3.2. Order PLATYHYPNIDIETALIA RUSCIFORMIS ( P h i l i p p i 1956)  A l l i a n c e Platyhypnidion rusciformis P h i l i p p i  Hygrohypnetum s m i t h i i a s s . nov.  prov.  1956  (Table XXIV).  An e v a l u a t i o n o f the s y n s y s t e m a t i c s communities" i n d i c a t e s P h i l i p p i ' s  o f the " a q u a t i c b r y o p h y t e  (1956) c l a s s i f i c a t i o n t o be  b e s t a p p l i c a b l e c l a s s i f i c a t i o n scheme o f stream a d j a c e n t  the  bryophyte  v e g e t a t i o n i n the study a r e a . T h i s system emphasizes a d i v i s i o n on b a s i s o f the h a b i t a t , r a t h e r than t h e f l o r i s t i c c o m p o s i t i o n  which shows  c o n s i d e r a b l e o v e r l a p between b r y o p h y t e assemblages w i t h s p e c i e s wide e c o l o g i c a l a m p l i t u d e The  ^.  newly d e s c r i b e d a s s o c i a t i o n Hygrohypnetum s m i t h i i  and d e s c r i b e d from s e v e r a l a l p i n e l o c a l i t i e s w i t h i n G a r b a l d i Park.  of  i s reported Provincial  I t o c c u r r e d on s o i l - c o v e r e d rocks d i r e c t l y a l o n g f a s t f l o w i n g  streams t h a t descend from l a t e snow a r e a s . The under i n f l u e n c e o f spray o r i s o c c a s i o n a l l y  substrate is constantly  inundated.  A s s o c i a t i o n s from t h i s type o f environment a r e c l a s s i f i e d Philippi  by  (1956) i n t o the a l l i a n c e P l a t y h y p n i d i o n r u s c i f o r m i s which i s  135  c o n t a i n e d i n the o r d e r P l a t y h y p n i d i e t a l i a r u s c i f o r n r i s . Both H e r t e l ( 1 9 7 4 ) , d e s c r i b i n g and p r o p o s i n g a new o c h r a c e i , and P h i l i p p i  a s s o c i a t i o n , t h e Hygrohypnetum  ( 1 9 6 5 ) , r e p o r t i n g on a Hygrohypnum ochraceum -  F i s s i d e n s c r a s s i p e s v a r . r u f i p e s - community, c l a s s i f y t h e s e s y n t a x a i n g the a l l i a n c e P l a t y h y p n i d i o n r u s c i f o r m i s P h i l i p p i However, G e i s s l e r ( 1 9 7 6 ) , employing c a t i o n , c l a s s i f i e s a newly proposed  .  K r a j i n a ' s (1933) c l a s s i f i -  a s s o c i a t i o n , t h e Solenostomo -  Hygrohypnetum, i n t o the a l l i a n c e Hygrohypniori which  1956  dilatati  Krajina  1933,  i s c o n t a i n e d i n the o r d e r H y g r o h y p n e t a l i a d i l a t a t i  Krajina  1933.  I t i s t h i s a s s o c i a t i o n which  shows c o n s i d e r a b l e f l o r i s t i c  (and  ecological)  a f f i n i t i e s t o t h e a s s o c i a t i o n Hygrohypnetum s m i t h i i , as d e s c r i b e d i n t h i s s t u d y . The c h a r a c t e r s p e c i e s o f G e i s s l e r ' s (1976) Solenostomo Hygrohypnetum a r e Solenostoma c o r d i f o l i u m (=Jungermannia c o r d i f o l i a ) , Scapania u n d u l a t a and Hygrohypnum s m i t h i i . The a s s o c i a t i o n c h a r a c t e r s p e c i e s o f t h e Hygrohypnetum s m i t h i i i n the p r e s e n t study a r e Hygrohypnum s m i t h i i , H_. ochraceum, Jungermannia e x s e r t i f o l i a . Constant companions were C h i l o s c y p h u s p o l y a n t h o s  and  Pel 1i a n e e s i a n a . The f o l l o w i n g bryophyte t a x a i n d i c a t e a d d i t o n a l a f f i n i t i e s , w i t h r e s p e c t t o f l o r i s t i c c o m p o s i t i o n , s p e c i e s ' abundance and  frequency,  between the p r e s e n t a s s o c i a t i o n Hygrohypnetum s m i t h i i and the Solenostomo Hygrohypnetum, as d e s c r i b e d by G e i s s l e r (1976): Scapania  undulata,  S c h i s t i d i u m r i v u l a r e , P h i l o n o t i s f o n t a n a , Jungermannia o b o v a t a , dontium pel 1ucidum and Bryum  Dicho-  pseudotriquetrum.  F u r t h e r r e s e a r c h , e s p e c i a l l y from s i m i l a r h a b i t a t s a t s u b a l p i n e  T a b l e XXIV. Hygrohypnetum s m i t h i i  a s s . nov.  Number o f r e l e v e s Releve-number Elevation(m) D i r e c t i o n ofgexposure Inclination( ) Total cover(%)  1 232 1487 SW 18 100  2 260 1492 E 56 100  3 234 1487 NW 38 100  4 327 2286 N-NE 68 100  5 328 2286 NE 84 100  6 233 1487 NW 22 100  Hygrohypnum s m i t h i i Chiloscyphus polyanthos Jungermannia e x s e r t i f o l i a Pel 1 i a n e e s i a n a Scapania subalpina Hygrohypnum ochraceum Blepharostoma t r i c h o p h y l l u m Scouleria aquatica Scapania undulata Bryum p a l l e s c e n s P h i l o n o t i s fontana Dichodontium p e l l u c i d u m Jungermannia obovata Cephalozia bicuspidata Brachythecium plumosum Hygrobiella l a x i f o l i a Jungermannia pumila Bryum p s e u d o t r i q u e t r u m Conocephalum conicum Drepanocladus u n c i n a t u s T r i t o m a r i a quinquedentata Bartramia i t h y p h y l l a Rhizomnium nudum  3.4 4.5 2.4 1.4  4.5 3.4 2.4 1.4 + .4 2.4  2.4 2.4 2.4 1 .4 1 .4 3.4 1.4  4.5 1.4 + .1 1.4 + .4  4.5 1.4 2.4 + .1 + .4  2.4 1 .4  1.4 + .1 1 .4  1.2  1.2 2.2  2.4 1.4 1.4 1.4 3.4 2.4 1.4 1.4 1.4 2.2 2.2  R e l e v e - l o c a t i o n : 232, 233, 234 B l a c k Tusk Meadows 260 Panorama Ridge 327, 328 W h i s t l e r Mountain  2.4 2.4 1 .4 3.3  1.4 2.2 1.2, 1 .2  1.4  1.4 1.4 1.4  1.4 1 .4 + .4 + .4 + .1  + .1 1.4 1.2  1.1 2.4 4.4 1.2 2.2  137  and a l p i n e v e g e t a t i o n s might shed more l i g h t on t h e i d e n t i t y o f t h i s v e g e t a t i o n type and i t s r e l a t i o n s h i p t o bryophyte  assemblages i n  s i m i l a r h a b i t a t s i n western North America and t o European s y n t a x a i n the same environment.  1. i . e . w i t h i n t h e R a c o m i t r i o n a c i c u l a r i s : D i c h o d o n t i e t u m p e l l u c i d i (Hubschmann 1967, Norr 1970, Neumayr 1971, M a r s t a l l e r 1973), B r a c h y t h e c i e t u m plumosi ( K r u s e n s t j e r n a 1945, Norr 1970, H e r t e l 1974, M a r s t a l l e r 1973, 1980), R a c o m i t r i e t u m a c i c u l a r i s ( P h i l i p p i 1965, Neumayr 1971). The f i r s t a s s o c i a t i o n was a l s o r e p o r t e d from Vancouver I s l a n d by Hubschmann (1978). W i t h i n t h e Scapanion undulatae: Scapanietum undulatae ( S c h w i c k e n r a t h 1944, P h i l i p p i 1956, Norr 1969, L e c o i n t e and P r o v o s t 1970, Dunk 1971, M a r s t a l l e r 1980). 2. Hubschmann ( 1 9 7 8 ) , s t u d y i n g t h e b r y o p h y t e v e g e t a t i o n on Vancouver I s l a n d , r e p o r t s and d e s c r i b e s t h e S c o u l e r i e t u m a q u a t i c a e i n which S c o u l e r i a a q u a t i c a i s t h e a s s o c i a t i o n c h a r a c t e r s p e c i e s and S c l e r o podium o b t u s i f o l i u m i s d e s i g n a t e d as t h e a l l i a n c e c h a r a c t e r s p e c i e s for the (provisional) Scleropodion o b t u s i f o l i a e . 3. Releves 66, 129, 274, 159. 235 and 236 ( C l u s t e r 2 ) , a l t h o u g h s t a n d i n g as an apparent u n i t i n t h e c l u s t e r a n a l y s i s (Appendix V ) , i s c o n s i d e r e d as a "fragment" o f t h e R a c o m i t r i o - S c o u l e r i e t u m a q u a t i c a e . T h i s i s because o f t h e t o t a l absence o f S c o u l e r i a a q u a t i c a and t h e lower frequency and c o n s t a n c y o f Brachytheciurn plumosum. F u r t h e r r e s e a r c h may s u b s t a n t i a t e i t s a f f i n i t i e s w i t h t h e a s s o c i a t i o n R a c o m i t r i e t u m a c i c u l a r i s P h i l i p p i 1965  4. S t r a s s e r (1971) and Mamczarz (1978) r e p o r t on an a s s o c i a t i o n , P e l l i o Conocephaletum c o n i c a e , synonymous t o t h e F e g a t e l l e t u m c o n i c a e Schade 1934. I n t h e i r a s s o c i a t i o n e i t h e r P e l l i a f a b b r o n i a n a o r Conocephalurn conicum a r e t h e dominant s p e c i e s . The companion s p e c i e s a r e c h a r a c t e r i s t i c f o r both s y n t a x a : Mnium punctatum, F i s s i d e n s a d i a n t h o i d e s and P l a g i o c h i l a a s p l e n i o i d e s . 5. A l s o H e r t e l (1974) d i s c u s s e s t h e s t a t u s o f t h i s syntaxon and s t a t e s t h a t t h e " F e g a t e l l e t u m " does not deserve t h e rank o f a s s o c i a t i o n .  138  Conocephalum conicum o c c u r s a f t e r the d e p o s i t i o n o f some m i n e r a l s o i l as a r e s u l t o f submergence. Because o f i t s r a p i d and v i g o r o u s growth the s p e c i e s can outcompete p l e u r o c a r p o u s mosses (Brachyt h e c i u m spp. and S t o k e s i e l l a s p p . ) . E v e n t u a l l y i t can become dominant and develop i n t o a "one-species-community" (Neumayr 1971). H e r t e l (1974) f u r t h e r s t a t e s t h a t Conocephalum conicum can become ' f a c i e s ' - b u i l d i n g w i t h i n s e v e r a l communities which proove the s p e c i e s n e i t h e r t o be f a i t h f u l nor t o be c o n s t a n t . Another b a s i c r e q u i r e m e n t t o a c h a r a c t e r s p e c i e s i s i t s confinement t o a narrow e c o l o g i c a l a m p l i t u d e . In o b v i o u s c o n t r a s t H e r t e l (1974) r e p o r t s Conocephalum conicum d o m i n a t i n g on humid t o wet s o i l , rock bases a d j a c e n t and d i r e c t above the water s u r f a c e . The s p e c i e s i s , a c c o r d i n g t o Boros (1968) i n d i f f e r e n t ; " A p i n i s (1939) r e p o r t s on a pH-range between s i t e s o f t h i s s p e c i e s ' o c c u r r e n c e from 5.0 t o 8.5. Hence, f o l l o w i n g the f l o r i s t i c a l method such v e g e t a t i o n s , dominated by one, e c o l o g i c a l l y u b i q u i t o u s s p e c i e s , s h o u l d be d e s i g n a t e d as 'facies'.  6. Under e x t e n s i v e spray-zone c o n d i t i o n s M u l l e r (1938) d e s c r i b e d an a s s o c i a t i o n Scapanietum paludosae i n w h i c h , b e s i d e s v a s c u l a r p l a n t s p e c i e s , the f o l l o w i n g mosses o c c u r r e d : P h i l o n o t i s s e r i a t a , Drepanoc l a d u s e x a n u l a t u s and D i c r a n e l l a s q u a r r o s a . 7. K r a j i n a (1933) e r e c t e d the o r d e r H y g r o h y p n e t a l i a which was t o c o n t a i n b r y o p h y t e communities from a l p i n e s i t e s Where streams o r i g i n a t e and from the banks o f the streams. However, as H e r t e l (1974) r i g h t l y s t a t e s , t h i s o r d e r i n c l u d e s groups o f a s s o c i a t i o n s o c c u r r i n g on c o n s t a n t l y i n u n d a t e d as w e l l as l o n g - l a s t i n g emerged s i t e s , a s s o c i a t i o n s from c a l c i u m - r i c h w a t e r s and streams o v e r s i l i c e o u s r o c k s . T h i s c l a s s i f i c a t i o n was a l s o i n c l u d e d i n Hubschmann's (1957) o v e r a l l s y n s y s t e m a t i c c l a s s i f i c a t i o n o f the " a q u a t i c b r y o p h y t e communities". His order F o n t i n a l e t a l i a a n t i p y r e t i c a e contains a l l i a n c e s of longl a s t i n g emerged and almost c o n s t a n t l y submerged b r y o p h y t e a s s o c i a t i o n s . F i n a l l y , Hubschmann (1957) proposes two c l a s s e s , between which o b v i o u s r e l a t i o n s e x i s t ( i . e . the a l l i a n c e s F o n t i n a l i o n and Scapanion both i n c l u d e o n l y o c c a s i o n a l l y emerged b r y o p h y t e a s s o c i a t i o n s , y e t are c l a s s i f i e d i n two d i f f e r e n t c l a s s e s ) .  8. F u r t h e r r e p o r t s on Hygrohypnum-dominated a s s o c i a t i o n s and subassociat i o n s are p r o v i d e d by Herzog ( 1 9 4 3 ) , P o e l t ( 1 9 5 4 ) , Hagel ( 1 9 6 6 ) , Neumayr ( 1 9 7 1 ) , P h i l i p p i ( 1 9 7 3 ) , Hubschmann (1973, 1974), G e i s s l e r ( 1 9 7 6 ) , Mamczarz (1978) and M a r s t a l l e r (1980).  139  8.4  BRYOPHYTE ASSOCIATIONS FROM CALCAREOUS ROCKS.  T a b l e XXV.  B r y o p h y t e a s s o c i a t i o n s from c a l c a r e o u s  A l l i a n c e ENCALYPTION PROCERAE a l l .  rocks.  nov.  C l u s t e r 1.  A s s o c i a t i o n Lophozio - B r y o e r y t h r o p h y l l e t u m r e c u r v i r o s t r i a s s .  C l u s t e r 2.  A s s o c i a t i o n Metaneckeretum m e n z i e s i i ass.  nov.  nov.  B a r b u l a v i n e a l i s - P I a t y d i c t y a j u n g e r m a n n i o i d e s - Jungermannia a t r o v i r e n s - community  C l u s t e r 3A.  "inside-grotto-type" with Distichium capillaceum  C l u s t e r 3B.  "outside-grotto-type"  and  w i t h S c h i s t i d i u m r i v u l a r e and  Fissidens bryoides  Bryum p s e u d o t r i q u e t r u m  141  8.4.  Bryophyte a s s o c i a t i o n s from c a l c a r e o u s r o c k s ( T a b l e  XXV).  8.4.1. S y n s y s t e m a t i c s .  The s u b s t r a t e s p e c i f i c i t y i s the most s i g n i f i c a n t f a c t o r i n d e l i m i t i n g the bryophyte assemblages van " c a l c a r e o u s r o c k s ^. ;  d e c i s i v e d i s t i n c t i o n between c a l c a r e o u s and s i l i c e o u s rock  The  bryophyte  communities f o c u s e s on t h e chemical c o m p o s i t i o n o f t h e rock s u r f a c e and/or t h e t h i n l a y e r o f d i s i n t e g r a t e d rock m a t e r i a l accumulated  om  t h a t s u r f a c e ( W i r t h 1972, H e r t e l 1974). C o n c e r n i n g the n a t u r e o f b r y o p h y t e communities on c a l c a r e o u s rocks i t i s e s s e n t i a l t o r e a l i z e t h a t t h e s e are r e f e r r e d t o as combin a t i o n s o f s p e c i e s which o c c u r p r e d o m i n a n t l y on l i m e s t o n e and show t h e i r o p t i m a l development on t h a t s u b s t r a t u m . When bryophyte  assemblages  a r e d e s c r i b e d a t t h e b o r d e r a r e a o r even o u t s i d e t h e i r e c o l o g i c a l range, fragments  o r l o c a l r e p r e s e n t a t i o n s can r e c e i v e a s s o c i a t i o n  These "communities"  are then not r e c o g n i z e d a c c o r d i n g t o the  rank.  Braun-  B l a n q u e t f l o r i s i c - s b c i o l o g i c a l method, but are based merely on dominance o f s e p a r a t e s p e c i e s . The t a x a become a s s o c i a t i o n c h a r a c t e r s p e c i e s a l t h o u g h most o f the time they are not f a i t h f u l  to the "community".  A s s o c i a t i o n s n e i t h e r can be based nor d e l i m i t e d on a s i n g l e dominant species. A f t e r an e x t e n s i v e d i s c u s s i o n , o f c l a s s i f i c a t i o n attempts  and  p r o p o s a l s by v a r i o u s b r y o - s o c i o l o g i s t s , H e r t e l (1974) p r o v i d e s a s a t i s f y i n g s y n s y s t e m a t i c h i e r a r c h y f o r t h e b r y o p h y t e communities on  142  c a l c a r e o u s rock s u b s t r a t a . Based on t h e f l o r i s t i c c o m p o s i t i o n o f t h e a s s o c i a t i o n s and supported by h a b i t a t c h a r a c t e r i s t i c s t h e a u t h o r t i n g u i s h e s (1) an o r d e r from exposed, dry r o c k s [ S c h i t i d i e t a l i a (Jezek and Vondracek 1962)  H e r t e l 1974],  disapocarpi  (2) an o r d e r from s h e l t e r e d ,  humid r o c k s ( N e c k e r e t a l i a complanatae Jezek and Vondracek 1962 an o r d e r from more o r l e s s s h e l t e r e d r o c k s and/or s o i l m o l l u s c i Hadac and Smarda 1944). S u b s e q u e n t l y ,  and  (3)  (Ctenidietalia  the a s s o c i a t i o n s w i t h i n  these o r d e r s a r e c l a s s i f i e d i n t o t h e a l l i a n c e s S c h i s t i d i o n a p o c a r p i Jezek and Vondracek 1962, N e c k e r i o n complanatae Hadac and Smarda  1944  2 and C t e n i d i o n m o l l u s c i S t e f u r e a c 1941,  The  p r e s e n t data show apparent  respectively  .  f l o r i s t i c and e c o l o g i c a l  affini-  t i e s t o a s s o c i a t i o n s d e s c r i b e d by b r y o - s o c i o l o g i s t s i n Europe and, p a r t i c u l a r l y , c l a s s i f i e d i n i l l u s t r a t i v e and r e v e a l i n g t r e a t m e n t s Hertel  by  (1974) and M a r s t a l l e r (1979, 1980). However, r e l a t i o n s h i p s a t  t h e l e v e l o f a l l i a n c e and o r d e r were not d e t e c t a b l e . In a d d i t i o n , t h e various t a x a , confined i n t h e i r d i s t r i b u t i o n to western  North A m e r i c a ,  and  t h e r e l a t i v e l y low number o f r e l e v e s per r e c o g n i z e d a s s o c i a t i o n , have c o n t r i b u t e d t o the p r e s e n t t r e a t m e n t o f t h e d a t a . Where p o s s i b l y supported  w i t h c h a r a c t e r s p e c i e s , new  s y n t a x a a r e proposed and  affinities  w i t h v e g e t a t i o n s i n o t h e r r e g i o n s i n t h e n o r t h e r n Hemisphere a r e A new  indicated.  a l l i a n c e i s proposed, the E n c a l y p t i o n procerae, best  c h a r a c t e r i z e d by E n c a l y p t a p r o c e r a , Homalothecium f u l g e s c e n s and  Porella  cordaeana. W i t h i n t h i s a l l i a n c e two a s s o c i a t i o n s have been d e s c r i b e d : (A) a Lophozio  - Bryoerythrophylletum r e c u r v i r o s t r i w i t h the c h a r a c t e r  143  species Bryoerythrophyl1um  r e c u r v i r o s t r u m , Lophozia  heterocolpos  and Gymnostomum r e c u r v i r o s t r u m and (B) a Metaneckeretum m e n z i e s i i w i t h t h e c h a r a c t e r s p e c i e s Metaneckera m e n z i e s i i . F i n a l l y , a t h i r d " v e g e t a t i o n t y p e " i s d e s c r i b e d , which i s c h a r a c t e r i z e d by s p e c i e s such as B a r b u l a v i n e a l i s , P l a t y d i c t y a j u n g e r mannioides and Jungermannia a t r o v i r e n s . However, t h e r e l e v e s o f t h i s 'community' l a c k c o n s t a n t (and abundant) and f a i t h f u l  s p e c i e s , hence,  cannot be g i v e n any syntaxonomic s t a t u s and be f u r t h e r c l a s s i f i e d .  8.4.2. P h y t o g e o g r a p h i c a l  setting.  The c o n t r i b u t i o n o f t h e c i r c u m b o r e a l element t o t h e f l o r i s t i c c o m p o s i t i o n o f t h e l i m e s t o n e r e l e v e s i s e x p r e s s e d by a s i g n i f i c a n t number o f s p e c i e s . Among t h e h e p a t i c s a r e Conocephalum conicum, Jungermannia a t r o v i r e n s , Lophozia h e t e r o c o l p o s and P l a g i o c h i l a des. The mosses B r y o e r y t h r o p h y l ! u m  asplenioi-  recurvirostrum, Plagiopus oederi  and t o a l e s s e r degree F i s s i d e n s b r y o i d e s , Gymnostomum r e c u r v i r o s t r u m and D i s t i c h i u m capi11aceum a l s o dominate t h e c i r c u m b o r e a l a s p e c t o f t h e bryophyte  v e g e t a t i o n on c a l c a r e o u s r o c k s i n t h e study a r e a .  A predominantly' 1imestone h e p a t i c , c i r c u m b o r e a l i n i t s d i s t r i b u t i o n , b u t m i s s i n g i n e a s t e r n A s i a , i s Lophozia  gillmanii.  Except f o r Homalothecium f u l g e s c e n s , which i s d e s i g n a t e d as one o f t h e c h a r a c t e r s p e c i e s f o r t h e a l l i a n c e E n c a l y p t i o n p r o c e r a e , t h e c o n t r i b u t i o n o f western stone bryophyte  North American endemics t o t h e o v e r a l l  lime-  v e g e t a t i o n i s 'expressed by l e s s f r e q u e n t companions.  144  Among t h e s e s p e c i e s a r e P o r e l l a r o e l l i i , B r a c h y t h e c i u m f r i q i d u m , Isothecium  s t o l o n i f e r u m and t o a l e s s e r e x t e n t L e u c o l e p i s m e n z i e s i i ,  Plagiomnium venustum, Claopodium b o l a n d e r i and P o r o t r i c h u m The  d i s j u n c t phytogeographical  bigelovii.  elements p l a y o n l y minor r o l e s  i n t h e i r c o n t r i b u t i o n t o t h e f l o r a and v e g e t a t i o n o f t h e c a l c a r e o u s r o c k s i n t h e study a r e a . The w e s t e r n North American - w e s t e r n Europe d i s j u n c t h e p a t i c , P o r e l l a cordaeana, and t h e w e s t e r n North American Japan d i s j u n c t moss, Hypnum subimponens, a r e f r e q u e n t companion s p e c i e s i n t h e a l l i a n c e E n c a l y p t i o n p r o c e r a e . B i p o l a r d i s j u n c t bryophytes among t h e l i m e s t o n e f l o r a here i n c l u d e s p e c i e s such as D i t r i c h u m f l e x i c a u l e and Brachythecium v e l t i t i n u m . -  8.4.3. A l l i a n c e E n c a l y p t i o n p r o c e r a e  Lophozio  - Bryoerythrophylletum  a l l . nov.  r e c u r v i r o s t r i a s s . nov. (Table  T h i s a s s o c i a t i o n i s d e s c r i b e d from exposed c a l c a r e o u s c h a r a c t e r i s t i c a l l y covered by a s u b s t a n t i a l s o i l s/lope  rocks,  l a y e r . Depending on t h e  o f t h e rock s u r f a c e where t h e sampling was done, t h i s  accumulation  XXVI).  soil  was measured from 0.5 t o 2 cm i n t h i c k n e s s .  The a s s o c i a t i o n ' s c h a r a c t e r s p e c i e s a r e B r y o e r y t h r o p h y l 1 urn r e c u r v i r o s t r u m and Lophozia h e t e r o c o l p o s . A l l i a n c e c h a r a c t e r s p e c i e s which a r e f r e q u e n t l y , and sometimes almost c o n s t a n t l y , a s s o c i a t e d w i t h t h e s e two t a x a a r e EncaTypta p r o c e r a and Homalothecium f u l g e s c e n s . A d d i t i o n a l s i g n i f i c a n t a s s o c i a t e s a r e P l a g i o p u s o e d e r i and Conocephalurn  Table XXVI. Lophozio-Bryoerythrophylletum  recurvirostri  Number o f r e l e v e s Releve-number Elevation(m) D i r e c t i o n ofgexposure Inclination( ) Total cover(%)  1 36 266 W 58 100  2 163 266 W-NW 38 100  Bryoerythrophyllum recurvirostrum Encalypta procera Plagiopus oederi Homalothecium f u l g e s c e n s Conocephalurn conicum Lophozia heterocolpos Racomitrium h e t e r o s t i c h u m var. heterostichum Hypnum subimponens P o r e l l a cordaeana Ditrichum f l e x i c a u l e Gymnostomum r e c u r v i r o s t r u m Brachythecium f r i g i d u m Mm* urn thorns on i i Geocalyx g r a v e d ens Barbula r e f l e x a S e l i g e r i a donniana Jungermannia a t r o v i r e n s  2,.3 2,.2 2..2  4 .3 3,.2  1,.4  3 164 258 W-NW 38 100  a s s . nov.  4 165 240 W-NW 36 100  5 153 266 W 39 100  6 166 266 N 74 60  7 167 258 NE 60 90  8 168 258 NE 68 75  2,.3 4,.3 2,.2 2..4 1,.4 3..4  2,.3 2,.2 2,.2 2,.4  2.3 1.2 2.2 2.4 1.4  2.3 3.2 1 .2 2.4  2.3  1,.4 2,.4 2,.4  3.3 3.2 2.2 2.4 3.4 2.4  1 .2 , 1 .4 , 2..4  1.2 1.4 2.4  1,.2 1,.4  3..3 1..4 3,.4 2.2 3.2 2.4 2.2  3.3 4.3 2.4  + ,.1  2.4 2.2 1.1 3.,4  1 .2 1.4 1.4 2.4  2.2 3.2 1 .4 1 .2 1.4 1.2 2.1 1.4  Found o n l y once: Gymnostomum aeruginosum, P l a g i o c h i l a a s p l e n i o i d e s , Dichodontium p e l l u c i d u m , Claopodium b o l a n d e r i , Bryum c a p i l l a r e , Oxystegus t e n u i r o s t r i s , Cratoneuron commutatum v a r . f a l c a t u m , T r i t o m a r i a e x s e c t i f o r m i s , Thamnobryum n e c k e r o i d e s , Lophozia g i l l m a n i i , C e p h a l o z i a b i c u s p i d a t a , S c h i s t i d i u m apocarpum, P o r e l l a n a v i c u l a r i s , Dryptodon patens. R e l e v e - l o c a t i o n : 166, 167, 168 S l e s s e Creek; o t h e r s : C h i l l i w a c k R i v e r V a l l e y .  146  conicum. O c c a s i o n a l l y , t h e p r e d o m i n a n t l y  "limestone bryophytes"  S e l i g e r i a donniana and Jungermannia a t r o v i r e n s , o c c u r w i t h i n t h e p r e s e n t association. The Lophozio - B r y o e r y t h r o p h y l ! e t u m r e c u r v i r o s t r i  occurs  w i t h i n t h e p r e s e n t d a t a s e t w i t h two " t y p e s " , o f which one o c c u r s on r o c k s , h a r d l y covered by s o i l m a t e r i a l , and i s c h a r a c t e r i z e d by s p e c i e s such as Racomitrium h e t e r o s t i c h u m v a r . a f f i n e and Hypnum subimponens. The second soil  " t y p e " i s a group o f s p e c i e s found on r o c k s w i t h e x t e n s i v e  accumulation. This combination o f s p e c i e s contains D i t r i c h u m  f l e x i c a u l e , Gymnostomum r e c u r v i r o s t r u m , Brachythecium  f r i g i d u m , Mnium  t h o m s o n i i , Geocalyx g r a v e o l e n s and B a r b u l a r e f l e x a . However, t h e p r e s e n t data f a i l  to indicate neither the bryosociological  syntaxonomic  s i g n i f i c a n c e nor the  r e l a t i o n of the vegetation "types" t o the synsystematic  t r e a t m e n t s o f c a l c a r e o u s rock b r y o p h y t e communities  of Hertel  (1974)  and M a r s t a l l e r (1979, 1980) . 3  Metaneckeretum m e n z i e s i i a s s . nov. ( T a b l e X X V I I ) .  The p r e s e n t syntaxon has been d e s c r i b e d from s h e l t e r e d and shaded l i m e s t o n e b o u l d e r s i n s i d e f o r e s t s on t h e s l o p e s o f t h e C h i l l i wack R i v e r V a l l e y near SI esse Creek adn Chipmunk Creek. Except f o r a few o c c a s i o n s , most o f t h e rock s u r f a c e s d i d not have much s o i l rial of  accumulated  these  logical  mate-  on them. T h i s , o b v i o u s l y , depends on t h e i n c l i n a t i o n  boulders,which i s i n d i c a t e d i n the a s s o c i a t i o n ' s phytosociotable.  Table, XXVII. Metaneckeretum m e n z i e s i i a s s . nov. Number o f r e l e v e s Releve-number Elevation(m) D i r e c t i o n ofgexposure Inclination( ) Total cover(%)  1 143 266 NW 64 100  2 144 266 NW 62 100  3 149 258 W-NW 82 100  4 145 230 NW 58 100  5 148 258 NW 58 100  6 150 258 SW 84 100  7 152 230 SW 64 100  Encalypta procera Metaneckera m e n z i e s i i Plagiopus oederi Homalothecium f u l g e s c e n s Eurhynchium p u l c h e l l u m Hypnum subimponens Isothecium s t o l o n i f e r u m Dichodontium p e l l u c i d u m Claopodium b o l a n d e r i Porotrichum b i g e l o v i i Gymnostomum aeruginosum Barbula v i n e a l i s P l a t y d i c t y a jungermmanioides P o h l i a cruda Blepharostoma t r i c h o p h y l 1 u m Plagiomnium venustum Porella r o e l l i i Claopodium c r i s p i l u m Bryoerythrophyllum recurvirostrum Mnium ambiguum Apometzgeria pubescens Barbula r e f l e x a Plagiomnium r o s t r a t u m  2.2 3.4  2.2  1.2 3.4 2.2 4.4 2.4  1.2 3.4 2.2 1.4  2.2 3.4 2.2 1.4  2.2  2.2  2.4  2.4  2.2  1.4 2.4 2.4  2.4 2.2  2.4 1.2 4.3  2.4 3.2  1.4 +.4 1.4 1.4 3.2 2.4  9 321 266 W 82 100  10 324 266 W 60 100  11 322 268 NW 78 100  12 323 268 W-NW 76 100  1.2 1.4 3.2 1.4  1 .2 3.4 1.2 1.4  + .2 1.4 1.2 1.4  1.2 1.4 2.2 1.4  + .4  +.4 2.2  1.2 +.2 2.4  2.4 4.3  8 151 • 230 SW 64 90  2.2 2.4 1.2  2.2 1.4 4.3 4.4 3.4 2.4  1.4 2.4 1.2 + .2 1.2  2.4 2,. 4 2.2 1.2 1.4 1 .2  + .4 1.4 3.2 1 .2  + .4 2.4 3.3 1 .2 2.4  1.2  1.2  T a b l e XXVII  Found o n l y once:  (continued).  Homalothecium n u t a l l i i , P l a g i o c h i l a a s p l e n i o i d e s , Racomitrium h e t e r o s t i c h u m v a r . a f f i n e , Bryum c a p i l l a r e , Conocephalum conicum, Mnium t h o m s o n i i , P o r e l l a cordaeana, I s o p t e r y g i u m e l e g a n s , S c h i s t i d i u m s t r i c t u m , Scapania americana, S t o k e s i e l l a p r a e l o n g a v a r . p r a e l o n g a , Timmia a u s t r i a c a  R e l e v e - l o c a t i o n s : 143,144,145,148,149  Chilliwack River Valley  150,151,152  Chipmunk Creek  321,322,323,324  S l e s s e Creek  149  The  a s s o c i a t i o n ' s c h a r a c t e r s p e c i e s i s Metaneckera m e n z i e s i i ,  which i s f r e q u e n t l y a s s o c i a t e d w i t h the a l l i a n c e c h a r a c t e r s p e c i e s E n c a l y p t a p r o c e r a , and Homalothecium f u l g e s c e n s . Other s i g n i f i c a n t accompanying s p e c i e s a r e P l a g i o p u s o e d e r i , Eurhynchium p u l c h e l 1 urn and Hypnum subimponens. W i t h i n the Metaneckeretum m e n z i e s i i two s u b a s s o c i a t i o n s have been d i s t i n g u i s h e d , based on s p e c i e s c o m p o s i t i o n The  and h a b i t a t d i f f e r e n c e s .  s u b a s s o c i a t i o n s Gymnostometosum aeruginosum o c c u r s on s t e e p l y  s l o p i n g , dry rocks s u r f a c e s ( e s p e c i a l l y t h e i r lower p o r t i o n s ) and i s c h a r a c t e r i z e d by the f o l l o w i n g d i f f e r e n t i a t e d s p e c i e s : Gymnostomum aeruginosum, B a r b u l a v i n e a l i s and P l a t y d i c t y a j u n g e r m a n n i o i d e s . s u b a s s o c i a t i o n , the P o r e l l e t o s u m r o e l l i i  A second  grows on g e n t l y as w e l l as  s t e e p l y s l o p i n g , humid r o c k s u r f a c e s where i t s l u s h v e g e t a t i o n i s c h a r a c t e r i z e d by the d i f f e r e n t i a t i n g s p e c i e s Pore!1 a r o e ! I i i , Claopodium c r i s p i f o l i u m , Bryoerythrophy!1 urn r e c u r v i r o s t r u m and Mnium ambiguum. Although  f l o r i s t i c and e c o l o g i c a l a f f i n i t i e s are apparent  between the p r e s e n t a s s o c i a t i o n Metaneckeretum m e n z i e s i i and  European  b r y o p h y t e s y n t a x a , the p r e s e n t data do not a l l o w i n t e g r a t i o n i n t h e s y n s y s t e m a t i c h i e r a r c h y o f b r y o p h y t e communities on c a l c a r e o u s r o c k s  as  c l a s s i f i e d by H e r t e l . (1974) and M a r s t a l l e r (1979, 1980), i n p a r t i c u l a r  4  .  F i n a l l y , a t h i r d t y p e o f b r y o p h y t e assemblage has been d e s c r i b e d : the B a r b u l a v i n e a l i s - P l a t y d i c t y a jungermannioides  - Jungermannia  a t r o v i r e n s - community (Table X X V I I I ) . T h i s c h a r a c t e r i s t i c  combination  o f s p e c i e s (extended w i t h the t h a l l o s e h e p a t i c Conocephalurn conicum)  T a b l e XXVIII B a r b u l a v i n e a l i s - P l a t y d i c t y a j u n g e r m a n n i o i d e s - Jungermannia Number o f r e l e v e s Rel eve-number El evation(m) D i r e c t i o n ofgexposure Inclination( ) Total cover(%)  1 146 266 NW no 40  2 147 266 NW 96 60  3 280 258 NW 60 50  Conocephalum conicum Barbula v i n e a l i s Leucolepis menziesii P l a t y d i c t y a jungermannioides  1 .4  1 .4  1.4 3.2 1 .2  Porel1 a r o e l I i i Schistidium rivulare Bryum p s e u d o t r i q u e t r u m Plagiochila asplenioides Racomitrium canescens B r y o e r y t h r o p h y l 1um r e c u r v i r o s t r u m Plagiopus oederi Plagiomnium r o s t r a t u m Ditrichum f l e x i c a u l e Gymnostomum aeruginosum Brachythecium r i v u l a r e Homalothecium f u l g e s c e n s Brachythecium velutinum Jungermannia a t r o v i r e n s Distichium capillaceum Fissidens bryoides Lophozia g i l l m a n i i S e l i g e r i a donniana Timmia a u s t r i a c a Tortula princeps Apometzgeria pubescens Gymnostomum r e c u r v i r o s t r u m  a t r o v i r e n s - community.  48 60  5 282 258 N-NW 52 50  6 277 240 NE 58 90  7 278 246 E 66 100  8 279 246 NW 54 90  9 283 262 M-NE 54 95  10 284 268 NE 42 100  11 285 250 N 18 100  1.4 1.2 1 .2  2.4 1 .2 + .2  2.4 1.2  3.4 + .2  3.4 1.2  1.4 1 .2  1.4  1.4  1.4 1 .2 3.3 2.4  + .2 2.2 1.4  1 .4 2.2 1.2 2.4 3.3 2.2 2.2  1 .4 1 .2 1 .2 1.4 1 .2 1 .2 1 .2  1 .2 1 .2  1 + 1 1  4 281 258 N-NW  2.4  2  .  1  1 .4 1 .2  1.4 2.2  1 .4 2.2 1.2 1.2  2.4 1.2 1.2 1.2  1.4 + .2 2.2  1 .4  2.4 1 .2  1.4 1.2  1.2  1.4 + .2 4.2  + .4  .2 .2 .4 .4  1.4 2.4 1 .2 + .2 1.4 1.2 1 .2 1 .2 1 .2  1.4 + .2 1.4 1 .2 2.2 1.2 1.2 + .2  1 .4 1 .2 4.3  3.2  + .2 1 .2  + .4 1.4  1.4 1 .4 1.4  Table XXVIII  Only found once:  (continued).  Dichodontium p e l l u c i d u m , C e p h a l o z i a b i c u s p i d a t a , Scapania u n d u l a t a , Mnium t h o m s o n i i .  R e l e v e - l o c a t i o n s : 146,147,283,284,285 280,281,282,277,278,279  S l e s s e Creek Chipmunk Creek  152  predominantly  o c c u r s under the very shaded and s h e l t e r e d c o n d i t i o n s o f  " r o c k - g r o t t o s " and t h e immediately  a d j a c e n t r o c k bases o u t s i d e  these  grottos. C o n f i n e d t o the dry i n t e r i o r o f the r o c k g r o t t o s was wing c o m b i n a t i o n  the  follo-  of species: Distichium capillaceum, Fissidens bryoides,  Timtnia a u s t r i a c a , T o r t u l a p r i n c e p s , Apometzgeria pubescens and Gymnostomum r e c u r v i r o s t r u m (1 i n T a b l e X X V I I I ) . The  humid t o moist  rock  bases d i r e c t l y o u t s i d e t h e s e g r o t t o s showed a more d i v e r s e and 1 u x u r i a n t v e g e t a t i o n (2 i n T a b l e X X V I I I ) . S p e c i e s  confined ( w i t h i n the  limestone  r e l e v e s ) t o t h i s type o f h a b i t a t were S c h i s t i d i u m r i v u l a r e , Bryum pseudotriquetrum,  Racomitrium c a n e s c e n s , Brachytheciurn r i v u l a r e , B_.  v e l u t i n u m and Plagiomnium  rostratum.  The a f f i n i t i e s between t h i s t h i r d community ( e s p e c i a l l y group o f s p e c i e s o u t s i d e the rock g r o t t o s ) and the a l l i a n c e p r o c e r a e are expressed  the  Encalyption  by s p e c i e s such as B r y o e r y t h r o p h y l l u m  recurvi-  rostrum, Plagiopus o e d e r i , Ditrichum f l e x i c a u l e , P o r e l l a r o e l l i i  and  Plagiochila asplenioides.  F u r t h e r r e s e a r c h i s needed i n o r d e r t o d i s t i n g u i s h  and  recognize  the nature o f the b r y o p h y t e v e g e t a t i o n on c a l c a r e o u s rock s u b s t r a t a i n w e s t e r n North A m e r i c a , i n general and i n B r i t i s h C o l u m b i a , i n p a r t i c u l a r . A l a r g e r set of releves w i l l  c e r t a i n l y enhance t h e d e s i g n a t i o n o f o r d e r ,  a l l i a n c e and a s s o c i a t i o n c h a r a c t e r s p e c i e s and w i l l  a l l o w a more r e l i a b l e  comparison w i t h data from l i m e s t o n e b r y o p h y t e v e g e t a t i o n s as d e s c r i b e d and c l a s s i f i e d i n Europe.  153  8.4.4. E v a l u a t i o n o f the o r d i n a t i o n  In o r d e r t o d i s p l a y and r e l e v e s from l i m e s t o n e nation techniques  v i s u a l i z e r e l a t i o n s h i p s between t h e  rocks, contained  i n major group number 4,  were a p p l i e d t o the data s e t . The  n a t i o n , namely, p r o v i d e s w i t h one  techniques.  a n o t h e r . The  use o f the  ordi-  ordi-  a means whereby a l l r e l e v e s can be compared  r e s u l t s o f the c e n t e r e d  A n a l y s i s are shown i n F i g u r e 8. The  Principal  Components  axes on t h i s o r d i n a t i o n i n d i c a t e  the p e r c e n t a g e d i s s i m i l a r i t y between the r e l e v e s which are a r r a n g e d a l o n g the X - a x i s and  Y-axis according  t o t h e i r sample s c o r e s  (listed  i n Appendix I V ) . A f i r s t group o f r e l e v e s ( t h e a s s o c i a t i o n Lophozio throphylletum  r e c u r v i r o s t r i ) , predominantly occurs  Bryoery-  i n the bottom r i g h t  c o r n e r o f the o r d i n a t i o n diagram ( F i g u r e 8 ) . A second group o f r e l e v e s ( t h e a s s o c i a t i o n Metaneckeretum m e n z i e s i i ) i s l o c a t e d towards the l e f t c o r n e r and a t h i r d group o f r e l e v e s , t h e B a r b u l a  top  vinealis -  P l a t y d i c t y a j u n g e r m a n n i o i d e s - Jungermannia a t r o v i r e n s - community, appears c o n f i n e d t o the bottom l e f t c o r n e r o f the o r d i n a t i o n diagram. A s c a t t e r diagram s u b s t a n t i a t e s t h i s arrangement i n F i g u r e 9, i n which the c e n t r o i d s and the s t a n d a r d  d e v i a t i o n s p l a c e t h e t h r e e groups o f  r e l e v e s i n the r e s p e c t i v e q u a d r a n t s . A comparison o f the r e s u l t o f t h i s c e n t e r e d A n a l y s i s w i t h the p a t t e r n r e s u l t i n g from the i n i t i a l  Principal  Components  cluster analysis,  shows a r e c o g n i t i o n and a s e p a r a t i o n o f the same groups. However, r e l e v e number 11 was but i n the c e n t e r e d  p a r t o f c l u s t e r 1 i n the c l u s t e r a n a l y s i s ( F i g u r e P.C.A. ( F i g u r e 8) i s grouped w i t h r e l e v e s 2, 3,  4,  11),  154  1 1 1 1 2 4 6 8 0 2 4 6  1 2 2 2 2 8  0  2 4 6  5 6 6 6 6 6 7 7 7 7 7 8 8 8 8 8 9 2 3 3 3 3 3 4 4 4 4 4 5 5 5 . 5 0 8 0 2 4 6 8 0 2 4 6 8 0 2 4 6 8 0 2 4 6 8 0 2 4 6 8 0 2 4 6 8  9 9 9 9 2 4 6 8  1 0 0  % dissimilarity F i g u r e 8. P r i n c i p a l components a n a l y s i s of t h e 27 l i m e s t o n e r e l e v e s . (A.=Metaneckeretum m e n z i e s i i ; • = L o p h o z i o - B r y o e r y t h r o p h y l l e t u m r e c u r v i r o s t r i ; Barbula v i n e a l i s - P l a t y d i c t y a jungermannioidesJungermannia a t r o v i r e n s ^ c o m m u m t y i f = m s i d e g r o t t o t y p e ; O = o u t s i d e grotto type.)  155  2  4  6  8  0  1 ( 2 2 2 2 2 3 3 3 3 3 4 4 4 4 4 5 S 5 5 5 6 6 6 6 6 7 7 7 7 7 8 8 8 8 8 9 9 9 9 9 0 4 6 8 O 2 4 6 R 0 2 4 6 8 0 2 4 6 8 0 2 4 6 8 O 2 4 6 8 O 2 4 6 8 0 6 8 0 2 4 6 8 0 2  %  dissimilarity  F i g u r e 9. S c a t t e r - d i a g r a m o f t h e 27 l i m e s t o n e r e l e v e s from t h e PCA-cent o r d i n a t i o n . l = L o p h o z i o - B r y o e r y t h r o p h y l l e t u m r e c u r v i r o s t r i ; 2= Metaneckeretum m e n z i e s i i ; 3=Barbula v i n e a l i s - P l a t y d i c t y a j u n germannioides-Jungermannia atrovirens-community. Standard d e v i a t i o n s a r e i n d i c a t e d by l i n e s .  156  7, 8 and  10  l y s i s had  (=cluster  included  2 ) . A manual t a b u l a t i o n  r e l e v e number 11  p r i o r t o the c l u s t e r ana-  i n the Metaneckeretum  w h i c h , a l s o , i s r e p r e s e n t e d by c l u s t e r 2. The  r e l a t i v e l y high cover  v a l u e o f Racomitrium h e t e r o s t i c h u m v a r . a f f i n e and Bryum c a p i l l a r e are s h a r e d w i t h  releves  the presence o f  numbers 12 and  However, the o c c u r r e n c e s o f I s o t h e c i u m s t o l o n i f e r u m , j u n g e r m a n n i o i d e s and  1,  respectively.  Platydictya  Gymnostomum aeruginosum i n d i c a t e i t s " a f f i n i t y "  the Metaneckeretum m e n z i e s i i p h y l l e t u m . The  menziesii,  to  r a t h e r than the Lophozio - B r y o e r y t h r o -  presence o f E n c a l y p t a p r o c e r a e x c l u d e s r e l e v e  number 11  from the t h i r d group o f r e l e v e s , the B a r b u l a v i n e a l i s - P l a t y d i c t y a 5 j u n g e r m a n n i o i d e s - Jungermannia a t r o v i r e n s The releves all  P r i n c i p a l Components A n a l y s i s  - community  .  shows t h i s t h i r d group o f  as a more o r l e s s " i s o l a t e d " c o l l e c t i o n o f l i m e s t o n e samples  r e s t r i c t e d t o the l e f t lower quadrant o f the o r d i n a t i o n  ( F i g u r e 9 ) . The i n t o two  cluster analysis  smaller  c l u s t e r 3A and  (Figure 3B.  The  11)  separated t h i s c l u s t e r 3  f l o r i s t i c and  r e l a t i o n s h i p s between t h e s e " i n s i d e - g r o t t o " and are d e s c r i b e d on page 152.  This separation  autonomous groups i s s u b s t a n t i a t e d  and  and  3B  (releves  19,  t h e i r d i s t r i b u t i o n a l o n g the  20,  more o r  by a subsequent p o l a r  10). Here, the d i s t i n c t i o n between 3A  24)  ecological  "outside-grotto"  i n t o two  (Figure  21, 25,  diagram  26 and  (releves 27)  types  less  ordination  5, 6, 22,  23  becomes a p p a r e n t i n  Y-axis.  With r e s p e c t t o the r e l a t i o n s h i p s between the r e l e v e s e n v i r o n m e n t s , the X - a x i s i n the p o l a r o r d i n a t i o n  and  suggests exposure  to be the environmental f a c t o r r e f l e c t e d i n the p o s i t i o n i n g o f  the  their  157  100 98 9E 94 92 90 88 86 84 82 80 78 76 74 72 70 68 66 64 62 60 >> 5 8 4-> 5 6 54  tl (0  »r-  E  •r-  i/i  to  •r-  52 50 48 46 44 42 40 38 36 34 32 30 28 26 24 22 20 18 16 14 12 10  10  7  O 2 7  12  3  20  S  2?,  °19  26  13 15 16 17 14  23.• •22 •, 24  IB  5 1  1 1 2  4  2 0  2 2  2 2 4 6  2 8  3 2  3 3 4 6  3 4 8 0  4 2  %  4 4 4 6  4 5 8 0  5 5 2 4  5 5 6 6 8 0  6 2  6 6  6 7 8 0  7 7 2 4  7 6  7 8 8 0  8 2  8 8 8 9 9 9 9 9 0 4 6 8 0 2 4 6 8 0  dissimilarity  F i g u r e 10, P o l a r o r d i n a t i o n o f t h e 27 l i m e s t o n e r e l e v e s (^=Metaneckeretum m e n z i e s i i ;•= L o p h o z i o - B r y o e r y t h r o p h y l l e t u m r e c u r v i r o s t r i ; B a r b u l a v i n e a l i s - P l a t y d i c t y a jungermannioides-Jungermannia a t r o v i r e s - c o m m u n i t y : a = i n s i d e g r o t t o t y p e ; 0 = o u t s i d e g r o t t o type.)  s T E P  2 2 2 2 2 6 5 4 3 2  2  1  2 1 1 1 1 1 1 1 1 1 0 9 8 7 G 5 4 3 2  0  9 8  7  6  5  4  3  2  1  --I I --I-  I-I-  -I I-I I---I  co  + -I + -I+ + -I i-B-  D I S T A N  7  7  8  8  8  1  1 2 2 3 3 4 4 5 5 G 7 7 7 7 8 1 1 9 3 7 5 7 4 7 3 9 1 1 2 3 7 5 1 3 2 5 0 0 8 2 4 5 0 0 0 3 7 0 5 0 6 7 2 5 0 0 1 5 5 1 4 2 8 . . 0 0 0 0 3 5 0 4 0 6 5 5 0 0 0 2 4 0 6 2 5 3 2 2 4  0  0  0  0  3  1  6  0  0  2  0  7  0  0  0  0  0  5  2  0  7  5  0  3  4  8  1 5 0 . 3  1 8 8 . 4  0  9  C E S  F i g u r e 1 1 . C l u s t e r a n a l y s i s o f the 27 limestone r e l e v e s . Symbols and numerical and a l p h a b e t i c c h a r a c t e r s are d i s c u s s e d i n the t e x t . Distance i s e u c l i d e a n .  159  r e l e v e s a l o n g t h i s a x i s . In a d d i t i o n , a l s o the measured of soil  a c c u m u l a t i o n on the s u b s t r a t a  X - a x i s . The show any  thickness  reveals a gradient along  the  g r o t t o r e l e v e s , on the l e f t p o r t i o n o f the a x i s , d i d  soil  not  a c c u m u l a t i o n , whereas the exposed b r y o p h y t e assemblages  o f the r e l e v e s , p o s i t i o n e d on the r i g h t s e c t i o n o f the X - a x i s , were growing on s o i l The gradient  depositions  on r o c k s up t o 2(3)  cm t h i c k .  d i s t r i b u t i o n o f the r e l e v e s a l o n g the Y - a x i s suggests a  from dry t o humid. The  inside-grotto bryophytes, c h a r a c t e r i s -  t i c a l l y , o c c u r r e d on d r y , c r u s t y l i m e s t o n e , Bryoerythrophylletum r e c u r v i r o s t r i  whereas the Lophozio -  grew on exposed, dry  soil-covered  r o c k s . Both groups o f r e l e v e s are p o s i t i o n e d a l o n g the lower p o r t i o n the Y - a x i s . The  outside-grotto  r e l e v e s and  the r e l e v e s c o n s t i t u t i n g  the a s s o c i a t i o n Metaneckeretum m e n z i e s i i , o c c u r r e d on s h e l t e r e d bases and  shaded b o u l d e r s i n s i d e f o r e s t v e g e t a t i o n s .  Their  rock  subsequent  p o s i t i o n i n g a l o n g the Y - a x i s i s more o r l e s s c o n c e n t r a t e d i n the p o r t i o n o f the o r d i n a t i o n diagram ( F i g u r e  To  nents A n a l y s i s . The Figure  1  upper  10).  f u r t h e r s u b s t a n t i a t e these r e s u l t s , a ' c l u s t e r a n a l y s i s  performed on the r e l e v e s  was  s c o r e s used i n the c e n t e r e d P r i n c i p a l Compo-  r e s u l t of t h i s c l u s t e r analysis i s presented i n  12. The  f i r s t cluster analysis  ( F i g u r e 11)  placed  r e l e v e number 11  i n the Lophozio - B r y o e r y t h r o p h y l l e t u m r e c u r v i r o s t r i . In t h i s second c l u s t e r a n a l y s i s , r e l e v e number 11 and  of  i s placed  i n c l u s t e r 2,  d e s i g n a t e d b e f o r e as the a s s o c i a t i o n Metaneckeretum  recognized  menziesii.  Distance 1 12 15 13 14 16 17 18 2 11 10 3 9 4 8 7 5 6 22 23 24 27 26 19 20 25 21  0  O0 N • • 0 0  oo r o t o oo i — . — i — o o o o • • • • o o o o o o o  CSJ  o  •si" (O  CO LO M J \  O O O  00  i  1—  1—  1—  • —  — o  OO 1—  1—  rCM  00  LO LO  OO O  LO  ^j- r\  r-~ 00  ro  I —  o  F i g u r e 12. C l u s t e r a n a l y s i s o f P.C.A.-scores (7 axes) o f the 27 limestone r e l e v e s . Distance i s Euclidean.  160  1. S i n c e l i m e s t o n e s u b s t r a t a a r e v e r y r e s t i c t e d i n t h e i r d i s t r i b u t i o n throughout the study a r e a , s a m p l i n g o f t h e b r y o p h y t e v e g e t a t i o n on c a l c a r e o u s r o c k s i s performed o n l y i n t h e C h i l l i w a c k R i v e r Valley. 2. S y n s y s t e m a t i c t r e a t m e n t o f t h e o r d e r C t e n i d i e t a l i a m o l l u s c i Hadac ans Smarda 1944 and t h e a l l i a n c e S c h i s t i d i o n a p o c a r p i J e z e k and Vondracek 1962 a r e g i v e n by M a r s t a l l e r (1979) and ( 1 9 8 0 ) , r e s p e c tively 3. S i g n i f i c a n t f l o r i s t i c a f f i n i t i e s can be d e t e c t e d between the p r e s e n t l y r e c o g n i z e d Lophozio - B r y o e r y t h r o p h y l 1 e t u m r e c u r v i r o s t r i and p h y t o s o c i o l o g i c a l t a b l e s p r o v i d e d by P o e l t ( 1 9 5 4 ) , Haybach ( 1 9 5 6 ) , Hb'fler ( 1 9 5 9 ) , P h i l i p p i ( 1 9 6 5 ) , Hagel ( 1 9 6 6 ) , Nagano ( 1 9 6 9 ) , Norr ( 1 9 7 0 ) , Neumayr ( 1 9 7 1 ) , S t r a s s e r ( 1 9 7 2 ) , H e r t e l ( 1 9 7 4 ) , S t u c h l y ( 1 9 7 6 ) , Hebrard ( 1 9 7 8 ) , Mamczarz (1978) and M a r s t a l l e r (1979, 1980a, 1980b). 4. F l o r i s t i c s i m i l a r i t i e s between t h e Metaneckeretum m e n z i e s i i and European l i m e s t o n e b r y o p h y t e v e g e t a t i o n s are found i n the f o l l o w i n g r e f e r e n c e s : Demaret ( 1 9 4 4 ) , P o e l t ( 1 9 5 4 ) , Koppe ( 1 9 5 5 ) , H o f l e r ( 1 9 5 9 ) , B r e u e r ( 1 9 6 2 ) , P h i l i p p i ( 1 9 6 5 ) , Hagel ( 1 9 6 6 ) , Hubschmann ( 1 9 6 7 ) , Norr ( 1 9 7 0 ) , Neumayr ( 1 9 7 1 ) , S t r a s s e r ( 1 9 7 2 ) , H e r t e l ( 1 9 7 4 ) , W i l c z y n s k a ( 1 9 7 4 ) , Mamczarz ( 1 9 7 8 ) , M a r s t a l l e r (1979, 1980). 5. A l s o the t y p e o f h a b i t a t , i n t h i s case shaded, humid b o u l d e r s i n s i d e f o r e s t s , i n d i c a t e t h e placement o f r e l e v e number 11 i n t h e Metaneckeretum m e n z i e s i i . The Lophozio - B r y o e r y t h r o p h y l 1 e t u m r e c u r v i r o s t r i i s an a s s o c i a t i o n from exposed, dry r o c k s .  161  9. SUMMARY.  In a study concerned w i t h t h e e p i l i t h i c i t i s important  bryophyte vegetation  t o r e a l i z e t h a t s u b s t r a t a such as b o u l d e r s  w a l l s u s u a l l y are "discontinuous" with the surrounding  and r o c k -  (vascular)  vegetation. Rocky s l o p e s i n s u b a l p i n e and a l p i n e areas u s u a l l y c o n t a i n a p i o n e e r stage i n which b r y o p h y t e s (and l i c h e n s ) a r e t h e s o l e i n h a b i t a n t s , and v a s c u l a r p l a n t s can e s t a b l i s h o n l y i n l a t e r stages o f s u c c e s s i o n . However, smooth rock s u r f a c e s , r e t a i n i n g any s o i l  i n which t r e e s  and shrubs may become a n c h o r e d , remain i n d e f i n i t e l y i n a s e m i - b a r r e n s t a t e c o n d i t i o n e d by t h e r a t e and amount o f s o i l  accumulation.  In most  i n s t a n c e s , exposed and s h e l t e r e d b o u l d e r s and rocks a r e u s u a l l y  covered  by a v e g e t a t i o n which i s , f l o r i s t i c a l l y , c o n s i d e r a b l y u n l i k e t h e p l a n t cover o f o t h e r w i s e wooded t e r r i t o r y . The v e g e t a t i o n growing on a bare r o c k s u r f a c e i s r e l a t e d p r i m a r i l y t o t h e k i n d o f r o c k , i t s chemical Philippi  composition  1961, Nagano 1972), t h e s t a t e o f w e a t h e r i n g  ( S h a c k l e t t e 1961, and s u r f a c e  micro-  s t r u c t u r e ( S j o g r e n 1964). I t i s a l s o i n f l u e n c e d by m a c r o c l i m a t e and m i c r o c l i m a t i c c o n d i t i o n s depending l a r g e l y on r e l i e f ,  i n c l i n a t i o n and  exposure ( K r u s e n s t j e r n a 1965). Hence, many e c o l o g i c a l f a c t o r s such as moisture,  light,  temperature and s o i l  bution of e p i l i t h i c  accumulation  determine t h e d i s t r i -  bryophytes and t h e i r communities. ( O o s t i n g and  Anderson 1947, Keever, O o s t i n g  and Anderson 1951). A l s o the r e l e v a n c e  o f b r y o p h y t e r h i z o f d s i n a t t a c h i n g t h e p l a n t s to t h e rock  substrate  162  i s o f major importance i n t h e e s t a b l i s h m e n t and development o f t h e b r y o p h y t e v e g e t a t i o n (Keever 1957, Hebrard 1970, Odu 1978). This " d i s c o n t i n u i t y " with the surrounding vegetation i s further reflected i n the variation i n f l o r i s t i c b r y o p h y t e assemblages  composition o f  from d i f f e r e n t e n v i r o n m e n t s . The b r y o p h y t e  communities from exposed, d r y r o c k s , from shaded, humid b o u l d e r s and from s t r e a m s i d e r o c k s a l l e x h i b i t " d i s c o n t i n u o u s " v a r i a t i o n i n t h e i r d e t a i l e d f l o r i s t i c c o m p o s i t i o n as compared t o t h e ( v a s c u l a r ) v e g e t a t i o n of  t h e i r immediate environment. T h i s o b s e r v a t i o n i m p l i e s some c o n s i d e -  r a b l e r e l a t i o n s h i p between t h e s p e c i e s o f a community w h i c h , i n growing t o g e t h e r , s u f f i c i e n t l y modify t h e environment t o form a r e c o g n i z a b l e and r e p e t i t i v e v e g e t a t i o n a l  g r o u p i n g (Anderson 1965).  Any procedure o f v e g e t a t i o n c l a s s i f i c a t i o n i n v o l v e s an a r r a n g e ment o f communities i n t o c l a s s e s : t h e members o f each c l a s s have i n common a c o n s t e l l a t i o n o f a t t r i b u t e s which s e r v e t o s e t them a p a r t from members o f a n o t h e r c l a s s . I m p l i c i t i n t h i s approach i s t h e s u g g e s t i o n t h a t t h e r e i s some d i s c o n t i n u i t y i n s p e c i e s ( a t t r i b u t e ) c o m p o s i t i o n between both c o n c r e t e samples o f v e g e t a t i o n ( r e l e v e s ) i n t h e f i e l d and t h e o r e t i c a l  u n i t s ( s y n t a x a ) a b s t r a c t e d from such f i e l d d a t a .  In t h e p r e s e n t attempt t o c l a s s i f y t h e e p i l i t h i c communities of  bryophyte  i n s o u t h w e s t e r n B r i t i s h Columbia, t h e Braun-Blanquet method  v e g e t a t i o n s t u d y has been a p p l i e d . T h i s approach i s based on t h e  concept o f e x i s t i n g d i s c o n t i n u o u s v a r i a t i o n w i t h i n t h e p l a n t c o v e r o f a g i v e n area and r e s u l t s i n t h e r e c o g n i t i o n o f more o r l e s s  readily  d e l i m i t e d c l e a r - c u t u n i t s . These u n i t s a r e then c l a s s i f i e d as a s s o c i a t i o n s  163  i n a h i e r a r c h i a l c l a s s i f i c a t i o n scheme o f s y n t a x a . In a d d i t i o n t o the t a b u l a t i o n t e c h n i q u e o f o r d e r i n g r e l e v e s , c l u s t e r a n a l y s e s have been used t o r e c o g n i z e data m a t r i x  patterns  and  i n the  o f each major group (groups from e c o l o g i c a l l y d i f f e r e n t  s i t e t y p e s ) . T h i s s e a r c h f o r groups o f s i m i l a r r e l e v e s has i d e n t i f i c a t i o n of separate a s s o c i a t i o n s , subassociations, f a c i e s , each c h a r a c t e r i z e d by t h e i r c h a r a c t e r In o r d e r t o r e c o g n i z e the p r e p a r a t i o n  o f raw  d i s t i n c t i v e patterns  t a b l e s and  of species  presence t a b l e s by t h e  computerized  d i s t r i b u t i o n t h r o u g h o u t the data s e t epilithic  lower s y n t a x o n o m i c a l u n i t s . However, above  a s s o c i a t i o n s i n t o the h i g h e r detects  and  been found s u c c e s s f u l . I t r e v e a l s  the a s s o c i a t i o n l e v e l , c l u s t e r a n a l y s i s d i d not p r o v i d e  analysis algorithm  variants  the  species.  p r o v i d e s an adequate b a s i s f o r the c l a s s i f i c a t i o n o f  b r y o p h y t e a s s o c i a t i o n s and  l e d to  these units a c l u s t e r a n a l y s i s , f o l l o w i n g  t a b u l a r s o r t i n g program VEGTAB, has  and  species  s y n t a x a o f a l l i a n c e s and  differences i n species  into higher  o r d e r s . The  composition  q u a n t i t i e s between r e l e v e s , r a t h e r than the v e g e t a t i o n c l a s s i f i c a t i o n of associations  a means t o combine  type.  cluster  and The  s y n t a x a , on the o t h e r hand,  i s based on t h e i r o c c u r r e n c e s i n , e c o l o g i c a l l y d i f f e r e n t h a b i t a t s . A l t h o u g h sometimes apparent f l o r i s t i c s i m i l a r i t i e s i n s p e c i e s and q u a n t i t i e s are d e t e c t e d  composition  by the c l u s t e r i n g a n a l y s i s a l g o r i t h m ,  this  i s not r e f l e c t e d i n the syntaxonomic c l a s s i f i c a t i o n schemes o f the major groups ( T a b l e s I I I , X I , XV and The  XXV)  l e v e l o f r e s o l u t i o n (the c u t - o f f p o i n t i n the c l u s t e r  analy-'  s i s ) used i n the r e c o g n i t i o n o f the s y n t a x a i s , t h u s , somewhat a r b i t r a r y .  164  However, attempts have been made t o d e f i n e b r y o p h y t e communities to correspond t o the e p i l i t h i c a s s o c i a t i o n s described  by v a r i o u s  European b r y o - s o c i o l o g i s t s . These workers m o s t l y a p p l i e d  tabulation  t e c h n i q u e s , s t a n d a r d w i t h i n the Braun-Blanquet method, t o t h e i r vegetation  d a t a . As s t a t e d b e f o r e , t h i s study used c l u s t e r a n a l y s e s  to a r r i v e at the separate vegetation  units.  I t i s t h i s comparison between data from European and  t h e p r e s e n t s t u d y ' s data m a t r i c e s , which r e v e a l e d  s i m i l a r i t i e s i n species and  vegetations  some remarkable  c o n t r i b u t i o n to the separate  associations  i n t h e m i c r o - s i t e s p e c i f i c i t y o f t h e b r y o p h y t e communities i n g e n e r a l .  Hence, where p o s s i b l e , t h e p r e s e n t l y d i s t i n g u i s h e d a s s o c i a t i o n s and lower syntaxonomical  u n i t s have been p l a c e d  i n already  f i c a t i o n schemes f o r b r y o p h y t e communities o c c u r r i n g  existing classi-  i n different  t y p e s o f environments.  The  a p p l i c a t i o n o f o r d i n a t i o n techniques, although l i m i t e d  i n t h e p r e s e n t s t u d y , has been welcomed as an a d d i t i o n a l means t o d i s p l a y and  i n t e r p r e t patterns  patterns  i n t h e d a t a m a t r i x . The r e c o g n i t i o n o f t h e s e  i s , then, r e f l e c t e d i n the p o s i t i o n i n g o f the releves  along  the X- and Y - a x i s o f t h e o r d i n a t i o n diagrams ( F i g u r e s 8 and 10). D i f f e rences i n the p o s i t i o n s o f t h e v e g e t a t i o n  samples have c o n t r i b u t e d t o  the i d e n t i f i c a t i o n o f "groups o f s i m i l a r r e l e v e s " , and a comparison w i t h t h e c l u s t e r a n a l y s i s ' arrangement The respect  followed.  P r i n c i p a l Components A n a l y s i s , which o r d i n a t e s  to a l l others simultaneously,  substantiated  releves  with  and s u p p o r t e d t h e r e c o g -  165  n i t i o n o f t h r e e l i m e s t o n e b r y o p h y t e v e g e t a t i o n u n i t s , each  charac-  t e r i z e d by i t s own d i a g n o s t i c s p e c i e s . A subsequent p o l a r o r d i n a t i o n , which o r d i n a t e s r e l e v e s on t h e b a s i s o f comparison w i t h two e n d p o i n t r e l e v e s , n o t o n l y r e v e a l e d d i f f e r e n c e s w i t h i n one o f t h e u n i t s (3A and 3B; F i g u r e 1 0 ) , b u t p r o v i d e d a means t o c o r r e l a t e t h e p o s i t i o n s o f t h e groups o f s i m i l a r r e l e v e s a l o n g t h e axes w i t h e c o l o g i c a l f a c t o r s such as e x p o s u r e , m o i s t u r e and s o i l  accumulation.  F i e l d data on e c o l o g i c a l s i t e c h a r a c t e r i s t i c s , i n a d d i t i o n to t a b u l a t i o n and c l u s t e r a n a l y s i s , a l r e a d y showed some r e l a t i o n s h i p s o f r e l e v e s t o one another and t o t h e environment. T h i s p r o v i d e d a b a s i s f o r i n t e r p r e t i n g the o r d i n a t i o n s , which, i n t u r n , f u r t h e r expressed and c l a r i f i e d those  relationships.  F u r t h e r r e s e a r c h on the e p i l i t h i c b r y o p h y t e v e g e t a t i o n i s needed. E s p e c i a l l y i n s u b a l p i n e and a l p i n e r e g i o n s , where rock s u b s t r a t a are i n abundant s u p p l y , t h e v a r i a t i o n i n s p e c i e s c o m p o s i t i o n  and t h e  m i c r o - s i t e s p e c i f i c i t y o f b r y o p h y t e assemblages need a d d i t i o n a l work. More r e l e v e s may s u b s t a n t i a t e f u r t h e r t h e sometimes apparent f l o r i s t i c and e c o l o g i c a l a f f i n i t i e s between t h e e p i l i t h i c b r y o p h y t e v e g e t a t i o n o f south-western  B r i t i s h Columbia and o t h e r areas i n t h e  P a c i f i c North West, i n g e n e r a l , and r e g i o n s elsewhere i n North and Europe. The knowledge on t h e s e v e g e t a t i o n s w i l l  gain i n understanding  and meaning when t h e s e methods a r e e x t e n s i v e l y a p p l i e d over areas i n t h e temperate r e g i o n s o f t h e N o r t h e r n On a more l o c a l  America  wider  Hemisphere.  s c a l e r e s e a r c h may be d i r e c t e d on t h e n a t u r e  166  o f t h e b r y o p h y t e assemblages. F i r s t , more d e t a i l e d measurements o f environmental  factors are required to characterize the ecological  d i f f e r e n c e s on and between t h e v a r i o u s r o c k s u b s t r a t a . Second, a u t ecological  ( p a r t i c u l a r l y p h y s i o l o g i c a l ) i n f o r m a t i o n i s needed f o r t h e  v a r i o u s s p e c i e s i n o r d e r t o p r o p e r l y examine t h e i r a b i l i t y and s u c c e s s to occupy t h e e p i l i t h i c h a b i t a t , and t o more a c c u r a t e l y d e s c r i b e n i c h e d i f f e r e n t i a t i o n . T h i r d , by s e t t i n g up permanent q u a d r a t s and u s i n g i n f o r m a t i o n from t h e p r e v i o u s two s u g g e s t i o n s  f o r further research,  the dynamic n a t u r e o f t h e e p i l i t h i c v e g e t a t i o n c o u l d be s t u d i e d and c o n t r a s t e d between e c o l o g i c a l l y d i f f e r e n t environments.  167 REFERENCES.  A n d e r s o n , D.J. 1965. C l a s s i f i c a t i o n and o r d i n a t i o n i n v e g e t a t i o n s c i e n c e : c o n t r o v e r s y over a n o n - e x i s t e n t problem? J . E c o l . 53: 521-526. A p i n i s , A. 1939. 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A s t a t i s t i c a l i n v e s t i g a t i o n i n t o the s t r u c t u r e and e c o l o g y o f some s a x i c o l o u s bryophyte and l i c h e n communities. Ph. D. T h e s i s , Univ. o f E x e t e r . 62 p. . 1967a. P r i n c i p a l components a n a l y s i s o f d a t a from s a x i c o l o u s b r y o p h y t e v e g e t a t i o n a t Steps B r i d g e , Devon. I . A q u a n t i t a t i v e assessment o f v a r i a t i o n i n the v e g e t a t i o n . Can. J . Bot. 45: 9.3-115. . 1967b. J . Bot. 45:  Ibid. I I . An experiment w i t h h e t e r o g e n e i t y . 229-247.  Can.  . 1967c. I b i d . I I I . C o r r e l a t i o n o f v a r i a t i o n i n the veget a t i o n w i t h environmental v a r i a b l e s . Can. J . Bot. 45: 249-258. . T967d. A q u a n t i t a t i v e study o f the b r y o p h y t e and macrol i c h e n v e g e t a t i o n o f the Dartmoor g r a n i t e . The L i c h e n o l o g i s t 3(3): 392-408.  179  APPENDIX I  180  ABUNDANCE SCALE ( a f t e r Frey 1933)  5  covering  1/2  (50  -  100%)  of the p l o t  4  covering  1/4 - 1/2  (25  -  50%)  o f the p l o t  3  covering  1/8 - 1/4  (12.5 -  25%)  of the p l o t  2  covering  1/16 - 1/8  ( 6  -  12.5%)  of the p l o t  1  covering  1/100 - 1/16  ( 1  -  6%)  o f the p l o t  +  covering  )  of the p l o t  <1/100  (  <1%  181  SOCIABILITY SCALE  5  compact mat -  20 cm  4  l o o s e mat o r l o o s e t u r f -  3  t u r f (cushion) -  3-8 cm  2  t u r f (cushion) -  1-3 cm  1  s o l i t a r y plants - t u r f between 1 cm i n a c r o c a r p o u s mosses and 3-4 (5) cm i n p l e u r o c a r p o u s mosses  8-20  cm  182  APPENDIX I I CHARACTER TAXA.  F i d e l i t y degree 5: E x c l u s i v e t a x a : t a x a c o m p l e t e l y  o r almost  completely  F i d e l i t y degree 4: S e l e c t i v e t a x a : t a x a o c c u r r i n g w i t h c l e a r  preference  c o n f i n e d t o one phytocoenon ( v e g e t a t i o n  unit);  f o r one phytocoenon b u t a l s o , though w i t h a c o n s i d e r a b l y presence d e g r e e , i n o t h e r  coena:  F i d e l i t y degree 3: P r e f e r e n t i a l perhaps w i t h about equal  lower  taxa: taxa present  i n several  coena,  p r e s e n c e d e g r e e , b u t w i t h a h i g h e r combined  e s t i m a t i o n v a l u e and ( o r ) w i t h a h i g h e r v i t a l i t y degree i n one particular  coenon;  COMPANIONS.  F i d e l i t y degree 2: I n d i f f e r e n t t a x a : t a x a w i t h o u t  pronounced  preference  f o r any coenon;  ACCIDENTALS.  F i d e l i t y degree 1: S t r a n g e t a x a : t a x a h a v i n g  a d e f i n i t e presence  degree optimum and m o s t l y a l s o a cover-abundance optimum o u t s i d e t h e considered bouring  coenon. These a r e o f t e n a c c i d e n t a l i n t r u d e r s from  neigh-  coena o r r e l i c s from a coenon t h a t preceded i n s u c c e s s i o n .  183  APPENDIX I I I  Bryophyte s p e c i e s  list  184  CLASS HEPATICAE  Family  Pseudolepicoleaceae BIepharostoma t r i c h o p h y l l u r n ( L . ) Dum.  Family  Herbertaceae Herbertus  Family  aduncus ( D i c k s . ) S. Gray  Ptilidiaceae P t i l i d i u m c a l i f o r n i c u m (Aust.) Underw. P_. c i l i a r e ( L . ) Hampe P_. pulcherrimum (G. Web.) V a i n .  Family  Lepidoziaceae B a z z a n i a denudata ( T o r r . ex G o t t . ) T r e v . B_. t r i c r e n a t a (Wahlenb.) L i n d b . i n B r o t h L e p i d o z i a r e p t a n s ( L . ) Dum.  Family  Calypogeiaceae C a l y p o g e i a i n t e g r i s t i p u l a Steph. C_. m u e l l e r i a n a ( S c h i f f n . ) K. M u e l l .  Family A n t h e l i a c e a e A n t h e l i a j u r a t z k a n a (Limpr.)  Family  Trev.  Cephaloziaceae C e p h a l o z i a b i c u s p i d a t a ( L . ) Dum.  185  Family Cephaloziaceae (continued) Hygrobiella l a x i f o l i a  (Hook.) Spruce  P l e u r o c l a d a a l b e s c e n s (Hook.) Spruce  Family C e p h a l o z i e l l a c e a e C e p h a l o z i e l l a byssacea (Roth.) Warnst. v a r . byssacea C;. byssacea v a r . a s p e r i f o l i a ( T a y l . ) Macv. C. p h y l l a c a n t h a (Mass. & C a r e s t . ) K. M u e l l . CL t u r n e r i (Hook.) K. M u e l l .  F a m i l y Geocalycaceae Geocalyx g r a v e o l e n s (Schrad.) Nees  Family  Lophocoleaceae C h i l o s c y p h u s p o l y a n t h o s (L.) Corda Lophocolea c u s p i d a t a (Nees) Limpr. L. h e t e r o p h i l a (Schrad.) Dum.  Family P l a g i o c h i l a c e a e P l a g i o c h i l a a s p l e n i o i d e s (L.)  Dum.  Family Gyrothyraceae G y r o t h y r a underwoodiana  Family  M.A.  Howe  Jungermanniaceae A n a s t r o p h y l l u m a s s i m i l e ( M i t t . ) Steph. A. minutum (Schreb.) S c h u s t . B a r b i l o p h o z i a f l o e r k e i (Web.  e t Mohr.) Loeske  186  F a m i l y Jungermanniaceae  (continued)  B a r b i l o p h o z i a h a t c h e r i . ( E v a n s ) Loeske B_. l y c o p o d i o i d e s ( W a l l r . ) Loeske Chandonanthus f i 1 i f o r m i s Steph. C. s e t i f o r m i s ( E h r h J L i n d b . J a m e s o n i e l l a autumnalis  (DC) Steph.  Jungermannia a t r o v i r e n s J_. c o n f e r t i s s i m a Nees J_. e x s e r t i f o l i a Steph. J_. obovata Nees J_. pumila With. L o p h o z i a e l o n g a t a Steph. L_. e x c i s a ( D i c k s . ) Dum. I. h e t e r o c o l p o s (Thed. ex Hartm.) M.A. Howe IL. i n c i s a (Schrad.) Dum. L_. s u d e t i c a (Nees ex Hueb.) G r o l l e L_. v e n t r i c o s a ( D i c k s . ) Dum. v a r . v e n t r i c o s a L_. wenzelii~("Nees) Steph. Mylia t a y l o r i  (Hook.) S. Gray  N a r d i a b r e i d l e r i (Limpr.) L i n d b . N_. geoscyphus (DeNot) L i n d b . N_. j a p o n i c a Steph. N_. s c a l a r i s . Gray T r i t o m a r i a e x s e c t i f o r m i s ( B r e i d l . ) S c h i f f n . ex Loeske I - P o ] i t a (Nees) J o e r g . J_. q u i n q u e d e n t a t a (Huds.) Buch F a m i l y Gymnomitriaceae Gymnomitrion concinnatum ( L i g h t f . ) Corda i n O p i z . G. obtusum ( l i n d b . ) P e a r s . M a r s u p e l l a b r e y i s s i m a (Dum.) G r o l l e M. condensata ( A n g s t r . ) S c h i f f n . M. emarginata (Ehrb.) Dum. M. s p a r s i f o l i a ( L i n d b . ) Dum. M. s p h a c e l a t a (Gieseke ex Lindenb.) Dum. M. s t a b l e r i Spruce  187  Family  Scapaniaceae D o u i n i a o v a t a ( D i c k s . ) Buch D i p l o p h y l l u m a l b i c a n s (L.) Dum. D. o b t u s i f o l i u m (Hook.) Dum. T). p l i c a t u m L i n d b . D. t a x i f o l i u r n (Wahlenb.) Dum. Scapania mucronata Buch S. paludosa (K. M u e l l . ) K. M u e l l . S_. s c a n d i c a (H. A r n . & Buch) Macv. S_. s u b a l p i n a (Nees ex Lindenb.) Dum. S. u l i g i n o s a (Sw. ex Lindenb.) Dum. undulata~~(L.) Dum.  F a m i l y Radulaceae Radula b o l a n d e r i G o t t . R. complanata (L.) Dum.  Family Porellaceae P o r e l l a cordaeana (Hueb.) Moore P. n a v i c u l a r i s (Lehm. & Lindenb.) L i n d b . F. r o e l l i i STeph.  Family F r u l l a n i a c e a e F r u l l a n i a t a m a r i s c i (L.) Dum.  ssp. n i s q u a l l e n s i s ( S u l l . ) Hatt.  Family P e l l i a c e a e P e l l i a neesiana  Family  (Gott.)  Limpr.  Metzgeriaceae Apometzgeria pubescens (Schrank) Metzgeria conjugata  Lindb.  Kuwah.  188  Family Blasiaceae B l a s i a p u s i l l a L.  F a m i l y Aneuraceae Aneura p i n g u i s (L.) Riccardia multifida  Dum. (L.) S. Gray  Family Aytoniaceae R e b o u l i a h e m i s p h a e r i c a (L.) Raddi  F a m i l y Conocephalaceae Conocephalurn conicum (L.) Dum.  Family Marchantiaceae P r e i s s i a q u a d r a t a (Scop.) Nees  ex L i n d b .  189  CLASS MUSCI  SUBCLASS SPHAGNIDAE  F a m i l y Sphagniceae  Sphagnum g i r g e n s o h n i i  Russ.  SUBCLASS ANDREAEIDAE  Family Andreaeaceae Andreaea b l y t t i i B.S.G. A. n i v a l i s Hook. A. r o t h i i Web. & Mohr. ft. r u p e s t r i s Hedw. SUBCLASS TETRAPHIDAE  Family Tetraphidaceae  T e t r a p h i s p e l l u c i d a Hedw.  SUBCLASS POLYTRICHIDAE  Family  Polytrichaceae Atrichum selwynii Aust. A. undulatum (Hedw.) P. Beauv. B a r t r a m i o p s i s Tescur'i'i (James) K i  190  Family P o l y t r i c h a c e a e ( c o n t i n u e d ) Oligotrichum aligerum M i t t , rj. hercynicum (Hedw.) Lam & DC 0. p a r a l l e l u m ( M i t t . ) K i n d b . Pogonatum contortum ( B r i d . ) L e s q . _P. urnigerum (Hedw.) P. Beauv. P o l y t r i c h u m a l p i n u m Hedw. v a r . a l p i n u m _P. alpinum v a r . macounii (Kindb.) S a i t o F\ formosum Hedw. F\ j urn p e r i num Hedw. £. p i l i f e r u m Hedw. _P. s e x a n g u l a r e F l o e r k e ex B r i d .  SUBCLASS BRYIDAE  Family D i t r i c h a c e a e Ceratodon purpureus  (Hedw.) B r i d .  D i s t i c h i u m c a p i l l a c e u m (Hedw.) B.S.G. D i t r i c h u m f l e x i c a u l e (Schwaegr.) Hampe D. heteromallum (Hedw.) B r i t t .  Family S e l i g e r i a c e a e B l i n d i a a c u t a (Hedw.) B.S.G. S e l i g e r i a donniana  (Sm.) C. M u e l l .  Family Dicranaceae Arctoa f u l v e l l a  ( D i c k s . ) B.S.G.  Campylopus a t r o y i r e n s De Not C. f r a g i l i s ( B r i d . ) B.S.G. £. paradoxus W i l s . i n Hardy Cynodontium j e n n e r i  (Schimp. i_n Howie) S t i r t .  191  F a m i l y Dicranaceae  (continued)  Dichodontium p e l l u c i d u m Ren. & Card. D. olympicum (Hedw.) Schimp. Dicranella  h e t e r o m a l l a (Hedw.) Schimp.  D i c r a n o w e i s i a c i r r a t a (Hedw.) L i n d b . i n M i l d e D. c r i s p u l a (Hedw.) L i n d b . i_n Mi 1 de Dicranum f u s c e s c e n s Turn JJ. scoparium Hedw. K i a e r i a b l y t t i i (B.S.G.) B r o t h . J<. f a l c a t a (Hedw.) Hag. K. s t a r k e i (Web. & Mohr.) Hag. Paraleucobryum enerve (Thed. ex C . J . Hartm.) Loeske  Family F i s s i d e n t a c e a e F i s s i d e n s b r y o i d e s Hedw. F. osmundoides Hedw.  Family  Encalyptaceae E n c a l y p t a c i l i a t a Hedw. E. p r o c e r a Bruch  Family P o t t i a c e a e Anoectangium a e s t i v u m Barbula r e f l e x a  (Hedw.) M i t t .  (.Brid.) B r i d .  Didymodon v i n e a l i s ( b r i d . ) Zander Gymnostomum aeruginosum Sm. G. r e c u r v i r o s t r u m Hedw. Oxystegus t e n u i r o s t r i s (Hook. & T a y l . ) A.J.E. Smith Tortella fragilis  (Hook f . & W i l s . )  T o r t u l a p r i n c e p s De Not  Limpr.  192  Family. Grimmiaceae Dryptodon patens (Hedw.) B r i d . Grimmia a l p e s t r i s  (Web. & Mohr.) S c h l e i c h . ex_ Nees var. a l p e s t r i s G. a l p e s t r i s v a r . h o l z i n g e r i ( C a r d . & Ther.) James i n Gro G. donniana~Sm. G. l e i b e r g i i B.S.G. I- Montana B.S.G. G. t o r q u a t a Hornsch. i n Grev. Racomitrium a c i c u l a r e (Hedw.) B r i d . R.. aquaticum ( B r i d . ex Schrad.) B r i d . R. " b r e v i p e s " Kindb. R. canescens (Hedw.) Kindb. R. f a s c i c u l a r e (Hedw.) B r i d . R. lanuginosum (Hedw.) B r i d . R. heterostTcFum (Hedw.) B r i d . v a r . h e t e r o s t i c h u m R. h e t e r o s t i c h u m v a r . a f f i n e (C. Jensen) Kindb. R. s u d e t i c u m (Funck) B.S.G. R. variurn ( M i t t . ) J a e g . & Sauerb. S c h i s t i d i u m apocarpum (Hedw.) B. & S. v a r . apocar.pum S_. apocarpum v a r . s t r i c t urn (Turn.) T a y l . S_. n v u l a r e ( B r i d . ) Podp. S c o u l e r i a a q u a t i c a Hook, i j i Drumm.  Family S c h i s t o s t e g a c e a e S c h i s t o s t e g a pennata (Hedw.) Web. & Mohr.  F a m i l y Bryaceae Bryum c a e s p i t i c i u m Hedw. B_. mini at urn Lesq. B_. p a l l e s c e n s S c h l e i c h . ex Schwaegr. B_. p s e u d o t r i q u e t r u m (Hedw.) Schwaegr. P o h l i a c a r d o t i i (Ren.,ex_Re. & Card.) B r o t h . P_. cruda (Hedw.) L i n d b . P_. l o n g i b r a c t e a t a B r o t h , vn R o e l l . P_. l u d w i g i i (Schwaegr.) B r o t h . P_. nutans (Hedw.) L i n d b . IP. p r o l i g e r a ( K i n d b . e_x Limpr.) B r o t h .  193  F a m i l y Mniaceae L e u c o l e p i s m e n z i e s i i (Hook.) S t e e r e i j i Koch Mnium ambiguum H. M u e l l . M T ~ 5 T y t t i i B.S.G. M. spinulosum B.S.G. M. thomsoniT~Schimp. Plagiomnium r o s t r a t u m (Schrad.) P_. venustum ( M i t t . ) Kop.  Kop.  Rhizomnium g l a b r e s c e n s (Kindb.) Kop. R. nudum~TBritt. & W i l l i a m s ) Kop.  F a m i l y Aulacomniaceae Aulacomnium androgynum  Family  (Hedw.) Schwaegr.  Bartramiaceae A n a c o l i a m e n z i e s i i (Turn.) P a r . Bartramia ithyphyl1 a B r i d . B_. pomiformis Hedw.~ P h i l o n o t i s a r n e l 1 i i Husn. P_. f o n t a n a (Hedw.) B r i d . v a r . f o n t a n a P l a g i o p u s o e d e r i ( B r i d . ) Limpr.  F a m i l y Timmiaceae Timmia a u s t r i a c a Hedw.  Family Orthotrichaceae Amphidium c a l i f o r n i c u m (C. M u e l l . ) B r o t h A. lapponicum (Hedw.T~Schimp. A. mougeotii (B.S.G.) Schimp.  194  Family  Hedwigiaceae Hedwigia c i l i a t a  (Hedw.) P. Beauv.  Pseudobraunia c a l i f o r n i c a (Lesq.) B r o t h .  Family  Neckeraceae Homalia t r i c h o m a n o i d e s (Hedw.) B.S.G. Metaneckera m e n z i e s i i  (Hook ijn Drumm.) S t e e r e  Porotrichuirnbigelovii  ( S u l l . ) Kindb.  Thamnobryum n e c k e r o i d e s (Hook.) Lawt.  Family  Leskeaceae Pseudoleskea a t r i c h a ( K i n d b . j_n Macoun & Kindb.) Kindb. P_. b a i l e y i Best & Grout j_n Grout P_. i n c u r v a t a (Hedw.) Loeske JP. patens ( L i n d b . ) Kindb. P_. r a d i c o s a ( M i t t . ) Macoun & Kindb. P_. s t e n o p h y l l a Ren. & Card, ex Roe! 1. P s e u d o l e s k e e l l a t e c t o r u m (Funck ex B r i d . ) Kindb. e_x B r o t h . Pterigynandrum f i l i f o r m e Hedw.  Family Thuidiaceae Claopodium b o l a n d e r i B e s t C_. c r i s p i f o l i u m (Hook.) Ren. & Card. C_. whippleanum ( S u l l . ) Ren. & Card. H e t e r o c l a d i u m dimorphum ( B r i d . ) B.S.G. H_. macouni i B e s t H_. p r o c u r r e n s ( M i t t . ) Rau & Herv.  Family A m b l y s t e g i a c e a e Cratoneuron commutatum (Hedw.) Roth. v a r . f a l c a t u m ( B r i d . ) Moenk.  195  Family A m b l y s t e g i a c e a e ( c o n t i n u e d ) Drepanocladus  u n c i n a t u s (Hedw.) Warnst.  Hygrohypnum b e s t i i (Ren. & Bryhn.) H o l z . H. 1uridurir(Hedw.) Jenn. H. ochraceum (Turn. j_n W i l s . ) Loeske TT. s m i t h i i ("Sw. vn L i l j . ) B r o t h . P l a t y d i c t y a j u n g e r m a n n i o i d e s ( B r i d . ) Crum  Family Brachytheciaceae Brachythecium a l b i c a n s (Hedw.) B.S.G. EL asperrimum ( M i t t . ) S u l l . B_. frigidum~TC. M u e l l . ) Besch. B_. p i umosum (Hedw.) B.S.G. B_. r e f l e x u m ( S t a r k e j_n Web. & Mohr.) B.S.G. B_. r i v u l a r e (Hedw.) B.S.G. B_. rutabulum B.S.G. B. s t a r k e i ( B r i d . ) B.S.G. B_. v e l u t i n u m (Hedw.) B.S.G. Eurhynchium  p u l c h e l l u m (Hedw.) Jenn.  Homalothecium f u l g e s c e n s ( M i t t , ex C. Muel1.) J a e g . H. nevadense (Lesq.) Ren. & Card. H_. n u t t a l l i i ( W i l s . ) Jaeg. & Sauerb. Isothecium s t o l o n i f e r u m B r i d . S c l e r o p o d i u m c e s p i t a n s (C. M u e l l . ) L. Koch S. o b t u s i f o l i u m (Hook.) Kindb. vn Macoun S t o k e s i e l l a oregana ( S u l l . ) Robins. S. prae1o~nga (Hedw.) Robins.  Family P l a g i o t h e c i a c e a e P l a g i o t h e c i u m c a y i f o l i u m ( B r i d . ) Iwats. P_. denticuTatum (Hedw.) B.S.G. P. laetum B.S.G. V. p i l i f e r u m (Sw. ex C.J. Hartm.) B.S.G. P. undulatum (HedwTJ B.S.G.  196  Family Hypnaceae Hypnum c i r c i n a l e Hook. l i - d i e c k i i Ren. & Card, ex Roe 1 1 . \\_. subimponens Lesq. I s o p t e r y g i u m elegans ( B r i d . ) L i n d b . J_. p u l c h e l l u m (Hedw.) Jaeg. & Sauerb.  Family  Rhytidiaceae Pleurozium schreberi (Brid.) M i t t . R h y t i d i a d e l p h u s l o r e u s (Hedw.) Warnst. R. s q u a r r o s u s (Hedw.) Warnst. R. t r i q u e t r u s (hedw.) Warnst.  197  APPENDIX  IV  O r d i n a t i o n data on t h e b r y o p h y t e v e g e t a t i o n from c a l c a r e o u s  rocks.  ORDINATION: AXIS 1 2 3 4 5 6 7 8 9 10 1 1 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27  ^CA-CENT EIGENVALUE  0.. 1 170347772E+03 0..8845344522E+02 0. 6636100796E+02 0. 6478132645E+02 0, 5392733340E+02 0. 4053179768E+02 0.. 3963538360E+02 0 . 3443028234E+02 0 .3260502171E+02 0. 3161708391E+02 0. 2669218426E+02 0.. 2080173402E+02 0. 1665683272E+02 0. 1584265435E+02 0. 1501971595E+02 0. 1271493798E+02 0. 8595393369E+01 0. 8348086760E+01 0. 6880249695E+01 0. 5611985637E+01 0. 5560991204E+01 0. 3052311112E+01 0. 2238079501E+01 0. 1854'169694E+01 0. 1659231780E+01 0. 1 186387531E+01 0. 3036854776E-11  %EV 16 .20772 12 .24960 9 .19010 8 .97134 7 .46820. 5 .61310 5 .48896 4 .76812 4 .51535 4 .37854 . 3 .69650 2 .88076 2 .30674 2 .19399 2 .08003 , 1 .76085 . 1 .19034 . 1 .. 15610 0. 95282 0. 77718 0. 77012 0. 42270 0. 30994 0. 25678 0. 22978 O. 16430 0. 00000  TRACE OF XX' = 0.7220924050278433E+03 AND SUM OF EV = O.7220924050278437E+03  SUM %EV 16 .20772 28 .45731 37 .64740 46 .61873 54 .08693 59 .70003 65 . 18898 69 .95709 74 .47244 78 .85097 82 .54747 85 .42822 87 .73495 89 .92894 92 .00896 93 .76979 94 .96013 96 , . 1 1621 97 .06903 . 97 .84621 . 98 .61632 99. 03902 99. 34895 99 .60573 99. 83549 99. 99979 99. 99979  SORT EV 0.. 1081826E+02 0. 940496BE+01 0, 8146227E+01 0. 8048685E+010, 7343522E+01 0. 6366458E+01 0. 6295663E+01 0. 5867731E+01 0. 5710080E+01 0. 5622907E+01 0. 5166447E+01 0. 4560891E+01 0. 4081278E+01 0. 3980283E+01 0. 3875527E+01 0. 3565801E+01 0. 2931790E+01 0. 2889305E+01 0. 2623023E+01 0. 2368963E+01 0. 2358175E+01 0. 1747087E+01 0. 1496021E+01 0. 1361679E+01 0. 1288112E+01 0. 1089214E+01 0. 1742657E-05  SCALE 99 .99998 86 .93604 75 .30069 74 .39905 67 .88078 58 .84917 58 . 19476 54 .23912 52 .78186 51 .97606 47 .75671 42 . 15918 37 .72581 36 .79225 35 82393 32 .96094 27 .10037 26.,70766 24 .24625 . 21 .89781 . 21 .79810 . 16 .14941 13. 82867 12. 58686 11 .90683 10.06829 0. 00002  ORDINATION: SAMPLES  N 1 2 3 4 5 6 7 8 9 10 1 1 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 MIN MAX ZERO  NAME B36 B143 B144 B145 B146 B147 B146 B149 B150 B151 B152 B153 B163 B164 B165 B166 B167 B168 B277 B278 B279 B280 B281 B282 B283 B284 B285  AXIS 66 26 3€ 50 21 24 44 61 36 3 37 80 93 98 94 83 100 68 33 31 26 9 19 14 33 0 8  1  2  35 59 71 80 31 26 100 71 81 63 61 58 36 38 52 1 1 9 OOOO 6938 0 3277 9 9602 18 3815 18 1590 26 1 176 27 6789 25 8506 18 17 0 9835 35 0030 2603 4258 4282 3299 4550 7655 2145 4539 5636 6696 8430 3895 9754 8663 5083  59 100 47 46 46 50 OOOO 67 64 4598 5256 17 9928 0 7573 37 7905 53 7245 69 1574 70 8111 62 5155 31 23 7192 28 0 59 7351 7065 53 4446 62 73 8562 3289 60 64 6669 7716 60 1301 60 5084 39 6590 1064 7089 4470 3806 9038  3 8346  OOOO 6094 9423 2422 2728 2378 0598 5231 0 1 138 3360 6514 3888 8172 0888 6481 0138 7458 4055 9720 6195 8359 3800 6374 0098 5620  26 87 53 94 54 54 100 65 38 14 7 0 3 1 1 20 89 97 95 32 40 32 60 53 53 35 36 31  SCORES  4  5  6  91 16 8431 5039 2649 5822 8831  8629 5316 9131 5486 4780 7983 8990 6609 4417 8954 8278 1447 3161 0537 2860 8592 9194 1735  OOOO  1 46 52 69 9 7 OOOO 69 3617 54 5738 0 3559 87 8 4050 36 0 1 193 43 57 7809 41 5123 0513 53 0443 42 5070 30 4831 100 6673 76 0479 75 7 8522 4068 13 1944 0 79 5916 4922 62 7584 57  100 0 53 55 62 56 52 67 4 43 26 43 26 31 32 38 1 1 50 OOOO 54 41 1382 36 8416 2431 29 2324 40 38 0 0765 39 33 5152 1403 30  PCA-CENT  RANKED  7  18 8461 27 9370 0 5233 58 2319 8857 100 OOOO 8040 72 2147 9247 59 1054 0397 51 1418 0627 5 5325 0432 0 0 6296 19 1456 78 6231 3210 1 148 67 1834 5457 47 3408 7446 55 7872 64 7493 2026 4869 40 8244 49 5281 73 16 49 2352 0632 27 4161 3182 3269 48 0 9 6 0 27 4426 3524 0414 40 8096 6389 43 1 162 •1 169 41 9849 1343 31 3225 2157 83 4620 9834 86 1852  26 10 27 22 24 23 5 6 2 21 20 19 25 3 9 1 1 7 4 8 1 18 12 16 13 15 14 17  B284 B151 B285 B280 B282 B281 B146 B147 B143 B279 B278 B277 B283 B144 B150 B152 B148 B145 B149 B36 B168 B153 B166 B163 B165 B164 B167  1  RANKED 2 0 3 8 9 14 19 21 24 26 26 31 33 33 36 36 37 44 50 61 66 68 80 83 93 94 98 100  0 5636 9835 1590 6789 1 176 3299 4550 2603 3815 9602 3277 8506 4258 4539 6696 7655 4282 2145 0030 6938 8430 5083 3895 8663 9754  OOOO  18 17 19 16 26 21 20 25 24 22 6 23 5 27 1 13 14 15 12 2 1 1 10 8 3 4 9 7  B168 B167 B277 B166 B284 B279 B278 B283 B282 B280 B147 B281 B146 B285 B36 B163 B164 B165 B153 B143 B152 B151 B149 B144 B145 B150 B148  0 9 9 1 1 17 18 18 18 25 26 26 27 31 35 35 36 38 52 58 59 61 63 71 71 80 81 100  -0.2822838E+00 0.3495352E+00  -0.3020527E+00 0.4462833E+00  -0.5052854E+00 0.4616321E+Q0  -0.3072802E+00 0.3352695E+00  -0.2924408E+00 0.3734319E+00  -0.3974504E+00 0.5787736E+00  -0.3868844E+00 0.4195840E+00  0.4467795E+02  0.4036324E+02  0.5225732E+02  0.4782201E+02  0.4391841E+02  0.4071303E+02  0.4797266E+02  0 7192 7351 5155 1301 4446 7065 7716 6669 8562 9038 3289 3806 5084 6590 7245 1574 8111 7905 1064 7573 9928 4598 7089 4470 5256  OOOO  ORDINATION: POLAR OF SAMPLES USING PD ENDPOINTS 26 AND 17 AUTOMATIC FIRST AXIS (BASED ON ORDINATION NUMBER BASELINE* 0.94737E+02 AND PD =100.00000 - PS  N 1 2 3 4 5 € 7 8 9 10 1 1 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27  NAME B36 B143 B144 B145 B146 8147 B148 B 149 B150 B151 B152 B153 B163 B164 B165 B166 B167 B168 B277 B278 B279 B280 B281 B282 B283 B284 B285  X  XREL 73 53 53 54 33 48 63 69 65 49 54 77 76 81 83 93 99 91 40 38 32 34 32 36 38 0 22  .37032 .14667 .15382 .85742 .59898 .3994 1 .43445 20155 .82291 .25737 40120 01830 74149 50082 53162 92409 99994 14244 51532 83049 98726 79553 80728 38588 87265 0 13565  0 0 0 0 0 0 0 0 0 0 0 0 O 0 0 0 0 0 0 0 0 0 0 0 0 0 0  6950874E+02 5034949E+02 5035628E+02 5197020E+02 3183063E+02 4585210E+02 6009581E+02 6555939E+02 6235857E+02 4666489E+02 5153799E+02 7296472E+02 7270248E+02 7721133E+02 7913524E+02 8898074E+02 9473682E+02 8634549E+02 3838295E+02 3678679E+02 3125110E+02 3296420E+02 3108060E+02 3447084E+02 3682674E+02 1 1 0 5 9 4 6 E - 10 2097063E+02  DI  E 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0  7189250E+02 7156238E+02 7106398E+02 6790128E+02 7773550E+02 7975952E+02 7265181E+02 7551135E+02 7817549E+02 8252521E+02 7819402E+02 6838237E+02 5918958E+02 5438899E+02 5026601E+02 3118045E+02 3053246E-04 3863406E+02 5169130E+02 4310268E+02 5285600E+02 7089256E+02 6825682E+02 7126794E+02 4819656E+02 4577635E-04 4986845E+02  0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 -0 0  10O000OE+O3 8749998E+02 8709676E+02 8550723E+02 8400000E+02 9199998E+02 9428571E+02 1000000E+03 1000000E+03 9480519E+02 9365079E+02 1000000E+03 9374998E+02 9444444E+02 9374998E+02 9428571E+02 9473683E+02 9459459E+02 6438356E+02 5666666E+02 6140349E+02 7818181E+02 7499998E+02 7916666E+02 6065573E+02 4577637E-04 5409834E+02  RANKED XREL  D2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 -0 0 0 0 0 0 0 . 0 0 0 0  7619048E+02 8421051E+02 8378378E+02 8024692E+02 1000000E+03 9354837E+02 8048781E+02 8095238E+02 8461537E+02 9550562E+02 8933333E+02 7176471E+02 6315788E+02 5714285E+02 5263156E+02 3170731E+02 3051758E-04 3953487E+02 7647058E+02 7222221E+02 8260869E+02 9402985E+02 9333333E+02 9333333E+02 7534247E+02 9473683E+02 8904109E+02  7)  26 27 23 21 5 22 24 20 25 19 6 10 2 3 1 1 4 7 9 8 1 13 12 14 15 18 16 17  B284 B285 B281 B279 B146 B280 B282 B278 B283 B277 B147 B151 B143 B144 B152 B145 B148 B150 B149 B36 B163 B153 B164 B165 B168 B166 B167  0 22 32 32 33 34 36 38 38 40 48 49 53 53 54 54 63 65 69 73 76 77 81 83 91 93 99  0 13565 80728 98726 59898 79553 38588 83049 87265 51532 39941 25737 14667 15382 40120 85742 43445 82291 20155 37032 74149 01830 50082 53162 14244 92409 99994  ro O O  201  IS £ 2  T 5 2™ininincop~Tro3aiinoiuJOOTPjp-r--oico  m  X  PI IX) 1 10 p~ in r- pi co co o — rt Pl p » 03 03 L0 CO CP CO 00 LO CO co CP oo * C Ol Ol Ol M OJ *~ »co 00 00 Ol CO 00 00 CQ 00 oo 00 00 00 m 00 00 m CO m  0) 01 co in Ol CO P~ in co in Ol Ol m 03 00 CO 00 00 00  in •<T CO OJ ro  _  10 CM  2 <  or  t  Ol  Ol  PI *-  CN CN  OOOOOOOOOO  in CO LO r~  0)  OJ *-  in o  o •— in in co m  00 Ol P) pOJ  Ol Ol OJ  o  Cl  oi 0 O 0 O oi oi O + + + + 0 O + LU U J + + U J UJ in o> 01 — LU oi 01 — LP 01 *r co io in T in O in r- TT 0 oi 0 oo in o CO OJ co ••- 0 — to •s- in •<r in in 00 00 01 01 01 01 in  CM oi OJ CM CO CO CO OJ o i OJ O O O O O O O ++ + + O O O + ++ U J U J UJ U J LU U J U J O 01 in m O O O CO 01 00 CO *T LO O O O — 01 CM ID — r- O O O p~ co in CM LO in O O O co o t~ O <T in co O O O r- co oi o CO OJ O O O co v CM in LO co o p*01 o CO 03 00 o o o O O o o O O O O o O O o o o  C M C O O I O I P I P I C M C M C M T Ol CM Ol Ol  + + + + + + + + +i  L U U I U J U J U J U J U J U J U J U J  inoo^OOcocoinoo OQOcoQQcoojoiin p-OQinoOcMcoOP~ t O O U O O C l B N -  K O O n o O M n i s i n i^OTinOOOOOlO Ol —  a.  CO CO CO CM OJ CM Ol OJ OJ OJ CM OJ OJ CM CM CM CM CM  in o  ~ Q  COOIOTPOCOOOJ  UJ  O o o O O O O ++ ++ +++  UJ  o o o *o o o O O O o O O O O o O O O o o o  O 11 UJ  v> a a.  --  CM OJ Ol CM CM TT OJ OJ OJ 1 CM Ol CM OJ OJ CM CM CM CM Ol CM CM CM CM Ol OJ Ol  0 o  O O O O O O 0 O O O O 0 O o o O O + + o oo o + ++ O O O + + + + + + + + ++ + + + O + ++ UJ UJ UJ U J U J U l UJ  £  UJ I P- CM — CO UJ  in co — in in i0 - o oi r~ 1 oo 01 in r- co in co CO co 1 L0 01 in — UJ ot CM CO CM CO cc 01 OJ LP co CO GO r* LP co p- p-  O Ul 1-  <  X  <  i/i O  o  oo o  O O O O + + + + UJ UJ Ul L0 01 pCO L0 •*r CO co CO T L0 O CO LO co O CO CM in CO LO 0) T—  UJ U J UJ UJ UJ U J L0 co in co un CO P- in OJ CO Ol P l Ol O oo 01 01 Ol p~ 01 oi oo o r- co CO ID CP in T 01 *T L0 L0 O in o oi oo Ol 01 O) L0 CO 01 01 pp- 01 Ol Ol  O O O CO O Q o 01 O O o 01 01 Q  OoiOinrooiOOi - o i O ^ n c i O O i  Z CJ a »—4 •-i z l/l i < - < 3  LL.  UJ  TcoOcocoinoOi TCiOcnoioiQOi  < o o o 6 z o o  C3  ui _i Cu  O O o o O O o + + + + + + UJ + UJ U J UJ UJ UJ CO 00 CO oo 01 —— CO 01 p~ O o 01 CM 01 01 in CO  U J U J U J U J U J U J U J U J I  x 5  a CL  C O C O ' - ' - C O O l L D' Pol 1  UJ UJ  Ul  UJ UJ  UJ U J  UJ UJ UJ UJ UJ UJ UJ  LP CO ID CO CM LP Pl LO n co CM Ol 1 01 - ID ro CO LP O O) T 01 r- in p- in co oi CM CO O CO co co in CM CO in — O - 01 in O oo O CP co O co in O in LP LO CO 03 p» ID OJ UJ O) O oo co — O 01 LP T 01 — P- CM in i CO 03 rr CO in oi O OJ in t- Ol tD O in UJ O in O -=r P» CD in oo oi O — CM *O UJ co co LO CO f - I- p- r- i - p» co ID P- P- o oi r- L0 p*- 03 P- LP  O  co ^  in *-  P~ P~ UJ  O O O O O O O O O O O O O O O O O O O O O O O O O O O  Pl  o  + or a U J < Z O _i < o o CL 10 O  OIOJCMOI —  O C M C M C M O I O I C M O I C M O I Ol Oi CM Ol  0 0 0 0 0 * - 0 0 0 0 0 0 0 0 0  *~  o i c o o i ^ c o c o o - i D O i o o p ^ o u * i c o i n 03 t - CM CO i n o i o i p - c o c o o c o o o i c o o c o c M c o CO CO CM in  o  c p o i o i i n i n c o o i c o i n o i T r o o i c o o i in oi •fl^ *uj^'-p~inO'-u5LooicMcop-inp03 *- o  Z o to UJ n M K I- Z Z z a _i >-i a. UJ OQin cc z < o U J aa  ,  U J C 0 P - O i n ' - r T 0 1 P ~ 0 1 P ~ O l C 0 P - C 0 p~ oo LP CM inoiTjoinoip-ooinoiLPoip-^p*  T rO n iOD iOn iOn o ujO i nO r -O - oOi uO> O u 3i^ ^ * r O Oi O  I  Ol CM CM CM CM CM CM  O O O O ' O O O O O O O + + + + + + + + + + + UJ I  + + + + + I + + + + + + + + + UJ UJ UJ U J U J U J U J U J U J U J U J U J U J U J U J U J U J U J  UJ UJ  UJ  Ol i p~ in pp~ I Pl CM I CO oo CO co i O CO Ol CO I LP in Ol co CM in i Ul CM I  ,  UJ  !-T  UJ  UJ U J  co in co 03 CO Ol Pl CM 01 UJ P- CM p- CO CM O) in •sr UJ p- in CM in *r LP  O o O OOO O  oi oo oi T in oi oi P~ in oi oi in in LP — ' - p ~ i n  p-Locncot^oincoincor-cMooinoiinTrp-ooinpj OC0O01C0Q00CMC0r>00CMinL0P~ — P-tcOOOCM o i o B i n o i T r o o i c o o i i n o^i '.j -^. p-oico^copicMoiLO — cpipoioioop-inp- — v - c i i t O i o n ^ u i S ( M i P C ^ O i n O T O i p - o i P - o i c o r - P i c o — p-ror-eoincMoip-cooi £ S J ^ 9 ^ ® ^ ^ ^ ^ 3 < c n c p c o » Q ^ ' ^ ^ ^ ^ n UJ in io in p. oi CP co * ^ ^ ^ ^ CM in in in CM CM OI in ? 3J UJ  m  < z  o  m  r  o  w  M  0  u  ao p- 03 UJ p~ rOl CM oo 00 00  CO 01  O  01 O CM CO r- CO oo CO 00 Ol CM OJ CM CM 00 00 00 00 00  l  f CO OJ m  in CO CM OQ  Ol CO T in UJ P~ CM CM CM Ol CM CM CM CM  202  APPENDIX V  D i s t r i b u t i o n maps o f the b r y o p h y t e s y n t a x a .  203  R a c o m i t r i o - Andreaeetum  petrophilae Frey  1922  204  205  206  207  208  209  210  211  212  213  214  215  216  217  219  220  221  222  223  224  225  APPENDIX VI  T a b u l a r s o r t i n g t a b l e s and  c l u s t e r a n a l y s e s f o r the f o u r major  groups o f b r y o p h y t e v e g e t a t i o n  releves.  226 BRYOPHYTE ASSOCIATIONS FROM EXPOSED, DRY ROCKS. Releve-numbers  "translation"  list.  ( i n the l e f t column are the numbers as given by the c l u s t e r a n a l y s i s ; i n the r i g h t column are the f i e l d releve-numbers as l i s t e d i n the s e p a r a t e association tables.)  1 3 6 52 26 53 10 62 64 63 115 117 83 84 85 21 8 116 51 36 37 54 55 56 20 128 126 129 127 2 69 58 5 75 78 79 22 33 35 34 57 77 67  = 9 = 11 — 15 — 179 = 119 = 181 26 = 243 = 245 — 244 = 331 = 333 — 288 = 289 — 290 = 65 zz 17 zz 332 = 178 = 134 — 135 = 182 — 222 — 223 = 46 zz 347 - 345 = 348 = 346 10 — 252 — 236 = 14 258 = 264 265 = 68 — 131 — 133 zz 132 — 235 — 263 = 248  68 71 72 74 76 4 9 11 12 13 40 41 81 82 89 91 90 92 93 94 123 125 124 98 100 99 101 7 17 16 19 18 86 88 87 109 110 111 112 113 14 15 38  = = = = = = = = = = = = = = = = = = = = = = = = = = = = = = = = = = = = = = = = = = =  249 254 255 257 262 12 22 27 31 32 157 158 286 287 294 296 295 297 300 301 341 343 342 305 307 306 308 16 38 37 44 43 291 293 292 316 317 318 319 320 33 35 154  39 23 24 42 43 29 30 47 48 46 65 66 59 60 61 49 50 118 119 122 120 121 104 105 108 106 107 25 31 102 114 103 32 80 70 95 96 97 27 28 73 44 45  = 155 = 116 = 117 = 169 = 170 = 122 = 123 = 174 = 175 = 173 = 246 = 247 = 240 = 241 = 242 = 176 = 177 = 334 = 335 = 338 = 336 = 337 = 311 = 312 = 315 = 313 = 314 = 118 = 124 = 309 = 330 = 310 = 125 = 266 = 253 = 302 = 303 = 304 = 120 = 121 = 256 = 171 = 172  227  BRYOPHYTE ASSOCIATIONS FROM SHADED, HUMID ROCKS. Rel eve-numbers " t r a n s l a t i o n "  list.  ( i n t h e l e f t column a r e t h e numbers as g i v e n by the c l u s t e r a n a l y s i s ; i n t h e r i g t h column a r e t h e f i e l d releve-numbers as l i s t e d i n t h e s e p a r a t e association tables.)  1 = 1 2 = 2 4 = 4 45 = 75 46 = 80 65 = 180 32 = 59 48 = 95 52 = 100 23 = 45 26 = 50 33 = 60 28 = 54 29 = 55 34 = 61 3 = 3 36 = 63 51 = 98 53 = 101 8 13 13 = 23 17 = 29 74 = 272 75 = 273 14 - 24 44 74 31 = 58 63 = 140 62 = 139 39 = 69 38 = 67 35 = 62 55 = 103 5 = 5 6 = 7 69 = 230 9 = 18 10 = 19 64 = 156 49 = 96 50 = 97 7 = 8  -  -  70 82 83 54 67 68 18 24 42 40 41 61 11 21 22 43 58 47 16 25 27 30 76 77 66 37 59 60 19 20 56 57 78 81 79 80 12 15 71 72 73  = 231 = 339 = 340 = 102 = 225 = 226 = 30 = 47 = 72 = 70 = 71 = 138 = 20 = 41 = 42 = 73 = 114 = 82 = 28 = 49 = 51 = 57 = 298 = 299 = 214 = 64 = 136 = 137 = 39 = 40 = 112 = 113 = 321 = 324 = 322 = 323 = 21 = 25 = 237 = 38 = 39  228 BRYOPHYTE ASSOCIATIONS FROM SILICEOUS ROCKS ALONG STREAMS. Releve-numbers " t r a n s l a t i o n "  list.  ( i n t h e l e f t column a r e t h e numbers as g i v e n by t h e c l u s t e r a n a l y s i s ; i n t h e r i g h t column a r e t h e f i e l d releve-numbers as l i s t e d i n t h e s e p a r a t e a s s o c i a t i o n t a b l e s . )  1 86 88 24 25 26 27 80 6 36 94 38 97 98 33 34 35 66 67 78 79 68 69 15 19 17 16 18 96 101 62 64 91 93 92 81 87 83 99 100 82 51 52 56  = 6 = 259 = 261 = 104 = 105 = 106 = 107 = 229 = 66 = 129 = 274 = 159 = 235 = 326 = 126 = 127 = 128 = 210 = 211 = 227 = 228 = 212 = 213 = 86 = 90 = 88 = 87 = 89 = 276 = 329 = 206 = 208 = 269 = 271 = 270 = 232 = 260 = 234 = 327 = 328 = 233 = 195 = 196 = 200  53 = 197 54 = 198 55 = 199 63 = 207 89 = 267 90 = 268 70 = 215 95 = 275 71 = 216 74 = 219 73 = 218 75 = 220 72 = 217 76 = 221 2 = 48 32 = 115 11 = 81 59 = 203 12 = 83 102 = 344 47 = 191 48 = 192 50 = 194 58 = 202 61 = 205 7 = 76 8 = 77 31 = 111 10 = 79 13 = 84 9 = 78 21 = 92 20 = 91 49 = 193 22 = 93 23 = 94 57 = 201 60 = 204 3 = 52 4 = 53 5 = 56 37 = 130 14 = 85 46 = 190  84 85 42 45 28 40 29 30 65 39 41 43 44 77  = = = = = = = = = =  250 251 186 189 108 184 109 no 209 183 185 = 187 = 188 224  229  BRYOPHYTE ASSOCIATIONS FROM CALCAREOUS ROCKS. Rel eve-numbers " t r a n s l a t i o n "  list.  ( i n t h e l e f t column a r e t h e numbers as g i v e n by t h e c l u s t e r a n a l y s i s ; i n t h e r i g h t column are t h e f i e l d releve-numbers as l i s t e d i n t h e separate association tables.)  1 13 14 15 11 12 16 17 18 2 3 8 4 7 9 10 5 6 22 23 24 19 20 21 25 26 27  = 36  = = = = = = = = = = =  = = = =  = = = = = = = = = =  163 164 165 152 153 166 167 168 143 144 149 145 148 150 151 146 147 280 281 283 277 278 279 283 284 285  230  1 RACOHETH POLYPIL  MARSEMA GYMNOBT CEPHTUR DICRCRI S-LEOBT DICHPEL POROBIG 10 POLYALF'A  11 MAR55PA 12 BRYUCAE 13 RACOCAN 14 CEPHBYSB 15 Of JOdEN 16 SCAPAME •i 7 I.OPHSUD -18 CAMPPAR 9 ANDRRUP :.0 POHLNUT ;:1 HYPNCIR :.2 HYPNSUB '.-3 CLAOBOL 24 DICRSCO 15 HERBADU S.6 l.OPHVENV 17 CHANSET 28 HEDWCIL 29 DICRCRI 30 CEPHBYSA 31 AULAAND 32 BARBFLO 33 POLYdJIM 34 CERAPUR 35 LOPHEXC 36 LOPHINC 37 PREIQUA 38 BRYUCAP 39 BARBHAT 40 RACOLAN 41 PLEUSCH 42 BARTPOM 43 BAZZDEN  44 ANASMIN 45 LOPHCUS 46 CAMPATR 47 ANASASS 48 CHANFIL 49 ANDRROT 50 PARAENE 51 DICRFUS 52 RACOBRE 53 "DIPLTAX 54 RACOHETA 55 GYMNCON 5G ANDRNIV 57 MARSSPH 58 OLIGHER 59 BLINACU 60 RACOACI 61 KIAESTA 62 RACOSUD 63 MARSSTA 64 LOPHWEN 65 ISOPELE 66 POHLCRU 67 PSEUSTE 68 POHLLUD 69 PSEUBAI 70 BARTITH 71 dUNGOBO 72 HYGROCH 73 PELLNEE 74 ANOEAES 75 GRIMALPH 76 SCAPSCA 77 CHILPOL 78 DITRHET 79 OLIGALI 80 PLAGOED 81 PSEURAD 82 SCHI PEN 83 DRYPPAT 84 PORENAV 85 GRIMTOR 86 DISTCAP 87 ENCAPRO 88 HOMANUT 89 SCHISTR 90 ATRIUND 91 RACOVAR 92 ANIDRBLY 93 GRIMMON 94 AMPHMOU 95 BRACSTA 96 PSEUINC 97 AMPHCAL 98 DICRHET 99 PTILCAL 100 RACOFAS 101 DIPLALB 102 BARBLYC 103 BAZZTRI 104 KIAEBLY 105 BRACALB 106 PSEUPAT 107 ANTHdUR 108 MARSCON 109 NARDdAP 110 TRITOUI 111 CEPHBIC 112 P H I L F O N 113 PLEUALB 114 MARSBRE 115 KIAEFAL 116 POLYSEX 117 POHLCAR 118 ANEUPIN 119 BARBVIN 120 AMPHLAP 12 1 RADUBOL 122 HETEDIM 123 DOUIOVA 124 ANACMEN 125 HOMANEV 126 PSEUCAL 127 LOPHHAT 128 PLAGPIL 129 PTERFIL 130 TORTPRI 131 PTILPUL 132 PORECOR 133 PSEUATR 134 GRIMDON 135 HYPNREV 136 SCAPPAL 137 SCAPULI 138 dUNGEXS 139 NARDBRE 140 HYPNDIC 141 GRIMALPA 142 CAMPFRA 143 ARCTFUL 144 dUNGCON 145 RHYTTRI 146 TIMMAUS 147 PTILCIL 148 PLAGASP 149 HETEMAC 150 POREROE 151 HETEPRO 152 ISOTSTO  1111111111  12345678901234567890  21-1 1  2  3 3  2  _  134  -323 43222-132443 _ S--323  1---  5-4--4--1 + 1 1 1 " 1 3 3-"1- + -++ — 2--S+ 2-3 + --3 + --22-1 1 2 — 21-42__ 1--3 4 1+ + -1 -51 -21 — -.  1-111 322111--  1  4-- " " " - — 1 4  —  3-121  --23-2  1211  43233321 1 1—2+1 1 1 + 1---2322 +31--1212  __  1-1-+1111  2  _  112  _-,  -+  - 4+13-2 -- + -— 3 + — 1-- + ---32-2-51344321 1 22 + 1-2- 11----+  +  +1 --++++-12++ 31-2-2 +1 1 2 1 131 --1 1 1  32 1 + ---+1135212-71 ----412+ 211--2 +  +1  ---32  1  —  1  22 — 322 2  t2  11- --11 —  21 1 1 1  11 1-  ---21-121  1-+-  1 14 + -1 2 _4*_  3  + 2  1  5  .1--  + 2  +  2  12+11  +1+111--+1--  —  2232  411  +11 -+  1—-  1211  1131 + -21122+12  _.  -  +  11 —  2122++21 + + -1++++--2  112—1  •1 •1-21-11  --1-+1-1312+-  13-3--1---  1 121 1 '  32--2+213  +1 12331 — 1 21 + - — 1 + 211 121 1 —  _323  •--31  1  H  1 +++ 1 + 1+1  31211 111 1-+1  1  1 13  11-  1--11 — 1231 11 11---222114-112  1 11-122 2 1 — 1-1-  31-2-2 42 — 3-1 ---2-32222 + 1_+1 __11 + ++ 1-2221 + 21-111 -4 —1-3 — ++  1+-  1  • -.  223-2+3  12-32-2 -21 43-1  1  .  ++21+  •1  111 1  ' -  -  1  124 + +12  3  —  -  1  111  --1-  1 — 1--  2-  2  12 4  -1---1 . +  —  •132 —  1  -  "  1  1 + 1 — 1 1-  --+1+  -+2-  +-—1111  4324 1--1 -11-1 -++-+ -2-22  3  - + --+2-  •13342-32-  41 12552233-4224312-232-11311--23121 + 21 11-3+1 1-  -43333-3 21-11111- •121  •1. 3 .1 -123231-1—2212131321-44 11- 1--2 — -13 — -1++-  •1+2-  -423 — -4411 — 1 1 1 - 2111-  -2423323-44-2144-  43__ _2+- + --24223333-3 —  -31 1  •12211•11-2221+•11 1+1 + -++-1-111+1411-+13313 21 1 1222-  2  22-24— 233-31 •12-3  23  21233 3223-311 11 — 1-21 — 1 4 — 1 1 4 11-  -1111221 112131 11-41 1122+-  •1-1-  •11 — 1-  -++1•1--+•11 + 1-  -31•1-  •1-1 —  •11-  •11-  •11• 1-•11•1-1222-  •1-  + 2•12-  -+-++1-  •141  —  •12-22-3-121111 — 1 -3 — -31-  11111 —-221-  ._  •113-  1 1 1 1-221  -31 — 1-  12111 1---11  -1 — -1 + -21-  •111-  -3-1--2-  •1 —  •111-  •11-  1+--1-  -211-  •1 1 - 2 : --2-2-12321 • 1  134•1 + 11  •1-  -221-  22++1+++12 — 31--- 11-  •1-1-1  -4333--  J  -11+3-1-2 — -1 + -  • — 2211-..-  —1++  —  — -1-1-  —  -31-1 — -1-•11-33•11-1-•13-22+ 313 -111 1 1 — ---1 -11-1-2 — --11 1 --11 :>.1211 132122—1211-+1-  -11-1211-1-+13 1311--2-4 1 1 +2— -  >0  •1211•1+-  •1-+-  •11+---+11-+1+-  0  °  p-  m  \  231  2 19 ANDRRUP 62 RACOSUD 2G LOPHVENV 53 DIPLTAX 3 MARSEMA 17 LOPHSUD 55 GYMNCON 20" POHLNUT 61 KIAESTA 16 SCAPAME 1 RACOHETH 52 RACOBRE 4 GYMNOBT 13 RACOCAN 10 POLYALPA 14 CEPHBYSB 32 BARBFLO 56 ANDRNIV 54 RACOHETA 81 PSEURAD 6 DICRCRI 22 HYPNSUB 76 SCAPSCA 2 POLYPIL 66 POHLCRU 83 DRYPPAT 42 BARTPOM 24 DICRSCO 15 CYNOdEN 23 CLAOBOL 85 GRIMTOR 107 ANTHUUR 30 CEPHBYSA 34 CERAPUR 11 MARSSPA 44 ANASMIN 49 ANDRROT 70 BARTITH 28 HEDWCIL 36 LOPHINC 51 DICRFU5 104 KIAEBLY 109 NARDUAP 125 HOMANEV 89 SCHISTR 94 AMPHMOU 98 DICRHET 112 PHILFON 133 PSEUATR 39 BARBHAT 69 PSEUBAI 99 PTILCAL 110 TRITOUI 128 PLAGPIL 21 HYPNCIR 33 POLYJUN 45 LOPHCUS 46 CAMPATR 48 CHANFIL 65 ISOPELE 101 DIPLALB 108 MARSCON 126 PSEUCAL 129 PTERFIL 148 PLAGASP 8 DICHPEL 38 BRYUCAP 68 POHLLUD 40 RACOLAN 41 PLEUSCH 50 PARAENE 63 MARSSTA 75 GRIMALPH 87 ENCAPRO 92 ANDRBLY 111 CEPHBIC 113 PLEUALB 120 AMPHLAP 130 TORTPRI 135 HYPNREV 137 SCAPULI 12 BRYUCAE 25 HERBADU, 47 ANASASS 72 HYGROCH 74 ANOEAES 124 ANACMEN 127 LOPHHAT 131 PTILPUL 134 GRIMDON 138 JUNGEXS 150 POREROE 151 HETEPRO 5 CEPHTUR 7 SCLEOBT 18 CAMPPAR 97 AMPHCAL 27 CHANSET 31 AULAAND 35 LOPHEXC 58 OLIGHER 59 BLINACU 60 RACOACI 64 LOPHWEN 105 BRACALB 106 PSEUPAT 71 UUNGOBO 78 DITRHET 79 OLIGALI 84 PORENAV 88 HOMANUT 91 RACOVAR 93* GRIMMON 115 KIAEFAL 121 RADUBOL 123 DOUIOVA 132 PORECOR 136 SCAPPAL 141 GRIMALPA 142 CAMPFRA 143 ARCTFUL 145 RHYTTRI 146 TIMMAUS 147 PTILCIL 9 POROBIG 29 DICRCRI 37 PREIOUA 43 BAZZDEN 57 MARSSPH 67 PSEUSTE 73 PELLNEE 77 CHILPOL 80 PLAGOED 82 SCHIPEN 86 DISTCAP 90 ATRIUND 95 BRACSTA 96 PSEUINC 139 NARDBRE 140 HYPNDIC 100 RACOFAS 102 BARBLYC 103 BAZZTRI 144 JUNGCON 114 MARSBRE 116 POLYSEX 117 POHLCAR 118 ANEUPIN 149 HETEMAC 119 BARBVIN 122 HETEDIM 152 ISOTSTO  11111111111111111111111 n 11111 1 1 1 1 1 1 1 1 112222222222333333333344444444445555555555G666666GG677777777778888888888999999999900000000001 111 11 1 1 1 12222P22222 12345G7890123456789012345678901234567890123456789012345678901234567890123456789012345G7890123456789012345678901 234567890123456789  31-2-2  3-1 2-32222 22--322 3121 1111 21111 111 21-121 1121 1 2122 + + 21 + 2232 31321-441 1-1 233-31-12-3 23 1-21--14--1 1 4 11 122 111213111-4 11 122 + 4 114+-1 13-3 1-+1-1312 + 1-21-11 11 12+11 1211 1 + 1131+-21122+12 11111 — 12 21 2-232-1 131 1--23 12 1 + 21 1 1-3+1 1 1- + 1 1 1 1 1 + 1 + - + + -1 21-42 22-1 21-1 1 32443 1-1 1 1 43233321 1 1--2+1 1 1 + 1 2322 4 12+ 21 1--2+ 2 1 — 1-1 1231 1 1 1 1 2221 14-1 12 21 + 1 + 21 1 121 1 43--1-2+- + ---2423323 1 423---44 1 1 — 1 1 1 --31 1 21 1 1222 13313 +1-+1++—11 12+ ++ 1-2221+21-1 1 1 2 1-+1 1 + -1+++ + --2 123231-1--22121 22-24-21233 3223-31 1 1 1 11 1 1-1 321 + +1 135212-21 . 1 1 1-122 1 1 . 1 323 1233 1 - - 1 5-4--4--1 + 1 11 13 3 1 43222 323 134 23-2 121 1 __ 4 3333 41 12552233-4224312 r-' 1 1-2221+ 1--3 4 1+ 33 -3--323 221 3221 1 1 • + 31 1212 32-2-51344321 1 1 21 131 1 11 1 1 1 13 --+ 3 4+13-2 +1 --++++- 12++ + -- - 1 + •-+ + --++ + -- 1----22+- •1- 2-11.u — ! : 11 + 1+++1 + -2+3•12 — 12-32-2 11-11 -1-++21+ -24223333-344-214 ! 2111 v 44 . • 33 -21 — -1+1411-+ •--3-121 1 1 1--1,_. -11111---221 -51-14 3-1211-1-+1111. -3-1-1-+1+111 12--1 , _ •1-1222- + -++1 •11•141 — -3+- -2-3•1•1-+2-2-++1-+1 11 •1 13•1111-221•11 + 1--1 1 +1 + • 132•11 — -32--2+21-+11-23-3-1-1 + 1•1-411 — 121111 11-4333--+ +•122++- + 2223 •1 + 1  42  2  3  1 +  2  4  1  1  •124 + -32-  •11-2 -—112  2-  -221  1221 1-  1 - in 11+3-1 —  -111 1 1---3 13 •111 3-1--2-211-  -2-2-12-  22 1 1 •11-1211-1111-  -11+1-321-  -23 1  11-  •1-1-  •134•11  212-  •1111++  -4324 -11-1  1-  •1 + 11  -+1+++•11-1-2 1 1  132-  • 1 1+ •112 -+12-  -31-2+-2-22  -111-33-  • 11 •122--1-  •1++•1311•11+21--1 - + 2-  •22+•111-  -21 1 -+13-+1 1 -+1 +  + -1-21-+1-  12-23-1  —  -----—'iT-11-13-  -+1--2-4-  •12•11• 1+-  KJ  ( !* c<-i'( >•••  o4"4  Do  ID  u  P\V>7 v >AI V  <CLUSTEI? MAX=*>  CLUSTER  OPTION=MINVAR  ANALYSIS  CASEWISE USING:  N=129 1.V1 .  18.V18. 34.V34. 50.V50, 66.V66, 82.V82. 98.V98,  CASE/* 1 3 6 52 26 53 10 62 64 63 1 15 t 17 83 84 85 21 8  CASES=1-129  VAR=1-152  CASES=CASE/f : 1- 129  2.V2,  3 . V3,  DI S T A N C E = E U C L I D E A N  4.V4,  5.V5,  6.V6,  7.V7,  8.V8.  9.V9,  20.V20. 21.V21 , 22.V22, 23.V23. 24.V24, 36.V36. 37.V37, 38.V38. 39.V39, 40.V40. 52.V52. 53.V53. 54.V54, 55.V55, 56.V56, 6 8 . V 6 8 , 6 9 . V 6 9 , 7 0 . V 7 0 . 71 .V7 1 , 7 2 . V 7 2 , 84.V84, 85.V85, 86.V86, 87.V87, 88.V88, 100.V100, 101.V101. 102.V102, 103.V103.  112.V112, 125.V125, 13B.V138. 151.V151,  1 1 3 . V 1 1 3, 126.V126, 139.V139. 152.V152  1  1  1  2 8  2 4  1 0 9 7  1 14.V 1 14, 127.V127, 140.V140,  115.V1 15. 128.V128. 14< . V 1 4 1 ,  10.V10.  12.V12.  13.V13.  14.V14.  116.V116, 129.V129, 142.V142,  117.V117. 130.V130, 143.V143,  118.V118. 131 . V 1 3 1 , 144.V144,  119.V119. 132.V132, 145.V145,  15.V15,  16.V16,  17.V17,  30.V30, 31.V31. 32.V32. 33.V33, 46.V46. 47.V47. 48.V48. 49.V49, 62.V62. 63.V63, 64.V64, 65.V65. 7 8 . V 7 8 , 7 9 . V 7 9 , 8 0 . V 8 0 , 8 1 .V8 1 , 94.V94, 95.V95, 96.V96. 97.V97, 108.V108. 109.V109. 110.V110,  120.V120, 133.V133, 146.V146,  121.V121. 134.V134, 147.V147,  122.V122, 135.V135. 148.V148,  123.V123. 136.V136, 149.V149,  1 9 8 9 7  8  8  4  2  8  7 0 7  7 0  6 6 4 2  6 1  5  4  4  1  9  8  3 9  3 7  3 6  3 5  2 0  1 1 9 8  1 4  1 1 1 3 2 1  9 8 7 6 5 4 3 2  4-~ +  - •  +  - •  +  - •  +  - •  4  I •I--  + + + + +  I  + • +  II  + I""+ -I + -I  I II-I -I  + +  +  J  +  I-  + + +  -I  +  •I-I I---I  I  +  j-  +  1  +  1-  ••-I I-  +  - T  4>  -  38 39 23 2 4 42 43 29 30 47 48 46 65 6S 59 60 61 49 50 1.13 1 19 122 "1.20  --I  Cn  --I-  +  + +  1-  ++++-  121 "104 105 108 106 107 25 31 102 1 14 103 32 80 70 95 96 97 27 28 73 44 45  - - i ii --i  1 1 -  CD  +-+--  D I S T A N C E S  1 4  5  9  1  1  2  4  O O O O  0 O  0 2  0 0 O. 0  0  1 6  1 7  1 8 4  1 9 5  2  2 5  2 7  2 8  2 9  2 1  3 3  3 4  3 6  3 8  3 1  4 2  4 5  4 9  4 0  2  5  4  2  7  0  5  8  4  0  4  1 8  0  0  1 5  5  5  0  0  0  0  4  2  7  0  0  7  0  0  0  0  0  5  2  8  3  4 0  <FINISH>  COMMAND  USES  LOADER  STORAGE  READ  1  .016  CLUSTER  1  .020  TOTALS Execut1on  $SIG  11.V11,  25.V25, 26.V26, 27.V27, 28.V28, 29.V29, 41.V41, 42.V42, 43.V43, 44.V44, 45.V45. 57.V57, 58.V58, 59.V59, 60.V60, 61.V61. 73.V73, 74.V74, 75.V75, 76.V76, 77.V77, 89.V89, 90.V90, 91.V91, 92.V92, 93.V93, 104.V104, 105.V105. 106.V106, 107.V107.  1  1 16 31 36 37 54 55 56 20 128 126 129 127 2 69 58 5 75 78 79 22 33 35 34 57 77 67 68 7 1 72 74 76 4 9 I 1 12 13 40 41 81 «!2 89 91 90 92 93 ' 34 123 12T. 124 98 100 £9 101 7 17 16 19 18 86 88 87 109 1 10 11 1 i12 1 13 14 — T 5  REL**  2  ALGORITHM-MINVAR  19.V19, 35.V35, 51.V51, 67.V67, 83.V83, 99.V99,  111.V111, 124.V124. 137.V137. 150.V150.  FUNCTION" EUCLIDEAN  .036 Term 1nated  MONITOR  VM(PGS)  CPU(SEC)  .084  135  649  .013  189  2. 3 0 6  . 1  $1  32  177  2. 9 5 5  . 1  $1  62  RC=0  $1 .65  .097 08:55:25  T=2.993  ELAPS(MIN) .0  COST  $ 30  5 2  5 8 8 2 5  5 9 8 9 6  5 0  6 2  6 7  6  6  7  8  0  0 6  2 5  7 O 6 6  7  0  8  4  0  7 6  0  7  8  2  5  3 8 5 7  1 6  8  7  1  8  8  9  8  2  7  6 7  4 4  5  0  7  9  1 1 1 1 1 1 1 1 1 2 2 2 3 4 5 0 3 3 4 5 5 5 8 3 0 2 3 4 6 0 6 6 8 7 0 4 6 5 9 2 1 4 9 2 1  7 7 8  5  3  9  5  7  4  7  1 3 4 7 4 0 7 3  7  1 9  3  4  5  7  6  7 3 5 7 1 0  233 11 • 1 11111 1 1 1 1 1| 1111 , 1 1 1 1 1 1 1 1 1 1| 1 1 1 52516661 18882 1533555222221 65 7 7 7 2 3 3 3 5 7 6 6 7 7 7 7 11 14488899999222PO90I 1 1 1 1 8 8 3 0 1 1 1 1 1 1 3 3 2 2 4 4 2 3 4 4 4 6 6 5 6 6 4 5 1 12220000CI230103879932274 1 3 6 2 6 3 0 2 4 3 5 7 3 4 5 1 8 6 1 6 7 4 5 6 0 8 6 9 7 E 2 9 8 5 5 8 9 2 3 5 4 7 7 7 8 1 2 4 6 4 9 1 2 3 0 1 129 1 0 2 3 4 3 5 4 B 0 9 1 | 7 7 6 9 8 6 8 7 9 0 12134589b42390786569019039201458G7|512432005677834  19 ANDRRUP 6 2 RACOSUD 2 6 LOPHVENV 5 3 DIPLTAX 3 MARSEMA 17 LOPHSUD 5 5 GYMNCON 2 0 POHLNUT 61 KIAESTA 16 SCAPAME 1 RACOHETH 5 2 RACOBRE 4 GYMNOBT 13 RACOCAN 10 POLYALPA 14 CEPHBYSB 3 2 BARBFLO 5 6 ANDRNIV 5 4 RACOHETA 8 1 PSEURAD 6 DICRCRI 2 2 HYPNSUB 7 6 SCAPSCA 2 POLYPIL 6 6 POHLCRU 8 3 DRYPPAT 4 2 BARTPOM 2 4 DICRSCD 15 CYNOdEN 2 3 CLAOBOL 8 5 GRIMTOR 1 0 7 ANTHdUR 3 0 CEPHBYSA 3 4 CERAPUR 11 MARSSPA 4 4 ANASMIN 4 9 ANDRROT 7 0 BARTITH 2 8 HEDWCIL 3 6 LOPHINC 51 DICRFUS 104 KIAEBLY 1 0 9 NARDdAP 1 2 5 HOMANEV 8 9 SCHISTR 9 4 AMPHMOU 9 8 DICRHET 112 PHILFON 1 3 3 PSEUATR 3 9 BARBHAT G9 PSEUBAI 99  r  "lT6  - - 1 - - - 2 3 2 1 1 2 1 1 1 2 2 2 2 2 2 3 2 2 4 3 2 2 2 |- 1 - 3 2 1 1 - 3 2 2 - 1 1 113 1 21 -13 -1-12 1 --4 1+221 311113 --1--11-+-112  2--221 1 1 21 11-212-1—323 --2 2+1 — 1-11 — - - 3 - - - 1 1 11 544433+3323213432 2 2 1 1 1 12|  -++--1-1  -+-  -+1 +-++)  12+1  111 —+1--1111-1-  3  1  -12-+ —  1  +  _ - _  3-321  1-+--1-  2-1+--3111-12 -312-  •112-1-  -32+++21-  13  •1 + 1 14-  1+  •1  1323-  31 —  --+1 +1 1 1+-++  14  -3-11-  •1-1-3 —  1--1  -+1-  •12 + •-11-  -111-  -32-  1  •11-1 •1+211-1-  -+11 1 —  331-  •  4433 2 2 1 — 1-  -121  1-1  22| -+2--1--++1-  2 11 -1-12C— 1+ 1 +2 1 2 - - -  -+1 -1 +  121 1 — -  12-  ++-+4+12| 1 1  +-  32121  1 1 + 22(344414442133213 1 1 1-222+1 + -  12-  •1•-11 •1  2++ + 2 1 2 1 1  1244433441-  1 121 1-1-  •1 + -  3  12 1 1 |33 12  - 12  i  22 21  -3-3—121-  -1 + -  1  111  •-  P--2-1 1  3--1- 1 644-32.1  _ . „  i  --21-31-21  211 1 1 1- - 1 2 - 2 2 2 1 2 3 1 3 2 1++-+1 1 - 2 - - 1 1 1 3 2 111422122—1 1 1 1 1+43323J 21-2+1 — 1 1 1 2 3 — 1 1+121 |43242+2?344423—1333111222] 11 1 + 2 + --12 ! 1--++2 1222- 1 — 4 3 3 1252133212 3 3 — 1 3 2 2 | 14-  -21-  •11311-  -23-  1-- + -  -41 1433  •122-  --223+-4443-1--3-+1-  1-3•13234  -31-  34+-  -4-+2-  21+4-  11-3212121-  •111-  1  -13  --1 1 3 3 1  •-- 1  •112-  221-  -31 131 — - 2 •1 + 1-2  -211  12-  •1-  •11  12121  -1  12l| 111-  -+1 + 1 1-  34-—,--1  TRITQUI  1  11-  -32-  •1 1 2 3 1 -  ---212--  -2-  --1-1-  31•1-21 •11-32443] -+-11  • 1+-  111 -+1 + •1-1-  11-132-  -11-  •1+1 •121 -13- + 2-  12^  -2222  1111  1-  •111-  • -a • 1-  -33 1 - 1 122131 12  1+ -+--1•13-+-11  •1-22•111-  -211-+13 -+11 -++1 + 1-1-  •1212-  5 8 OLIGHER 5 9 BLINACU 6 0 RACOACI 6 4 LOPHWEN 1 0 5 BRACALB 1 0 6 PSEUPAT 7 1 dUNGOBO 7 8 DITRHET 79 OLIGALI 8 4 PORENAV 8 8 HOMANUT  23-  12f  -23 -31-  -21--2-  RACOVAR  9 3 GRIMMON 1 15 K I A E F A L 12 1 RADUBOL 1 2 3 DOUIOVA 1 3 2 PORECOR 1 3 6 SCAPPAL 1 4 1 GRIMALPA 142 CAMP FRA 1 4 3 ARCTFUL1 4 5 RHYTTRI 1 4 6 TIMMAUS 147 P T I L C I L 9 POROBIG 2 9 DICRCRI 3 7 PREIOUA 4 3 BAZZDEN 5 7 MARSSPH 6 7 PSEUSTE 7 3 PELLNEE 77 CHILPOL 8 0 PLAGOED 82 SCHIPEN 8 6 DISTCAP 9 0 ATRIUND 9 5 BRACSTA 9 6 PSEUINC 1 3 9 NARDBRE 1 4 0 HYPNDIC 1 0 0 RACOFAS 1 0 2 BARBLYC 103 BAZZTRI 1 4 4 dUNGCON 1 1 4 MARSBRE 1 1 6 POLYSEX 1 1 7 POHLCAR 1 18 ANEUPIN 1 4 9 HETEMAC 1 1 9 BARBVIN 122 HETEDIM 152 ISOTSTO  --2-  - - +++  - -23+1-11-21  128 PLAGP IL  91  L _ _  1+-1+  H21-231343344553343 - - - 2 1 2 + 1 3 2 2 + 4 3 3 3 3 3 2 3 1 11221 11•1•12C53443111131-  1  1  1 11  ---1 11-2+11+ 1-21311— --1 1 1 + --1—3-2 + - 1 - 1-2 44322223| — 1-1 — 1 1 111 +2 --4412 —-2-1--221 -I 11 1 1 — + 2 + - 1 - 1 + 1+ +-1 + 1 222  PTILCAL  2 1 HYPNCIR 33 POLYuu^ 4 5 LOPHCUS 4 6 CAMPATR 4 8 CHANFIL 6 5 ISOPELE 101 D I P L A L B 1 0 8 MARSCON 1 2 6 PSEUCAL 129 PTERFIL 1 4 8 PLAGASP 8 DICHPEL 3 8 BRYUCAP 6 8 POHLLUD 4 0 RACOLAN 4 1 PLEUSCH 5 0 PARAENE 6 3 MARSSTA 7 5 GRIMALPH 8 7 ENCAPRO 9 2 ANDRBLY 111 C E P H B I C 113 PLEUALB 1 2 0 AMPHLAP 1 3 0 TORTPRI 1 3 5 HYPNREV 137 SCAPULI 12 BRYUCAE 2 5 HERBADU 4 7 ANASASS 7 2 HYGROCH 7 4 ANOEAES 1 2 4 ANACMEN 1 2 7 LOPHHAT 131 P T I L P U L 134 GRIMDON 1 3 8 dUNGEXS 1 5 0 POREROE 151 HETEPRO 5 CEPHTUR 7 SCLEOBT 18 CAMPPAR 9 7 AMPHCAL 2 7 CHANSET• 3 1 AULAAND 3 5 LOPHEXC  -11122111-  •11-  •13b41-2-  •11• 1+• 1 +-  RC=0  $5.12  1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 3 3 3 3 3 3 3 3 3 3 4 4 4 4 4 4 4 4 4 4 5 5 5 5 5 5 5 5 5 5 6 6 6 6 6 8 6 6 6 6 7 7 7 7 7 7 7 7 7 7 8 888 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 3 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 123  1 DIPLALB 2 RACOHETH 3 ISOPELE 4,.RHI ZGLA 5 POGOCON 6' DICRHET BARTPOM . 8 .L E P I R E P 9:;NARDSCA 10 MARSEMA ' 11 SCAPAME '-,1 2 CLAOBOL 13 POLYALPA 14 POHLNUT 15 HYPNSUB 16 PLAGASP 17 STOKPRAP 18 ISOTSTO 19 HETEMAC 20 METZCON 21 P L A G L A E 22 D I C R F U S 23 RACOCAN 24 LOPHHET 25 DICRSCO 26 LOPHCUS 27 LOPHVENV 28 SCAPSCA 29 PLAGUND 30 SCAPMUC 31 HYPNCIR 32 CEPHBYSB 33 CYNOJEN 34 LOPHINC 35 PLAGDEN 36 PORECOR 37 PLAGOED 38 PLAGINS 39 CLAOCRI 40 PORENAV 4 1 POREROE 42 CEPHBYSA 43 BARBFLO 44 POLYFOR 45 B L E P T R I 46 D I P L T A X 47 BAZZDEN 48 POHLCRU 49 C E P H B I C 50 A N A S M I N 51 HERBADU 52 S C A P U N D 53 GYMNOBT 54 ANACMEN 55 DOUIOVA 56 GRIMTOR 57 TRITQUI 58 P L A G P I L 59 FRULTAMN 60 EURHPUL 61 ANDRRUP 62 CHANFIL 63 MARSSPA 64 UAMEAUT 65 MYLITAY 66 B A Z Z T R I 67 CALYMUL 68 RACOBRE 69 RACOHETA 70 HYPNREV 7 1 AMPHCAL 72 APOMPUB 73 HOMANUT 74 METAMEN 75 POGOURN 76 DIPI..OBT 77 POHLPRO 78 CEPHPHY 79 C A L Y T R I 8 0 GYROUND 81 O L I G A L I 82 POLYLON 83 RHYTLOR 84 POLYALPM 85 LOPHGIL 86 RACOACI 87 BRACPLU 88 A T R I S E L 89 P H I L F O N 90 PELLNEE 91 D I C H P E L 92 BRACFRI 93 JUNGOBO 94 D I P L P L I 95 RICCMUL 96 GRIMHAR 97 RAOUCOM 98 BART ITH 99 DRYPPAT 100 GYMNCON 101 POROBIG 102 CLAOWHI 103 RADUBOL 104 PSEUINC 105 PLAGROE 106 BARBLYC 107 MNIUSPI 108 P S E U T E C 109 DREPUNC 1 10 B A R B H A T 1 1 1 RACOSUD 1 12 K I A E B L Y 113 DICRCRI 1 14 P T I L C I L 1 15 STOKPRAS 1 16 HOMATRI 1 17 PSEURAD 1 18 ENCAPRO 1 19 BARBREF 120 MNIUAMB 12 1 BRYOREC 122 THAMNEC 123 HOMAFUL 124 PLAGROS 125 TIMMAUS 126 S C H I S T R 127 E N C A C I L 128 B R Y U C A P  •T,  5432 1 332-12-2 4-+1 - - 1 ++ 2—1-1-243--1 + 21 + -- 1 ++--2++ +  2- + --+5S--2 2++22-333-42+ + +-+  •--44-22+433+ 3-+--1-214 32+24 1+1-2+2+11 1--1--12--•-- + +1-342 2++ +  --11--11-  +2  +2  + - -2--+-  -441++31++4 +13-1 25 -23 14 +1-3+31 1 + -32-2--1++1 1 122-•11- 1 + 3-243412333 -2532 + 32 43132+1 1 -33 1 121 43-1-323 34 2+ 2++2--4423--1--1-  -+41 1 + -2-1--341 — 3 113-1142+311--- 1--3--2 ---1 1 + 2 1  +  21 + -3-- --4--1-11--1+---1 132 -3-- 1-2 1 11 11 '  2  53 3+- -32--2+13+- ++1+++12 •1--- + 3+ --121-22-+ -+--S+- 3-+++-1-2---1 + ----1-  - + 3+  -31  -++2++-21-  -241 1 1-  3412  1-21 12  1  2  -5-- 111---1 1 + 1-+11- -22- 1 4-15-++3-++-  -21-  •1  143122-  -43-  • —  U  .++--+ -11 + 1 + +-3 -+13131-4-43 -_3__ + 1  + +  214  ++- +  ---1 + 2+-31 1 1—  2  1+  2  + 11--11 ++ + --43+-1 + 1 121 14221+  +  1--33  ++1  +1 2-++  i  '  111  + 21  ;  -3  :  2  4 121-++ 1R  .  —  2 23  -.__  111 +  :._  +  4 1 -3-++ _  1  1  1  : 324 1  22-332  1-12-1 4  2  1  -3 . -3  + -1  2 + + +1  ,  1  22  111  +  ______  1  2—3 +  2  +  +  :  +  '  3  ++_  3  331 +  2 11 +  1  2  2  , i __  3  1  3  4-1  ++  1, i 2  '  2  513  2 1  .  :  1  .  2  1  21-1- +  3  1  :—  4--1 + 22-1-22 12-1 22  12  -- +  :::::::.;:::::::::::±:: -1-21 +  !  _  ,n  1  3  L__  •1 32-----1 1+11-1--1-+11,1-1332--  •1  1.- — • -12 •1-1-1-  I  1 1 1 1 1 1 1 1 1 12222222222 ' 3 3 3 3 3 3 3 3 3 4 4 4 4 4 4 4 4 4 4 5 5 5 5 5 5 5 5 5 5 6 6 6 6 6 6 6 6 6 6 7 7 7 7 7 7 7 7 7 7 8 888 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 9 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 123  11 SCAPAME 10 MARSEMA 16 PLAGASP 2 RACOHETH 3 ISOPELE 12 CLAOBOL 18 ISOTSTO 46 D I P L T A X 19 HETEMAC 1 DIPLALB 4 RHIZGLA 61 ANDRRUP 27 LOPHVENV 41 POREROE 47 BAZZDEN 59 FRULTAMN 15 HYPNSUB 26 LOPHCUS 45 B L E P T R I 48 POHLCRU 49 C E P H B I C 55 DOUIOVA 7 BARTPOM 9 NARDSCA 13 POLYALPA 29 PLAGUND 31 HYPNCIR 35 PLAGDEN 37 PLAGOED 17 STOKPRAP 50 ANASMIN 58 P L A G P I L 72 APOMPUB 1.4 POHLNUT 20 METZCON 25 DICRSCO 28 SCAPSCA 32 CEPHBYSB 39 CLAOCRI 42 CEPHBYSA 51 HERBADU 103 RADUBOL 6 DICRHET 2 1 PLAGLAE 33 CYNOJEN 74 METAMEN 98 B A R T I T H 104 P S E U I N C 8 LEPIREP 22 D I C R F U S 34 LOPHINC 40 PORENAV 60 EURHPUL 62 CHANFIL 66 BAZZTRI 67 CALYMUL 75 POGOURN 101 POROBIG 107 MNIUSPI 118 ENCAPRO 120 MNIUAMB 121 BRYOREC 63 MARSSPA 56 GRIMTOR 65 MYLITAY 7 1 AMPHCAL 84 POLYALPM 105 PLAGROE 109 DREPUNC 111 RACOSUD 123 HOMAFUL 5 POGOCON 23 RACOCAN 24 LOPHHET 44 POLY FOR 76 D I P L O B T 52 SCAPUND 54 ANACMEN 68 RACOBRE 80 GYROUND 69 RACOHETA 70 HYPNREV 83 RHYTLOR 87 BRACPLU 88 A T R I S E L 89 P H I L F O N 106 BARBLYC 112 K I A E B L Y 116 HOMATRI 119 BARBREF 124 PLAGROS 126 SCHISTR 93 J U N G O B O 94 D I P L P L I 95 RICCMUL 96 GRIMHAR 97 RADUCOM 30 SCAPMUC 99 DRYPPAT 100 GYMNCON 36 PORECOR 102 CLAOWHI 38 P L A G I N S 43 BARBFLO 53 GYMNOBT 57 T R I T Q U I 64 JAMEAUT 108 P S E U T E C 73 HOMANUT 110 BARBHAT 7 7 POHLPRO 7 8 CEPHPHY 113 DICRCRI 114 P T I L C I L 115 STOKPRAS 79 C A L Y T R I 1 17 PSEURAD 81 O L I G A L I 82 POLYLON 85 LOPHGIL 86 RACOACI 122 THAMNEC 90 P E L L N E E 91 D I C H P E L 125 TIMMAUS 92 BRACFRI 127 E N C A C I L 128 BRYUCAP  --31 13- 1 142+31 1-3 1 1- 1 + 3-2434 12333-2532 + 32 43132+1 1 33 1 121 43 -+41 1 + -2- 1 - - £ 4 1 -3 -- + 1 25-23 14 +1-3+31 1 + 32-2--1++1 1 122 3 + -32--2+13 + -++1 +++12 1 + + 3+ ++ 2++ 132-3 — 1-21 — - 1 1 1 1 332-12-2 4-+1--1 ++ 2++22-333-42 + --1-214 32 + 24 11 2--1-1-243--1 + 21 + 1 + 1-2 + 2+1 1 1--1--12 11 1--3--2 3 •- 1 323 34 2+ 2 + + 2--4423-- 1 - - 1 12 1-22- + 3 1 — 21 241 1 1 34 12 1-21 1 1+ 1+ + -3 + --43 + -1 + 1 12 1 14221 + 1 + _ •) +__3 _3_ _ 1-2--3 1 1--2- + 2 21 12 5432 1 2- + --+53--2 44-22 + 4 33+ 3 + --1 ++--2 + + + -+ +-+ ++1-342 2++ + • 2+++1 1 11 +  +  +  1  j-31 j--1 '  +  1  1  4_ .. 15  +  +  4  +  1  2  3 12  2  +13131-4-43 1-3-++  ,  1  +  214 22-332  1  + __2 _1 1 1+2  22 ++-- +  +  +  1+  ++3_ + +  ,  :  21-1 21 + --2-1 1  1-12-1  1 +  4  21 + 1-----1  +  1 2 2  .  2--212'2--++  21 131 1---  2-++ :  2  +  2  _  + _+  1+ _2  __  1  2  +  . _-4  +  +  1+  5--1 1 1  • 312 1-2-j ' -22 211 +  +1  2  +  :  + 2  2  2  i  2+  4 • 1-- +  ++  :  4  +  :  11  + 21 1 1 1+ 3--21 31++4 +13 + 2- + 2- + . + 2-- + 32  2--1--3 + +1 + -- + 3+ 1+2+  111 + + 2- + j i 4--1-1 1--1 + -|--1  1  _ — 43  3  3  ++1  23  1 1  4  3 2-2 _-! 1  --•  :++- + 121- + + 441++  2  324 1  53 11 + 1-+11  1  2  21-  1  t, -3  +  1  +  ++ -++  11 l  21-1- + 22-1-22-  l 3  --  1+  2  •1  1--1 2 -34  2 1--  :-4  ' •  + -1>  14 -2-2 .-1+++  1 1 + --1 + 1 1-+11 1-133 2-  1_.---1  1  —  J  -1  --32 _  2--++--12-1 1-21--•13  -211-  .4  2  2 1-  -3  -•13•12-22•11-32  1-12  1 .  1 2  :  2  .__1 .__2  -• 2  " —  I::-::::::::::::::::::::::::::::::::::  u>  __„,____!„_ 2 +  ._ +  \  •_  5  + 2 :  --  1-1 _  •!_  1RC=0  $1.  <CLUSTER OPTION-MINVAR MAX=*>  FUNCTION = EUCLIDEAN  REL =* C A S E S M - 8 3  VAR=1-128  LO  CLUSTER A N A L Y S I S CASES = CASE/>': 1-83 CASEWISE N = 83 ALGORITHM=MINVAR DI ST ANCE = EUCL. I DE AN  USING: 1 . V 1 . 2V2, 3.V3, 4.V4 5.V5, 6 .V6. 7.V7, 8.V8, 9.V9, 10.V10, 11.V11, 12.V12, 13.V13, 14.V14. 15.V15, 16.V16 17 V17 18.V18 19.V19, 2 0 . V 2 0 , 21.V21 22.V22, 2 3 . V 2 3 , 24.V24, 25.V25, 2 6 . V 2 6 . 27.V2"? 28.V28 29.V29, 30.V30, 31.V31 32.V32. 33.V33, 34.V34 3 5 . V 3 5 , 36.V36 37.V37 3 8 . V 3 8 , 39.V39, 4 0 . V 4 0 , 4 1 . V 4 1 , 42.V42, 43.V43 44 . V44 45.V45, 4 6 . V 4 6 . 47.V47 48.V48, 49.V49, 50.V50 51 . V51 , 5 2 . V 5 2 53.V53 54.V54 , 55.V55, 56.V56, 57.V57, 58.V58. 59.V59 60.V60 61.V61, 62.V62 63.V63, 64.V64, 6 5 . V 6 5 , 66.V66 6 7 . V 6 7 . 68.V68 69.V69 7 0 . V 7 0 , 7 1 . V 7 1 , 72.V72, 73.V73, 74.V74, 75.V75 76 . V76 77.V77, 78.V78 79.V79, 8 0 . V 8 0 , 81.V81 , 82 . V82 8 3 . V 8 3 , 84.V84 85.V85 86 . V86, 8 7 . V 8 7 , 88.V88, 89.V89, 9 0 . V 9 0 , 91.V91 92 . V92 93.V93, 94.V94, 95.V95, 96.V96, 9 7 . V 9 7 , 98.V98 9 9 . V 9 9 , 100.V100, 101.V101, 102 V102. 103.V103, 104.V104, 105.V105, 106.V106, 107.V107, 108.V108 109V109 110V110 1 1 I .V1 1 1 , 112.V1 12, 113.V1 13, 114.V1 14, 115.V115 , 116.V116, 117.V117, 118.V118, 119.V119, 120.V120, 121 .V121 , 122.V122, 123.V123, 124.V124, 125.V125, 126.V126, 127.V127, 128.V128  8 8 7 7 7 7 7 6 6 2 0 9 7 6 2 0 8 7  CASE* 1 2 4 45 46 65 32 48 52 23 26 33 28 29 34 3 36 51 53 8 13 17 74 75 14 44 31 63 62 39 38 35 55 5 6 :69 9 10 64 49 50 7 70 82 83 54 67 68 18 24 42 40 4 1 6 1 1 1 21 22 43 58 47 16 25 27 30 76 77 66 37 59 60 19 20 56 57 78 8 1 79 80 12 15 71 72 73  6 5  5 5 4 2 0 9  4 4 3 3 3 0 9 6  ++-  + + +  1  +  1  +  +  + -I  +++++++++-  —  -  I  -  1  0 I  3 6 8 9  s  T A N C E S  1 1 1 1 1 1 1 1 2 2 2 2 2 2 3 3 3 3 3 3 4 4 4 4 4 4 5 5 5 6 6 6 7 7 7 8 8 9 9 1 1 1 1 1 1 1 1 1 2 2 0 2 3 4 5 6 6 9 0 1 2 3 6 8 0 1 2 6 8 9 1 3 4 5 6 7 3 6 9 2 4 6 3 5 6 3 5 3 9 0 0 1 3 4 5 6 8 8 3 3 4 6 3 6 1 1 1 0 9 6 5 5 0 2 0 7 5 5 0 2 5 7 1 8 O 0 0 5 8 5 0 0 0 5 0 5 9 0 5 3 6 4 7 6 5 O O O O 3 O 0 O O O O 0 6 0 8 O 6 O O 3 0  <FINISH>  COMMAND  USES LOADER  STORAGE  MONITOR VM(PGS) C P U ( S E C ) E L A P S ( M I N ) COST  READ  1  .016  .055  1 10  .376  .0  $ . 10  CLUSTER  1  .020  .012  135  .867  .0  $.30  .036  .067  128  1 . 243  . 1  $.40  TOTALS  9 8 7 6 5 4 3 2  +++-  4 0 9 9 3 1 9 9  1  4463452232231 355 1177 1436633351 6 1645 7885661244461122454 1 2 2 3 7 7 6 3 5 6 1 2 5 5 7 8 7 8 1 1 777 1 2 4 5 6 5 2 8 2 3 6 3 8 9 4 P 6 1 3 8 3 7 4 5 4 4 1 3 2 9 8 5 5 B 6 9 9 0 4 9 0 7 0 2 3 4 7 8 8 4 2 0 1 1|1 1 2 3 8 7 6 5 7 0 6 7 6 7 9 0 9 0 6 7 8 1 9 0 2 5 123  11 SCAPAME 10 MARSEMA 16 PLAGASP 2 RACOHETH 3 ISOPELE 12 CLAOBOL 18 ISOTSTO 46 D I P L T A X 19 HETEMAC 1 DIPLALB 4 RHIZGLA 61 ANDRRUP 27 LOPHVENV 41 POREROE 47 BAZZDEN 59 FRULTAMN 15 HYPNSUB 26 LOPHCUS 45 B L E P T R I 48 POHLCRU 49 C E P H B I C 55 DOUIOVA 1 BARTPOM 9 NARDSCA 13 POLYALPA 29 PLAGUND 31 HYPNCIR 35 PLAGDEN 37 PLAGOED 17 STOKPRAP 50 ANASMIN 58 P L A G P I L 72 APOMPUB 14 POHLNUT 20 METZCON 25 DICRSCO 28 SCAPSCA 32 CEPHBYSB 39 CLAOCRI 42 CEPHBYSA 51 HERBADU 103 RADUBOL 6 DICRHET 21 PLAGLAE . 33 CYNOJEN 74 METAMEN . 98 BART I T H 104 P S E U I N C 8 LEPIREP 22 D I C R F U S 34 LOPHINC 4 0 PORENAV 6 0 EURHPUL 62 CHANFIL 66 B A Z Z T R I 67 CALYMUL 75 POGOURN 101 POROBIG 107 MNIUSPI 118 ENCAPRO 120 MNIUAMB 121 BRYOREC 63 MARSSPA 56 GRIMTOR 65 MYLITAY 71 AMPHCAL . 84. POLYALPM 105 PLAGROE 109 DREPUNC 111 RACOSUD 123 HOMAFUL 5 POGOCON 23 RACOCAN 24 LOPHHET 44 POLYFOR 76 D I P L O B T 52 SCAPUND 54 ANACMEN 68 RACOBRE 80 GYROUND 69 RACOHETA 70 HYPNREV 83 RHYTLOR 87 BRACPLU 8 LEPIREP 8 LEPIREP 89 P H I L F O N 106 BARBLYC 1 12 K I A E B L Y 1 16 HOMATRI 1 19 BARBREF 124 PLAGROS 126 SCHISTR 93 JUNGOBO 94 D I P L P L I 95 RICCMUL 96 GRIMHAR 97 RADUCOM 30 SCAPMUC 99 DRYPPAT 100 GYMNCON 36 PORECOR 102 CLAOWHI 38 P L A G I N S 43 BARBFLO 53 GYMNOBT 57 T R I T O U I 64 JAMEAUT 108 P S E U T E C 73 HOMANUT 110 BARBHAT 77 POHLPRO 78 CEPHPHY 1 13 DICRCRI 114 P T I L C I L 115 STOKPRAS 79 CAL Y TRI 117 PSEURAD 81 O L I G A L I 82 POLYLON 85 LOPHGIL 86 RACOACI 122 THAMNEC 90 PELLNEE 91 D I C H P E L 125 TIMMAUS 92 BRACFRI 127 E N C A C I L  +!  + -3 -- 1 142 32243333+ 13343124335231 13214-31-1--1 11-1 + 32-|2 1 13 f 1 1— 1 |45332332244 123322 1.-+1 - - 2 - + + + +2+-2+11+-1-2 -3232^ + + + 1 1-- + + + 3311+ ++2 3 1 -- 1 1 1+ + 1--1 + + 32 33 2 + - + - 2+323122--- 1 1 + -2422434|12 1-2 1-1 1+++1-1-4312+ 211-2 1 1 2- 1 1 + 1 -: ++ +i - 2 - 2 2 C 4 3 3 3 124431332 2|i -- 1 22- 1234- + -- 1 1| 1 1 1 — 3 if1 1-22 21--241  II 1 1 1 1 4 1,  t  1--- + - -  + HI  +  1+  1 y -1-2-++ 5 4 2 4 4 3 5 2 3 - 2 3 + H•I3243 + - 1+ + + + --2--2 +111- + 1 1  +  1 __  3  1--1 .  422  1 1 143  1  +  +-+ 1 1+- 1 1 1  •++  1-1-2-1+ 1--2++ + + 34-  1 12  31•13 +  •1|-  +  +  - 1 1  ---+-3-  1- + +  2 1+  • 1  --334  1-  -1-  ^32-2 -1  •11- - + -2  + 1 1 1 45 143 12 + +  21  -2334  P34--3- 1 + 2-  +++ -!2+  - + + --422312J  •1-2•1 1 1 - - - 1 + 1 -  23  • 1-  - 3 4 2 4 3 1|  +  312312  22++333+ 221 1 4421+1-  1 + - -+--2-  --1+2-  D43 P  r - 2 - - -  1  2121-  -+11- 1--.+  332-'-2-  -2 +  -31123+-  . 1  h 1+  122-12 1+  2  1-  4  + -2  •1-2-  -2-21 + 1 — 11-  M +  2-  1  -51-  122 -44 • -221211-3-2•124-  + 2111311  •1--1-  --2-22-  -22  -2 + -  - 1 -  1 -  -2  4--  •1344 -323-2++112• 1+-11+1-+1 1 1-1233- 1 1 -  •1 •1--3-  1-  -2-133  •1  122 1-  -+13111-  + 2-  -42-21-3+-  -2 + -2+-22•1 1  -32•1111-  •12-  o  yj  ---— \> "i  —  —  -  RC=0  CM  $3.  238  111  11 1 1 1111 1 122222222223333333333444444.44445555555_-556G6666G66677777777778 8 8 8 8 8 8 8 8 8 9 9 9 9 9 9 9 9 9 9 0 0 0 1234.678301234567890123456789012345678^01234567890123456789012345G789012345G7890 1234567890123456789012  1 SCOUAQU 2 HYGROCH 3 DICHPE1. 4 SCAPUND 5 R/ICOACX 6 UL NGPUM 7 PELLNEE 6 NARDSCA 9 CEPIIBIC 10 HYPNDIC 11 SCAPSLIB 12 EI.INACU 13 RHIZGLA 14 SC/PAME 15 MARSEMA 16 BRACPLU 17 POGOURN 1B PLAGASP 13 PI.AGROE 23 B L E P T R I 21 BARTPOM 22 RACOHETH 23 POLYALPA 21 CERAPUR 23 D I P L A L B 23 HYGRLAX 27 ANASMIN 2B RACOBRE 29 A T R I S E L 30 UUNGOBO 31 DITRHFT 32 D I P L P L I 33 0LIGA1 I 34 RACOHETA 35 RACOFAS 36 CONOCON 37 RICCMUL 38 P 0 R 0 8 I G 39 STOKPRAP 4 0 GEOCGRA 4 1 POHLCRU 42 AMPHCAL 43 METZCON 44 BRACFRI 45 CYNOJEN 46 HETEMAC 47 POHLLON 48 POHLNUT 49 I S O P E L E  1  3223 21 312 + -: 223 2 +31 124 +- - +2 2121 132 23122 + 3+21 122-2 2 2 3 2 2 2 2 12 1 - - 1 221 1 + 1 - 2 4 3 - 3 12 11 1 1+2 1 1-- + - 2 2 2 2 3 122 — -23-33 + 4 4 3 3 3 4 3 - 2 - 2 4 143- - 1 1 1 + + - - 2 2 2 - - 1 1 13 1 2- + 2 1 +++ + - - 2 - 3 2 2 3 - 2 22 321 1 2221 + 3 - 1 3 3 -22 + 222-1 21 12--2-1 +1 + +2 + --1 +- + +12- + 2 2 - 2 - 2 2 - 1++1 1 1+1 3++1-1 1- 111-111 + 1 1 + -1 -4 2+4 1352 4332 3 21334 223221232322 ' 2 _1_4 1- + +1 1 +1 1 122 1 1 1- 1-1221 2 2 2 2 - 2 - -1 1 1 +++ + - 1 4 3 2 + 31 1 + 23- 4 + 2 + - - 2 1 1 2 4 3 3 2 - 2 2 3 3 2 1 2 4 - 2 2 2 - 2 2 4 2 4 1 12 1 + 1+42-1 212+1 45 3-12 + - - 1 3 -3--31 322122 1-2-3343 . - - 1 - 1 4 4 + - 3 1 - 1 1+1 12+ --1 _1 1 +1 121 1 222 14 1 1-4 1-44444 34-1 1 11 44 14111--+1  4 3 42--+  ;  245  3344  2-1  — -  2+  • '  2  1  J  2  -1 1 2 3 - + - - 2 - 2 - - 123+1 121 1 1-1--1 12223 -21-1 •1-1 -33424524213+-3344 12 + 121+3 23 2 22223-111 4331231+-232--2 •1--1-  121-1  2  11- 1  -21121-  1--21  111-1-  22+1  • 1+1 + - -  -2121-  1 1 + 111 + 2 1 + - 1 1 1 -  -2 + 111 •11-1--•-2 1-  • 1 ++-  -13 - - 12-2-  .-2 — 2 - 2 1 2 + 32121-1-22 1334442-1-2--  -2+1-  2 + + -31 + 1 1 1-  - + 3 2 - 3 2 — 22 — 22-  -2-22-2  •1  -+1-1  1-1 1+4-  1-1-  21234-  --2  -4 1-2-- -  1-  11-  -43335-2---1-  -+1++--  -212113-4 1 2 — 1 -211-11-1 -1-  •12  -+4-1 1-  -211-+-  -232-  -21 1-  •1221112-  ---2• 1•11-  -__ —  •12-  13  2  -2--2-  31  •1 1 1-  5 0 JAMfcAUT*  5 1 RACOAOU 52 SCLEOBT 53 HYPNSUB 54 D I P L T A X 55 PHILFON 56 CHILPOL 57 BART ITH 58 POHLPRO 59 SCAPSCA GO RACOVAR 61 PLEUALB 62 OLIGPAR 63 B A Z Z T R I 64 CLAOBOL 65 ANEUPIN 66 BARTLES 67 DICRHET 68 SPHAGIR G9 OXYSTEN 7 0 PHILARN 71 JUNGATR 72 ATRIUND 73 RADUBOL 74 ISOPPUL 75 T R I T Q U I 7G MYLITAY 77 P L A G L A E • 78 DREPUNC 79 RHYTSOA 80 BRYUPAL 8 1 RACOLAN 82 SCHISTR 83 S C H I R I V 84 BRACVEL 85 GRIMTOR 86 PREIQUA 87 SCAPPAL 88 FISSOSM 89 JUNGEXS 9 0 DRYPPAT 91 ANOEAES 92 PSEUPAT 93 BRYUPSE 94 JUNGCON 95 ISOTSTO 96 F I S S B R Y 97 POREROE 98 PLAGROS 99 HYGRSMI 100 RHIZNUD 101 PORECOR 102 MNIUBLY 103 P L A T J U N 104 LOPHHET 105 BARBLYC 10G TORTFRA 107 HYGRLUR 108 BARBREF 10? BRYUCAP 110 SCAPMUC 111 BRYUMIN 112 STOKPRAS 113 DICRCRI 114 T R I T P O L 115 P L A G P I L 116 BLASPUS 117 LOPHSUD  -2++--12-23-  -2-  -4 — 21-21-22-  .+ 21-1 2-434- 1 1 1•111 1 4 1 2 - - 1 3 1 — 3 + - 1 - - - 1 1 11-1--  -2+-  1-  •11-  -21-  -2-1112 2--1131+2--2 2--22  -+1+14-  1-112++3-  -22 1-22--212--  1-121-  1-1-  -2-21-21 1 + 1  322-1-  •1-  1-121212-22-  •1-1-  -322 1 12 + - 13244 12--131 1+12-  1+•212-1222--1  •+1---+2-  -+11 1•22-  •322---4-  -44-  •1-•1111-  -2-3•11-11-  -++— 2-  ii-  li § I  i  Do  r-  x3  1  1  1  I f ^ "  ci,'  ft-  239 111 1 1 1 1 1 1 1 1 1 122222222223333333333444444444455555555556666G666G6777777777788888888889999999999000 1234567890123456789012345G789012345G78901234567890123456789012345678901234567890 123456789012345G789012  10 HYPNDIC 3 DICHPEL 4 SCAPUND 7 PELLNEE 12 B L I N A C U 15 MARSEMA 3 0 UUNG0B0 9 CEPHBIC 5 RACOACI 13 RHIZGLA 11 SCAPSUB 8 NARDSCA 16 BRACPLU 20 B L E P T R I 2G HYGRLAX 1 SCOUAQU 37 RICCMUL 25 D I P L A L B 56 C H I L P O L 18 PLAGASP 6 JUNGPUM 55 P H I L F O N 17 POGOURN 8 0 BRYUPAL 89 JUNGEXS 39 STOKPRAP 62 OLIGPAR 3E POROBIG 44 BRACFRI 65 ANEUPIN 71 UUNGATR 83 S C H I R I V 87 SCAPPAL 14 SCAPAME 36 CONOCON 52 SCLEOBT 2 HYGROCH 4 0 GEOCGRA 78 DREPUNC 86 PREIOUA 24 CERAPUR 75 TRITQUI 88 FISSOSM 99 HYGRSMI 6 1 PLEUALB 29 A T R I S E L 33 O L I G A L I 48 POHLNUT 51 RACOAQU 53 HYPNSUB 54 D I P L T A X 57 B A R T I T H 84 BRACVEL 93 BRYUPSE 94 JUNGCON 23 POLYALPA 27 ANASMIN 35 RACOFAS 59 SCAPSCA GO RACOVAR 4 1 POHLCRU 43 METZCON 49 I S O P E L E 64 CLAOBOL 68 SPHAGIR 69 OXYSTEN 82 SCHISTR 104 LOPHHET 109 BRYUCAP 32 D I P L P L I 34 RACOHETA 46 HETEMAC 58 POHLPRO 74 ISOPPUL 63 B A Z Z T R I 66 B A R T L E S 81 RACOLAN 9 0 DRYPPAT 95 ISOTSTO 102 MNIUBLY 103 P L A T J U N 107 HYGRLUR 108 BARBREF 110 SCAPMUC 114 T R I T P O L 19 PLAGROE 2 1 BARTPOM' 22 RACOHETH 28 RACOBRE 31 DITRHET 91 ANOEAES 92 PSEUPAT 42 AMPHCAL 45 CYNOJEN 47 POHLLON 9G F I S S B R Y 97 POREROE 98 PLAGROS 50 UAMEAUT 100 RHIZNUD 101 PORECOR G7 DICRHET 70 PHILARN 72 ATRIUND 105 BARBLYC 10G TORTFRA 73 RADUBOL 76 MYLITAY 77 P L A G L A E 79 RHYTSQA 111 BRYUMIN 1 12 STOKPRAS 113 DICRCRI 85 GRIMTOR 115 P L A G P I L 116 BLASPUS 117 LOPHSUD  -1432+31 1+23+4 + 2 + - - 2 1 1 2 4 3 3 2 - 2 2 3 3 2 1 2 4 - 2 2 2 - 2 2 4 2 4 1 1 2 1 + 1 + 4 2 - 1 - - - 2 1 2+ 1 - -45 3-12+--13 42-- + 31 124 + - - t-2 2 12 1 132 23122 + 3 + 2 1 122-2 2232222 12 1 - - 1 221 1 + 1 - 2 4 3 - 3 12111 11+2 1 1 - - + - 2 2 2 2 3 122 23-33 + 4433343-2-24 143--1 1 1--- + + - - 2 2 2 - - 1 1 13 +- - 1 +- + +12- + 2 2 - 2 - 2 2 - 1 + +1 1 1+1 3++ 1- 1 1 - 1 1 1 - 1 1 1 + 1 1 + -1 -1 2 1+1 121 1 222 14 1 1-4 1-4444434-1 1 11 44 14 1 1 1--+1 - - 3 3 4 2 4 5 2 4 2 13+ 3344 12+121+3 23 2---22223 21 12 1 2 12-2 2 + + -31+1 1 1 +32-32--22 --22 2-22-2 21234 1-1 +- + 2 -1- + 1- + +1 1 +1 1 1 22 1 1 1-1-122 1 2 2 2 2 - 2 - -1 1 1 '+ + + + 1----2- + 2 1+++ + - - 2 - 3 2 2 3 - 2 22 32 11 2221 + 3-133--22 -1 123- + - - 2 - 2 - - 1 2 3 + 1 12 1 1 1-1 — 1 12223 121-1 2 1 1-1 -3--31 322122 1-2-3343 1- 144 + - 3 1 - 1 1 + 1 12++- 1 -4 2+4 1352 4 332 3 2 133422322 1232322 2 --1 1 1 4331231 + - 2 3 2 - - 2 1 1- 1 2 1 - 3 - - - 1 1 1-1 1+1+ -+ , 1--21 22+1 1 1+ 1 1 1 + 2 1 + - 1 1 1 2121- 1 ,-1 1-22 1334 4 4 2 - 1 - 2 1 1-1 1+4 +1-1 4 3344 245 2-1 2+ 3223 21 312 »--2 1 2121 13 1221 1 12 21 1 232 + 111 1++ 2+1 2--2-212 + 32121 •! t 4 1 2 - - 13 1 3+ - 1 11 111-1-22•1-222-1112--2-1-2 1-21-1 2-434- 1 1 1-2 + •1--1-2 11+1-322-1 1-212-1222 +1 + 2--+4-11 21 1-11 2 - - 1 131 + -+1+1412--+1++412--1 13 31 4' 2 - -2 1-1 12 ++3-22--212--11-121 ... 11-1212121324442 • 1+-21-1•1 + • 1-4333512-2-2 4--211-+ -223 2-21 1- + 1• ! -2-21--1•-322 1 12 + -2+-4-+1•-1-1--1311+1-322--44-2-1 112•13+  2  1  n  2  2  2  •111-2++-21-2--22-  •1-1-  •1111-21-11-  -2--22-221-  •11-41--1-  1  i-  - 2 - - 1--22-•1111-2-3•11•11-  -3  -+-1-  0 °  IT)  1  1  •'  vjyt  ,1 c  C) ^ ^  <CLUSTER OPTION-MINVAR MAX=*>  FUNCTION'EUCLIDEAN  REL = " CASES=1- 102  VAR=1-117  CLUSTER ANALYSIS CASES = CASEX': 1 - 102 CASEWISE N=102 AIGC1RITHM = MINVAR DISTANCE=EUCLIDEAN USING: 1.V1, 2.V2, 3.V3, 4.V4. 5.V5. 6.V6. 7.V7. 8.V8, 9.V9, 10.V10. 11.V11, 12.V12, 13.V13, 14 . V 14 , 15 . V15 . 16.V16, 17.V17, 18.V18. 19.V19. 2 0 . V 2 0 . 2 1 . V 2 1 , 2 2 . V 2 2 , 2 3 . V 2 3 , 24.V24, 2 5 . V 2 5 , 26.V26, 27.V27, 28.V28. 29.V29, 30.V30, 3 1 . V 3 1 , 32.V32, 3 3 . V 3 3 . 3 4 . V 3 4 . 3 5 . V 3 5 , 36.V36, 3 7 . V 3 7 . 3 8 . V 3 8 , 3 9 . V 3 9 . 4 0 . V 4 0 , 4 1 . V 4 1 . 42.V42, 43.V43, 44.V44, 4 5 . V 4 5 . 46.V46, 4 7 . V 4 7 , 4 8 . V 4 8 , 4 9 . V 4 9 , 5 0 . V 5 0 , 5 1 . V 5 1 . 5 2 . V 5 2 , 5 3 . V 5 3 . 5 4 . V 5 4 , 55.V55, 5 6 . V 5 6 . 57.V57, 5 8 . V 5 8 . 59.V59, 6 0 . V 6 0 . 6 1 . V 6 1 . 62.V62, 6 3 . V 6 3 . 64.V64, 6 5 . V 6 5 , 6 6 . V 6 6 , 6 7 . V 6 7 . 6 8 . V 6 8 , 6 9 . V 6 9 . 7 0 . V 7 0 , 7 1 . V 7 1 , 72 . V72. 73 . V73, 74.V74, 75.V75, 7 6 . V 7 6 . 77.V77, 78.V78, 79.V79, 8 0 . V 8 0 , 81 . V8 1 , 8 2 . V 8 2 , 8 3 . V 8 3 , 8 4 . V 8 4 . 8 5 . V 8 5 . 8 6 . V 8 6 , 8 7 . V 8 7 , 8 8 . V 8 8 , 8 9 . V 8 9 , 9 0 . V 9 0 , 9 1 . V 9 1 , 9 2 . V 9 2 , 9 3 . V 9 3 , 94.V94, 9 5 . V 9 5 . 9 6 . V 9 6 , 9 7 . V 9 7 , 9 8 . V 9 8 . 9 9 . V 9 9 , 100.V100, 1 0 1 . V 1 0 1 . 102.V102. 103.V103. 104.V104, 105.V105, 106.V106, 107.V107, 108.V108, 109.V109, 110.V110, 1 1 1 . V 1 1 1 . 112.V112. 113.V113, 114.V114, 115.V115, 116.V116, 117.V117  S T E P  CASE* 1 66 88 24 25 26 27 80 6 36 94 38 97 98 33 34 35 66 67 78 79 68 69 15 19 17 16 18 3o 101 62 64 91 93 92 8 1 87 83 99 100 82 51 52 56 53 54 55 63 89 90 70 95 71 74 73 75 72 76 2 32 1 1 59 12 102 47 48 50 58 61 7 8 31 10 13 9 21 20 49 22 23 57 60 3 4 5 37 14 46 84 85 42 45 28 40 29 30 65 39 41 43 44 77  0 9 9 8 8 7 7 6 6 6 5 5 5 5 5 5 4 4 4 4 4 3 3 3 3 3 3 3 2 2 2 2 2 2 2 2 2 1 1 1 1 0 7 0 3 2 5 1 8 5 2 8 7 6 5 2 0 7 5 4 2 1 9 8 7 5 4 3 1 9 7 6 5 4 3 2 1 0 9 8 7 6 +  1 1 3 1  9 8 7 6 5 4 3 2 1  1  + ---•£• --I I TI-I  +  ro 1  T  I I  1  ;  1 j  -p. + -I + -I-  + + -I  +  •toco:  o + + + + + + +  I I-I-I 1 Ij  + + +  1 1  + + _.  + -.  D I S T A N C  2 5 8 1 1 1 . 0 1 5 6 8 9 0 7 2 . 0 5 5 6 2 0 5 6 7 0 0 0 6 5 0 0 6 0 0 0 7 0 0 0 7 0 1  1  1 2 2 2 2 2 2 2 3 3 3 3 3 3 4 4 4 4 4 4 5 5 5 5 5 6 6 6 7 8 8 9 9 9 1 2 3 3 4 5 7 0 3 4 5 6 7 0 1 2 3 6 8 0 2 5 6 7 4 8 9 6 2 4 0 2 5  1 1 1 1 1 1 2 2 2 3 5 5 5 1 1 2 3 7 8 2 6 9 2 1 9 0 1 5 9 8 0 5 8 5 7 3 2 0 0 0 0 0 6 5 8 5 1 8 3 0 3 0 4 0 5 2 5 3 5 9 0 1 6 1 6 2 8 2 0 9 18 3 5 0 8 0 6 0 0 0 2 0 7 0 3 0 5 8 0 0 8 8 0 2 8 8 6 7 6 8 5 7 0 7 7 9 0 9 1 5 6 1 0 4 7 17 8. 0 3 0 7 0 0 0 5 0 5 0 3 0 0 3 0 8 3 3 0 5 3 1 7 7 4 3 4 9 0 2 7 7 2 8 7 8 7 3 0 3 5 5 7 6 8  241 1 1 1 8822228 39399]333667766|1 1 1 1 19066|99d888908l55555 6897SJ777777J 3151044556 31 1 22422 53 31488442423634447 16845670fe64878l345678989B97686124 133173902112634 09005)143526122 19227808178103910923 7034574645258090591347  10 HYPNDIC 3 DICHPEL A SCAPUND 7 PELLNEE 12 BLINACU 15 MARSEMA 30 JUNGOBO 9 CEPHBIC 5 RACOACI 13 RHIZGLA 1 1 SCAPSUB 8 NARDSCA 16 BRACPLU 20 BLEPTRI 26 HYGRLAX 1 SCOUAOU 37 RICCMUL 25 DIPLALB 56 CHILPOL 18 PLAGASP 6 dUNGPUM 55 PHILFON 17 P0G0U3N 80 BRYUPAL 89 dUNGEXS 39 STOKPRAP 62 OLIGPAR 33 POROBIG 44 BRACFRI 65 ANEUPIN 7 1 d'JNGATR 83 SCHIRIV 87 SCAPPAL 14 SCAPAME 36 C'.NOCON 52 SCLEORT 2 HYGROCH 40 GcOCGRA 711 DREPUMC 8G PRE I QUA 2<l CERAPUR 7!i TRITOUI 8!! FISSOSM 99 HYGRSMI 6 PLEUALB 29 ATRISEL 33 OLIGALI 48 POHLNUT 51 RACOAOU 53 HYPNSUB 54 DIPLTAX 57 BART ITH 84 BRACVEL 9 3 BRYUPSE 94  t  u  2  2  1  1 + 1344422 1  \--2-  B5333-  121  -42  1223-  b+11-  — 1--1 1 1 1-31  1342442!  1--1-  -21  21 122 +  •12+;  1  21 12-  •1  ---2 2++  2  --+1-  -+ 3 - 1  1 12-  -223--1-  -22-22  dljmCON  2 3 POLYALPA 27 ANASMIN 3 5 RACOFAS 551 SCAPSCA 6 0 RACOVAR 4• POHLCRU 4C. METZCON 4<ISOPELE 6 4 CLAOBOL GE SPHAGIR 6£ OXYSTEN SCHISTR 104 LOPHHET 109 BRYUCAP 3 2 DIPLPLI 34 RACOHETA 4 6 HETEMAC 5 8 POHLPRO 7 4 ISOPPUL 6 3 BAZZTRI 6 6 BARTLES 8 1 RACOLAN 9 0 DRYPPAT 9 5 ISOTSTO 102 MNIUBLY 103 FLATdUN 107 HYGRLUR 108 BARBREF 1 1 0 SCAPMUC 1 14 TRITPOL 19 PLAGROE 2 1 BARTPOM 2 2 RACOHETH 28 RACOBRE 31 DITRHET 91 ANOtAES 92  21 12 H-3312422312--+1I 1 1 1112 1 1-223324441312++1+22--+ 4322424 52-443322-224 1 + -222-- + + -- + 2212+1 22131 22(34 1 12-122|234|-1-12 |3 1322+1 1-3 |222 1 22(22- 1 1+-2222 32 123-3331 I-1--12144 1 - - 11|444333| 1 + 223-- + -2- 121- 1 + -3 : !1--+ 1 + -122+212-2--21 11111 + + - + 3 + 1J2 + - 1- 1 1 + 11 1 + -- 1 221214^4 4444443441 1 11 1 • + 1+' J2- 1-1 1| H 11 f 1 1-2 1--] -12+1 --21122231 145343233244 223343 + 2 -2-2-1+ +-2++1--12--34 1- 1-- 222-331+2+232- 122-222-22-- 1 12 1212 __1 \ 12+21 + 1-222- 12+1 1+1 2 + 1 -- 1 +++1(23323332232221 1 12+ -2222 +2 1-2 1-1- 1 1 + 1--22321122--3322+1 1 21-- 11-1-121 1 -111 1-212322334443 | , + 1 ++ -_;, , 1 ^33352332222+-12434432 3 232221 -1 1433 -21+1232 1 1 1-1 1 1--- 1 1--2-31 2 • -1 + --2 1 + 121-^ 11-12 2 + 11+11 •12 1 -1 12 2 111 + 1 — 11—11—1• -+ 1032444 43233443 1245-222+ _ 2 1 -3 &111 1--22213PI" f 1 : 222 1 1 r-22--2+ •1--21+21 1 1 1--1 1-2 + + 32I 1 •111 1 h + 31432 1 1 _i 11+ + 122- 1 + + 212(2122611--2 •1-2 + • 1- i a ++H-+--+ 1 3-2--2| 11 + 21 -22222 + 2 • 1 2 1 1 1-31-14-2112 42-1H--+1+---2 2 12 --433+ 1++32 -2111-21 -22--21-11-21-  •11•11 -421 --11 + 2-  ei-  1 1-  -22-  -22-  •11•11-  -23-  1 1•1 1  PSESJPAT  42 AMPHCAL 4 5 CYNO.JEN 4 7 POHLLCN 9 S FISSBRY 9 7 POREROE 9 8 PLAGROS. bO dAMEAUT 100 RHIZNIJD 101 PORECOR 6 7 DICRHET 7 0 PHILARN 7 2 ATRIUND 105 BARBLYC 106 TORTFRA 7 3 RA3UB0L 7 6 MYLITAY 7 7 PLAGLAE 7 9 RHYTSQA 111 BRYUMIN 1 12 STOKPRAS 113 DICRCRI 8 5 GRIMT! R 1 1 5 PLAGPIL 116 BLASPUS 1 1 7 LOPHSUD  N  i RC=0  $2.11  U^ . v  Ik l ? C M , .  UlM.  'f  L63 67  \ i 4  ^ > H v  242  1 1 PLAGOED 2 OXYSTEN 3 CRATCOMF 4 BRYOREC 5 LOPHHET 6 TRITEXI 7 dUNGATR 8 MNIUTHO 9 ENCAPRO 10 PLAGASP l l BRYUCAP 12 THAMNEC 13 METAMEN 14 BARBVIN 15 PLAGVEN 16 GYMNAER 17 CO.MOCON 18 POROBIG 19 P L A T J U N 20 BRACREF 21 LOPHGIL 22 HOMAFUL 23 HYPNSUB 24 D I S T C A P 25 TIMMAUS 26 TORTPRI 27 S E L I D O N 28 POHLCRU 29 BL E PTR I 30 EURHPUL 31 PORECOR 32 HOMANUT 33 CLAOBOL 34 ISOTSTO 35 DICHPEL 36 POREROE 37 CLAOCRI 38 I S O P E L E 39 SCAPAME 4 0 RACOHETA 41 S C H I S T R 42 PORENAV 43 CRYPPAT 44 TRI CC Y L 45 BRACFRI 46 CEPHBIC 47 D I T R F L E 48 BARBREF 49 GEOCGRA 50 SCHIAPO 51 S C H I R I V 52 RACOCAN 53 BRYUPSE 54 BRACRIV 55 PLAGROS 56 LEUCMEN 57 APOMPUB 58 F I S S B R Y 59 GYMNREC 6 0 SCAPUND 61 BRACVEL  11111 1 1 1 1 122222222 1234 567890123456789012341367  '  2-22--22 2-2221 121 1 1— 2 3 2 2 243222221 1 2 — 1 223 3 3 11 1 + --111 + 2 2-1 1 2221--212-2233413 + 3 211 I'+ 3 + - 3 - 3 — 33 _ _+-1 + 131111 + -41 -13 2--1 + 1+1 -2 1 231 1 1 12231 1 131 1 --2 --222 1 1 1 121 — 3 2 — 3-23 + 1-1 212 2 4 - - 1 - 1 1 - 1 1 2222 1 - 1 2— 2 1 1 1 1 12 +-+ 11 11 21 1 2 42 24 2-2 2 322 4 22-1 212 321 3 3 1 12 1 12 2 + 331 1 1 1 2 2 343 221 2--1 2321 1 +----21 — --21 -2 211 13 31 1+1 1+ + 41 +1 1 1111 + - 3 2 21 214 1 1 + — 11 4  .  •  •  2  1  243  111111111122222222 123456789012345G78901234567  17 C0N0C0N 1 PLAGOED S ENCAPRO 4 BRYOREC 22 HOMAFUL 14 BARBVIN 19 P L A T J U N 7 JUNGATR 16 GYMNAER 20 BRACREF 10 PLAGASP 23 HYPNSUB 36 POREROE 47 D I T R F L E 8 MNIUTHO 40 RACOHETA 51 S C H I R I V 53 BRYUPSE 56 LEUCMEN 5 LOPHHET 21 LOPHGIL 13 MET AMEN 24 D I S T C A P 27 S E L I D O N 31 PORECOR 35 D I C H P E L 52 RACOCAN 33 CLAOBOL 34 ISOTSTO 44 T RI CC YL 45 BRACFRI 54 BRACRIV 55 PLAGROS 58 F I S S B R Y 11 BRYUCAP 15 PLAGVEN 25 TIMMAUS 26 TORTPRI 28 POHLCRU 29 B L E P T R I 30 EURHPUL 46 C E P H B I C 48 BARBREF 49 GEOCGRA 57 APOMPUB 59 GYMNREC 61 BRACVEL 2 OXYSTEN  .  3 CRATCOMF 5 0 SCHIAPO 6 TRITEXI 12 THAMNEC 18 POROBIG 32 HOMANUT 37 CLAOCRI 38 I S O P E L E 39 SCAPAME 41 SCHISTR 42 PORENAV 60 SCAPUND 43 DRYPPAT  -2 1 2311 1 12231 1 131 1 2-22 — 22 2-2221 121 1 1 — 2221--212-2233413 2 2 243222221 1 2 — 4 - - 1 - 1 1 - 1 12222 1 - 1 -4- + --2 1+131111 + - - 222 1 1 1 121 3 1 1 1 + --1 1 1 -13 2—1! 1+1 --32--3-23+ 1-1 + 3 21 1 1 — 2--2 1 1 1 1 3 1 12 112321 1 +-+ 2 2-1 1 331 1 1 211 13 1 1+ +41 1 + -32 1 223 ---212 2 - 3 - 3 — 33 12 +-+ 21 1 2 2 322 321 3 31 1 +22-1 212 343 221 1 +1 1 1 1214 + 3 -41 11 1 1 42 24 2-2 2--1 21 21 21 1 1 11 2  3  -  -2  3 + --2 4  •  2 2 + 1 2 + — 2  1  244  GO  ^ 9 a i - 0 - A _ u c j i o ) 0 - > i J i ( n _ u _ s i i j ) u  o  -----t. 01 J> 03 ->• C/l  o i / i - n z > - ( ( / i H O O O 01 M  -J  + •••- + + + + + + + + + + + + + + -t- + + -i- + + -i- + -i- + 01 M  O O O ui o o o o  CD  o o o o  03  CO OJ U 00  ._. ro CJ M  CO  O cn --i •  _.  O O O '  01 —  M A 01 • CO O O O '  01 M  M M -- M  -  0 M CO -  CJ M  CS —  •J en  CT> • M O u i s • 01 CJ  -» 01 - -  O 01 M • ^1 CJ  01 —  J-  O O 01 • O O O '  -«1 J>  O O O -  CJ Ol  -  _. — Cd —  u—  Ol M _» • CO 01 M J - 01 • O O 01 •  01 -J  - J 01 — • M -1 01 M J - •  CJ - J  O H M -  -J  -  CJ CJ 00 • Ol CD  4>  —  CS M - -> CJ — M • M O CJ • O Ol CO J— • 09 CO  CO  O CO 00  01 Ol JCJ M  -  245  11111111 1 222J122222 134512678 2384 79056234,901567  17 CONOCON 1 PLAGOED 9 ENCAPRO 4 BRYOREC 22 HOMAFUL 14 BARBV IN 19 P L A T J U N 7 JUNGATR 16 GYMNAER 3 0 EURHPUL 10 P L A G A S P 23 HYPNSUB 36 PORFROE 47 DITIJFLE 8 MNIUTHO 4 0 RACOHETA 51 S C H I R I V 53 BRYUPSE 56 LEUCMEN 5 LOPHHET 21 LOPHGIL 13 METAMEN 24 O I S T C A P 27 SE LI DON 31 PORECOR 35 D I C H P E L 52 RACCCAN 33 C L A C B O L 34 I S O T S T O 44 T R I C C Y L 45 BRACFRI 54 BRACRIV 55 PLAGROS 58 F I S S B R Y 1 1 BRYUCAP 15 PLAGVEN 25 TIMMAUS 26 TORTPRI 28 POHLCRU 29 B L E P T R I 30 EURHPUL 46 C E P H B I C 48 BARB REF 49 GEOCGRA 57 APOMPUB 59 GYMNREC 61 BRACVEL 2 OX Y ST EN 3 CRATCOMF 50 SCHIAPO 6 T R I T EX I 12 THAMNEC 18 POROBIG 32 HOMANUT 37 CLAOCRI 38 I S C P E L E 39 SCAPAME 41 S C H I S T R 42 PORENAV 60 SCAPUND 43 DRYPPAT  -111 1|22331 1 - 2 3 1 - - 1 1 1|2 12 1 1 1 - 2 - 2 2 - 2 2 1 12222--I| 233422 1 3 - |22 1 122-| 2 1 12-2432-2222| 1 - 122-1221 -4111 "T 1 3 1 1i 1 + 1 1 1 + - + 2-1 •? 11 1-121 -2-21111 11+ 3 1 '1 + 1 - - 1 + - - - 1322  -i2 1 1 1 —  -3i-22-  •111-1---  -|1 12 11--11 1- + - - - 1-J  -2 32| -2- 1 -2-• 1 1 133-  I  I ,211-13 11 1+ + 4 1 1+1 32 I  1223-2-I3-333-- 1 -22--321-  1212 + -+, &1—•  2  I -22 -2 1  1-343 -22 1  -214-  Hi-2-4-  -2 1 -2 1 -21-11-  246  222,122222 i n u i 134512570238479056234,901567  1 PLAGOED  9 ENCAPh'j-  4 BRYOREC 22 HQMAFUL 17 CONOCON 14 B A R B V I N 19 P L A T c U N 16 GYMNAER 7 JUNGATR 8 MNIUTHO 10 PLAGASP 36 POREROE 23 HYPNSUB 47 D I T R F L E 56 LEUCMEN 21 LOPHGIL 27 SELIOON 31 PORECOR 35 D I C H P E L 33 CLAOBOL 34 ISOTSTO  46 CEPhEIC  4 0 RACOHETA 5 LOPHHET 44 T R I C C Y L 45 BRACFRI 11 3RYUCAP 48 BARBREF 49 GEOCGRA 2 OXYSTEN 3 CRATCOMF 50 5CHIAP0 6 TRI TEX I 12 THAMNEC 41 S C H I S T R 42 PORENAV 43 DRYPPAT 30 EURHPUL 13 METAMEN 15 PLAGVEN 28 POHLCRU 29 B L E P T R I 30 EURHPUL 18 POROBIG 32 HOMANUT 37 CLAOCRI 38 I S O P E L E 39 SCAPAME 24 D I S T C A P 58 F I S S B R Y 25 TIMMAUS 26 TORTPRI 57 APOMPUB 59 GYMNREC 51 S C H I R I V 53 BRYUPSE 52 RACOCAN 54 B R A C R I V 5 5 PLAGRJS 61 B R A C V E L 60  5CAPUND  b  2 - 2 2 - 2 2 1 1|2 2 2 2 - 1 1 1-23342213- 221122 -12 1 1 2 - 2432-222.' -2 - 1 2 2 - 122 1. - - 4 1 1 1 1 • 1 - 2 3 1 - - 1 1 2l| 1 1 1 12233 1 1 -3111+111+ ,4-- + 2,11-121 - 2 - 2 - - (2 1+ 1 --,1+1--132 1 1 1 1 1t+ 31-1 +2 -2 +12 1 1 1 - 3| 3 11 121 122--111-1I J11- + - -2?2 1 1+t 32 -2 , 12---I 2 PI—I --1 2 - -I - 2 2 - - 3 - - - -23j 2 1----22! 1 • ---2 1 •-2 - 11 1331223---343| -22 1  1_.  -2 1 -2 1  2 2| [3-3J3 -2-4-  12+-+- -  214i-  11---11 1 - - -,--21H-  12- 1 1 - 1 3 -li 1+ + 4!31- 1 + 1--+1 -111I 1 1 I  

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