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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 State 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 accept t h i s t h e s i s as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA Kandidaats' degree May 1981 © Johannes Petrus van Velzen, 1981 In presenting t h i s thesis i n p a r t i a l f u l f i l m e n t of the requirements for an advanced degree at the University of B r i t i s h Columbia, I agree that the Library s h a l l make i t f r e e l y a v a i l a b l e f o r reference and study. I further agree that permission for extensive copying of t h i s thesis for scholarly purposes may be granted by the head of my department or by his or her representatives. I t i s understood that copying or pu b l i c a t i o n of t h i s thesis for f i n a n c i a l gain s h a l l not be allowed without my written permission. Department of J$>crt(*nij The University of B r i t i s h Columbia 2075 Wesbrook Place Vancouver, Canada V6T 1W5 Date (2/79) through a succession of sinuous rocky banks and of c l i f f s and moss-covered rocks I gl ided along Liu Tsung-yuan (773-819). i i i 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 of bryophyte assemblages from rocky habitats that ranged in elevation from sea level to subalpine and alpine areas. Small scale variation was expressed in the segregation of various types of micro-environments on rocks, a l l characterized by th e i r own bryophyte vegetation. On a larger scale, 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) siliceous rocks along streams and (4) calcareous rocks. Within these major groups cluster analyses and tabular sorting, combined, were u t i l i z e d to detect patterns in the data set. The subse-quent recognition of groups of simi l a r releves formed the basis for 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 further characterized by micro-site 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 integration in already existing 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 sociological a f f i n i t i e s assigned to the e p i l i t h i c bryophyte associations in south i v western B r i t i s h Columbia were r e l a t e d to those i n Europe. Where necessary and a p p r o p r i a t e , new syntaxa were d i s t i n g u i s h e d and defined. F i n a l l y , the a p p l i c a t i o n of o r d i n a t i o n techniques, although l i m i t e d i n the present study, was u t i l i z e d as an a d d i t i o n a l means to d i s p l a y and i n t e r p r e t patterns i n the data s e t . The p r i o r c l a s s i f i c a t i o n , p a r a l l e l e d with 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 , already showed some r e l a t i o n s h i p between r e l e v e s . This provided a basis 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 r e l a t i o n s h i p s o f releves to one another and to the environ-ment. 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. Landforms 8 3.1.2. G l a c i a t i o n i n B r i t i s h C olumbia 12 3.1.3. V o l c a n i s m 14 3.2. Geology 16 3.2.1. Bedrock t y p e s 16 I n t r u s i v e igneous r o c k 17 Se d i m e n t a r y r o c k 18 V o l c a n i c r o c k 19 Metamorphic r o c k . . 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 Zone... • 27 5.2. The C o a s t a l Western Hemlock Zone 28 5.3. The S u b a l p i n e M ountain Hemlock Zone 30 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 31 5.5. The I n t e r i o r D o u g l a s - F i r Zone. 32 5.6. The A l p i n e Tundra Zone 33 v i 6. METHODS , 36 6.1. D e l i m i t a t i o n of e p i l i t h i c bryophyte 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 quadrat-size 38 6.2.3. Estimation of species 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 the 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. Tabular comparison 46 6.3.3. Numerical techniques 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. Ordination 54 7. SYNSYSTEMATICS 56 7.1. Terminology and nomenclature 56 8. ANALYSIS AND DESCRIPTION OF THE EPILITHIC BRYOPHYTE VEGETATION 60 8.1. Bryophyte a s s o c i a t i o n s from exposed, dry rocks 62 8.1.1. Synsystematics 65 8.1.2. Phytogeographical 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 rocks 88 8.2.1. Synsystematics 90 8.2.2. Phytogeographical s e t t i n g 91 8.2.3.1. Order LEPIDOZIETALIA REPTANTIS 93 v i i 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 rocks along streams 104 8.3.1. Synsystematics 107 8.3.2. Phytogeographical s e t t i n g 110 8.3.3.1. Order BRACHYTHECIETALIA PLUMOSI I l l 8.3.3.2. Order PLATYHYPNIDIETALIA RUSCIFORMIS 134 8.4. Bryophyte a s s o c i a t i o n s from calcareous rocks 139 8.4.1. Synsystematics 141 8.4.2. Phytogeographical s e t t i n g 143 8.4.3. A l l i a n c e E ncalyption procerae 144 8.4.4. Evaluation of the o r d i n a t i o n techniques 153 9. SUMMARY 161 REFERENCES. 167 APPENDIX I. Abundance s c a l e 180 S o c i a b i l i t y s c a l e 181 APPENDIX I I . F i d e l i t y degrees 182 APPENDIX I I I . Bryophyte species l i s t : Class Hepaticae. 184 Class Musci 189 APPENDIX IV. O r d i n a t i o n data on the bryophyte vegetation from calcareous rocks 197 APPENDIX V. D i s t r i b u t i o n maps of the bryophyte syntaxa 202 APPENDIX VI. Tabular s o r t i n g t a b l e s and c l u s t e r analyses f o r the four major groups of bryophyte vegetation r e l e v e s . . . 225 v i i i LIST OF TABLES TABLE I S i m p l i f i e d o u t l i n e of 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 Climate data from weather s t a t i o n i n the study area 24 I I I Bryophyte a s s o c i a t i o n s from exposed, dry rocks.. 63 IV Racomitrio - Andreaeetum p e t r o p h i l a e 69 V Campy!opus a t r o v i r e n s - Paraleucobryum enerve - f a c i e s 72 VI "Racomitrietum b r e v i p i s " 74 VII Gymnomitrietum conc i n n a t i 77 VII I Racomitrietum s u d e t i c i 80 IX Racomitrio - Polytrichetum p i 1 i f e r i 83 X Scapanietum americanae 86 XI Bryophyte a s s o c i a t i o n s from shaded, humid rocks 89 XII Diplophylletum a l b i c a n s 94 XIII Marsupello - Scapanietum americanae 96 XIV Claopodietum bolanderi 101 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 along streams. 105 XVI Racomitrio - Seoulerietum aquaticae 113 Dichodontietum p e l l u c i d i XVII Brachythecio - Conocephaletosum conicae 117 XVIII R i c c a r d i o - 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 i x TABLE XXII Marsupello - 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 calcareous rocks 140 XXVI Lophozio - Bryoerythrophylletum r e c u r v i r o s t r i 145 XXVII Metaneckeretum men z i e s i i 147 XXVIII Barbula v i n e a l i s - P l a t y d i c t y a jungermannioides - Junger-mannia a t r o v i r e n s - community 150 X LIST OF FIGURES Figure 1. Reference map f o r the study area 7 Figure 2. Physiographic 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 of the study area 10 Figure 3. Annual p r e c i p i t a t i o n i n south-western B r i t i s h Columbia, given i n centimeters 22 Figure 4. Vegetation zones i n south-western B r i t i s h Columbia... 26 Figure 5. Nested p l o t technique, used f o r the determination of a minimum r e l e v e ( p l o t ) - s i z e 41 Figure 6. Species-area curve 42 Figure 7. Categories of rock surface h a b i t a t s 44 Figure 8. P r i n c i p a l Components A n a l y s i s of the 27 limestone releves 154 Figure 9. Scatter-diagram of the 27 limestone r e l e v e s from the P.C.A.-cent 155 Figure 10. P o l a r o r d i n a t i o n of the 27 limestone releves 157 Figure 11. C l u s t e r a n a l y s i s of the 27 limestone r e l e v e s 158 Figure 12. C l u s t e r a n a l y s i s of P.C.A.-scores.(7 axes) o f the 27 limestone releves 160 x i ACKNOWLEDGEMENTS I am very g r a t e f u l to 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 of my b r y o - s o c i o l o g i c a l study, i n arranging the finance f o r my f i e l d w o r k and research and i n enhancing my knowledge and understanding of northern hemisphere bryophytes. I o f f e r s i n c e r e a p p r e c i a t i o n f o r h i s good advice during the term of my study and i n the w r i t i n g of t h i s t h e s i s . I am a l s o indebted to him the use of many unpublished B r i t i s h Columbia d i s t r i b u t i o n maps of 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 the use of 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 valuable help i n the computer analyses o f the vegetation data. A l s o , I thank Rob Scagel f o r his time and e f f o r t to help me c a r r y i n g out these analyses and d i s c u s s i n g t h e i r r e s u l t s . 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 appre-c i a t i o n f o r h i s valuable help and pleasant companionship i n the f i e l d . I am indebted to my parents, Mr. and Mrs. J . van Velzen-van Meer, f o r t h e i r encouragement throughout the years of my academic education. In p a r t i c u l a r , g r a t e f u l l y recognized i s the time I spent a t the I n s t i t u t e f o r Systematic Botany of the State U n i v e r s i t y of Ut r e c h t , Holland where my i n t e r e s t i n bryology was born and grew under the e l e c t r i f y i n g i n f l u e n c e of Dr. S. R. Gradstein. I thank the f o l l o w i n g persons who helped i n the t y p i n g o f the manuscript and the f i n a l t h e s i s : Eta Thiessen, Dr. R. Turkington. x i i I thank Deanne Brethower, O l i v i a Lee and John Pinder-Moss of the Herbarium of the U n i v e r s i t y of B r i t i s h Columbia, f o r t h e i r help i n typing the l a b e l s f o r the herbarium specimens. To many people I would l i k e to express my s i n c e r e thanks and a p p r e c i a t i o n f o r he l p i n g me, m a t e r i a l l y or morally: V i j k o Lukkien, Paul van Westendorp, A l l e n Banner, S h e i l a Humphrey, Leni G e l t e n , Nechemjah Cohen, Benito Tan, Ross H i l l , Terry Mcintosh, John Wehr, Debby Donaldson, Trevor Goward, W i l f N i c h o l l s , Tim and Suzanne Thompson, Michael Dunn. 1 1. INTRODUCTION. The o b j e c t i v e s of t h i s study are to sample, analyze and c l a s s i f y the e p i l i t h i c bryophyte communities as they occur i n south-western B r i t i s h Columbia. D e s c r i p t i o n s of the separate vegetation u n i t s (syntaxa) are concentrated on the r e c o g n i t i o n of d i a g n o s t i c species (character species and d i f f e r e n t i a l s p e c i e s ) , m i c r o - s i t e s p e c i f i c i t y of species and communities and t h e i r geographic d i s t r i -b ution throughout the study area. Secondly, 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 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 described i n Europe (a.o. Hertel 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 with the expansion o f knowledge o f e p i l i t h i c bryophyte a s s o c i a t i o n s of North America and temperate A s i a . I t i s evident t h a t i n North America ( i n c l u d i n g the west coast) 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 are 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 techniques of 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 of vege t a t i o n . In the study area and the adjacent regions of the present study, some e c o l o g i c a l research has been done on the bryophyte (and l i c h e n ) vegetation of Vancouver I s l a n d (Szczawinski 1953, Hubschmann 1978) and the Olympic Peninsula i n Washington, U.S.A. (Hoffman and Kazmierski 1969). Szczawinski (1953), i n an extensive study on t h e c o r t i c o l o u s 2 and 1 i g n i c o l o u s plant communities i n the f o r e s t a s s o c i a t i o n s of the Douglas F i r f o r e s t on Vancouver I s l a n d , described r e l a t i o n s between occurrence, abundance, dominance, constancy and.vigour of bryophytes and l i c h e n s and he d i s t i n g u i s h e d ten s o c i a t i o n s of c o r t i c o l o u s and 1 i g n i c o l o u s ( = e p i x y l i c ) cryptogams. Hubschmann (1978), 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 the Ziirich-Montpel 1 i e r School method of vegetation c l a s s i f i c a t i o n . Beside a q u a t i c , t e r r e s t r i a l and e p i p h y t i c bryophyte communities, he d i s t i n g u i s h e d and described seven e p i l i t h i c bryophyte a s s o c i a t i o n s and provided some inf o r m a t i o n on physiognomy and d i s -t r i b u t i o n of these vegetation u n i t s . Knowledge on the ecology of bryophytes i s scant i n comparison with that of higher p l a n t s . Studies of bryophytes up to about 1930 were mainly l i s t s of species i n c e r t a i n areas and h a b i t a t s , sometimes with comments on the environmental r e l a t i o n s or m i c r o - h a b i t a t s . Later works contain data on abundance, cover and frequency of species. E s p e c i a l l y i n Europe, and to a l e s s e r degree i n Japan, the a p p l i c a t i o n of the BraurfBlariquet r e l e v e technique l e d to a c l a s s i f i -catory system of bryophyte communities on a v a r i e t y of s u b s t r a t a such as t r e e s (Barkman 1969, Iwatsuki 1960), rocks (Hertel 1974, Sjogren 1964), decaying wood ( P h i l i p p i 1965) and s o i l (Hubschmann 1975, Wald-heim 1947). North American b r y o - e c o l o g i c a l research, e s p e c i a l l y r e c e n t l y , has concentrated mainly on s t a t i s t i c a l and m u l t i v a r i a t e a n a l y s i s of community s t r u c t u r e and composition, d i s t r i b u t i o n and e c o l o g i c a l r e l a -3 t i o n s h i p s between bryophytes, t h e i r s ubstrate and environment (a.o. Redfearn 1957, 1960, Foote 1963, 1966, Slack 1971, Culberson 1955, Hale 1955). E a r l i e r b r y o l o g i c a l 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 the r e c o g n i t i o n of more or l e s s d i s c r e t e vegetation u n i t s , e s p e c i a l l y on t r e e s . Cain and Sharp (1938), u t i l i z i n g coverage and presence of bryophytes i n c e r t a i n f o r e s t types of the Great Smokey Mountains, were able to c l a s s i f y c o r t i c o l o u s (=bark-inhabiting) commu-n i t i e s . In another study from the same geographic 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 of c o r t i c o -lous bryophyte communities. These workers described "unions" of bryophy-t e s , dominated by one or two s p e c i e s . At Mountain Lake, V i r g i n i a , Patterson (1940) recognized 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 the dominant species i n quadrats on random mature t r e e s . The occurrence and d i s t r i b u t i o n o f these s o c i e t i e s was found to depend upon d i f f e r e n t l e v e l s of evaporational s t r e s s and d i f f e r e n t moisture holding c a p a c i t y of 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) recognized a s s o c i a t i o n s of bark-i n h a b i t i n g bryophytes i n three regions of the State o f Michigan. The a s s o c i a t i o n s were based on and named a f t e r the dominant species (those with highest coverage, constancy and v i t a l i t y ) . A major Canadian study employing the same types of methods as the previous works i s the i n v e s t i g a t i o n by LeBlanc (1962) on the ecology and phytosociology of cryptogamic epiphytes i n southern Quebec. 4 Coverage and dominance w i t h i n quadrats l e d to the d i s t i n c t i o n and r e c o g n i t i o n o f e p i p h y t i c s o c i e t i e s of bryophytes 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 to recognize and describe e p i l i t h i c bryophyte assemblages. Major a t t e n t i o n i s paid to the i n v e s t i g a t i o n of the s i g n i f i c a n c e of using the t a b u l a t i o n technique, e s s e n t i a l i n the Braun-Blanquet method of vegetation c l a s s i f i -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 m u l t i v a r i a t e techniques. In the hope th a t some c o o r d i n a t i o n can be achieved between these approaches i n t h e i r a p p l i c a t i o n to bryo-ecology, the present study has been undertaken. 5 2. LOCATION OF THE STUDY AREA. The area of the present study i s l o c a t e d i n south-western B r i t i s h Columbia, Canada's westernmost province. I t s l o c a t i o n i s shown i n Figure 1. Boundaries of the area are e s s e n t i a l l y the mainland coast-l i n e on the west; the Canada-U.S.A. border on the south; the moun-t a i n range south o f the 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 to the south-east; and, f i n a l l y , Pemberton to the north. To provide f a m i l i a r i t y w i t h t h i s region and a basis 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 v e g e t a t i o n , the physio-graphy, geology, c l i m a t e and vegetation o f the study area are described i n the f o l l o w i n g chapters. Regional climate (macroclimate), physiography, and geology i n f l u e n c e the (v a s c u l a r ) vegetation which develops i n a re g i o n . A l l four elements determine the a v a i l a b i l i t y of h a b i t a t s f o r bryophytes, i n ge n e r a l , and the development of e p i l i t h i c bryophyte assemblages, i n p a r t i c u l a r . The proximity to the P a c i f i c Ocean and the v a r i e d topography of the study area, c r e a t i n g a great d i v e r s i t y of 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 region f o r a study of e p i l i t h i c bryophyte communities. 7 Figure 1. Reference map f o r the study area. 1. Black Tusk Meadows 2. Brandywine F a l l s 3. B r i d a l V e i l F a l l s 4. B r i t t a n i a Beach 5. Callaghan Creek 6. Cheakamus River V a l l e y 7. C h i l l i w a c k Lake 8. Chipmunk Creek 9. Diamond Head 10. Ford Mountain 11. G a r i b a l d i Lake 12. Cypress Bowl 13. Lighthouse Park 14. Lonetree Creek 15. Lynn Canyon 16. Mount Seymour 17. Murrin P r o v i n c i a l Park 18. Nairn F a l l s 19. Panorama Ridge 20. Porteau Camp 21. Shannon F a l l s 22. Slesse Creek 23. Stoney Creek 24. Sunset Creek 25. W h i s t l e r Mountain 26. White C l i f f Park, Horseshoe Bay 27. Sumas Mountain 28. Agassiz Mountain 8 3. PHYSIOGRAPHY AND GEOLOGY OF THE STUDY AREA. 3.1. Physiography. 3.1.1. Landforms. The area of t h i s study l i e s w i t h i n the C o r d i l l e r a n Region, one of the f i v e major physiographic s u b d i v i s i o n s recognized f o r Canada (Holland 1976). An extreme v a r i e t y i n topography i s shown i n the presence of an impressive 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 the several B r i t i s h Columbian physiographic s u b d i v i s i o n s (Holland 1976), two are represented i n the study area: the Coastal Trough and the Coast Mountain Area which encompasses the Coast Mountains and the Northern Cascade Mountains. To the east the study area i s bounded by the I n t e r i o r Plateau (Figure 2 ). In the study area the Coastal Trough (1) i s represented by a minute s e c t i o n of the Georgia Lowland (a narrow s t r i p on the mainland southeastward from Horseshoe Bay) and by the Fraser Lowland. The Georgia Lowland i s underlain 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 eros i o n . The Fraser Lowland i s a part o f the Georgia Lowland, from which i t d i f f e r s i n being a l o w - l y i n g area of 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 Point Grey f o r 110 km to Laidlow and southeastward to the coast at Bellingham, Washington, U.S.A. The Fraser Lowland c o n s i s t s of 10 Figure 2. Physiographic subdiv i s ions and genera l ized geology of the study area. LEGEND. Physiographic subd iv i s ions : 1. Coastal Trough 2. I n te r io r Plateau 3. Coast Mountain Area A. Coast Mountains B. Cascade Mountains Height of Mountains: A below 800 m over 800 m General ized geology: Fo l i a ted metamorphic rock. ^ V A . A Intrus ive igneous rock, Late Mesozoic or T e r t i a r y Folded and fau l ted vo lcan ic and sedimentary rock c h i e f l y Mesozoic F l a t or gently d ipping sedimentary rock. Cretaceous and younger 11 extensive low h i l l s ranging i n e l e v a t i o n from 15 to 300 meters separa-ted by wide, flat-bottomed v a l l e y s (Armstrong 1954). The d e l t a of the Fraser River has been the s i t e o f sedimentary d e p o s i t i o n s i n c e the l a t e Cretaceous. Deep d r i l l i n g has shown th a t the g r a n i t e basement i s o v e r l a i n by Cretaceous, T e r t i a r y and Quaternary sedimentary rocks. Recent d e p o s i t s , s t i l l i n the process of formation, c o n s i s t of d e l t a i c and f l o o d - p l a i n deposits of the Fraser River as i t b u i l d s i t s d e l t a seaward at the r a t e of about 8.5 m a year (Holland 1976). The major physiographic s u b d i v i s i o n (Holland 1976) represented i n the study area i s the Coast Mountain Area (3) which encompasses two regions, the Coast Mountains (3A) and the Cascade Mountains (3B). As an unbroken mountain chain the Coast Mountains extend from t h e i r south end at the Fraser River northwestward along the mainland c o a s t l i n e o f B r i t i s h Columbia i n t o the Yukon. Of these Coast Mountains the P a c i f i c Ranges, making up the major part of the area of the present study, comprise the e s s e n t i a l l y g r a n i t i c mountains extending southeastward from the B e l l a Coola R i v e r f o r about 480 km to the Fraser R i v e r mouth. The P a c i f i c Ranges contain the highest peaks i n the Coast Mountains: Mount Haddington ( e l e v a t i o n 4017 m) l i e s north of the study area. High mountains c l o s e , or w i t h i n , the study area includ e Mount G i l b e r t (3110 m) and Mount G a r i b a l d i (2679). The Coast Mountains comprise sedimantary and v o l c a n i c rocks (middle 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 rocks that are e s s e n t i a l l y g r a n o d i o r i t e and quartz d i o r i t e . The Cascade Mountains extend northward from the northern border 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 of Paleozoic 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 intruded by g r a n i t e b a t h o l i t h s . The summits of the peaks and ridges a t t a i n an approximately uniform e l e v a t i o n , with the exception of some of the highest peaks i n Oregon and Washington such as Mount Baker (3286 m) and G l a c i e r Peak (3180 m), Mount St. Helens (3225 m) and Mount Hood (3745 m). A l l are v o l c a n i c cones b u i l t up by the accumulation of the p l e i s t o c e n e and recent lava and ash upon the T e r t i a r y erosion surface. Rich i n mountain features such as hanging v a l l e y s , tarns and g l a c i a t e d rock s u r f a c e s , the Cascades' rugged peaks, ridges and cirques give evidence of extensive a l p i n e g l a c i a t i o n . To the east the study area i s bounded by another physiographic s u b d i v i s i o n (Holland 1976) termed here the I n t e r i o r Plateau ( 2 ) . Most of t h i s area i s drained by the Fraser R i v e r and i t s t r i b u t a r i e s . I t s f l a t or gently r o l l i n g s u r f a c e , which l i e s f o r most part below 900 m, i s covered with g l a c i a l d r i f t and has few exposures of bedrock. 3.1.2. G l a c i a t i o n i n B r i t i s h Columbia. A b a s i c knowledge of g l a c i a t i o n i s important i n the c o n s i d e r a t i o n of landforms i n B r i t i s h Columbia, because the Province was i n t e n s e l y g l a c i a t e d during the P l e i s t o c e n e . The topography everywhere shows the e f f e c t s of g l a c i a l erosion and d e p o s i t i o n , almost e n t i r e l y of the Wis-consinan stage. Holland (1976) s t a t e s t h a t g l a c i a t i o n i n mountain regions i s considered to f o l l o w a c y c l e . I t begins with the formation of small 13 cir q u e g l a c i e r s which, by continued growth, expand i n t o mountain and v a l l e y g l a c i e r s , then formation of a mountain ice-cap whose movement i s c o n t r o l l e d by underlying topography, and u l t i m a t e l y , at the maximum stage, by a regional ice-sheet c o n t r o l l e d f o r the most part by c l i m a t e . As the i c e cover diminishes there i s r e g r e s s i o n to the mountain i c e -cap w h i l e lobes of the main ice-sheet p e r s i s t longer i n c o l d e r , protec-ted v a l l e y s (Mathews 1951). The l a s t i c e to remain i s i n small c i r q u e g l a c i e r s as i n the beginning of the c y c l e . The C o r d i l l e r a n Region i n B r i t i s h Columbia was covered e n t i r e l y by the C o r d i l l e r a n i c e - s h e e t . The P l e i s t o c e n e began with the accumu-l a t i o n o f i c e at several major centres and numerous subordinate ones (Holland 1976). The d i s t r i b u t i o n o f present day g l a c i e r s , to some-extent i n d i c a t e major, centres.;;of: formation of P l e i s t o c e n e i c e . With respect to the present study area these were i n the P a c i f i c Ranges of the Coast Mountain area between B e l l a Cool a and G a r i b a l d i . Ice moved across B r i t i s h Columbia from the Coast Mountains i n a general e a s t e r l y d i r e c t i o n across the Fraser Plateau. Elsewhere i n the v i c i n i t y o f the present study area the i c e has recorded the d i r e c t i o n of i t s movement by the landforms i t created. The i c e moved westward from the mainland across the northern end of Vancouver I s l a n d , southward and southeastward down the S t r a i t o f Georgia and southwestward across the southern end of 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 to i n d i c a t e t h a t there were at l e a s t two major advances of the C o r d i l l e r a n i c e - s h e e t , separated by an i n t e r -g l a c i a l stage. Late P l e i s t o c e n e events i n south-western B r i t i s h Columbia 14 are summarized i n Table I. The P l e i s t o c e n e i c e , w i t h a thickness of as much as 2400 m i n some areas, depressed the land with respect to i t s e a r l i e r l e v e l . Since disappearance of the i c e - s h e e t , the land surface has r i s e n , and regained i t s mountainous topography from before the onset of the P l e i s t o c e n e . 3.1.3. Volcanism. 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 occurred s i n c e mid-T e r t i a r y time. With respect to the present study area several centres of P l e i s t o c e n e and recent v o l c a n i c a c t i v i t y extend i n a l i n e northward from Mount G a r i b a l d i . There i s evidence of some a c t i v i t y w i t h i n the l a s t few hundred y e a r s , much of i t having taken place s u f f i c i e n t l y recent that the c h a r a c t e r i s t i c v o l c a n i c landforms, such as cones and l a v a p l a i n s have been l i t t l e modified by erosion and g l a c i a t i o n (McKee 1972). Within h i s t o r i c times there have been a c t i v e volcanoes i n the Cascade Mountains of Washington and Oregon. Of these p l e i s t o c e n e s t r a t o -volcanoes, Mount Baker emitted steam and t o x i c fumes during 1975 which melted g l a c i a l 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 with a major eruption i n May 1980 and a subsequent decrease i n s i z e . 15 Table I. S i m p l i f i e d o u t l i n e of 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). ERA AGE (1 0 6 y r s ) PERIOD (Epoch) GEOLOGICAL EVENTS (Recent) P o s t - g l a c i a l u p l i f t of land. ca. 2.5 27 QUATERNARY (PIeistocene) o o >-Di cu c OJ cm o cu G l a c i a l e r o s i o n ; deposits of d r i f t , outwash; ice-marginal 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 Mountains; increased steam e r o s i o n ; formation of mountainous topography. Depositions o f sediments w i t h i n c o a s t a l trough and on western s i d e of Is l a n d Mtns reduction o f land surface to one of low r e l i e f ; volcanism of southern Vancouver I s l a n d . 65 cu cu o cu to Q_ Ancient erosion land surface u p l i f t e d , 2 axes of greatest u p l i f t were along Islan d and Coast Mtns. (separated by an anc e s t r a l trough). o I—I o M O OO CRETACEOUS Land surface o f low r e l i e f ; e r o s i o n of (Upper) mountains l e d to the formation of s e d i -ments which were deposited i n marine and b r a c k i s h water b a s i n s . 110 16 3.2. Geology. The Coast Range orogeny, which l a s t e d from e a r l i e s t J u r a s s i c time through post Lower Cretaceous, together with extensive 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 of the study area. Much of t h i s g e o l o g i c a l h i s t o r y i s recorded by landforms many of which are s c u l p t u r e d on bedrock by the forces of er o s i o n (Holland 1976). Since t h i s study deals with e p i l i t h i c bryophyte vegetations i t i s important to t r e a t the d i f f e r e n t bedrock types on the b a s i s o f the phys 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 the o v e r - a l l form of the topography. Subsequently, these landform types r e f l e c t the a v a i -l a b i l i t y o f rock s u b s t r a t a and h a b i t a t s and determine the development of v ascular p l a n t v e g e t a t i o n , on which, i n t u r n , some e p i l i t h i c bryophyte assemblages are e c o l o g i c a l l y dependent. 3.2.1. Bedrock types. The landforms w i t h i n the physiographic regions discussed before are commonly underlain by r e l a t e d bedrock types and have undergone the same geomorphic h i s t o r y . The character of these landforms i s i n f l u e n c e d by the r e a c t i o n of bedrock to weatering and e r o s i o n . The r a t e o f physical weathering or mechanical d e s i n t e g r a t i o n of a rock i s i n f l u e n c e d by the presence of Tines of weakness and pore spaces t h a t allow water, . a i r and temperature changes to penetrate beneath the s u r f a c e . Thus, densely j o i n t e d rocks such as b a s a l t , or very porous material such as 17 sandstone, are more r e a d i l y f r a c t u r e d than massive, s p a r s e l y j o i n t e d g r a n i t e s (Ryder 1978). The m i n e r a l o g i c a l composition o f a rock determines i t s suscep-t i b i l i t y to chemical weathering, s i n c e minerals d i f f e r i n t h e i r r e s i s -tance to chemical a t t a c k . I n t r u s i v e igneous rock (see Figure 2) These are t y p i c a l l y coarse c r y s t a l l i n e rocks 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 types, to d i o r i t e . They are r e l a t i v e l y r e s i s t a n t to weathering s i n c e they are composed l a r g e l y o f durable minerals ( q u a r t z , hornblende, f e l d s p a r ) arranged as a cohesive f a b r i c of i n t e r l o c k i n g c r y s t a l s . As a r e s u l t , slopes on these rocks are g e n e r a l l y steep 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 to weathering, some d i s i n t e g r a t i o n and decomposition o f t h i s type o f bedrock i n e v i t a b l y occurs. Mechanical breakdown through processes such as f r o s t s h a t t e r i n g tends to produce r e l a t i v e l y l a r g e fragments. Extremely coarse rubble on c o l l u v i a l slopes and la r g e boulders i n t i l l c h a r a c t e r i z e t h i s t e r r a i n type. A combination of phys 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 causes granular d i s i n t e g r a t i o n (Ryder 1978). Components l i k e c a l c i c f e l d s p a r are most s u s c e p t i b l e to at t a c k by chemical weathering; c r y s t a l s o f quartz and potash and sodic f e l d s p a r are released to form a sandy or g r i t t y r esidue. T i l l s of t h i s residue and much sandy outwash are thus a s s o c i a t e d with areas of coarse c r y s t a l l i n e rocks. 18 Sedimentary rocks (see Figure 2) The r e s i s t a n c e o f sedimentary rocks to weathering and er 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 composition, degree of i n d u r a t i o n , i . e . the hardening through cementation, pressure and heat. Poorly indurated 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 are e a s i l y eroded. Well in d u r a t e d , massive sandstone ( e.g. depositions on the Islands i n the S t r a i t of Juan de Fuca and Georgia 2 S t r a i t as w e l l as the Sumas Escarpment), limestones ( o c c u r r i n g w i t h i n the study area i n the C h i l l i w a c k R i v e r v a l l e y ) and conglomerates are r e l a t i v e l y r e s i s t a n t to e r o s i o n . P h y s i c a l d i s i n t e g r a t i o n o f sedimentary rocks produces fragments of a s i z e r e l a t e d to the spacing of l i n e s of weakness i n the bedrock. Thus s h a l e s , which are 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 that 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 other extreme conglomerates shed l a r g e blocks which s u r v i v e i n t a c t during g l a c i a l t r a n s p o r t . Besides t h i s p hysical 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 the cementing agent by s o l u t i o n o r other 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 dipping sedimentary rocks shows a strong r e l a t i o n s h i p between rock type, s t r u c t u r e and topography. I n d i v i d u a l ridges or mountains are commonly asymmetric with steeper scarp and g e n t l e r dip s l o p e s ; dips o f over 45^ produce extremely jagged 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 with f a u l t planes or steep bedding planes (Ryder 1978). 19 Volcanic rocks (see Figure 2) Topography upon these rock types tends to be s i m i l a r to th 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 lava flows form plateaus 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 B r i t i s h Columbia. Metamorphic rocks (see Figure 2). No p a r t i c u l a r type o f t e r r a i n can be a s c r i b e d to metamorphic rocks i n general. T h e i r r e s i s t a n c e to weathering and erosion depends upon t h e i r i n d i v i d u a l c h a r a c t e r i s t i c s . Many metamorphic rocks such as gneiss and s c h i s t are f o l i a t e d , t hat i s , the minerals are segregated i n t o bands w i t h i n the rock. Meta-morphic q u a r t z i t e s are extremely r e s i s t a n t to weathering. In g e n e r a l , weathering products are s i m i l a r to those o f coarse textured igneous rocks of the same composition. 1. This i s the c a l c u l a t e d r a t e of advance at a depth o f 90 m. 2. The limestones o c c u r r i n g i n the study area are of Carboniferous age and are as s o c i a t e d with q u a r t z i t e and a r g i l l i t e i n ste e p l y dipping d e p o s i t s . The limestone as sampled and analyzed by Goudge (1944) contained over 3% of s i l i c a , some had over 10%. The magnesium carbonate content, however, i s low as i t ranged from 1.3 to 2.5%. 20 4. CLIMATE OF THE STUDY AREA. On a macro-scale, c l i m a t e a s s o c i a t e d with the major landscape region i s a product of geographical l o c a t i o n and large s c a l e topography. On the m i c r o - s c a l e , c o n d i t i o n s at the earth's surface ( i . e . rock sur-face) are modified by l o c a l physiographic f a c t o r s such as s l o p e , aspect, e l e v a t i o n and surrounding vegetation (Schaefer 1978). Two f a c t o r s play a dominant r o l e i n determining the climate of the study area: (1) the P a c i f i c Ocean and (2) the Coast Mountains (Kendrew and Kerr 1955). Onshore westerly oceanic a i r masses impinge upon successive 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 d i r e c -t i o n . Since these are roughly perpendicular to the mean winds a l o f t , they determine to a major extent the o v e r a l l d i s t r i b u t i o n of 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 westerly winds a l o f t drop considerable moisture over extensive areas, p a r t i c u l a r l y w esterly exposures and at higher elevations.- East f a c i n g slopes and lowlands r e c e i v e s u b s t a n t i a l l y l e s s . Snowfall accounts f o r a small f r a c t i o n o f the annual p r e c i p i t a t i o n near sea l e v e l . In s h o r t , the c l i m a t e of 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 of temperature extremes, high amount of p r e c i p i t a t i o n and frequent cloud-cover and fog. Climate data (from Environment Canada 1970) as obtained 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 given i n t a b l e I I . 71 22 Figure 3. Annual p r e c i p i t a t i o n i n southwestern B r i t i s h Columbia, given i n centimeters. Measurements i n centimeters converted from measurements i n inches given by Chapman and Turner (1956). LEGEND. over 381 101-152 254-381 76-101 A 7v 152-254 23 Annual p r e c i p i t a t i o n i s shown i n Figure 3. The map was sketched f o l l o w i n g a map compiled by the D i v i s i o n of Geography, U n i v e r s i t y Of B r i t i s h Columbia (Chapman and Turner 1956). Table I I . Climate data from weather s t a t i o n s i n the study area (from Environment Canada 1970). c o •c ta cu E Q-• "OSt •r™• ta > Annual mean rainfal r— "(0 <1 1—' LO 4-S t a t i o n L a t i t u d e Longitude Annual mean rainfal Annuc mean snowl • ta o C 4-> CU ,— ,—, c C r — 03 ta -c -a +-> UJ Annual mean rainfal Annuc mean snowl O >-«t +-> Q. QJ ("0 =t*= s PEMBERTON 50 27 N 122 56 W 223 742 2824 1024 7.2 128 GARIBALDI 49 59 N 123 08 W 366 1301 4229 1724 SQUAMISH 49 42 N 123 09 W 1.5 1916 1455 2061 8.9 84 WOODFIBRE 49 40 N 123 16 W 6 2856 1466 3003 HOLLYBURN RIDGE 49 22 N 123 12 W 951 2128 8108 2939 5.1 164 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 1068 9.8 57 SUMAS CANAL 49 07 N 122 07 W 6 1702 1034 1805 10.9 36 ABOTTSFORD 49 01 N 122 22 W 60 1425 782 1503 9.4 76 BRIDAL VEIL FALLS 49 12 N 121 45 W 61 846 1648 10.2 CHILLIWACK RIVER (Mt. Thurston) 49 05 N 121 46 W 198 1276 998 1396 25 5. VEGETATION OF THE STUDY AREA. Related to q u i t e d i s t i n c t seasons, c l i m a t e s and a wide v a r i e t y of t e r r a i n and s o i l 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-t a t i o n . The d e s c r i p t i o n of the vegetation of the present 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 the province i n t o 4 b i o g e o c l i m a t i c formations comprising 7 b i o g e o c l i m a t i c regions which include 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 (vegetation and fauna) and s o i l s , being the products of g e o l o g i c a l parent m a t e r i a l s , s o i l animal a c t i v i t y , topography, anf the e f f e c t s o f c l i m a t e and time upon each of these. Generally each b i o g e o c l i m a t i c zone (=geographic area having c h a r a c t e r i s t i c vegetation with a s s o c i a t e d animals, s o i l s and climate) can be broken i n t o subzones which are d i s t i n g u i s h e d by the climax p l a n t a s s o c i a t i o n (=grouping of s i m i l a r p l a n t communities) on a mesic s i t e . This s i t e i s the mid-point of the n u t r i e n t and moisture range o c c u r r i n g w i t h i n the subzone; i t i s t h e r e f o r e not subject to excessive drainage nor to seepage waters and hence the vegetation best r e f l e c t s the macro-climate (Jones and Annas 1978). Each subzone possesses a s e r i e s of communities i n h a b i t a t s which, other than by macro-climate, are i n f l u e n c e d by f a c t o r s such as d i f f e r e n t s o i l s , drainage, moisture regime, s l o p e , aspect and micro-climate. The 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 present study area, are shown on the map i n Figure 4 which i s derived from K r a j i n a ' s map, " B i o g e o c l i m a t i c Zones of B r i t i s h Columbia", published by the B r i t i s h 26 Figure 4. Vegetation zones i n south-western B r i t i s h Columbia. VEGETATION ZONES IN SW BRITISH COLUMB IA . ] Coastal Douglas F i r Coastal Western Hemlock Subalpine Mountain Hemlock Subalpine Engelmann Spruce-Subalpine F i r I n t e r i o r Douglas F i r Alpine Tundra Gl a c i e r s 27 Columbia E c o l o g i c a l Reserves Committee. These vegetation zones and c h a r a c t e r i s t i c vascular plants are discussed below. Since t h i s study focuses on bryophyte vegetation w i t h i n some major vegetation zones, l i v e r w o r t s and mosses are mentioned t h a t are c h a r a c t e r i s t i c taxa o c c u r r i n g i n these zones. 5.1. The Coastal Douglas-Fir Zone. This zone [termed Boreomeridional zone by Hamet-Ahti (1965)] i s s i t u a t e d along the south coastal region i n the rainshadow of the Vancou-ver I s l a n d and the Olympic Mountains of 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 to 450 m on the east coast Of Vancouver I s l a n d and from sea l e v e l to 150 m on the mainland and Gulf I s l a n d s . .. 2 In the t r e e stratum, the dominant t r e e , Pseudotsuga m e n z i e s n , occurs together w i t h species such as Tsuga h e t e r o p h y l l a , Abies grandis 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 understory vegetation includes taxa such as G a u l t h e r i a s h a l l o n , Berberis nervosa, Ribes sanguineum, Vaccinium  p a r v i f o l i u m , Polystichum 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 the Coastal Douglas-Fir Zone i n t o two subzones. The wetter 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 contorta and Thuja p l i c a t a . The vegetation of the d r i e r subzone includes species such as Quercus garryana, 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 , Brachythecium 28 asperrimum, Buxbaumia pi p e r i , Calypogeia m u l l e r 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 , 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 , Dicranoweisia c i r r a t a , Dicranum fuscescens, JJ. scoparium, Douinia ovata, Drepanocladus aduncus, Homalothecium n u t a l l i i , Hypnum subimponens, Isopterygium elegans, Lophocolea c u s p i d a t a , L. h e t e r o p h y l l a , Metzgeria conjugata, Metaneckera m e n z i e s i i , Neckera dou-gl a s i i , Orthotrichum c o n s i m i l e , 0. l y e l l i i , P h i l o n o t i s fontana, P l a g i o c h i l a  a s p l e n i o i d e s , PIagiothecium denticulatum, Polytrichum alpinum, P_. j u n i -perinum, P_. p i l i ferum, 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 , Porotrichum b i g e l o v i i , P t i 1 idium c a l i f o r n i c u m , P_. pulcherrimum, Radula  b o l a n d e r i , Racomitrium canescens, R. heterostichum, Rhytidiadelphus  squarrosus, R. t r i q u e t r u s , Scapania americana, S_. b o l a n d e r i , Scieropodium  caespitans, Sphagnum capillaceum, 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_. tenel 1 um, w a r n s t o r f i i , Tetraphis pel l u c i d a . 5.2. The Coastal Western Hemlock Zone. Within the study area t h i s zone [enclosed i n the Hemiboreal zone by Hamet-Ahti (1965)] encompasses productive temperate maritime, 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 the d r i e r Coastal Douglas-Fir Zone, but below the Subalpine 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 are hemlock-moss communities ( K r a j i n a 1965) recognized by a combination of 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  splendens, Isothecium s t o l o n i f e r u m , PIagiothecium undulatum and 29 Rhytidiadelphus 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 successional t r e e species to the climax Tsuga h e t e r o p h y l l a . In the wetter subzone, t h i s l a t t e r species i s o f t e n accompanied by Abies a m a b i l i s . Plant communities w i t h i n the Coastal Western Hemlock Zone vary with l o c a l topography and slope p o s i t i o n . Rock outcrops and areas with we l l drained s o i l s , p a r t i c u l a r l y those with 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 carpet of l i c h e n s and bryophytes. Depressional areas and lower s l o p e s , where there i s permanent seepage water, maintain a mixed f o r e s t of Tsuga h e t e r o p h y l l a and Thuja p i i c a t a with an extensive understory of f e r n s . Bryophytes: Andreaea r u p e s t r i s , Apometzgeria pubescens, Bazzania 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 , Calypogeia neesiana, 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  fuscescens, D_. scoparium, F r u l l a n i a t a m a r i s e r ssp. n i s q u a l l e n s i s , Heterocladium macounii, H_. procurrens, Hookeria 1 ucens, Hylocomi um splen- dens, Hypnum subimponens, L e p i d o z i a reptans, 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 , Metzgeria conjugata, Mylia t a y l o r i , 0 1igotrichum 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, P h i l o n o t i s  fontana, 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 , Plagiothecium  denticulatum, P_. laetum, Polytrichum alpinum, 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. complanata, Racomitrium heterostichum, R. lanuginosum, Rhizomnium glabrescens, I*, punctatum, R. venustum, Rhytidiadelphus t r i q u e t r u s , Sphagnum f i m b r i a turn, S_. f us cum, S_. imbricatum, S_. mageTlanicum, S^ . russowi , Tetraphis p e l l u c i d a , Ulota megalospora. 30 5.3. The Subalpine Mountain Hemlock Zone. Occurring above the Coastal Western Hemlock Zone and below the A l p i n e Zone, t h i s zone includes t e r r a i n along the P a c i f i c Coast of B r i t i s h Columbia at a l t i t u d e s from 915 to 1680 m. The Subalpine Mountain Hemlock Zone [i n c l u d e d i n Hamet-Ahti's (1965) Upper Oroboreal zone] i s c h a r a c t e r i z e d by rugged topography, with rock outcrops, r i d g e s , t a l u s s l o p e s , narrow steep-sided v a l l e y s , f a s t f l o w i n g mountain streams and heavy snow accumulation during w i n t e r time. Within the Subalpine Mountain Hemlock Zone, two subzones are d i s t i n g u i s h e d : the Forest Subzone and the Parkland Subzone. Forest Subzone. In t h i s lower subzone (at e l e v a t i o n s of approximately 900 - 1100 m) the major t r e e species are Tsuga mertensiana, Abies  amabilis and Chamaecyparis nootkatensis, with the l a t t e r g e n e r a l l y occupying h a b i t a t s w i t h abundant moisture. Dominant shrubs on mesic s i t e s are Vaccinium alaskaense, V_. membranaceum and Menziesia 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 wel l developed with only 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 robusta dominate u s u a l l y extensive moss l a y e r of mesic s i t e s . On lower and middle slope p o s i t i o n s , seepage water is p l e n t i f u l expressed by an understory of Streptopus  roseus, 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 u n i f o l i a t a . Parkland Subzone. In t h i s subzone, o c c u r r i n g at e l e v a t i o n s of approxi-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 the dur a t i o n of the snow-lie r e s u l t i n a vegetation which i s broken i n t o a mosaic of f o r e s t patches, shrubby areas and meadows. These mea-dows, formed on s i t e s w i t h longer snow duration and i r r i g a t i o n from snow-melt c r e a t i n g h y g r i c and hydr 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 d r i e r flower (forb) meadows and moist heather (heath) meadows (Brooke et a l . 1970, Brink 1959, Archer 1963). Frequent species i n these f l o r i s t i c a l l y r i c h flower meadows include Caltha l e p t o s e p a l a , A r n i c a 1 a t i f o l i a , C a s t i l l e j a m i n i a t a , Erigeron peregrinus, Leptarrhena 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 vegetation o f moist heather meadows i s c h a r a c t e r i z e d by Phyllodoce empetriformis, Vacciniurn d e l i - ciosum, Cassiope mertensiana and Luetkea p e c t i n a t a . Bryophytes: 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 f l o e r k e i , Bazzania denudata, Dicranum fuscescens, D_. p a l l i d i s e t u m , Hookeria 1 ucens, Hylocomium splendens, Hypnum c i r c i n a l e , Mniurn spinulosum, Oligotrichum  aligerum, 0_. hercynicum, P h i l o n o t i s fontana, Plagiothecium denticulatum, P_. undulatum, Pleurozium s c h r e b e r i , Pseudoleskea b a i l e y i , P t i l i d i u m  pulcherrimum, Racomitrium sudeticum, Rhytidiadelphus l o r e u s , R h y t i d i o p s i s  robusta, Scapania b o l a n d e r i , S_. g i r g e n s o h n i i , S_. magellanicum, S.. squar-rosum, S_. t e r e s . 5.4. The Subalpine Engelmann Spruce-Subalpine F i r Zone. As i s shown i n Figure 4, t h i s zone occurs toward the northern part of the study area ( G a r i b a l d i Park). The Subalpine Engelmann Spruce-Subalpine F i r Zone [part of the Middle Oroboreal zone (Hamet-Ahti 1965)] 32 encompasses vegetation of a more c o n t i n e n t a l c h a r a c t e r : subalpine coniferous f o r e s t , r e p l a c i n g the Mountain Hemlock f o r e s t of coastal 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 subalpine are i n d i c a t e d by lower annual temperature, l e s s t o t a l annual p r e c i p i t a t i o n and annual s n o w f a l l , and higher p r o b a b i l i t y o f summer drought and frozen ground i n w i n t e r ( K r a j i n a 1965, Fraser 1970). Throughout the e n t i r e zone the dominant t r e e species are Picea engelmannii, Abies l a s i o c a r p a , Pinus contorta var. l a t i f o l i a ( a f t e r f i r e s ! ) and Pinus a l b i c a u l i s . Productive seepage s i t e s o f t e n show presence of p l a n t species 1ike Alnus incana, 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 productive parts of t h i s zone many species 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 occur f r e q u e n t l y e.g. Arctostaphylos  u v a - u r s i , Calamagrostis rubescens and Aster conspicuus. Bryophytes: 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 fuscescens, JJ. scoparium, Mnium spinulosum, Pogonatum urnigerum, Polytrichum alpinum, P_. juniperinum, Racomitrium sudeticum, R h y t i d i o p s i s robusta, Sphagnum  capill a c e u m , S_. fimbriatum, fuscum, S_. recurvum, S_. rubel 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 Douglas-Fir Zone. Although not encountered i n the present study, t h i s zone [termed Hemiboreal zone by Hamet-Ahti (1965)] marks the northeastern boundary of the study area. Located i n the shadow of the Coast Mountains, the I n t e r i o r 33 Douglas-Fir Zone occupies e l e v a t i o n s of 610 - 1340 m i n south eastern B r i t i s h Columbia C h a r a c t e r i s t i c species of open f o r e s t s at 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 spicatum. In these areas grazing a c t i v i t y and f i r e h i s t o r y tend to determine the composition and p a t t e r n o f present day vegetation (Jones and Annas 1978). In moister and more closed f o r e s t s , at higher e l e v a t i o n s of the zone, the two dominant tre e s p e c i e s , Pinus ponderosa and Pseudotsuga  menziesi i , 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 , Picea  engelmannii and Pinus c o n t o r t a . Closed f o r e s t species Paxistima m y r s i n i t e s and Chimaphila umbellata are more frequent i n wetter parts of 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 , Brachythecium  a l b i c a n s , Bryum canariense, Ceratodon purpureus, Picranurn fuscescens, Drepanocladus uncinates, Hylocomium splendens, Mnium spinulosum, P o h l i a  nutans, P t i l i d i u m pulcherrimum, RhytidiadeTphus t r i q u e t r u s , Timmia  a u s t r i a c a , T o r t u l a r u r a l i s . 5.6. The A l p i n e Tundra Zone. This zone [ i n c l u d e d by Hamet-Ahti (1965) i n the Orohemiarctic zone] contains t e r r a i n at higher e l e v a t i o n s , l y i n g above continuous f o r e s t , parkland and meadows, and below g l a c i e r s or permanent snowfields. As encountered i n the study area, some landscape features i n the A l p i n e Zone are depressions 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 , boulder-f i e l d s , bare rock outcrops, rock w a l l s , exposed summits and ridges with 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 ormation, type of v e g e t a t i o n , and p l a n t succession i s snow i . e . presence, depth, and movement of snow and snow p e r s i s t e n c e . Vegetation i s composed of herbs, bryophytes and l i c h e n s and only very low shrubs. No trees 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 plants develop a dwarf or stunted growth form at the lowest e l e v a t i o n s i n the zone. Among them are Tsuga merten- s i a n a , Abies 1asiocarpa and Pinus a l b i c a u l i s . The vegetative season i s very short. Bryophytes: Amphidium lapponicum, Anastrophyllurn 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 , Arctoa f u l v e l l a , Bartramia i t h y p h y l l a , BI i n d i a acuta, Bryum alpinum, B_. w e i g e l i i , Desmatodon  l a t i f o l i u s , Dichodontium Olympicurn, Dicranodontiurn denudatum, Dicrano- w e i s i a c r i s p u l a , Diplophyl1 urn t a x i f o l i u m , Grimmia a l p e s t r i s , (3. donniana, 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 , K_. s t a r k e i , Lophozia v e n t r i c o s a , Marsupella emarginata, M. s p h a c e l a t a , Oligotrichum  hercynicum, Paral eucobryum enerve, Pohl i a c a r d o t i i , P_. l u d w i g i i , Poly- 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 sudeticum, Scapania 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, T r i t o m a r i a quinquedentata. 1. Hamet-Ahti (1965) has described the vegetation zones of Western Canada by applying the f o r e s t s i t e type system, commonly used i n Scandinavia. This one i s not purely 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 . This author distinguishes': s i x 1 a t i -35 tud.ional zones, which are a l s o recognized i n mountains from peak to 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 zonations are combined under the same u n i t s . 2. Nomenclature f o r v a s c u l a r plants i s t h a t of Hitchcock and Cronquist 1973; t h a t f o r bryophytes f o l l o w s I r e l a n d , B i r d , Brassard, S c h o f i e l d and V i t t 1980. 36 6. METHODS. 6.1. D e l i m i t a t i o n of e p i l i t h i c bryophyte communities. Bryophytes seldom grow d i r e c t l y on rock s u r f a c e s . They occupy cr a c k s , depressions and occur along 'micro-ridges'. Moreover, bryophytes do not c o l o n i z e rocks not i n f l u e n c e d by weathering, but can invade the p h y s i c a l l y and chemically d i s i n t e g r a t e d rock surface. Because t r a n s i t i o n s between rock surface and mineral ' s o i l ' (product of weathering) are often gradual, i t i s d i f f i c u l t to determine whether bryophyte communities are e p i g e i c ( = t e r r e s t r i a l ) on mineral s o i l or e p i l i t h i c on the weathered rock surface. With time, humus ma t e r i a l w i l l accumulate beneath the bryophyte cover. E s p e c i a l l y under the extensive bryophyte cushions and carpets of progressive successional stages humus l a y e r s of several centimeters tend to develop. In t h i s case the d e l i m i t a t i o n from e p i g e i c bryophyte 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 Rhytidiadelphus spp., Pleurozium s c h r e b e r i , R h y t i d i o p s i s robusta, S t o k e s i e l 1 a oregana and Plagiothecium denticulatum. These e p i g e i c bryophyte species can com-p l e t e l y cover smaller stones (up to 20 centimeter i n diameter). Where they occur on more extensive rock s u r f a c e s , however, the c o n t r i b u t i o n of predominantly e p i l i t h i c bryophytes i s high (Sjogren 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 (1964) di v i d e s the mosses of South Sweden i n t o categories r e l a t e d to t h e i r 37 substrate preference. With respect to e p i l i t h i c bryophytes he d i s t i n -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 amphisubstratic species with t r a n s i t i o n s between the groups. In regions where the species i s very common, the substratum preference i s u s u a l l y weaker than where i t i s r a r e . Hence, diaspore input i s e s s e n t i a l i n the judgement of substrate preference (Hertel 1974). This change i n s u b s t r a t e , f o l l o w i n g weakened su b s t r a t e preference, becomes apparent i n comparison of several f l o r i s t i c areas. Sjogren (1964) presents observations comparing the bryophyte f l o r a s of South Sweden and West Europe. Gams (1932) i n d i c a t e s the change from e p i p h y t i c to e p i l i t h i c s ubstrate preference i n subalpine and a l p i n e areas. Herzog (1926) des-c r i b e s the t r a n s i t i o n from e p i l i t h i c i n t o e p i g e i c c o n d i t i o n s i n a r c t i c regions. In the present study sampling was confined to rock surfaces which were covered by a l a y e r of mineral m a t e r i a l that o r i g i n a t e d from d i s i n t e g r a t e d rock, or a mixture of deposited organic and i n i t i a l mineral ' s o i l ' up to a thickness of 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 before t a k i n g any releves i s the search f o r and f a m i l i a r i z a t i o n with p o t e n t i a l r e l e v e - s i t e s . The reconnaissance (meaning " p r e l i m i n a r y examination o f a given area") i n the present study 38 included an assessment of: a. a v a i l a b i l i t y of rock substrate types (boulders, outcrops and rockwalls) b. i n f l u e n c e o f topography, vegetation and macro-climate on the rock substrates present (exposure, moisture, accumulated s o i l ) c. the bryophyte f l o r a ; i n i t i a l c o l l e c t i o n s were made and specimens i d e n t i f i e d d. minimum area measurement f o r releves e. geographic d i s t r i b u t i o n of taxa to i d e n t i f y p o s s i b l e regional v a r i a t i o n 6.2.2. Choice of 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 the choice of the r e l e v e s i t e s i s the homogeneity o f the quadrat. Only rock surfaces with uniform c o n d i t i o n s and homogeneous vegetation 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 of d e c i s i v e i n f l u e n c e . Since several (environmental) f a c t o r s c o i n c i d e on the s i t e , c o n d i t i o n s on rock surfaces are f r e q u e n t l y v a r i a b l e . Surfaces w i t h changing i n c l i n a t i o n and exposure were excluded from sampling, as were rocks with i n s i d e - p l o t -changes i n rock s u r f a c e roughness, amount of accumulated s o i l or changing moisture c o n d i t i o n s (seepage water). In g e n e r a l , quadrats were taken i n homogeneously developed e p i l i t h i c bryophyte assemblages which cover a rock surface of apparently uniform e c o l o g i c a l c o n d i t i o n s of at l e a s t 1/2 m (Homogeneous r e f e r s to 38a LEAP 39 OMITTED IN NUMBERING 40 vegetation large enough to reach i t s f u l l f l o r i s t i c 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 vegetation has s u f f i c i e n t time to become w e l l developed). Types of s i t e s chosen include boulders, detached pieces of stone of at l e a s t 1 meter i n diameter, outcrops, parts of bedrock p r o j e c t i n g from s o i l and r o c k - w a l l s which encompass steep rock s u r f a c e s . For t a b u l a r comparison and synthesis of a s s o c i a t i o n s i t i s important to take only one r e l e v e at each s i t e . This avoids"over-r e p r e s e n t a t i o n of a c c i d e n t a l l y l o c a l t a x a , fragments \ or characters (e.g. dominance/abundance, species combinations) i n the t a b l e s . This requirement could 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 w e l l developed, more rel e v e s were taken (see Wirth 1972, Hertel 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 o f 625 cm (25 x 25 cm) was found appropriate to encompass the c h a r a c t e r i s t i c combination of s p e c i e s . This i s i n d i c a t e d i n the t a b l e s of the separate bryophyte syntaxa. As a c o n t r o l f o r the quadrat-size the Nested P l o t Technique was a p p l i e d (Figure 5). The species 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 species/area curve (Figure 6 ) , a f t e r which the d e c i s i o n i s made on the quadrat s i z e . 6.2.3. Estimation of species q u a n t i t y and s o c i a b i l i t y . In order to estimate the cover of bryophyte species w i t h i n the 41 cn o 10 CM ro o O 40 CM Figure 5. Nested p lo t technique, used fo r the determination o f a minimum r e l e v e ( p 1 o t ) - s i z e . 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 other cryptogam-(lichen- and bryophyte-) s o c i o l o g i s t s , In t h i s s c a l e , cover values below 50%, e s p e c i a l l y , are 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 are made to estimate the degree of s o c i a b i l i t y . Although 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 reveal information on the c o n t r i b u t i o n of the species to the o v e r a l l community s t r u c t u r e (physiognomy). For the est i m a t i o n of the 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 of the r e l e v e - s i t e . The l o c a t i o n on the rock surface was recorded f o l l o w i n g Frey's (1922) types of rock surface h a b i t a t s (Figure 7). To obt a i n information on environmental f a c t o r s determining and c o n t r o l l i n g the bryophyte assemblages on rocks, data 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 geographical maps a l t i t u d e : measured with an a l t i m e t e r d i r e c t i o n of slope (exposure): measured w i t h Bezard Compass i n c l i n a t i o n of slope: s o i l accumulation: d i v i d e d i n two c l a s s e s 0-1 cm 1-2 cm roughness of rock surface: s u b j e c t i v e l y assessed 44 1 Figure 7. Categories o f rock surface h a b i t a t s . 1. top surface or nearly l e v e l (0-20°) 2. gentle slope (20-70°) 3. steep slope (70-90°) 4. overhanging slope 5. g r o t t o 45 exposure: l i g h t was estimated as e i t h e r b r i g h t s u n l i g h t or shade and a s i t e was considered shaded only i f i t was shaded by t r e e s (or other rocks) f o r one-half or more of each d a y l i g h t period moisture: expressed as e i t h e r wet or dry. The wet s i t e s i n c l u d e only those which are v i s i b l y moist from r u n - o f f water, tem p o r a r i l y submerged or e f f e c t e d by spray 6.3. Data a n a l y s i s . 6.3.1. Bryophyte i d e n t i f i c a t i o n . In the 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 of the bryophyte specimens from the r e l e v e s , a s e r i e s of f l o r a s and s p e c i a l generic and s p e c i e s -complex treatments were used. For the Hepaticae these were: Flowers (1961), Frye and Clark (1937-1947), Godfrey (1977), Macvicar (1926), M u l l e r (1957) and Schuster (1966, 1969, 1974). For the Musci the f o l l o w i n g f l o r a s were used: Crum (1976), Flowers (1973), Grout (1928-1940), Lawton (1971), Nyholm (1954-1969) and Smith (1978). I d e n t i f i c a t i o n of the genus Plagiotheciurn followed I r e l a n d (1969), that of Hygrohypnum followed Jamieson (1976), that of Dicranum followed Peterson (1979) and t h a t of S e l i g e r i a f o l l owed V i t t (1976). F i n a l l y , Koponen (1974) was used i n the i d e n t i f i c a t i o n o f the family Mniaceae and Hoisington (1979) was consulted i n the r e c o g n i t i o n 46 of taxa out of the Brachythecium asperrimum - f r i g i d u m complex. B a s i c a l l y , the nomenclature f o l l o w s Crum, Steere, and Anderson (1973) and the species l i s t (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 bryophyte specimens were examined, the i d e n t i -f i c a t i o n o f which formed the basis f o r the bryophyte a s s o c i a t i o n t a b l e s . This p l a n t m a t e r i a l has been deposited i n the Herbarium of the U n i v e r s i t y of B r i t i s h Columbia. Information on the phytogeographical s e t t i n g o f the e p i l i t h i c bryophyte vegetations of south-western B r i t i s h Columbia was derived from Godfrey (1977), S c h o f i e l d (1968a, 1968b, 1969, 1976, 1980) and Worley (1972). 6.3.2. Tabular comparison. The raw data set was subdivided i n t o four major groups which c o n s t i t u t e c o l l e c t i o n s of releves from the f o l l o w i n g h a b i t a t types: 1. exposed, dry rocks (129 r e l e v e s , 152 species) sandstone and i n t r u s i v e ( s i l i c e o u s ) rocks 2. shaded, humid rocks (83 r e l e v e s , 128 species) 3. s i l i c e o u s rocks on and along streambanks (102 r e l e v e s , 117 species) 4. limestone, both exposed and.shaded (27 r e l e v e s , 61 species) A c r u c i a l procedure i n the a n a l y s i s of p h y t o s o c i o l o g i c a l data i s 47 the development of t a b l e s . Braun-Blanquet (1951a,b) wrote ( t r a n s l . from Westhoff and Maarel 1973): "Appropriately elaborated a s s o c i a t i o n t a b l e s are comparable with a thorough species d i a g n o s i s . By means of the t a b l e s the considerably 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 of 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 evaluable. From the t a b l e i t also shows whether one has worked s e r i o u s l y and r e l i a b l y ; the t a b l e s are the proper touch-stone of 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 (1956), Knapp (1971), Shimwell (1971) and M u e l l e r -Dombois and Ellenberg (1974) presented d e t a i l e d accounts of the t a b l e technique which was performed on the releves of each major group i n t h i s study. I t s procedure can be summarized as f o l l o w s : a. Entrance of the r e l e v e data i n t o a raw t a b l e . Species values (cover and s o c i a b i l i t y ) are recorded i n rows, re l e v e s i n columns, w h i l e the order of entry i s i n c o n s e q u e n t i a l . b. Count of presence of species and c a l c u l a t i n g t h e i r constancy value, ranked i n classes from I to V. In t h i s presence t a b l e i t becomes apparent whether releves vary i r r e g u l a r l y , or whether c e r t a i n combination of species are found r e c u r r e n t l y . In the l a t t e r cases such groups are more or l e s s mutually e x c l u s i v e and serve as groups of d i f f e r e n t i a t i n g s p e c i e s . Those are not l i k e l y to be found i n the presence c l a s s e s V and I ; they w i l l mostly belong to the intermediate c l a s s e s I I , I 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 are p r o v i -s i o n a l sets of d i a g n o s t i c s p e c i e s , are used to rearrange the t a b l e . 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 species and using them to rearrange rows i n t h e i r corresponding groups, and the columns i n t h e i r corresponding r e l e v e s . This rearrangement seeks to show a diagonal order of species groups from the l e f t to the r i g h t of 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 t a b l e . d. Replacement of p a r t i a l t a b l e by a column i n which, f o r each p a r t i c i p a t i n g s p e c i e s , the presence degree i s i n d i c a t e d . A f t e r comparison of t h i s synoptic t a b l e (Westhoff and Maarel '1973) with those from other types of vegetation from the same r e g i o n , an idea i s formed about the l o c a l d i a g n o s t i c species groups i n the t a b l e under study. With these species as we l l as the constant t a x a , a phytocoenon [=vegetation u n i t (Westhoff and Maarel (1973)] i s then i d e n t i f i e d and described. e. Comparison of the sy n o p t i c t a b l e with those a v a i l a b l e i n the l i t e r a t u r e and with t a b l e s from s i m i l a r vegetation types from other r e g i o n s , r e s u l t s i n the design a t i o n of 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 of t a b l e s covering the vegetation of a given area makes i t p o s s i b l e to d i s c e r n character species on c r i t e r i a of d i s t r i b u t i o n of the species i n d i f f e r e n t phytocoena. This e n t a i l s the determination of f i d e l i t y of the taxa ( f i d e l i t y i s the degree to which a taxon i s concentrated i n one phytocoenon 49 vs. dispersed with more even occurrence i n several phytocoena). Five degrees have been d i s t i n g u i s h e d (Braun-Blanquet 1964) i n the r e l a t i o n of a species to a given phytocoenon (Appendix I I ) . The ' c l a s s i c a l ' arrangement technique by manual o r d e r i n g o f taxa and releves i s a time consuming procedure and a p o t e n t i a l source of e r r o r . A number of computerized techniques f o r t a b l e arrangement have been developed of which the more recent ones have been reported to 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, Maarel, Janssen and Louppen 1978, Meulen, Morris and W e s t f a l l 1978, Louppen and Maarel 1979). In order to s i m p l i f y and mechanize the procedure of the tabu-l a t i o n technique a computer program was used, which was designed to s o r t the rows and columns of 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 . This program, 'VEGTAB'. developed by Dr. G. B r a d f i e l d , Department o f Botany of the U n i v e r s i t y of B r i t i s h Columbia, takes care of steps a, b and c of the t a b u l a t i o n procedure: producing a raw t a b l e , transforming t h i s i n t o a presence t a b l e , and, f i n a l l y , rearranging t h i s l a t t e r one i n t o a d i f f e r e n t i a t e d t a b l e . 6.3.3. Numerical techniques. A d d i t i o n a l to the t r a d i t i o n a l methods of the Zu'rich-Montpellier s c h o o l , q u a n t i t a t i v e techniques have been developed and a p p l i e d to study bryophyte vegetation on a v a r i e t y of s u b s t r a t a . 50 In most studies concerning the q u a n t i t a t i v e ecology of bryo-phytes (and l i c h e n s ) , workers have used random sampling techniques. T h e i r s t a t i s t i c a l analyses of the vegetation of quadrats and the e c o l o g i c a l f a c t o r s w i t h i n them, mostly showed veg e t a t i o n a l continua r e l a t e d to some environmental gradient (a.o. Jesberger 1973, Lambert and Maycock 1968, Culberson 1955, Hale 1955). With respect 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 e p i l i t h i c 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 to show r e l a t i o n s h i p s w i t h environmental parameters (Foote 1963, 1966, Yarranton 1967a,b,c). Other workers have endeavoured to i n t e r p r e t q u a n t i t a t i v e data i n an attempt to recognize more or l e s s d i s c r e t e vegetation u n i t s (Redfearn 1957, N o r r i s 1964, West and S t o t l e r 1977). Foote (1963, 1966) s t u d i e d sandstone and limestone outcrops i n south-western Wisconsin by means of random sampling technique. Ordinations of bryophyte "stands" d i s p l a y e d a continuum o f the h a b i t a t s from x e r i c to moist s i t e s . The moisture r e l a t i o n s of the i n d i v i d u a l stands appeared to most i n f l u e n c e the frequency of the s p e c i e s . Other e c o l o g i c a l f a c t o r s 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 of the rock slope 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 of these . 2 species. . Yarranton (1967a,b,c) produced a q u a n t i t a t i v e a n a l y s i s ( P r i n c i p a l Components A n a l y s i s ) of v a r i a t i o n i n the e p i l i t h i c bryophyte vegetation at Steps B r i d g e , Devon (England). Results of the 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 environmental 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) stu d i e d the e p i l i t h i c vegetation o f a Snowdonian c l i f f on the western seaboard o f B r i t a i n . The o r d i n a t i o n s , constructed on frequency data from quadrat-vegetations, showed r e l a t i o n s h i p s between species d i s t r i b u t i o n s and a major environmental gradient from dry o l i g o t r o p h i c to wet eutrophic c o n d i t i o n s . Redfearn (1957), i n h i s quadrat s t u d i e s of the bryophyte vege-t a t i o n on exposed limestone i n c e n t r a l and northern F l o r i d a , U.S.A., used high frequency species i n d e l i m i t i n g d i f f e r e n t bryophyte assem-blages. C a l c u l a t i o n s o f Cole's Index of A s s o c i a t i o n between the most frequent species made i t p o s s i b l e to recognize fourteen bryophyte "unions". Moisture was the predominant environmental 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 the bryophytes and t h e i r unions. Norn's' (1964) study of e p i l i t h i c h a b i t a t s i n the Appalachian Spruce-Zone r e s u l t e d i n an arrangement of quadrats i n t o a number o f bryophyte communities or "unions". Groups o f quadrats ( i n which coverage of species was estimated i n increments of ten percent) were sorted i n a way as to minimize the number of s i g n i f i c a n t l y a s s o c i a t e d species p a i r s . S i g n i f i c a n c e of a s s o c i a t i o n i s judged by the Chi square method f o r each p o s s i b l e species p a i r . The author used t h i s o b j e c t i v e c l a s s i f i c a t i o n of the vegetation 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 with h a b i t a t s between p l o t s o f s i m i l a r species composition than between p l o t s of s i m i l a r species dominance" ( N o r r i s 1964, p. 8 ) . His study r e s u l t e d i n the r e c o g n i t i o n of 12 e p i l i t h i c bryophyte unions, 8 o f which were confined to t h i s type of sub s t r a t e and 4 had a wider 52 e c o l o g i c a l range ( s o i l , t r e e s ) . Major f a c t o r s i n the d i s t r i b u t i o n o f bryophytes and t h e i r unions included source o f water, frequency and duration of drought and l i g h t i n t e n s i t y . West and S t o t l e r (1977) analyzed the bryophyte - macro-lichen f l o r a o f sandstone canyons i n southern I l l i n o i s by using quadrats i n which r e l a t i v e cover and frequency was determined f o r each species. In t h e i r data analyses they computed the importance values 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 the cover and frequency values and again from presence/absence data. However, the data and the authors' difcussion of the r e s u l t s f a i l to d e l i m i t a s s o c i a t i o n s and recognize d i s t i n c t groupings. In the present study, c l u s t e r a n a l y s i s was used to 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 the e p i l i t h i c bryophyte vegetation i n the study area. Furthermore, attempts were made to or d i n a t e some releves to 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 of a s s o c i a t i o n s and lower syntaxa to one another and to the 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 of the four major groups, using the MIDAS s t a t i s t i c a l package (Fox and Guire 1976). C l u s t e r a n a l y s i s i s an agglomerative method i n which the c l u s t e r i n g algorithm "proceeds from the 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 of decreasing i n t e r n a l s i m i l a r i t y u n t i l the whole set of samples i s combined" (Goodall 1973). 53 The c l u s t e r algorithm used here to d e l i m i t the bryophyte vegetation at the various s i t e s i s the procedure described by Ward (1963). I t i s a h i e r a c h i a l method (using Euclidean distance measure) grouping, i n t h i s case, releves i n t o c l u s t e r s , based on cover values of species i n the r e l e v e s . The t o t a l decrease 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 the t o t a l sum of squared d e v i a t i o n s of every point from the mean of the c l u s t e r to which i t belongs. At each step i n the a n a l y s i s , union of every p o s s i b l e p a i r of c l u s t e r s i s considered 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 the minimum increase i n the e r r o r sum of squares are combined ( E v e r i t t 1974). The e r r o r sum of squares (E.S.S.) i s given by N p -j y E.S.S. = S X, - -rj- (S x.) i = l N The aim of these c l u s t e r analyses i s to explore and subdivide the l a r g e sets of releves i n t o a number of c l u s t e r s , each of which i s c h a r a c t e r i z e d by one or more constant s p e c i e s . These c l u s t e r s are then compared to the grouping of releves a r r i v e d at through the tabu-l a t i o n technique of the Braun-Blanquet method. In t h i s comparison (which was only c a r r i e d out on the group of releves from calcareous rocks) connec-are sought between the value of s i m i l a r i t y index (given as a 54 Euclidean distance measure) and syntaxonomic l e v e l ( a l l i a n c e s , 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 ) . 6.3.3.2. O r d i n a t i o n . O r d i n a t i o n techniques are used t o search f o r patterns i n a data matrix (Gaugh 1977) and r e s u l t i n an arrangement of e n t i t i e s ( r e l e v e s ) i n a u ni- or multidimensional order. In vegetations with a s s o c i a t i o n s g e n e r a l l y discontinuous with one another and the environment, o r d i n a t i o n has been a p p l i e d to under-stand the r e l a t i o n s of a s s o c i a t i o n s and lower syntaxa to one another between the r e l e v e s (Matuszkiewicz and Traczyk 1958, F a l i n s k i 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 to the r e l a t i o n s of a s s o c i a t i o n s and higher syntaxa as wholes, to one another and the e n v i -ronment (Westhoff and Maarel 1973). In the present study a major f u n c t i o n of o r d i n a t i o n i s to d i s -play groups of s i m i l a r releves i n the c o l l e c t i o n of limestone vegetation samples (27 r e l e v e s , 61 s p e c i e s ) . This a p p l i c a t i o n of o r d i n a t i o n tech-niques to the data seeks to enhance the c l a s s i f i c a t i o n by f u r t h e r expressing and c l a r i f y i n g r e l a t i o n s h i p s of a s s o c i a t i o n s to one another and the environment. The analyses were performed using the ORDIFLEX computer program developed by Gaugh (1977) at C o r n e l l U n i v e r s i t y , Ithaca,. New York, U.S.A. 1. "A community can be c a l l e d fragmentary i f i t l a c k s species t h a t are u s u a l l y present i n the r e c u r r i n g p l a n t assemblages of t h i s k i n d " 55 (Mueller-Dombois and E l l e n b e r g 1974). Neither 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 are omitted i n the present study, s i n c e they may be important i n the understanding of 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 syntaxonomical p o s i t i o n o f species groups and vegetation types (Barkman 1969, Hale 1955, Culberson 1955, Hertel 1974) 2. Within the frame work and set of o b j e c t i v e s i n the present study, i t i s i n t e r e s t i n g to note the f o l l o w i n g f i n d i n g s of Foote (1963): "Several stands on portions of the continuum are discussed and the vegetations of these stands compared to a s s o c i a t i o n s recognized i n Europe. Many of the European a s s o c i a t i o n s are s i m i l a r to the vegetation of 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 that the bryophyte and l i c h e n vegetation of middle Europe i s probably a continuum s i m i l a r to that found i n the present study. Because the European method of sam-p l i n g includes only " t y p i c a l " stands, intermediate vegetations are ignored. Such studies r e s u l t i n a c l a s s i f i c a t i o n of separate, w e l l defined a s s o c i a t i o n s r a t h e r than a continuum of s p e c i e s " . 56 7. SYNSYSTEMATICS. 7.1. Terminology and nomenclature. Centering on the nature 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 important on the one hand to recognize the heterogeneity of species d i s t r i b u t i o n , y e t to emphasize on the other hand the i n t e r a c t i o n s between p l a n t s i n the community. In t h i s r e s p e c t , the plant community or phytocoenose has a c e r t a i n i n d i v i d u a l i t y because of 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 the f i e l d . This a p p l i e s , e s p e c i a l l y , to 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 discontinuous i n space. Moreover, the i n d i v i d u a l rock s i t e s ( b o u l d e r s , outcrops, r o c k w a l l s ) d i f f e r widely i n form and s i z e . Each s i t e , i n t u r n , may be very heterogeneous i n e c o l o g i c a l features (aspect, i n c l i n a t i o n , exposure, moisture). Within the context of t h i s t h e s i s phytocoenose and phytocoenon are defined as f o l l o w s ( a f t e r Westhoff and Maarel 1973): phytocoenose: a part of a vegetation c o n s i s t i n g of i n t e r a c t i n g populations growing i n a uniform environment and showing a f l o r i s t i c composition 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 uniform and d i s t i n c t from the surrounding vegetation phytocoenon: a type of phytocoenose derived from the charac-t e r i z a t i o n of a group of phytocoenoses c o r r e s -57 ponding with each other i n a l l characters that are considered t y p o l o g i c a l l y r e l e v a n t In applying the Z i i r i c h - M o n t p e l l i e r school method of vegetation a n a l y s i s , the present study seeks to c l a s s i f y e p i l i t h i c bryophyte assemblages, as phytocoena, i n t o synsystematic u n i t s ( C l a s s , Order, A l l i a n c e , A s s o c i a t i o n , etc.) based on constancy and f i d e l i t y o f taxa and defined by character s p e c i e s , d i f f e r e n t i a t i n g species and companions. The fundamental u n i t of the .hierarchy i s the a s s o c i a t i o n , a un i t t h a t corresponds i n f u n c t i o n to the species as the fundamental u n i t i n idiotaxonomy, or the c l a s s i f i c a t i o n of 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 defined as "a phytocoenose i d e n t i f i e d by i t s charac-t e r i s t i c species combination, i n c l u d i n g one or more character-taxa or d i f f e r e n t i a t i n g t axa". 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 to samples of a given 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 or more character species are used f o r the charac-t e r i z a t i o n of any syntaxon from the a s s o c i a t i o n to the c l a s s . Degrees o f r e s t r i c t i o n to a given taxon are expressed by the degree of f i d e l i t y (Appendix I I ) . Hence, character species c h a r a c t e r i z e syntaxa by t h e i r normal (or sometimes high) occurrence i n phytocoenose of that syntaxon, contrasted with t h e i r absence or l e s s frequent occurrence i n phytocoenoses of other syntaxa. I t i s p o s s i b l e a l s o to 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 syntaxa by the presence and absence of c e r t a i n species without concern about the broader d i s t r i b u t i o n o f those species i n other phytocoenoses. As such, two 58 subassociations of 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 of 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 of d i a g n o s t i c species with d i a g n o s t i c value are the companions which occur i n most releves of a syntaxon but are not designated 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 to the character taxa to form the " c h a r a c t e r i s t i c species combination" f o r a s s o c i a t i o n s and higher syntaxa. With respect to e p i l i t h i c bryophyte assemblages, various authors have described a large number of bryophyte communities without proper a s s o c i a t i o n r e c o g n i t i o n . These bryophyte communities are based on dominance r a t h e r than f i d e l i t y o f taxa. As Westhoff and Maarel (1973) s t a t e "there i s no necessary r e l a t i o n between f i d e l i t y and dominance; character taxa can as w e l l be minor as major ones". In the present study attempts are made to f o l l o w as s t r i n g e n t l y as p o s s i b l e the p r i n c i p l e s of the Z i i r i c h - M o n t p e l l i e r school i n the d e f i n i t i o n o f syntaxa by means of character and d i f f e r e n t i a t i n g taxa. Where p o s s i b l e , the synsystematic 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 study are incorporated i n e x i s t i n g syntaxa, many of them described from e p i l i -t h i c bryophyte assemblages i n Europe. In accordance to nomenclature proposals as those of Neuhausl (1968) and name-assigning r u l e s of Bach, Knock and Moor (1962) new u n i t s are described. For the s t a n d a r d i z a t i o n of syntaxon names, the f o l l o w i n g procedures are used. To the generic part of the names of one or two c h a r a c t e r i s t i c taxa of a syntaxon a s u f f i x i s added. These s u f f i x e s are 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 - e t a l i a f o r order - i o n f o r a l l i a n c e -etum f o r a s s o c i a t i o n -etosum f o r su b a s s o c i a t i o n When de a l i n g with an a s s o c i a t i o n or 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 dominant charac-t e r taxon), the genus name of the taxon i s used with the appropriate s u f f i x (-etum, - i o n ) , followed by the species name i n the g e n i t i v e . When a syntaxon was to be named a f t e r two character t a x a , the name of the second taxon was provided with the appropriate s u f f i x , while the name of the f i r s t taxon was j o i n e d with the second one by the s u f f i x -eto. (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 the second name represents the charac t e r taxon considered d i a g n o s t i c a l l y most important). 60 8. ANALYSIS AND DESCRIPTION OF THE EPILITHIC BRYOPHYTE VEGETATION. The raw data set, containing a l l releves in columns and a l l species in rows, i s subdivided into the following major groups: 1. releves from exposed, dry rocks 2. releves from shaded, humid rocks 3. releves from siliceous rocks along streams 4. releves from calcareous rocks, both exposed and shaded A cluster analysis i s performed on each of these major groups, in order to recognize groups of "similarr.eTe.ves". In analysing the phyto-sociological data these clusters function as the star t i n g point i n the preparation of the association tables. After the preparation of the raw table and the presence table, VEGTAB i s given the command to rearrange the order of re-1 eves according to the sequence shown in the cluster analysis. This results in the organization of a "differentiated" table, in which, subsequently, the clusters are drawn in by v e r t i c a l l y separating columns into groups of releves (see Appendix Vi),. The f l o r i s t i c composition of the clusters i s studied and related 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 cha r a c t e r i s t i c s . Then, within these c l u s t e r s , attempts are made to recognize diagnostic species: recognition of order, a l l i a n c e and associ-ation character species and, where possible and appropriate, d i s t i n c t i o n 61 of 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 designation 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 are shown i n Table I I I , XI, XV, XXV and Appendix V ( w i t h i n t h i s Appendix a l i s t i s provided f o r each major group, which " t r a n s l a t e s " the releve-number i n the c l u s t e r analyses i n t o the releve-number as taken i n the f i e l d and given i n the separate a s s o c i a t i o n t a b l e s ) . F i n a l l y , the c l u s t e r analyses may serve to i n d i c a t e syntaxono-mic r e l a t i o n s h i p s at the various l e v e l s of the c l a s s i f i c a t i o n h i e r a r c h y and between syntaxa at the same l e v e l . To enhance and c l a r i f y these r e l a t i o n s h i p s references are made to f l o r i s t i c composition and h a b i t a t of the bryophyte assemblages. The c l u s t e r a n a l y s i s procedure, i t s r e s u l t i n each major group, are shown and discussed on the f o l l o w i n g pages. In the d i s c u s s i o n of major group number 4, the a p p l i c a t i o n of o r d i n a t i o n techniques on the data w i l l a l s o be evaluated. These methods are used to enhance the c l a s s i f i c a t i o n as achieved by the t a b u l a t i o n technique and c l u s t e r ana-l y s i s . They a l s o may c l a r i f y and f u r t h e r express r e l a t i o n s h i p s between groups of releves and t h e i r a f f i n i t i e s to the environment. 62 8.1 BRYOPHYTE ASSOCIATIONS FROM EXPOSED, DRY ROCKS. Table I I I . Bryophyte a s s o c i a t i o n s from exposed, dry rocks. 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 1944 C l u s t e r 1. A s s o c i a t i o n Racomitrio ( h e t e r o s t i c h i ) - Andreaeetum petrophilae 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. A s s o c i a t i o n "Racomitrietum b r e v i p i s " ass. 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 - v a r i a n t Dicranum fuscescens - B a r b i l o p h o z i a f l o e r k e i - v a r i a n t Andreaea r o t h i i - v a r i a n t Table I I I (continued). C l u s t e r 6. A s s o c i a t i o n Racomitrietum 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. A s s o c i a t i o n 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 ass. nov. 65 8.1. Bryophyte a s s o c i a t i o n s from exposed, dry rocks (Table I I I ) . 8.1.1. Synsystematics. Within the a l l i a n c e Andreaeion p e t r o p h i l a e Hadac and K l i k a 1944, P h i l i p p i (1956) d i s t i n g u i s h e d three a s s o c i a t i o n s : the Racomitrio -Andreaeetum p e t r o p h i l a e Frey 1922, the Andreaeetum r o t h i i P h i l i p p i 1956 and the Gymnomitrietum conci n n a t i Herzog 1943. This l a s t a s s o c i a t i o n was o r i g i n a l l y described as two separate vegetation u n i t s by Herzog (1943): a Racomitrium sudeticum- 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 the present study, these two vegetation types are recognized s e p a r a t e l y again and are given the status of a s s o c i a t i o n , based on the dominance and f i d e l i t y of t h e i r r e s p e c t i v e character 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 " . of which various a l l i a n c e character species (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 sudetica) show i t s a f f i l i a t i o n with the other three 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 Andreaeion p e t r o p h i l a e . Both, the Racomitrio - Andreaeetum p e t r o p h i l a e and the "Raco-mitrietum b r e v i p i s " show a wide e l e v a t i o n a l range. The f i r s t a s s o c i a t i o n occurs predominantly at lower e l e v a t i o n s , whereas the l a t t e r one shows i t s best development i n subalpine areas. On exposed a l p i n e rocks these a s s o c i a t i o n s occur only r a r e l y : here, the Gymnomitrietum c o n c i n n a t i and 66 the Racomitrietum s u d e t i c i are the most frequent and dominant a s s o c i a t i o n s . L i k e Neumayr (1971) and Hertel (1974), t h i s study, synsystema-t i c a l l y , places the a l l i a n c e Andreaeion p e t r o p h i l a e i n the order Raco-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. From exposed r o c k s , a second group of a s s o c i a t i o n s has been described, which are c l a s s i f i e d i n the order 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. The two syntaxa, d i s t i n g u i s h e d i n the present study are confined to low e l e v a t i o n s and c h a r a c t e r i s t i c a l l y grow on h o r i z o n t a l and s l i g h t l y s l o p i n g rock surfaces o v e r l a i n by a mixture of mineral s o i l and humus. The f i r s t a s s o c i a t i o n , the Racomitrio - P o l y t r i c h e t u m p i l i f e r i Herzog 1943 has been described from several l o c a t i o n s i n Europe (Herzog 1943, Norr 1969, Neumayr 1971, Hubschmann 1975) and i s a l s o reported 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 the a s s o c i a t i o n has been placed i n the a l T v P o l y t r i c h i o n p i l i f e r i by Norr (1969). In the present study, a newly proposed a s s o c i a t i o n , the Scapa-nietum americanae, i s described from s i m i l a r h a b i t a t s and e c o l o g i c a l c o n d i t i o n s as the previous a s s o c i a t i o n . I t s own i d e n t i t y , however, i s based on character species as w e l l as t y p i c a l accompanying s p e c i e s . I t s syntaxonomic a f f i n i t y to the Racomitrio - P o l y t r i c h e t u m p i l i f e r i and i t s subsequent placement i n the same a l l i a n c e and order i s expressed by a d d i t i o n a l companion species. 67 8.1.2. Phytogeographical s e t t i n g . Both orders 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 circumboreal bryophyte species designated as a s s o c i a -t i o n character species or important companions. Among these the most frequent and abundant are Diplophyllum t a x i f o l i u m , Gymnomitrion concin- natum, Marsupel1 a emarginata, Racomitrium heterostichum, R. canescens, 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 of s p e c i e s , endemic to western North America i s of minor importance i n most bryophyte a s s o c i a t i o n s from exposed, dry rocks. An obvious exception i s "Racomitrium b r e v i p e s " , which i s t r e a t e d as a separate taxonomical e n t i t y i n t h i s study. Because of i t s constancy, t h i s western North American endemic i s designated as the character species f o r the newly proposed a s s o c i a t i o n "Racomitrietum b r e v i p i s " . In a d d i t i o n , other s i g n i f i c a n t l y accompanying endemics i n releves from low e l e v a t i o n s are Scapania americana and Hypnum c i r c i n a l e . Frequently o c c u r r i n g b i p o l a r d i s j u n c t bryophytes are species such as B a r b i l o p h o z i a f l o e r k e i , B_. hatch e r i and Bartramia pomiformis. 8.1.3.1. Order RACOMITRIETALIA HETEROSTICHI P h i l i p p i 1956 A l l i a n c e Andreaeion p e t r o p h i l a e Hadac and K l i k a 1944. Racomitrio ( 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 (Table IV). In the study area t h i s a s s o c i a t i o n i s found and described from 68 e l e v a t i o n s ranging from sea l e v e l (Lighthouse Park) to a l p i n e areas. This 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 lowest 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 subalpine to a l p i n e areas. Reports and documentation on the present syntaxon are given by a.o. Frey (1922), Herzog (1943), P h i l i p p i (1956, 1965), Neumayr (1971) and Hertel (1974). From Vancouver I s l a n d Hubschmann (1978) described small fragments of t h i s a s s o c i a t i o n . Horikawa, Ando and Kawai (1961) describe Andreaea r u p e s t r i s var. f a u r i e r i - and Racomitrium heterostichum -communities from the a l p i n e zone of Mount Hakusan, Japan. Although 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 , the 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 Columbia and Europe. The Racomitrio - Andreaeetum i n the present study shows a wide geographic d i s t r i b u t i o n w i t h i n the Coast Mountains area of 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 north of Vancouver to Nairn F a l l s (Figure 1). Since both character s p e c i e s , Racomitrium heterostichum and Andreaea r u p e s t r i s , occur over a much l a r g e r area throughout the province ( S c h o f i e l d 1976, Tan 1980), the a s s o c i a t i o n i s l i k e l y to be more wide-spread than encountered i n the present study. The a s s o c i a t i o n predominantly occurs on dry and exposed r o c k s , where i t forms lush bryophyte assemblages. At lower e l e v a t i o n s these rocks are mostly i n t r u s i v e i n nature; i n a l p i n e areas the a s s o c i a t i o n i s a l s o described from metamorphic rocks. Frequent a s s o c i a t e s of the character species are bryophyte taxa Table IV. Racomitrio (heterostichi) - Andreaeetum petrophilae Frey 1922 Number of releves 1 2 3 4 5 6 7 8 9 10 11 12 Releve-number Elevation(m) Direction ofQexposure Inclination ) 78 30 46 68_ 32 84~ ~66~ 32 46 24 24" 32 Total cover(%) 80 75 80 85 100 100 85 75 80 90 80 90  2    8    9 11 15 179 119 181 26 243 245 244 331 36 61 61 1433 610 1433 18 1492 1492 1492 610 E-SE E-SE S-SW E-SE N E-SE E-NE NW E NM N NW            5.5 4.4 4.4 4.5 3.4 3.4 1.2 3.4 3.4 2.4 3.4 1.4 1.2 2.2 3.2 2.4 1.4 1.4 2.4 1.4 2.4 + .4 1.2 1.2 1.2 2.4 + .4 1.4 1.4 1.2 4.4 1.4 + .4 2.4 2.4 + .1 2.2 + .1 1.1 1.1 + .1 1.2 2.2 2.2 2.4 1.2 1.2 1.2 3.4 2.4 1.4 1.4 1.4 1.4 1.4 1.4 2.4 1.2 3.3 1.2 Racomitrium heterostichum var. heterostichum .4 . .5 .    2.4 Andreaea rupestris .  .     2.4 Gymnomitrion obtusum 2.4 -  .4 1.4 Diplophyll urn taxi folium .  .4  +.4 Lophozia ventricosa var. ventricosa .4  + -4 2.4 2.4 1.4 Pohlla nutans .1 .1 .1 +.1 Kiaeria starkei -* - - 1-2 Marsupella emarginata  + .4 Lophozia sudetica  1.4 Scapania americana Racomitrium sudeticum Polytrichum alpinum var. macounii +.1 Polytrichum piliferum 2.2 1-1 + . l Grimmia torquata + -2 Cephaloziella byssacea var. asperifolia 2.4 2.4 3.4 +.4 Anastrophyllum minutum +.4 1.4 2.4 +.4 Dicranella heteromalla 2.2 2.2 1.2 2.3 Oryptodon patens ]-2 1.2 3.4 Homalothecium nevadense 1-4 1-4 1.4 Cephaloziella byssacea var. byssacea +.4 1.4 4.4 Racomitrium heterostichum var. affine 2.4 1.2 Scapania scandica '.2 2.4 13 14 15 16 17 18 19 20 21 22 23 24 25 288 289 290 65 17 332 178 134 135 182 222 223 21 610 610 610 61 61 610 1433 427 427 1433 610 610 18 E E E NW N NW NW E SE N NW E SE W W N 76 102 98 66 64 74 72 50 62 88 70 68 84 75 90 90 90 60 100 90 100 100 100 70 80 90 1.4 3.4 4.4 3.4 3.4 3.4 2.4 2.4 2.4 4.4 1.2 1.2 2.4 2.4 2.2 2.4 2.4 2.4 2.4 2.4 3.4 2.2 2.4 3.4 2.4 2.4 + .4 4.4 3.4 3.4 1.4 3.4 3.4 2.4 2.4 1.4 3.4 3.4 1.4 1.4 3.4 1.4 1.4 + .1 1.4 1.4 1.2 2.2 3.4 + .1 1.2 4.4 2.2 1.4 1.4 1.2 + .4 2.4 2.4 4.5 4.5 3.4 2.4 + .4 1.4 2.4 + .1 1 .2 + .1 1.4 1 .4 1.1 1.4 + .1 + .4 * ° 1.1 + .1 1.2 1.2 1.2 Pohlia cruda Bartramia pomiformis • 1.4 1.4 + .1 +.1 Dicranum scoparium 1.2 1.2 Cynodontium jenneri 3.3 1 -2 Anthelia juratzkana +,4 +.4 1 - 2 Marsupella sparsifolla 1.4 3.4 Hypnum circinale 1.4 4 4 Isopterygium elegans 1.4 +.1 Marsupella condensata +,4 +.4 Pseudobraunia californica ' 1.4 1.4 Plagiochila asplenioldes +.1 ' ' +1 Bryum caespiticium +.1 1.2 Herbertus aduncus 1 1 3 2 Cephaloziella turnerl +.1 1.1 ' Chandonanthus setiformis i 4 -> « Ptilidium ci l iare 1.4 +.4 4 Table IV (continued). Found only 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 , Dicranoweisia c i r r a t a , Hypnum subimponens, Claopodium  b o l a n d e r i , Ceratodon purpureus, Dicranum fuscescens, P h i l o n o t i s fontana, B a r b i l o p h o z i a 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 , Polytrichum juniperinum, Chandonanthus f i 1 i f o r m i s , Diplophyllum a l b i c a n s , Racomitrium lanuqinosum, Anacolia 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 , Lophozia 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 culare, Barbi1ophozia lycopodi oi des, Bazzania  t r i c r e n a t a , Heterocladium 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 Murrin P r o v i n c i a l Park Mount Seymour Cypress Bowl Lighthouse Park G a r i b a l d i Lake Callaghan Creek Nairn F a l l s Shannon F a l l s Brandywine F a l l s Cheakamus River V a l l e y Black Tusk Meadows 71 such as Gymnomitrion obtusum, Diplophyl1um t a x i f o l i u m and Lophozia  v e n t r i c o s a var. v e n t r i c o s a . Rock i n c l i n a t i o n up to about 70^ permits some s o i l and humus accumulation underneath the bryophyte veg e t a t i o n . In such s i t u a t i o n s frequent accompanying species are 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 rock s u r f a c e s , where of t e n a considerable depo-s i t i o n o f humus can be found, the two character species o f the present a s s o c i a t i o n w i l l e v e n t u a l l y be outcompeted by Racomitrium canescens, Polytrichum p i l i f e r u m and Dicranum fuscescens, the f i r s t two taxa being the c haracter species of the Racomitrio - Polyt r i c h e t u m p i l i f e r i . Under exposed c o n d i t i o n s with only a s l i g h t humus l a y e r or no s o i l accumu-l a t i o n whatsoever, Gymnomitrion obtusum, Marsupel1 a emarginata and C e p h a l o z i e l l a byssacea are important accompanying s p e c i e s . At higher e l e v a t i o n several s p e c i e s , character taxa of other 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 of these are Racomitrium sudeticum, Gymnomitrion concinnatum, K i a e r i a b l y t t i i and Pseudoleskea i n c u r v a t a . Accompanying taxa with r a r e occurrences ( a c c i d e n t a l species) i n these reTeves from s u b a l p i n e - a l p i n e s i t e are Marsupella condensata, 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 . F i n a l l y , a l o c a l " f a c i e s " i s recognized from steep, wet rocks at Shannon F a l l s ^. At places where some s o i l had accumulated - i n and around cracks and toward the base of these rocks - major species from the Racomitrio - Andreaeetum p e t r o p h i l a e had e s t a b l i s h e d (Andreaea  r u p e s t r i s , A. r o t h i i , Marsupel1 a emarginata, Racomitri um heterostichum Table V. R a c o m i t r i o ( h e t e r o s t i c h i ) - Andreaeetutn pet 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 of releves Releve-number Elevation(m) D i r e c t i o n ofgexposure I n c l i n a t i o n ( ) Total cover(%) 1 2 3 4 5 46 347 345 348 346 61 61 61 61 61 NW N-NW NW NW N-NW 60 72 58 43 62 75 80 75 85 80 3.4 2.4 4.4 4.4 3.4 4.4 3.4 2.4 2.4 2.4 2.4 2.4 2.4 2.4 3.4 1.2 1.2 1.2 1.2 2.3 1.4 1.4 1.4 2.4 2.4 + .4 T.4 + .4 + .4 + .4 2.2 2.2 2.2 2.2 + .4 1.4 1.4 1.4 1.4 1.4 1.4 + .4 + .2 + .2 + .2 + .2 + .2 1.2 1.2 Campylopus a t r o v i r e n s Andreaea r u p e s t r i s Marsupella emarginata Racomitrium heterostichum var. heterostichum Gymnomitrioni obtusum C e p h a l o z i e l l a byssacea var. byssacea Paraleucobryum enerve Chandonanthus f i l i f o r m e Diplophyllum a l b i c a n s 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 + .4 Location of a l l r e l e v e s : .Shannon F a l l s . 73 Gymnomitrion obtusion). However, under these c o n d i t i o n s , a combination of spec i e s , 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 the f o l l o w i n g taxa: Campy!opus a t r o v i r e n s , Paraleuco- bryum enerve, Anastrophyl1 urn a s s i m i l e , Diplophyllurn a l b i c a n s , Chandonan- thus 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). Since most species 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 considerably beyond the present study area, f u r t h e r research i s needed i n order to designate t h i s ' a c c i d e n t a l ' combination of species as a suba s s o c i a t i o n of the Racomitrio - Andreaeetum p e t r o p h i l a e , or 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 " ass. nov. (Table V I ) . Problems have a r i s e n here around the taxon designated as character species f o r the 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 to R. heterostichum, but d i f f e r s from i t by the mixture of long and short a p i c a l l e a f c e l l s and the ridged (resembling p a p i l l a e ) c r o s s w a l l s i n l e a f cross 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 , the name "Racomitrium b r e v i p e s " Kindb. appears to be synonymous with R. sudeticum (Funck)~B.S.G. based on the apparent type. According to Lawton (1971) t h i s type c o l l e c t i o n of R. brevipes 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) suggestions and Banu's (1969) concepts, "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 separate taxonomical 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) the name "R_. brevipes" Kindb. i s used f o r convenience, since the western North American material i s c l e a r l y not i d e n t i c a l to the European concept of JR. heterostichum. The geographic d i s t r i b u t i o n of "Racomitrium brevipes" extends considerably beyond the present study area: from the west coast o f Vancouver I s l a n d to the Queen C h a r l o t t e I s l a n d s , a l l along the north coast and eastward i n t o the P u r c e l l Mountains i n the Kootenays (Tan 1980). Table VI. "Racomitrietum brevipis" ass. nov. Number of releves 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Releve-number 252 236 258 264 265 131 133 132 235 263 248 249 254 255 257 262 Elevation(m) 1487. 1487 1492 1487 1487 457 457 457 1487 1487 1487 1487 1492 1492 1492 1487 Direction ofnexposure S-SE NE NE S-SW S-SW SW NW E NW W E-SE E-SE NE NE NW NW Inclinat1on( ) 68 38 50 68 52 38 52 30 44 62 26 28 80 64 76 78 Total cover(%) 100 100 90 100 90 100 100 80 90 100 90 100 100 80 90 90 "Racomitrium brevipes" 2.4 1.2 2.4 3.4 1.4 4.4 3.4 3.4 4.4 4.4 5.5 5.5 3.4 3.4 4.4 2.2 Andreaea rupestris 1.2 2.4 1.2 1.2 3.4 2.2 2.2 1.2 1.2 1.4 1.2 Kiaeria starkei 1.4 1.2 2.4 1.2 2.4 1.4 3.4 2.4 3.4 3.4 Lophozia sudetica 4.4 1.4 2.6 2.4 1.4 2.4 2.4 1.4 Dicranoweisia crispula 4.5 4.4 3.4 2.4 1.4 1.4 1.2 1.4 Pseudoleskea radicosa 3.4 1.4 1.4 1.4 1.4 2.4 1.4 Lophozia ventricosa 1.4 var. ventricosa 1.4 1.4 1.4 + .2 1.4 1.4 2.4 Barbilophozia floerkei 1.2 2.4 3.4 + .2 1.2 1.4 1.2 Racomitrium sudeticum 1.2 3.4 1.2 1.2 2.2 1.2 3.4 Diplophyllum taxifolium 1.4 1.4 1.2 + .4 1.4 Scapania americana 1.2 1.2 Polytrichum piliferum 1.1 + .1 + .1 Pohlia nutans 1.2 2.2 2.2 2.2 1.4 1.1 Racomitrium canescens 1.4 Gymnomitrion concinnatum 1.4 1.4 1.4 1.4 1.4 Ptilidium californicum 3.4 4.4 1.2 1.2 Andreaea nivalis 2.2 Marsupella stableri 1.4 1.4 1.4 Polytrichum alpinum + .1 + .1 + .1 var. macounii Hypnum circinale 3.4 1.1 2.2 Kiaeria blytt i i 2.2 ^1 -p. Found only once: 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 75 Hence, the a s s o c i a t i o n i s l i k e l y to occur more widespread than encoun-tered i n the present study. As such i t i s , apart from low e l e v a t i o n l o c a l i t i e s such as Murrin P r o v i n c i a l Park and Brandywine F a l l s , des-c r i b e d from subalpine 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 rare occurrences i n other a s s o c i a t i o n s "Racomitrium  brevipes" i s h i g h l y constant i n , hence, a good character 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 to the three.other syntaxa of the Andreaeion p e t r o p h i l a e i s i n d i c a t e d by the frequency of the a l l i a n c e character species 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 Lophozia  s u d e t i c a . The "Racomitrietum b r e v i p i s " forms lush vegetations on gently s l o p i n g surfaces of exposed i n t r u s i v e rocks. The predominant (sub)alpine character 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 the presence of accompa-nying species such as Racomitrium sudeticum, 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 Marsupella s t a b l e r i Gymnomitrietum co n c i n n a t i Herzog 1943 (Table V I I ) . This a s s o c i a t i o n has been reported and described from subalpine and a l p i n e areas i n the southern p o r t i o n of West Germany (Herzog 1943, P h i l i p p i 1956 and Hertel 1974) . Theirs as we l l as the present data show that the Gymnomitrietum conci n n a t i replaces the Racomitrio -Andreaeetum p e t r o p h i l a e at these higher e l e v a t i o n s . Likewise i t s character species - Gymnomitrion concinnatum -, t h i s a s s o c i a t i o n i s 76 s t r i c t l y confined to subalpine and alpine s i t e s . I t forms a low but extensive bryophyte cover over steeply sloping exposed rocks surfaces. The association's most frquent and abundant accompanying species are Diplophyllum taxi folium, Marsupella emargi nata and Lophozia sudetica. Important a l l i a n c e character species are Lophozia ventricosa var. ventricosa and Andreaea rupestris. This s t r i c t subalpine-alpine charac-ter of the association i s also reflected 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 b a i l e y i , • Ahthelia juratzkana, Marsupella condensata, Kiaeria b l y t t i i and Bartramia ithy-3 phylla. Additional associates constitute three local 'variants' The Gymnomitrietum concinnati - typicum (1 i n Table VII) predominantly occurs i n Cypress Bowl Provincial Park and Mount Seymour (1433 m). Most frequent and abundant accompanying species in these areas are Andreaea n i v a l i s and Kiaeria s t a r k e i . Along Garibaldi Lake (1492 m), Tritomaria quinquedentata, Anthelia  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 constitutes the Tritomaria quinquedentata-Andreaea b l y t t i i -variant (2). In the Diamond Head area (1524 m), in the southwest corner of Garibaldi Provincial Park, Barbilophozia f l o e r k e i , Dicranum fuscescens, Dryptodon patens and Racomitriurn heterostichum var. af f i n e were frequent accompanying species. This group of species constitutes the Dicranum  fuscescens-Barbilophozia floerkei - variant (3), which, c h a r a c t e r i s t i c a l l y , occurs on gently sloping rock surfaces with considerable s o i l accumu-Table VII. Gymnomitrietumconcinnati Herzog 1943 Number of releves 1 Releve-number 116 Elevation(m) 610 Direction ofnexposure N-NW Inclination(u) 106 Total coverU) 100 2 3 4 5 6 7 117 169 170 122 123 174 610 1433 1433 610 610 1433 N-NW NW NW N N NW 100 72 76 74 64 80 90 95 100 80 100 80 8 9 10 11 12 13 175 173 246 247 240 241 1433 1433 1487 1487 1492 1492 N-NW NE N N SW SW 62 68 54 60 58 48 100 100 100 100 100 90 14 15 16 17 18 19 20 242 176 177 334 335 338 336 1492 1433 1433 1524 1524 1524 1524 SW E E NW N NE NW S 60 76 84 32 52 58 58 100 80 100 95 80 100 80 21 22 23 24 25 26 337 311 312 315 313 314 1524 1219 1219 121f 1219 1219 SE N NW NE N NE E NE C NE 34 62 40 74 78 66 90 100 90 100 100 100 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 lytt i i Marsupella condensata Cephaloziella byssacea var. byssacea Scapania scandica Barbilophozia floerkei Dicranum fuscescens Dryptodon patens Racomitrium heterostichum var. affine Cynodontium jenneri Pt111di urn californicum Andreaea rothii Amphidium californicum Campylopus fragilis Arctoa fulvella Kiaeria blytt i i 4.4 3.4 2.4 4.5 2.4 + .4 2.4 3.4 3.4 4.5 4.5 4.5 2.4 3.4 1.2 3.4 3.4 1.2 1.2 1 .2 2.4 2.4 2.4 2.4 1.4 2.4 3.4 1.4 3.4 2.4 1.2 + .2 + .2 4.5 2.4 2.4 1.4 2.4 2.4 1.4 1.4 1.4 1.4 2.4 1.4 2.4 + .4 1.4 1.4 1.4 1.4 2.4 3.4 1.4 2.4 1.2 3.4 1.2 2.2 1.2 1.2 2.4 1.4 1.4 1.4 1.4 2.4 2.4 2.4 1.2 1.2 2.2 + .2 + .2 + .2 1.2 1.2 1.2 + .2 + .2 2.3 3.4 1.4 + .2 2.2 1.2 2.4 + .1 + .1 1.2 1.2 2.2 1.1 + .1 + .1 1.2 1.1 1.2 4.5 4.5 2 4 2.4 2.4 1.4 1.2 4.4 3.4 3 4 2.4 3.4 1.4 + .2 1 4 2.4 1.4 1.2 1.2 + 2 2.4 2.4 + .2 2.2 1.2 1.2 1.2 1 -2 + .1 + 1 + .1 1.1 2.4 4.4 4.4 4.4 3.4 3.4 4.4 4.4 1.2 1.2 2.2 2.2 2.2 1.2 4.4 +.4 1.4 +.4 1.4 1.4 1.4 +.2 1.4 2.2 3.3 2.3 3.3 2.4 1.4 +.2 1.2 3.4 3.4 2 1.4 1.4 2.4 3.4 1.4 1.4 2.4 2.4 1.4 1.4 2.4 1.2 1.2 1.2 +.4 1.4 +.4 1.4 1.4 1.4 1.4 2.4 1.4 3 3.4 2.4 +.4 +.4 +.4 2.4 1.4 3.3 2.2 1.2 2.3 1.2 2.2 1.2 1.2 1.2 3.3 1.2 1.2 1.2 +.2 1.2 1.2 4.4 2.2 3.3 3.4 2.4 4 1.2 3.4 2.4 3.4 4.4 1.2 +.2 1.2 1.2 1.2 1.2 +.2 1.2 +.2 Table VII (continued). Found only 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 , Marsupella sphacelata, Pel 1 i a neesiana, Brachytheciurn s t a r k e i , Pseudoleskea i n c u r v a t a , Jungermannia c o n f e r t i s s i m a . R e l e v e - l o c a t i o n s : 116,117,122,123 Cypress Bowl 169,170,173,174,175,176,177 Mount Seymour oJS'oi? B 1 a c k T u s k Meadows 240,241,242 G a r i b a l d i Lake 311,312,313,314,315 Wh i s t l e r Mountain 334,335,336,337,338 Diamond Head 79 l a t i o n . On Mount Whisler (2286 m) Andreaea r o t h i i becomes an important a s s o c i a t e of the 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 these l o c a l companions can be designated as d i f f e r e n -t i a t i n g species to d i s t i n g u i s h subassociations needs f u r t h e r i n v e s t i -g a t i o n . E s p e c i a l l y , because a l l taxa have a d i s t r i b u t i o n which extends beyond the l i m i t s of the present study area, t h i s cannot be done w i t h -out more research. Racomitrietum s u d e t i c i Herzog 1943 (Table V I I I ) . Herzog (1943) reports and describes a Racomitrium sudeticum-a s s o c i a t i o n from the Black Forest region i n southern West Germany. In the study area the Racomitrietum s u d e t i c i occurs on exposed rocks at subalpine and a l p i n e e l e v a t i o n s of Cypress Bowl P r o v i n c i a l Park, 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. In c o n t r a s t to the Gymnomitrietum c o n c i n n a t i , which occurs on steep (or even over-hanging) rock s l o p e s , the Racomitrietum s u d e t i c i predominantly grows on gently s l o p i n g rock s u r f a c e s . G e n e r a l l y , more humus and mineral s o i l has been formed underneath the lush bryophyte assemblages i n which the a s s o c i a t i o n character s p e c i e s , Racomitrium sudeticum, i s the dominant taxon. Most frequent accompanying species are Gymnomitrion concinnatum, Andreaea n i v a l i s , K i a e r i a s t a r k e i and Dicranoweisia c r i s p u l a . Although they were found only a few times, three taxa out of the genus Grimmia Table VIII. Racomitrietum sudetici Herzog 1943 Number of releves 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Releve-number 118 124 309 330 310 125 266 253 302 303 304 120 121 256 171 172 Elevation(m) 610 610 1219 1219 1219 610 1492 1492 1219 1219 1219 610 610 1492 1433 1433 Direction ofnexposure Inclination(u) W S N E-SE E-NE SE W-NW W NW N W S-SW S NW W-NW NW 58 26 112 60 108 78 62 14 24 48 52 36 70 58 42 84 Total cover(!fc) 60 .100 75 85 85 100 85 100 90 80 80 90 100 100 40 90 3.4 4.4 4.4 4.5 1.4 4.4 4.5 4.4 2.4 1.4 3.4 3.4 2.4 1.4 3.4 3.4 2.4 1.2 + .2 3.4 1.2 1.2 1.2 1.2 2.4 3.4 3.4 2.4 1.2 2.4 1.2 1.2 2.4 1.2 1.2 2.4 3.4 3.4 3.4 1.4 2.4 3.3 1.2 3.2 1.2 2.2 2.2 1.2 2.3 1.2 1.2 1.2 1.4 1.4 1.4 + .4 1.4 4.4 4.4 3.4 3.4 3.4 2.4 2.4 1.4 1.4 1.2 2.3 2.2 1.2 1.4 1.4 1.4 2.4 1.4 2.4 1.4 1.4 2.4 1.4 3.4 2.2 1.2 1.2 + .2 1.2 3.2 1.4 1.4 1.4 Racomitrium sudeticum Gymnomitrion concinnatum Andreaea rupestris Andreaea nival 1s Kiaeria starkei Dicranoweisia crispula Diplophyllum taxifollum Racomitrium heterostichum var. heterostichum Pseudoleskea radicosa Kiaeria blytti i Grimmia alpestris var. holzingerl Lophozia sudetica Marsupella sparsifolia Amphidium californicum Grlnrnia donniana Pseudoleskea atricha Pohlia cruda 2.2 1.1 Grimmia alpestris var. alpestris 1.2 1.2 Racomitrium canescens 1.2 2.2 Lophozia ventricosa var. ventricosa 1.4 3.4 Lophozia indsa 1.4 1.4 Marsupel la emarginata. 1.4 1.4 Grimmia montana 1-2 1.2 Pohlia nutans +.2 1.2 Found only once: Andreaea b ly 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 171,172 Mount Seymour 302,303,304,309,310,330 Whistler Mountain 253,256,266 Panorama Ridge 81 were confined to 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 var. a l p e s t r i s and var. h o l z i n g e r i and Grimmia donniana. Herzog (1943), i n h i s o r i g i n a l d e s c r i p t i o n of t h i s syntaxon from southern West Germany, mentions Grimmia commutata and G_. elongata 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 , together 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 s p a r s i f o l i a r e f l e c t the s u b a l p i n e - a l p i n e character of the Racomitrietum s u d e t i c i . 8.1.3.2. Order POLYTRICHETALIA PILIFERI v. Hubschmann 1967 A l l i a n c e P o l y t r i c h i o n p i l i f e r i Norr 1969. Racomitrio - Pol y t r i c h e t u m p i l i f e r i Herzog 1943 (Table I X ) . The Racomitrio - Polytrichetum p i l i f e r i was f i r s t reported by Herzog (1943) from the Black Forest region i n southern West Germany. The author described 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 (1955), Norr (1969), Neumayr(1971) and Hubschmann (1967, 1975). The l a s t author a l s o reported the syntaxon from Vancouver I s l a n d , again from sandy s o i l (Hubschmann 1978). Separate a s s o c i a t i o n s (Poly-trichetum p i l i f e r i and Racomitrietum canescentis) have been described f o r both character s p e c i e s , Polytrichum piliferum and Racomitrium  canescens, r e s p e c t i v e l y (Krusenstjerna 1945, F r e g r i 1933, Smarda 1947, 82 Giacomini 1951, Dunk 1971). In the present study t h i s predominantly 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 surfaces at low e l e v a t i o n s only. In both m i c r o - s i t e types the amount of accumulated humus and/or mineral s o i l c o n s t a n t l y ranged from 1 - 2 cm. The character s p e c i e s , Polytrichum p i l i f e r u m and Racomitrium  canescens, are commonly accompanied by other ' t e r r e s t r i a l ' species such as Ceratodon purpureus and Polytrichum j u n i p e r i num. Other frequent a s s o c i a t e s , which o f t e n occur on rocks with a t h i n n e r s o i l l a y e r are species such as C e p h a l o z i e l l a byssacea var. byssacea, Hedwigia c i l i a t a and S c h i s t i d i u m apocarpum var. s t r i c t u m . In the d i f f e r e n t i a t e d t a b l e of the f i r s t major group, a c o l l e c -t i o n of four releves (305 to 308) i s s p l i t o f f from the releves c o n s t i -t u t i n g the present a s s o c i a t i o n . These four releves taken at Mount W h i s t l e r , are discussed f u r t h e r i n the d i s c u s s i o n of major group number 3 (8.3.3.1 Ass. Dichodontietum p e l l u c i d i ; subass. B l i n d i o - Scapanie-tosum paludosae). Scapanietum americanae ass. nov. (Table X). The d i s t r i b u t i o n of 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 the same pattern as that of the Racomitrio - Polyt r i c h e t u m p i l i f e r i . With respect to i t s h a b i t a t i t a l s o occurs on h o r i z o n t a l or s l i g h t l y s l o p i n g rock surfaces o v e r l a i n by s o i l and/or humus. The character species of the a s s o c i a t i o n i s Scapania americana. a western North American endemic. I t Table IX. Racomitrio - Polytrichetum pi l i fer i Herzoa 1943 Number of releves Releve-number Elevation(m) Direction ofQexposure Inclination( ) Total cover(%) 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 12 22 27 68 31 32 157 158 286 287 294 296 295 357 358 359 36 - 18 18 18 61 61 266 266 610 610 610 610 610 354 354 354 E-SE S-SW SE U-NU S-SW N-NE SW S E-NE E E E-NE E-NE E E-SE E 54 38 52 46 38 54 56 54 48' 36 42 68 30 52 44 62 60 95 80 85 90 100 90 90 60 60 80 100 100 85 100 90 3.4 5.4 3.4 4.4 4.4 3.4 1.2 1.2 1.2 1.2 3.4 1.4 2.4 3.4 4.4 3.4 3.3 2.2 3.3 1.2 2.2 1.2 1.2 2.2 1.2 + .2 1 .2 1.2 2.2 1.2 2.3 2.2 2.3 1.2 1.2 1.2 1.2 1.2 3.4 3.4 1.4 2.4 1.4 2.4 2.4 3.4 1.2 1.2 3.4 1.2 1.2 1.2 2.4 2.4 1.4 1.4 1.4 1.4 1.4 1.2 1.2 1.2 1.2 1.2 1.2 2.4 + .2 + .2 + .2 1.2 2.3 1.2 2.3 3.4 4.4 + .2 5.4 1.4 2.4 1.2 + .2 2.3 1.2 1.2 1.2 1.4 + .4 1.4 + .4 1.4 2.4 1.1 + .1 + .1 1.1 1.1 2.2 + .4 1.4 + .4 1.4 1.4 2.4 1.2 1.2 3.4 3.4 1.2 1.2 1.2 1.2 2.4 2.4 2.3 2.2 1.2 1.2 1.2 1.4 2.4 1.4 1.4 1.4 3.3 3.3 1.2 1.2 1.2 2.2 2.2 1 .2 Racomitrium canescens Polytrichum piliferum Ceratodon purpureus Hypnum -subimponens Schistidium strictum Cephaloziella byssacea var. byssacea Racomitrium heterostichum var. heterostichum Pohlia nutans Hedwigia c i l iata Grimmia torquata Lophozia ventricosa var. ventricosa Polytrichum juniperinum Scapania americana Homalothecium nevadense Racomitrium sudeticum Aulacomnium androgynum Barbilophozia floerkei Amphidium lapponicum Tortula princeps Dryptodon patens 3.4 1.2 1.2 Homalothecium nutalli l 2.3 1.2 _^ 1.2 Claopodium bolanderi 1.4 1.4 1.4 Pseudoleskea radicosa 1-4 1.4 1.4 "Racomitrium brevipes" 2.2 3.2 Porella navicularis 3.4 1.4 Douinia ovata 1.4 3.4 Anacolia menziesii 2.3 2.3 Reboulla hemisphaerica 1-4 2.4 Cephaloziella byssacea var. asperifolia +.4 2.4 Lophocolea cuspidata 1.4 1.4 Radula bolanderi 1.4 1.4 Encalypta ci l iata 1.2 +.2 Found only once: 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 (continued). Rel e v e - l o c a t i o n s : 12 22,27 68 31,32 157,158 286,287,294,295,296 357,358,359 Horseshoe Bay Lighthouse Park B r i t t a n i a Beach H i s t o r i c S i t e B r i d a l V e i l F a l l s C h i l l i w a c k River V a l l e y Nairn F a l l s Agassiz Mountain 85 i s o ften a s s o c i a t e d with Polytrichum alpinum var. macounii, P o h l i a  nutans and Bartramia pomiformis. Other frequent companions are Lophozia v e n t r i c o s a var. v e n t r i c o s a , Scapania scandica and Dicranum  scoparium. The syntaxonomically c l o s e r e l a t i o n s h i p of the Scapanietum americanae to the p r e v i o u s l y described syntaxon i s expressed by species such as Racomitrium canescens, Hypnum subimponens and Ceratodon purpu- reus. However, i n t h i s a s s o c i a t i o n , Polytrichum p i l i f e r u m , P_. j u n i - perinum and species l i k e C e p h a l o z i e l l a byssacea var. byssacea and Hedwigia c i l i a t a occur as rare a s s o c i a t e s . In c o n t r a s t to the Racomitrio - Polytrichetum p i l i f e r i , which, with no ex c e p t i o n , occurs i n exposed h a b i t a t s , the current a s s o c i a t i o n i s sometimes found on rock surfaces shaded or somewhat protected by surrounding low vegetation or other boulders. In these s i t u a t i o n s , the character species as w e l l as the otherwise frequent companions, can be associated with species such as Claopodium b o l a n d e r i , Plagiothecium  p i l i f e r u m and Pterigynandrum f i l i f o r m e . These taxa play a major r o l e i n bryophyte assemblages described from shaded and protected rocks. 1. A ' f a c i e s ' i s a more or l e s s a c c i d e n t a l combination of species w i t h i n an a s s o c i a t i o n or s u b a s s o c i a t i o n . 2. Hebrard (1978) reports on an a s s o c i a t i o n with 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 new-l 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 species composition, to the present data as w e l l as to phytosocio-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 Hertel (1974). The low constancy of Andreaea r u p e s t r i s and the presence of species l i k e Racomi t r i um  sudeticum, Gymnomitrion concinnatum, Dicranoweisia c r i s p u l a , Bart-Table X. Scapanietum americanae ass, nov. Number of releves Rel eve-number Elevation(m) Direction ofQexposure Incl ination( ) Total cover(X) 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 ci l iata Plagiothecium piliferum Pseudobraunia californica Pterigynandrum filiforme Lophocolea cuspidata Claopodium bolanderi 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 16 38 37 44 43 291 293 292 316 317 318 319 33 35 154 155 61 61 61 61 61 610 610 610 266 266 266 266 61 61 266 266 N-NW E S-SW NE NE E E E NW N NW N-NW E-NE N-NW N NE 32 48 38 48 26 50 60 66 44 56 32 64 60 56 50 40 100 100 90 100 100 90 80 90 100 100 100 100 100 90 90 90 1.4 2.4 5.4 2.4 1.4 3.4 3.4 2.4 1.4 2.4 3.4 3.4 1.4 3.4 2.4 2.4 3.2 3.2 1.2 2.2 + .1 + .1 4.3 + .1 1.2 2.2 1.2 1.2 1.2 + .1 2.2 1.2 2.2 1.2 1.2 2.2 1.2 3.4 1.4 1.4 1.4 3.4 1.4 1.4 2.4 2.4 1.4 1.4 1.4 3.4 2.4 1.4 1.4 1.2 1.2 + .2 4.3 4.3 4.3 3.3 2.3 1.2 1.2 3.3 2.2 1.2 1.2 1.2 + .2 2.3 1.2 2.3 1.2 1.2 1.4 1.4- + .4 + .4 + .4 2.4 2.4 1.4 + .2 1.4 + .2 1.4 1.4 1.4 2.4 1.4 + .4 4.4 2.4 2.4 2.4 1.4 2.4 1.4 1.4 1.4 + .4 2.4 1.1 2.4 3.2 3.2 + .1 2.2 1.2 2.2 1.2 2.2 +.2 1.4 1.4 1.4 1.4 1.4 1.1 1.2 3.4 1.4 +.4 + .4 1.4 1.2 1.2 1.4 1.4 1.4 1.4 + .2 2.4 2.4 + .4 1.4 + .4 2.2 1.1 1.4 2.2 1.2 2.2 2.2 1.4 Found only once: 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 Murrin Provincial Park Bridal Veil Falls Nairn Falls Chilliwack River Valley SI esse Creek C O 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 des-c r i b e d a s s o c i a t i o n ' s nature as fragments of high e l e v a t i o n s syntaxa as the Gymnomitrietum co n c i n n a t i Herzog 1943 and Racomitrietum 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 with one p a r t i c u l a r taxon being dominant, which i s absent or only o c c a s i o n a l l y (and with 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 - "typicum". 88 8.2 BRYOPHYTE ASSOCIATIONS FROM SHADED, HUMID ROCKS. Table XI. Bryophyte a s s o c i a t i o n s from shaded, humid rocks. Order LEPIDOZIETALIA REPTANTIS ( P h i l i p p i 1956) Hertel 1974 A l l i a n c e DIPLOPHYLLION ALBICANTIS ( P h i l i p p i 1956) Hertel 1974 C l u s t e r 1. A s s o c i a t i o n Diplophylletum 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 Marsupello - Scapanietum americanae ass. nov. C l u s t e r 3. A s s o c i a t i o n Claopodietum bolanderi A s s o c i a t i o n - typicum Subassociation Isopterygietosum e l e g a n t i s subass. nov. Subassociation Porelletosum r o e l l i i subass. nov. 90 8.2. Bryophyte a s s o c i a t i o n from shaded, humid rocks (Table X I ) . 8.2.1. Synsystematics. P h i l i p p i (1956) proposed the order 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 to contain bryophyte 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. Within t h i s order he a l s o proposed 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 , i n c l u d i n g a.o. the a s s o c i a t i o n Diplophylletum a l b i c a n t i s Krusen-s t j e r n a 1945. This l a t t e r syntaxon had been reported and described e a r l i e r by Schade (1923), hence, as s t a t e d by Hertel (1974). i t should be termed Diplophylletum a l b i c a n t i s Schade 1923. However, the order 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 c o n f l i c t s w i t h the order 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 P h i l i p p i 1963, described from mineral s o i l . Although P h i l i p p i s t r e s s e s the d i s s i m i l a r i t i e s between these two syntaxa, Hertel (1974) r i g h t l y s t a t e d t h a t a species can not be a c h a r a c t e r species f o r two orders. Mineral s o i l communities with Diplophyllum a l b i c a n s show a species combination s i m i l a r to samples from rocky h a b i t a t s . Hence, they should, on f l o r i s t i c c r i t e r i a , be grouped i n one category. This 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 contains a s s o c i a t i o n s from humid, shaded rocks as w e l l as mineral s o i l . Together with the T e t r a -phidion p e l l u c i d a e Krusenstjerna 1945 (also 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 ( P h i l i p p i 1956) em. i s placed i n the order 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 1963) em., f o l l o w i n g Hertel (1974). In the present study, a new a l l i a n c e i s proposed, the Scapanion 91 americanae, c o n t a i n i n g two newly described a s s o c i a t i o n s Marsupello -Scapanietum americanae and Claopodietum b o l a n d e r i . Based on the f o l l o -wing c r i t e r i a these syntaxa are c l a s s i f i e d i n the order 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 1963) Her t e l 1974: (1) a l l three a s s o c i a t i o n s share a considerable number of s p e c i e s ; (2) they occur i n the same types of h a b i t a t s and (3) many species (character species as w e l l as companions) w i t h i n the releves are known from mineral s o i l and humus-covered-rocks. As Neumayr (1971) and Hertel (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 mineral 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 l e v e l s . 8.2.2. Phytogeographical s e t t i n g . 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 ameri-canae, the presence of endemic and circumboreal taxa i s predominant. Although only represented by l e s s frequent companions, two other phytogeographical elements are shared by these syntaxa: b i p o l a r d i s j u n c t s (Blepharostoma t r i c h o p h y l l u m , Polytrichum alpinum var. macounii, Bart- ramia pomiformis) and circumboreal-missing-in-Japan (Lophozia v e n t r i c o s a var. v e n t r i c o s a , Isopterygium elegans). In the Diplophylletum a l b i c a n t i s the circumboreal character s p e c i e s , Diplophyllum a l b i c a n s , i s f r e q u e n t l y accompanied by a western North American endemic, Scapania americana (which replaces Scapania  nemorosa front European samples of t h i s a s s o c i a t i o n . Other endemic taxa 92 are Rhizomnium glabrescens and the more infrequent Diplophyllum t a x i - f o l i u m . The most important circumboreal companions of the Diplophylletum a l b i c a n t i s i n the present study are Marsupella emarginata, Cephalozia  b i c u s p i d a t a , Racomitrium hetrostichum var. heterostichum and Polytrichum  alpinum ( i n f a c t the var. macounii, an endemic taxon). The a l l i a n c e , Scapanion americanae, r e f l e c t s a notable increase i n c o n t r i b u t i o n by western North American endemics. For i n s t a n c e , Scapania americana and Claopodium bolanderi are the r e s p e c t i v e character species 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 . Impor-t a n t companions are Isothecium s t o l o n i f e r u m , Rhizomnium glabrescens, 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 ssp. n i s q u a l l e n s i s . However, there s t i l l i s a large c o n t r i b u t i o n by circumboreal species i n c l u d i n g 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 a l b i c a n s and to a l e s s e r extent by Racomitrium heterostichum var. heterostichum and Polytrichum alpinum var. 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 of both a s s o c i a t i o n s of the Scapanion americanae are r e p r e s e n t a t i v e s from three a d d i t i o n a l phytogeo-graphical elements: western North America - western Europe - d i s j u n c t s (Scapania scandica, Plagiothecium p i l i f e r u m ) , a western 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 (Polytrichum  alpinum, Bartramia pomiformis, S t o k e s i e l l a praelonga var. praelonga and Plagiothecium denticulatum 93 8.2.3. Order LEPIDOZIETALIA REPTANTIS ( P h i l i p p i 1956) He r t e l 1974 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 ( P h i l i p p i 1956) Hertel 1974. Diplophylletum a l b i c a n t i s Schade 1923 (Table X I I ) . Bryophyte communities c o n t a i n i n g Diplophyllum a l b i c a n s occur on mineral s o i l as w e l l as on rock. This oceanic a s s o c i a t i o n (Gams 1952) i s widely d i s t r i b u t e d over c e n t r a l and northern Europe. Frequent reports from shaded e p i l i t h i c h a b i t a t s are given by Schade (1923', 1934), Herzog (1943), P h i l i p p i (1956, 1965), Norr (1969), Neumayr (1971), Dunk (1971), M a r s t a l l e r (1973) and Hertel (1974). In the study area the Diplophylletum a l b i c a n t i s occurs" wide-spread at predominantly low but sometimes at subalpine e l e v a t i o n s . I t s h a b i t a t i s confined to shaded r o c k s , adjacent to o r i n the d i r e c t neighbourhood of water, on humid ro c k w a l l s i n canyons or at the base of b i g boulders 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 lush assemblages c h a r a c t e r i z e d by mostly low-growing bryophytes i n f r e q u e n t l y mixed with l u x u r i o u s growth of some mosses and l i v e r w o r t s . The c h a r a c t e r species i s Diplophyllum a l b i c a n s , which i s most fre q u e n t l y accompanied by Scapania americana, a western North American endemic which replaces Scapania nemorosa from the Diplophylletum a l b i -c a n t i s i n Europe ^. Another species used as character species f o r the l a t t e r syntaxon i s Heterocladium heteropterum ( P h i l i p p i 1956, Norr 1969). Present data show Heterocladium macounii w i t h only occasional occurrences Table XII. Diplophylletum albicantis Schade 1923 Number of releves 1 2 3 4 5 6 7 8 9 10 11 12 13 1.4 15 Releve-number 1 2 4 75 80 180 59 95 100 45 50 60 54 55 61 Elevation(m) 1150 1150 30 18 61 1433 61 30 610 61 61 30 61 61 61 Direction ofQexposure NE NE SW NE E E-SE N SW SW NW N N-NW N-NW N N-NE Inclination ) 85 40 64 76 52 86 70 56 64 78 74 56 76 78 86 Total cover(%) 90 85 80 90 90 100 100 100 100 100 95 100 70 80 60 Diplophyllum albicans 5.5 • 4.5 2.4 4.4 4.5 3.4 5.4 2.4 3.4 2.4 3.4 + .4 Scapania americana 1.4 1.4 4.4 2.4 3.4 2.4 2.4 4.4 3.4 Racomitrium heterostichum var. heterostichum 3.3 3.3 2.2 + .2 + .1 2.2 + .1 3.3 Bazzania denudata + .4 3.4 4,5 IA 4.4 3,1 1-4 Marsupella emarginata + .4 1.4 2.4 + .4 + .4 + .4 Rhizomnlum glabrescens + .1 1.2 2.2 + .1 + .1 + .1 Isopteryglum elegans 2.4 1.4 1.4 + .4 1.4 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 2.2 + .2 + .4 + .1 1.4 + .1 2.2 1.1 +.1 2.2 +.4 +.4 1.4 2.2 +.4 +.4 2.4 1.1 1.4 3.4 4.4 4.5 1.2 + .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 Mount Seymour 4,60,95 Lynn Canyon 75 Lighthouse Park 61,80 Bridal Veil Falls 45,50,54,55,59 Shannon Falls 100 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 species f o r the 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 to the European representa-t i o n of the Diplophylletum 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 are: Marsupella emarginata, Isopterygium elegans, Cephalozia b i c u s p i d a t a , Bartramia pomiformis and Lophozia v e n t r i c o s a var. v e n t r i c o s a . However, D i c r a n e l l a heteromalla, a frequent and important character species f o r the a l l i a n c e and order i n Europe occurs i n the study area only o c c a s i o n a l l y . Within the Diplophylletum a l b i c a n t i s the present data d i s t i n -guish a Bazzania denudata - v a r i a n t , which, i n a d d i t i o n , can be charac-t e r i z e d by species such as Chandonanthus f i l i f o r m i s and to a l e s s e r extent by Herbertus aduncus. I t s d i s t r i b u t i o n i s confined to very shaded and protected steep boulders w i t h i n f o r e s t vegetation around Shannon F a l l s and B r i d a l V e i l F a l l s . Releves 59, 95 and 100 i n d i c a t e a Bazzania t r i c r e n a t a - C a l y p o g e i a muelleriana - f a c i e s through the a c c i d e n t a l species combination of these two bryophyte taxa. A l l i a n c e Scapanion americanae a l l . nov. Marsupello - Scapanietum americanae ass. nov. (Table X I I I ) . The a s s o c i a t i o n a l s o occurs on rocks and boulders, not exposed but shaded and s h e l t e r e d by overhanging and/or surrounding v e g e t a t i o n . I t forms lush assemblages on gently s l o p i n g rock surfaces w i t h only a Table XIII. Marsupello - Scapanietum americanae ass. nov. Number of releves 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 Releve-number 3 63 98 101 13 23 29 272 273 24 74 58 140 139 69 67 62 103 Elevation(m) 1150 61 30 • 30 36 18 61 266 266 18 18 61 30 30 18 30 61 30 Direction of0exposure NE W NW SE SE S-SE S-SW W W E-NE NW N S-SE S-SE S-SI' E-SF NW SE Inclination( ) 74 74 52 72 36 82 72 54 52 62 58 70 64 78 52 78 70 24 Total cover(%) 80 100 100 100 95 95 80 90 90 90 80 85 100 100 100 90 100 80 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 2.4 5.5 3.4 3.4 + .1 3.4 2.4 1.4 + .2 1.4 + .1 3.4 3.4 2.2 1.4 3.4 + .4 1.2 2.4 2.4 +.1 1.4 4.5 3.4 2.4 + .4 2.2 2.4 1.4 2.4 2.4 5.3 1.4 1.4 3.4 3.4 2.4 1.4 + .1 + .1 3.4 3.4 + .1 2.2 + .4 +.4 + .2 1.2 + .1 1.4 + .4 2.4 4.4 1.2 1.4 1.4 1.4 1.4 1.2 1.4 1.4 2.4 3.4 4.5 1.4 1.2 +.2 1.4 +.1 1.4 +.4 + .1 + .4 1.2 1.4 1.4 1.4 1.4 + .4 + .1 4.4 2.4 + .4 1.4 + .1 1.4 2.4 4.5 3.4 2.4 1.4 + .4 + .4 + .1 4.4 1.4 1.4 1.2 1.4 + .4 1.4 + .4 3.4 3.4 2.4 .2.4 1.4 3.4 5.5 1.2 +.2 1.4 1.4 + .2 2.2 + .4 2.2 2.4 1.4 2.4 2.4 2.4 3.4 3.4 1.2 2.4 3.4 1.4 + .1 1.4 1.4 2.4 4.4 3.4 4.4 1.4 2.2 + .4 + .4 1.2 Found only once: 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 (continued). R e l e v e - l o c a t i o n s : 3 58,63 98,101,103 13 23,24 29,62 272,273 74 67,139,140 69 Mount Seymour Shannon F a l l s Lynn Canyon Horseshoe Bay Lighthouse Park B r i d a l V e i l F a l l s ^ C h i l l i w a c k R i v e r V a l l e y Sunset Creek Murrin P r o v i n c i a l Park Porteau Camp 98 t h i n l a y e r of humus or s o i l accumulation. I t s d i s t r i b u t i o n , as recognized by the present data, i s confined to the southern p o r t i o n o f the study area and ranges from Lighthouse Park i n t o the C h i l l i w a c k River V a l l e y . I t s a f f i l i a t i o n to the previous a s s o c i a t i o n i s , aside 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 the presence and abundance of major species w i t h i n the r e l e v e s . The character species o f the present asso-c i a t i o n are Scapania americana and, co n s t a n t l y a s s o c i a t e d w i t h i t , Marsupella emarginata. Other frequent companions are such species as Racomitrium heterostichum 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 and Isopterygium elegans. Less frequent accompanying species that s t i l l 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 Diplophyllurn a l b i c a n s and Rhizomnium glabrescens. A l l i a n c e character s p e c i e s , which are confined to the Scapanion americanae and, thus, d i f f e r e n t i a t e i t from the 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 cuspidata and Plagiothecium denticulatum. Species which were, despite t h e i r few occurrences, found only i n t h i s a s s o c i a t i o n were Dicranum scoparium and Cynodontium j e n n e r i . However, they cannot be designated as chara c t e r species f o r the Marsupello - Scapanietum americanae, since they occur f r e q u e n t l y as major ass o c i a t e s i n vegetation types from open and exposed s i l i c e o u s r o c k s , 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 bolanderi ass. 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 c h a r a c t e r i s t i c a l l y on steep rock surfaces of rock w a l l s i n s h e l t e r e d creek canyons, and on s l i g h t l y s l o p i n g surfaces of shaded boulders which are protected by surrounding vegetation. Character species f o r the 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 character species f r e q u e n t l y a s s o c i a t e d with 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 , Scapania americana and Isopterygium elegans. Hubschmann (1978) reports on a newly described 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 boulders i n s i d e some Vancouver I s l a n d f o r e s t v egetations. Besides Porotrichum b i g e l o v i i and Claopodium c r i s p i f o l i u m as the most abundant and constant character s p e c i e s , a l s o Claopodium bolanderi 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 to 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 , there 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 Claopodio - Porotrichetum b i g e l o v i i and the Claopodietum b o l a n d e r i , as a newly described a s s o c i a -t i o n i n t h i s study. These s i m i l a r i t i e s are expressed i n a s s o c i a t e d species such as Isothecium s t o l o n i f e r u m , Rhizomnium glabrescens, S t o k e s i e l l a  praelonga var. praelonga, Apometzgeria pubescens and Metaneckera menzie- s i i . The present data d i s t i n g u i s h a Claopodietum bolanderi - t y p i -cum and f u r t h e r d i f f e r e n t i a t e two subassociations w i t h i n t h i s syntaxon. The a s s o c i a t i o n - typicum (1 i n Table XIV) i s c h a r a c t e r i z e d by species 100 such as Isothecium s t o l o n i f e r u m , Plagiothecium p i l i f e r u m , Pseudo-leskea i n c u r v a t a , Hypnum c i r c i n a l e and to 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 Claopodietum 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 americana, Marsupel!a emarginata and P l a g i o c h i l a a s p l e n i o - ides i s minor. These bryophyte taxa play a more important r o l e i n the o v e r a l l f l o r i s t i c composition of the 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 . The f i r s t s u b a s s o c i a t i o n , the Isopterygietosum e l e g a n t i s ( 2 ) , i s defined by the 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 : Isopterygium  elegans, Hypnum subimponens, Rhizomnium glabrescens and Heterocladiurn  macounii. Other species c o n t r i b u t i n g to t h i s sub-unit 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 , are Polytrichum alpinum var. macounii, Plagiothecium undulatum and P_. 1 aetum. This 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 occupies lower portions o f boulders and rockwalls where moisture and r e l a t i v e humidity are found to be higher from s o i l i n f l u e n c e (Sjogren 1964). The second s u b a s s o c i a t i o n o f the Claopodietum bolanderi occurs on very steep to overhanging rock surfaces without any s o i l accumulation 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 species f o r t h i s syntaxon Porelletosum r o e l l i i (3) are Pore!la r o e l l i i , F r u l l a n i a  t a m a r i s c i ssp. 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 bryophytes occasional i n t h i s type o f micro-habitat are Anacol i a m e n z i e s i i , Douinia ovata and Plagiopus o e d e r i . As i n d i c a t e d i n the c l u s t e r a n a l y s i s and the subsequent " d i f f e r -Table XIV. Claopodietum bolanderi ass, nov. Number of releves Rel eve-number Elevation(m) Direction ofQexposure Inclination( ) Total cover(iS) 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 5 7 230 18 19 156 96 97 8 231 339 340 102 225 226 30 47 72 70 71 138 30 30 610 18 18 266 30 30 30 610 1524 1524 30 610 610 61 61 18 18 18 30 SW N SW N N NE N N NW W NW W NW SE SW SW NW E N NE S SW N W Nil W NW W NW E 20 40 66 84 76 68 54 64 68 56 82 108 26 68 68 100 96 84 92 84 94 80 80 100 90 90 100 100 80 100 100 100 100 100 100 100 80 90 80 90 90 80 2.4 2.4 3.4 2.4 3.4 2.4 1.4 3.4 2.4 1.4 4.5 1.1 1.1 1.1 1.4 + .4 + .4 1.4 1.1 2.4 1.1 1.2 + .1 3.4 4.5 3.4 3.4 3.4 1.4 2.4 4.5 4.4 3.4 1.4 3.4 3.4 2.4 2 .4 + .4 + .4 + .4 1.4 1.4 + .4 + .4 + .4 3.4 3 .4 3.4 1.4 1.4 1.4 1.4 1.4 1.4 3.4 2.4 .1 2.4 2.4 1.1 2.4 3.4 4.4 4.4 + .1 1.1 1, 1.4 + .4 1.4 1.4 1.1 1.1 + .1 + .1 1.1 1.1 + .1 2.2 1.4 + .4 + .4 2.4 Claopodium bolanderi 2.4 Plagiochila asplenioldes 3.4 Scapania americana Isothecium stoloniferum Marsupella emarginata Racomitrium heterostichum var. heterostichum Pohlia cruda Apometzgeria pubescens 1.4 Polytrichum alpinum var. macounii .1 +.1 2.2 Diplophyllum albicans 1.4 2.4 +.4 +.4 Mnium spinulosum +.1 +.1 l - 2 + - ' Bartramia ithyphylla 1.2 1.2 2.2 Diplophyllum taxifolium 2.4 1.4 +-4 Cephaloziella byssacea var. asperifolla v +-4 +.4 2.4 Metzgeria conjugata 1.4 1.4 1.4 Radula bolanderi 1.4 +.4 +.4 Plagiothecium denticulatum +.4 2.4 1.4 / Plagiothecium laetum 2.4 2.4 Grimnia torquata I-2 1 , 2 Blepharostoma trlchophyllum 1.4 1.4 Lophocolea heterophylla +-4 1.4 Plagiothecium piliferum 1-4 2.4 2.4 1.4 2.4 Pseudoleskea incurvata 1.4 2.4 2.4 1.4 2.4 1.4 1.4 1.4 2.4 Hypnum circinale Plagiothecium roeseanum Racomitrium sudeticum +.1 1.4 3.4 Isopterygium elegans Hypnum subimponens 5.5 3.4 4.5 ^ 3.4 +.4 +.4 Rhizomnium glabrescens Heterocladium macounii Lophocolea cuspidata Plagiothecium undulatum Porella roel l i i Frullania tamarisci var. nisquallensls Amphidium californicum Anacolia menziesii. Plagiopus oederl Douinia ovata 1.4 4.4 3.4 2.4 2.4 + .4 .  + .1 + .1 + .1 3.3 4.3 1.4 1.4 2.4 1.4 1.4 + .4 2.4 + .1 + .1 2.4 4.5 2.1 2.4 3.4 1.4 2.4 2.4 3.4 3.4 4.5 1.2 3.3 3.3 4.5 2.4 2.3 +.1 1.4 1.4 Table XIV (continued) Found only 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 var. praelonga, Lophozia v e n t r i c o s a var. v e n t r i c o s a , Andreaea r u p e s t r i s , Bazzania denudata, Cephalozia b i c u s p i d a t a , Anastrophyl1um minutum, P o h l i a  nutans, Scapania scandica, C e p h a l o z i e l l a byssacea var. byssacea, Herbertus  aduncus, Metaneckera m e n z i e s i i , Dicranum fuscscens, P o r e l l a n a v i c u l a r i s , Pogonatum urnigerum, Drepanocladus uncinatus, Racomitrium canescens, Racomitrium heterostichum var. a f f i n e , Rhytidiadelphus l o r e u s , Brachythecium  plumosum, Grimmia' anomala ,'. Radula complanata, Scapania mucronata, B a r b i l o p h o z i a f l o e r k e i , T r i t o m a r i a quinquedentata, Pseudoleskeella  tectorum, 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 Lighthouse Park C h i l l i w a c k River V a l l e y Cheakamus River V a l l e y Diamond Head B r i d a l V e i l F a l l s Lonetree Creek Murrin Creek 103 e n t i a t e d " t a b l e (Appendix V ), the rest-group of re l e v e s from shaded, humid rocks i s too fragmentary and too heterogeneous to allow any arrangement. This r e s u l t s from e i t h e r l a c k of r e l i a b l e d i a g n o s t i c species or too few releves w i t h i n c l u s t e r s to designate as separate syntaxa. Hence, a f f i n i t i e s with already recognized a s s o c i a t i o n s were considered too vague and obscure these p o s s i b l e f l o r i s t i c and phyto-s o c i o l o g i c a l r e l a t i o n s h i p s . As such, t h i s rest-group i s not f u r t h e r discussed here. Addi-t i o n a l research may reveal t h e i r a f f i n i t i e s to already e s t a b l i s h e d syntaxa, both i n B r i t i s h Columbia and i n Europe, or t h e i r "own" nature, based on which they may become designated as separate syntaxa. 1. Reports on communities, i n which both species are designated as the co-dominant character species are given by Dunk (1971), Wilczynska (1974), 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 along streams. Order BRACHYTHECIETALIA PLUMOSI ( P h i l i p p i 1956) prov. A l l i a n c e RACOMITRION ACICULARIS Krusenstjerna 1945 C l u s t e r 1 + 3. A s s o c i a t i o n Racomitrio - Scoulerietum aquaticae ass. 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. Subassociation Brachythecio - Conocephaletosum conicae subass. nov C l u s t e r 7. Subassociation R i c c a r d i o - Hygrobielletosum l a x i f o l i a e subass. nov C l u s t e r 9. Subassociation B l i n d i o - Scapanietosum paludosae subass. nov. 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 Marsupello - Nardietum s c a l a r i s ass. nov. Table XV (continued). C l u s t e r 11. A s s o c i a t i o n Hypnetum d i e c k i i ass. nov. A s s o c i a t i o n - typicum Subassociation 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 ass. 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 rocks along streams (Table XV). 8.3.1. Synsystematics. Bryophyte assemblages growing on boulders and rocks along streams, e i t h e r i n open areas or 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 incorporated i n the "aquatic bryophyte communities" ("Wassermoosgesell-schaf t e n " ; a.o. P h i l i p p i 1956, 1965, Neumayr 1971 , Hertel 1974). Hubschmann (1957) has proposed a synsystematic c l a s s i f i c a t i o n of the aquatic bryophyte communities. This treatment 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 are grouped together, and other vegetation 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 c o n d i t i o n s are c l a s s i f i e d i n t o n o n - a l l i e d syntaxa. The present study f o l l o w s the proposals by P h i l i p p i (1956), who emphasizes more c o n s i s t e n t l y the r e l a t i o n between h a b i t a t and t h e i r bryophyte assemblages, and c l a s s i f i e s these vegetations a c c o r d i n g l y . The bryophyte vegetations from stream adjacent rocks, because o f t h e i r unique species composition and species combi-n a t i o n s , can har d l y be l i n k e d with bryophyte assemblages from rocks not i n f l u e n c e d by surrounding or adjacent water bodies. The y e a r l y p e r i o d i c i t y i n water i n - f l u x '(varying water l e v e l through snow melt 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 the development of stream-accompanying bryophyte assemblages. This phenomenon causes a zonation pattern which includes vegetations inundated f o r most of the ye a r , vegetation which l a y dry f o r s h o r t e r or longer periods o f time and, f i n a l l y , bryophyte assemblages only o c c a s i o n a l l y submerged at 108 the time of the year's highest water l e v e l s . The many a s s o c i a t i o n s described by European b r y o - s o c i o l o g i s t s are based mostly on dominant species (designated as character species) w i t h i n these zones. Dominance i n and f i d e l i t y to these 'communities' f a i l e d to u n i t e them i n t o a s o c i o l o g i c a l system, and f o r each of them higher synsystematic u n i t s were erected. Hence, e c o l o g i c a l c r i t e r i a were used i n the proposal f o r a c l a s s i f i c a t i o n of the "aquatic bryophyte 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 various b r y o - s o c i o l o g i s t s , Hertel (1974) provides the most useful ( f o r t h i s study) synsystematic h i e r a r c h y . In the present study, predominantly, those aquatic bryophyte communities were sampled, which are emerged f o r most of the year and those which are only a f f e c t e d ( o c c a s i o n a l l y submerged) at the times o f the year's highest water l e v e l s . These assemblages, "amphibische g e s e l l -s c h a f t e n " ( H e r t e l 1974), united i n the a l l i a n c e Racomitrion a c i c u l a r i s K r u senstjerna, are contained w i t h i n the order B r a c h y t h e c i e t a l i a plumosi, as proposed by Krusenstjerna (1945) and accepted by P h i l i p p i (1956) and Hertel (1974). This order includes a l s o the a l l i a n c e o f submerged bryophyte a s s o c i a t i o n s , the Scapanion undulatae P h i l i p p i 1956, as such not studied here. Another a s s o c i a t i o n has been recognized and proposed as a new syntaxon. I t i s the 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 rocks along 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 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 1956 and the order 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 m i s 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 of t h i s newly proposed syntaxon. The two a l l i a n c e s , Racomitrion a c i c u l a r i s and Scapanion undulatae, cont a i n species f o r which European b r y o - s o c i o l o g i s t s have erected separate a s s o c i a t i o n s ^. Although the present data show apparent r e l a t i o n -ships ( f l o r i s t i c a l l y and e c o l o g i c a l l y ) with these syntaxa, most of these species (Dichodontium pellucidum, Brachythecium plumosum, Racomitrium  a c i c u l a r i s , Scapania undulata) have been designated as a l l i a n c e or as order character species. Order character species f o r the 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 neesiana. A l l i a n c e character species f o r the.Racomitrion a c i c u l a r i s Krusenstjerna 1945 i s Racomitrium a c i c u l a r i s . The f o l l o w i n g a s s o c i a t i o n s and subassociations have been recog-nized w i t h i n the a l l i a n c e Racomitrion a c i c u l a r i s : (1) Ass. Racomitrio - Scoulerietum aquaticae ass. nov. (2) Ass. Dichodontietum p e l l u c i d i v. Hubschmann 1967 Subass. Brachythecio - Conocephaletosum conicae Subass. R i c c a r d i o - Hygrobielletosum l a x i f o l i a e Subass. B l i n d i o - Scapanietosum paludosae (3) Ass. Marsupello - Nardietum s c a l a r i s ass. nov. (4) Ass. Hypnetum d i e c k i i ass. nov. In a d d i t i o n , two.facies have been recognized 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 fontana f a c i e s and a B l i n d i a acuta - f a c i e s . no 8.3.2. Phytogeographical s e t t i n g . The phytogeographical 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 syntaxa o f the order B r a c h y t h e c i e t a l i a plumosi, 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 study f o r B r i t i s h Columbia. This i s , predomi-n 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 the o v e r a l l species composition of the syntaxa. Among the 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 of the circum-boreal element, are Scapania undulata, Pel 1 i a neesiana, Marsupella  emarginata, Jungermannia obovata and Nardia s c a l a r i s . T h e i r important c o n t r i b u t i o n to the syntaxa of t h i s order i s obviously r e f l e c t e d i n t h e i r designation as character s p e c i e s . Also H y g r o b i e l l a l a x i f o l i a i s a frequent hepatic o f stream accompanying bryophyte v e g e t a t i o n , mostly as an a s s o c i a t e , but foremost as the d i f f e r e n t i a t i n g species o f the su b a s s o c i a t i o n R i c c a r d i o -Hygrobielletosum l a x i f o l i a e . The species has a broken-circumboreal d i s -t r i b u t i o n , o c c u r r i n g i n oceanic areas i n Europe, Japan and North America and a l s o f r e q u e n t l y found i n (sub)alpine areas o f Central Europe. Among major companions are l i v e r w o r t species such as Chiloscyphus  polyanthos, P l a g i o c h i l a a s p l e n i o i d e s , Diplophyllurn a l b i c a n s and Cepjha-l o z i a b i c u s p i d a t a . Mosses, i n d i c a t i n g the predominantly circumboreal aspect o f the syntaxa w i t h i n the B r a c h y t h e c i e t a l i a plumosi are Dichodontium pellucidum, B l i n d i a acuta, Racomitrium a c i c u l a r e , Brachythecium piumosum and, to a l e s s e r degree, Pogonatum urnigerum. As western North American endemics only S c o u l e r i a aquatica and i n Rhizomnium glabrescens play a f l o r i s t i c a l l y important r o l e . Porotrichum b i g e l o v i i and Scleropodium o b t u s i f o l i u m occur only as o c c a s i o n a l l y 2 accompanying species 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 phytogeographical element only a few bryo-phyte species stand out 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 plumosi. 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 m u l t i f i d a and Blepharostoma t r i c h o p h y l l u m . A western 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 constant companion i n most a s s o c i a t i o n s and the character species f o r the Hypnetum d i e c k i i . In t h i s syntaxon, and i n the Marsupello - Nardietum s c a l a r i s , another r e p r e s e n t a t i v e of the l a t t e r phytogeographical element, the P o l y t r i -chaceous moss OTigotrichum p a r a l l e l u m occurs as an a d d i t i o n a l companion. 8.3.3.1. Order BRACHYTHECIETALIA PLUMOSI ( P h i l i p p i 1956) prov. A l l i a n c e Racomitrion a c i c u l a r i s Krusenstjerna 1945 Racomitrio - Scoulerietum aquaticae ass. hov. (Table XVI). This a s s o c i a t i o n occurs over a wide geographical range w i t h i n the study area, from Lynn Canyon to Brandywine F a l l s . I t was found most f r e q u e n t l y at Tow e l e v a t i o n s and only a few releves were taken from a l p i n e s i t e s w i t h i n the Black Tusk Meadows area. Both character 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 a c i c u l a r e , are widespread throughout B r i t i s h Columbia, i n c l u d i n g Vancouver I s l a n d and the 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 of the a s s o c i a t i o n i s l i k e l y to extend beyond the range of the present study area. Hubschmann (1978) has described a Scoulerietum aquaticae from Vancouver I s l a n d . A constant companion was 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 a l s o f r e q u e n t l y o c c u r r i n g . The short s i t e d e s c r i p t i o n s show that the author has taken h i s releves from streamside rocks which are f r e q u e n t l y inundated. Under such c o n d i t i o n s s o l i d stands of S c o u l e r i a aquatica develop with Scleropodium o b t u s i f o l i u m as a frequent a s s o c i a t e . However, f o r t h i s study the releves from bryophyte vegetations a s s o c i a t e d with streams were taken on rocks emerged f o r most part of the year or only a f f e c t e d by spray i n periods of high water l e v e l s . 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 , although sometimes becoming the dominant bryophyte s p e c i e s , was o f t e n a s s o c i a t e d w i t h major compa-nions such as Racomitrium a c i c u l a r e , Scapania s u b a l p i n a , Brachythecium 3 plumosum and.Hypnum d i e c k i i . Other order character 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 obovata, Dichodontium p e l l u - cidum, Chiloscyphus polyanthos and Scapania undulata. The s i m i l a r i t i e s i n f l o r i s t i c composition (except f o r the endemic character species S c o u l e r i a aquatica) and i n h a b i t a t , i n d i c a t e the a s s o c i a t i o n ' s synsystematic a f f i n i t i e s to the a l l i a n c e Racomitrion a c i c u l a r i s . Subsequently, as suggested by Hubschmann (1978) - who pro-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 syntaxon Table XVI. Racomitrio - Seoulerietum aquaticae ass. nov. Number of releves 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 Releve-number 6 259 261 104 105 106 107 229 224 126 127 128 210 211 227 228 212 213 Elevation(m) 30 1487 1487 30 30 30 30 610 610 457 457 457 457 457 610 610 457 457 Direction ofgexposure Inclination! ) NE S N-NE N W-SW S NE SE N SU SW SW E NE SW SW E S 48 80 10 58 22 54 50 62 26 30 22 56 24 26 66 64 12 48 Total cover(%) 95 100 90 95 100 95 100 90 100 100 90 100 100 90 90 90 100 100 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 3.4 2.4 3.4 3.4 3.4 1.2 1.2 + .2 + .2 1.4 1.4 4.5 3.4 3.4 1.4 1.4 2.4 1.4 1.4 2.4 + .4 + .4 1.4 + .4 2.4 2.4 2.4 4.3 1.2 + .1 1.4 1.4 + .4 1.4 + .4 2.2 2.2 1.4 2.4 + .4 + .4 + .4 1.4 1.4 2.2 + .2 3.4 2.4 3.4 2:4 2.4 1.4 4.5 4.5 3.4 3.4 2.4 4.5 + .2 1.2 2.3 3.3 2.3 2.2 1.4 3.4 2.4 2.4 3.4 2.4 + .4 1.4 3.4 2.4 2.4 2.4 2.4 2.4 3.4 1.4 2.4 1.2 1.2 + .4 + .4 1.4 1.4 1.4 2.2 1.4 2.4 1.4 2.4 + .2 2.4 2.4 2.4 2.3 2.3 2.4 1.2 2.2 + .1 5.5 2.4 2.4 2. ,4 + .2 3.3 2.2 2.3 2.3 1. .2 1.2 1.4 1.4 1.4 1.4 3.4 3.4 4.5- 4.5 4. .5 3.4 1.4 2.4 + , .4 1.4 1.4 2.4 3. .4 4.5 1. .4 2. .3 2.3 + .4 1.4 2.4 1.4 1.4 1.4 1.4 1.4 2.2 1.4 2.4 2.4 1.4 1, .4 2.4 2.4 1.4 2, .4 1 .4 1.2 1.2 1, .2 2. .4 2.4 2.4 2.4 2.2 + .2 1.4 1.4 \ 1.1 + .1 1.2 + .1 1.2 + .1 Table XVI (continued). Found only 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 praelonga var. praelonga, Aneura pingu  Conocephalum conicum, Ceratodon purpureus, Scapania scandica, Metzgeria  conjugata, Heterocladium macounii, Isopterygium pulchellum, Anoectangium  aestivum, Pseudoleskea patens, F i s s i d e n s bryoides, P o r e l l a r o e l l i i , Plagiomnium rostratum, Grimmia torquata. Rel eve-locations:6,104,105,106,107 259,261 224,227,228,229 126,127,128,210,211,212,213 Lynn Canyon Black Tusk Meadows Cheakamus River V a l l e y Brandywine F a l l s 115 should be c l a s s i f i e d i n the order 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. < Dichodontietum p e l l u c i d i v. Hubschmann 1967 This a s s o c i a t i o n with Dichodontium pellucidum as i t s character s p e c i e s , i s described by Hubschmann (1967) from streamside r o c k s , covered with s o i l but submerged f o r most of the year. From s i m i l a r h a b i t a t s i t has been reported and documented by Norr (1970), Neumayr (1971), M a r s t a l l e r (1973) and as a su b a s s o c i a t i o n o f the Brachythecietum plumosi Krusenstjerna 1945 by M a r s t a l l e r (1980). Hubschmann (1978) a l s o described the a s s o c i a t i o n from Vancouver I s l a n d . Frequent a s s o c i a t e s i n the Dichodontietum p e l l u c i d i i n Europe are Mnium punctatum, Conocephalum com'cum, Eurhynchium s c h w a r t z i i , Platyhypnidium r i p a r i o i d e s . Further a s s o c i a t e d s p e c i e s , c o n t r i b u t i n g to the present syntaxon both i n Europe and the Vancouver I s l a n d vegetations documented by Hubschmann (1978), are bryophytes such as Brachythecium  r i v u l a r e , Chiloscyphus polyanthos, S t o k e s i e l l a praelonga and Brachy- thecium plumosum. The present data do not contain any r e p r e s e n t a t i o n of the Dichodontietum p e l l u c i d i - "typicum". Despite i t s constancy the chara c t e r species u s u a l l y occurred with low cover v a l u e s , with the exception o f releves 90 and 92. In comparison to the type of h a b i t a t , described i n European reports on t h i s syntaxon and the 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 of the Dichodontietum 116 p e l l u c i d i i n the present treatment were taken from d i f f e r e n t micro-h a b i t a t s . These w i l l be elaborated on i n the d i s c u s s i o n s of 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 . Within the Dichodontietum p e l l u c i d i t h i s study recognizes three subassociations and two f a c i e s . Brachythecio - Conocephaletosum conicae subass. nov. (Table XVII). This s u b a s s o c i a t i o n was found at several s i t e s i n Lynn Canyon and occurs a l s o northward at Brandywine F a l l s , a t the base of Whisler Mountain and, f i n a l l y , was described from the C h i l l i w a c k River V a l l e y . I t s h a b i t a t i s bases of boulders and steep r o c k w a l l s , where, under shady and s h e l t e r e d c o n d i t i o n s , the bryophyte vegetation never becomes inundated but i s e x t e n s i v e l y a f f e c t e d by spray, e i t h e r from adjacent streams or permanent seepage. The accumulation o f sandy-loamy s o i l on the rock surface was 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 with Conocephal urn com'cum as the dominant s p e c i e s , have been described as a s s o c i a t i o n s from calcareous rocks (Herzog and H o f l e r 1944, Koppe 1955, H o f l e r 1959, Smarda 1947, Jezek and Vondracek 1962, P h i l i p p i 1965, Hebrard 1975). As such t h e i r f l o r i s t i c composition 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 that e x h i b i t e d i n the present data, which i n d i c a t e close a f f i n i t i e s to the Conocephalurn conicum - community described by P o e l t (1954), the Brachythecium r i v u l a r e - Conocephal urn conicum - community of Hagel (1966) and the Fegatelletum conicae Schade 1934, reported on Table XVII. Dichodontietum p e l l u c i d i v. Hubschmann 1967: Brachythecio-Conocephaletosum conicae subass. nov. Number of releves 1 2 3 4 5 6 7 8 9 Releve-number 86 90 88 87 89 276 329 206 208 E l e v a t i o n (m) 30 30 30 30 30 266 1219 457 457 D i r e c t i o n o f ^exposure NE E E SE E-NE W SW NE S-SW I n c l i n a t i o n ( ) 74 72 84 78 76 56 48 48 56 Total cover {%) 90 100 80 90 90 100 90 100 100 Dichodontium pellucidum 3.3 4.3 1 .2 1 .2 2.2 1.2 2.2 2.2 Scapania undulata 1.4 1.4 2.4 1.4 4.4 4.4 Conocephalurn conicum 4.5 5.5 3.4 3.4 3.4 Geocalyx g r a v e d ens 2.4 + .4 1.4 1 .4 1.4 Jungermannia pumila 2.4 1.4 2.4 2.4 S t o k e s i e l l a praelonga var. praelonga + .4 1.4 4.5 1.4 P l a g i o c h i l a a s p l e n i o i d e s + .4 + .4 2.4 2.4 Porotrichum b i g e l o v i i + .4 + .4 1.4 + .4 Brachythecium f r i g i d u m 4.5 3.4 3.4 Hypnum d i e c k i i + .1 1.4 1.4 B l i n d i a acuta + .1 1.2 + .1 R i c c a r d i a m u V t i f i d a . 2.4 1.4 Scapania subalpina 1.4 2.4 Rhizomnium glabrescens 1.1 2.2 Racomitrium a c i c u l a r e 1.2 2.3 Metzgeria conjugata + .4 2.4 Pel 1 i a neesiana + .1 + .1 Found only once: Jungermannia obovata, Cephalozia b i c u s p i d a t a , Blepharostoma tr i c h o p h y l 1 urn, Chiloscyphus polyanthos, P h i l o n o t i s fontana, 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 , Scapania americana, Scleropodium obtusum, Hygrohypnum ochraceum. Ceratodon purpureus, T r i t o m a r i a quinquedentata, Brachythecium velutinum, P o h l i a cruda, Amphidium c a l i f o r n i c u m , Cynodontium j e n n e r i , Atrichum undulatum, Mylia t a y l o r i , S t o k e s i e T l a praelonga var. s t o k e s i i , B l a s i a p u s i l l a , Lophozia sudetica Table XVII (continued). Releve - locat ions: 86. 87. 88, 89, 90 276 329 206, 208 Lynn Canyon Chi l l iwack River Vai l Whistler Mountain Brandywine Fa l l s 119 by Hiibschtnann (1967), 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 expressed by the two major d i f f e r e n t i a t i n g species 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 . Further i n d i c a t i o n i s found i n the presence of species such as Stoke-s i e l l a praelonga var. praelonga, P l a g i o c h i l a a s p l e n i o i d e s and Rhizo-mnium glabrescens, a western North American endemic. However, the releves comprising the present 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 composition i n comparison to European b r y o - s o c i o l o g i c a l s t u d i e s mentioned before. Further research may show t h i s to be i n accordance to M a r s t a l l e r ' s (1973) statement t h a t "the Fegatelletum conicae Schade 1934, due to i t s wide h a b i t a t amplitude,^has to be subdivided i n numerous su b a s s o c i a t i o n s " As encountered i n the present study t h i s c o l l e c t i o n o f releves has been given s u b a s s o c i a t i o n rank based on two main reasons: (1) Although Conocephalum conicum and Brachythecium f r i g i d u m are domi-nant taxa, the releves do not contain 'one-species-vegetations 1 but w e l l developed combinations of s p e c i e s ; (2) The c o l l e c t i o n o f releves i s not confined to one s e c t i o n o f the study area but i s widespread. The Brachythecio - Conocephaletosum conicae J s , besides i t s character s p e c i e s , f u r t h e r defined by species such as Scapania undulata, Jungermannia pumila, Geocalyx graveolens. Less frequent bryophytes 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 Porotrichum b i g e l o v i i . 120 R i c c a r d i o - Hygrobielletosum l a x i f o l i a e subass. nov. (Table X V I I I ) . This s u b a s s o c i a t i o n i s described from several s i t e s i n Lynn Canyon where i t occurred 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 shaded rocks which become o c c a s i o n a l l y inundated at high water l e v e l s r e s u l t i n g from snow melt or extremely high p r e c i p i t a t i o n . I t s m i c r o - h a b i t a t i s h o r i z o n t a l and gently s l o p i n g rock surfaces covered with only a t h i n l a y e r of deposited s i l t , c o n t a i n i n g 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 these releves were taken from one geographic a r e a , the r e c u r r i n g combination of s p e c i e s , as opposed to "one-species-communities", was 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 of t h e i r constancy and near f i d e l i t y to t h i s set of 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 designated as 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 s p e c i e s . Further constant, and sometimes h i g h l y abundant, d i f f e r e n t i a t i n g taxa a s s o c i a t e d with the two previous species are Scapania undulata, Nardia 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 Cephalozia b i c u s p i d a t a . Constant, but not d i f f e r e n t i a t i n g , companions are Dichodontium  pellucidum and B I i n d i a acuta. B l i n d i o - Scapanietosum paludosae subass. nov. (Table XIX). The present s u b a s s o c i a t i o n i s described from several s i t e s i n f r o n t of 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 occurred on h o r i z o n t a l and gently s l o p i n g boulder surfaces with only a t h i n l a y e r of accumulated Table XVIII. Dichodontietum p e l l u c i d i v. Hubschmann 1967: Riccardio-Hygrobielletosum l a x i f o l i a e subass. nov. Number of releves Releve-number Elevation(m) D i r e c t i o n ofpexposure I n c l i n a t i o n ( ) Total cover 1 2 3 4 5 6 195 196 200 197 198 199 30 30 30 30 30 30 SE S-SW SE SE SE S 48 46 24 58 70 28 100 100 100 100 90 90 Scapania undulata 4.5 4.4 4.5 3.4 3.4 3.4 H y g r o b i e l l a l a x i f o l i a 3.4 3.4 2.4 4.4 4.4 4.4 Dichodontium pellucidum 3.3 1.2 3.3 2.2 2.2 + .2 Nardia s c a l a r i s 2.4 3.4 2.4 2.4 2.4 1 .4 B l i n d i a acuta 2.3 2.3 1.2 2.2 1 .2 4.3 R i c c a r d i a m u l t i f i d a 2.4 2.4 2.4 1 .4 1.4 1 .4 Hypnum d i e c k i i 1.4 1.4 1.4 2.4 1.4' + .1 Jungermannia obovata 2.4 2.4 2.4 Cephalozia b i c u s p i d a t a 2.4 1.4 2.4 Jungermannia pumila 2.4 1.4 Oxystegus t e n u i r o s t r i s 2.2 1.2 Location of a l l r e l e v e s : Lynn Canyon. 122 ., 6 s o i l 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 s u b a s s o c i a t i o n are Scapania paludosa and B l i n d i a acuta. Other bryophytes, which, because of t h e i r near confinement to t h i s s u b a s s o c i a t i o n , are h i g h l y characte-r i s t i c f o r i t are P r e i s s i a quadrata and F i s s i d e n s osmundoides. Constant companions to the B l i n d i o - Scapanietosum paludosae are species such as Dichodontium pellucidum, Blepharostoma trichophylTurn, 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 Hygro-b i e l l a l a x i f o l i a . G e i s s l e r (1976), i n her study of the h y g r o p h i l i c bryophyte vegetation of the eastern Swiss A l p s , proposes a new a l l i a n c e , the Marsupello - Scapanion. A s s o c i a t i o n s w i t h i n t h i s syntaxon occur on s i t e s where streams o r i g i n a t e i n subalpine and a l p i n e areas. Predominant environmental f a c t o r s f o r the bryophyte vegetation are (1) the water volume and i t s low cu r r e n t v e l o c i t y and (2) the impermeability of the (mainly 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 with high moisture s t a t u s and high a i r humidity. The present s u b a s s o c i a t i o n shows apparent f l o r i s t i c r e l a t i o n s h i p s to the a s s o c i a t i o n s , B l i n d i o - Scapanietum undulatae and B l i n d i o -Scapanietum uliginosum, as described by G e i s s l e r (1976) i n the a l l i a n c e Marsupello - Scapanion. Further research might s u b s t a n t i a t e t h i s . A p a r t i c u l a r l y a l l i e d [to G e i s s l e r ' s (1976) data] bryophyte assemblage was found at 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 described by t h i s Swiss b r y o l o g i s t . Is i s represented by the releves 305 to 308, taken from a l p i n e s i t e s on W h i s t l e r Mountain. These assemblages are predomi-Tab'le XIX. Dichodontietum p e l l u c i d i v. Hubschmann 1967: BIindio-Scapanietosum paludosae ass, nov. Number of r e l e v e s 1 2 3 4 5 6 Releve-number 216 219 218 220 217 221 Elevation(m) 457 457 457 457 457 457 D i r e c t i o n ofgexposure I n c l i n a t i o n ( ) W SW W S E S 26 40 32 68 40 40 Total cover{%) 100 100 100 100 100 100 B I i n d i a acuta 4.4 4.4 4.5 3.4 4.5 4.4 Scapania paludosa 3.4 4.5 4.5 4.5 2.4 2.4 Dichodontium pellucidum 2.3 2.2 2.3 1.2 2.2 2.2 P r e i s s t a quadrata 2.4 1.4 1.4 2.4 2.4 + .1 F i s s i d e n s osmundoides 1.2 1.2 1 .2 + .1 3.3 1.2 BIepharostoma t r i c h o p h y l l u r n 1.4 1.4 + .4 1.4 1.4 1 .4 H y g r o b i e l l a l a x i f o l i a 1.4 1.4 1 .4 + .4 4.5 Aneura pinguis 1.4 + .4 + .4 3.4 2.4 1 .4 Jungermannia a t r o v i r e n s 1.4 1.4 2.4 1.4 Bryum pallescens 1.2 1 .2 + .2 2.3 1 .2 Pel 1 i a neesiana + .1 3.4 + .1 Jungermannia pumila 1.4 1.1 2.4 Rhizomnium glabrescens 1.1 Jungermannia e x s e r t i f o l i a 2.4 Scapania subalpina 1.4 Location of a l l r e l e v e s : Brandywine F a l l s 124 nantly 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 frequent companion species i n the a s s o c i a t i o n Nardietum compressae described by G e i s s l e r (1976) and c l a s s i f i e d i n the a l l i a n c e Marsupello - Scapanion. Further character species of the bryophyte vegetation of releves 305 to 308 are bryophytessuch as P h i l o n o t i s fontana, Pseudoleskea a t r i c h a , Hypnum subimponens and Scapania 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 are Racomitrium sudeticum, Lophozia s u d e t i c a , K i a e r i a s t a r k e i , Dicranoweisia  c r i s p u l a , Bartramia i t h y p h y l l a , P o h l i a l u d w i g i i , Pleuroclada albescens, Jungermannia e x s e r t i f o l i a and Scapania paludosa. The releves 305 to 308 a l s o contain 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 - Brachytheci um g l a c i a l e - v a r i a n t and the Racomitri um  sudeticum - v a r i a n t o f the a s s o c i a t i o n Dermatocarpetum riv u l o r u m as newly proposed and described by G e i s s l e r (1976). Further research, e s p e c i a l l y i n subalpine and a l p i n e areas o f western North America, may elaborate on the s t r u c t u r e and nature of these vegetations 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 with European vegetations. In a d d i t i o n to the three subassociations w i t h i n the a s s o c i a t i o n Dichodontietum p e l l u c i d i , the 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 the presence of two groups of r e l e v e s , recognized here as ' f a c i e s ' . These are bryophyte 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 'parent' a s s o c i a t i o n but are dominated by an a c c i d e n t a l combination of 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 recognized. Extremes are expressed by "one-species-communities". As such, the present data d i s t i n g u i s h a P h i l o n o t i s fontana - 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 fontana - f a c i e s . Number of releves 1 2 3 Releve-number 269 271 270 Elevation(m) 305 305 305 D i r e c t i o n of nexposure SW W W I n c l i n a t i o n ^ ) 52 38 60 Total cover(%) 100 100 100 P h i l o n o t i s fontana 4.3 4.3 3.3 Dichodontium pellucidum 2.2 3.3 4.3 R i c c a r d i a m u T t i f i d a 2.4 2.4 3.4 P o h l i a nutans 1 . 2 1 . 2 1 . 2 Cephalozia b i c u s p i d a t a +.4 +.4 +.4 Chiloscyphus polyanthos Scapania undulata +.1 +.1 3.4 Location of a l l r e l e v e s : C h i l l i w a c k Lake Table XXI. Dichodontietum p e l l u c i d i v. Hubschmann 1967: B l i n d i a acuta - f a c i e s . Number of releves 1 2 3 4 5 Releve-number 207 267 268 215 275 Elevation(m) 457 305 305 457 305 D i r e c t i o n ofgexposure I n c l i n a t i o n ( ) E SW S NW NW 104 58 32 26 32 Total cover(%) 60 90 95 100 100 B l i n d i a acuta 4.5 4.5 4.5 4.5 4.5 Dichodontium pellucidum 1.2 3.3 3.3 Bryum c a p i l l a r e 1.2 1 .2 1.2 Scapania paludosa 1 .4 + .1 1.4 Scapania undulata 1.4 2.4 P h i l o n o t i s fontana 2.2 1.2 Barbula r e f l e x a 1 .2 1.2 Scapania mucronata + .1 + .1 Radula bolanderi 3.4 P r e i s s i a quadrata 3.4 Jungermannia a t r o v i r e n s 2.4 Bryum miniatum 2.2 P e l l i a neesiana 1.1 Rhizomnium glabrescens 1.1 Chiloscyphus polyanthos 1.4 Jungermannia e x s e r t i f o l i a 1.4 Brachythecium f r i g i d u m 1.1 Aneura pinguis 1.4 Isopterygium elegans 1 .4 Isopterygium pulchellum 1.2 P l a g i o c h i l a a s p l e n i o i d e s + .1 Racomitrium a c i c u l a r e + .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 127 and a B l i n d i a acuta - f a c i e s . The f i r s t f a c i e s i s described from s o i l - c o v e r e d rocks at e x t e n s i v e , open seepage areas along C h i l l i w a c k Lake. Under these c o n d i t i o n s P h i l o n o t i s fontana was c o n s t a n t l y a s s o c i a t e d with 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 b i c u s p i d a t a . The second f a c i e s i s reported from the same area as the P h i l o n o t i s fontana - f a c i e s but recurred under the same c o n d i t i o n s at Brandywine F a l l s . Frequent companions i n t h i s B l i n d i a acuta - f a c i e s were Dichodontiurn pel 1ucidum, Scapania paludosa and Bryum c a p i l l a r e . Marsupello - Nardietum s c a l a r i s ass. nov. (Table X X I I ) . This a s s o c i a t i o n i s described from rocks adjacent to streams which do not become inundated but are a f f e c t e d only by spray at times of highest water l e v e l s . The Marsupello - Nardietum s c a l a r i s c h a r a c t e r i s -t i c a l l y occurs on h o r i z o n t a l top surfaces and gently s l o p i n g parts of rocks 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 bryophyte species were found growing on only a t h i n l a y e r o f s o i l that had accumulated on these rocks. Geographically, the present a s s o c i a t i o n i s widely d i s t r i b u t e d throughout the study area. I t i s documented from streamside rock h a b i t a t s i n Lynn Canyon, Murrin Creek and Stoney Creek canyons along the Squamish Highway, northward from the Cheakamus R i v e r banks and Brandywine F a l l s and, f i n a l l y , eastward from s i t e s along the C h i l l i w a c k R i v e r . Although 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 environment, the 128 Marsupello - Nardietum s c a l a r i s shows some a f f i n i t i e s to the Nardietum s c a l a r i s P h i l i p p i 1956, an a s s o c i a t i o n reported 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, Norr * 1969, Neumayr 1971, Dunk 1971, M a r s t a l l e r 1973, 1980). These authors a l l describe the Nardietum s c a l a r i s as a t e r r e s t r i a l a s s o c i a t i o n from open, d i s t u r b e d s i t e s where i t occurs on loamy-sandy s o i l along roads, t r a i l s and uprooted t r e e s . This syntaxon i s contained i n the a l l i a n c e Pogonation u r n i g e r i Krusenstjerna 1945 and c l a s s i f i e d i n the order D i c r a n e l l e t a l i a heteromallae P h i l i p p i 1956, an order from a c i d l y r e a c t i n g s o i l . This i s r e f l e c t e d i n the major companions, designated as order and a l l i a n c e character s p e c i e s : D i c r a n e l l a heteromalla, Ditrichum  heteromal1 urn, Atrichum undulatum, Cephalozia b i c u s p i d a t a , Isopterygiurn  elegans, Pogonatum urnigerum. In the present study, a l l these species occur e i t h e r o c c a s i o n a l l y (except 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 constant companion i n t h i s a s s o c i a t i o n ) or are not present at a l l . Present bryophyte 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 the "European" syntaxon and the Marsupello - Nardietum s c a l a r i s recog-nized i n t h i s study are: Diplophyllum a l b i c a n s , O l i gotrichum p a r a l l e l urn, 0_. al igerum and Atrichum selwyni i . Despite these a f f i n i t i e s i n f l o r i s t i c composition, two major reasons f o r the r e c o g n i t i o n of the Marsupello - Nardietum s c a l a r i s as a separate syntaxon and i t s syntaxonomic p o s i t i o n w i t h i n the Raco-m i t r i o n a c i c u l a r i s Krusenstjerna 1945 are (1) the h a b i t a t of the present syntaxon i s d e c i s i v e l y d i f f e r e n t from the ' t e r r e s t r i a l ' Nardietum s c a l a r i s P h i l i p p i 1956 and Table XXII. Marsupello - Nardietum scalaris ass. nov. Number of releves 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 Releve-number 48 115 81 203 83 344 191 192 194 202 205 76 77 111 79 84 78 92 91 193 93 94 201 204 Elevation(m) 61 61 30 610 30 61 457 457 457 610 610 266 266 30 30 30 30 30 30 457 30 30 610 610 Direction ofgexposure NW N-NW NE SE N N-NW S-SE S-SW S SE S-SE NE NE NW NW NE NE W NE S NE SW SE E-SE Inclination! ) 12 82 52 62 34 32 52 48 68 44 82 18 26 50 28 64 36 62 62 50 30 52 82 68 Total cover(%) 90 80 95 100 100 80 90 100 100 100 100 95 90 100 100 .100 100 95 95 95 100 80 100 100 Nardia scalaris 4.4 3.4 3.4 3.4 5.5 2.4 3.4 3.4 2.4 2.4 2.4 2.4 + .4 1.4 2.4 4.5 3.4 4.4 4.5 3.4 2.4 3.4 2.4 Marsupella emarginata 2.4 1.4 1.4 2.4 2.4 2.4 3.4 1.4 1.4 4.5 5.5 3.4 4.4 3.4 2.4 3.4 3.4 2.4 4.5 4.5 2.4 2.4 Hypnum dieckii 1.4 2.4 2.4 3.4 3.4 2.4 4.5 4.4 4.5 1.4 3.4 1.4 2.4 + .4 + .4 1.4 + .4 2.4 2.4 + .4 Rhizomnium glabrescens 1.2 1.2 + .1 1.2 2.2 2.3 3.3 2.2 1.2 1.2 2.3 2.2 3.3 3.3 2.2 2.3 + .1 1.2 1.2 Cephalozia bicuspidata 1.4 2.4 + .4 2.4 1.4 + .4 1.4 2.4 2.4 2.4 1.4 2.4 + .4 1.4 1.4 + .4 1.4 2.4 Diplophyllum albicans 1.4 2.4 1.4 + .4 2.4 1.4 1.4 1.4 1.4 1.4 1.4 2.4 + .4 + .4 3.4 2.4 Pellia neesiana 2.4 +.1 1.4 1.4 1.4 + .1 1.4 1.4 1.4 + .1 1.4 + .1 + .1 Scapania undulata + .1 2.4 2.4 3.4 + .1 2.4 1.4 2.4 1.4 Jungermannia obovata 1.4 2.4 2.4 2.4 2.4 2.4 2.4 2.4 1.4 1.4 2.4 Dichodontium pelluddum 2.3 2.3 1.2 2.3 2.2 2.3 + .1 + .1 1.2 Blindia acuta 1.2 1.2 1.2 1.2 1.2 1.2 2.3 1.2 Oligotrichum parallelum + .1 3.3 1.2 1.2 4.3 + .1 + .1 1.2 Hygrobiella laxifolla 1.4 1.4 1.4 1.4 1.4 1.4 2.4 Racomitrium aciculare 2.3 2.3 2.3 2.3 + .2 2.3 Pogonatum urnlgerum + .1 1.2 + .1 + .1 Atrichum selwynii + .1 3.3 1.2 Oligotrichum allgerum 2.2 1.2 1.2 Anastrophyllum minutum 1.4 2.4 1.4 Scapania subalplna 3.4 1.4 + .4 Brachythecium frigidum 2.4 1.4 Scapania americana 2.4 1.4 Riccardia mul'tifida + .1 1.4 Found only once: Brachythecium plumosum, Plagiochila asplenioides, Philonotis fontana, Pohlia nutans, Diplophyl1um  taxifolium. Isopterygium elegans , Oxystegus tenuirostris, Diplophyllum piicatum, Heterocladium macounli, Bartramiopsis lescuri i, "Racomitrium brevipes", Ditrichum heteromallum, Pohlia longebracteata, Jamesoniella autumnal is, Philonotis arnelli i Releve-locations: 48,344 Shannon Falls 115 Murrin Creek 78,79,81,83,84,91.92,93,94 Lynn Canyon 201,202.203,204,205 Cheakamus River Valley 191 ,192,193,194 Brandywine Falls 111 Stoney Creek 76,77 Chilliwack River Valley 130 (2) the presence and frequency of a l l i a n c e character species as major companions. Because of t h e i r constancy and abundance, Nardia s c a l a r i s and Marsupella emarginata are designated as the character species o f the present 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 , with major occurrence i n t h i s syntaxon i n c l u d e Cephalozia b i c u s p i d a t a , Rhizomnium  glabrescens, Diplophyllum a l b i c a n s and Oligotrichum p a r a l l e l u m . Order and a l l i a n c e character species as s i g n i f i c a n t a s s o c i a t e s are Scapania undulata, P e l l i a neesiana, Dichdontium pellucidum, 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 ass. nov. (Table X X I I I ) . The a s s o c i a t i o n i s reported and documented from Lynn Canyon, i n the southern p o r t i o n of the study area, to Brandywine F a l l s i n the north, where the syntaxon i s a l s o described from two subalpine s i t e s i n the Black Tusk Meadows area. I t s h a b i t a t i s h o r i z o n t a l to s l i g h t l y s l o p i n g rock surfaces with mineral s o i l accumulation up to 1 cm i n thickness ( i n only a few instances t h i s was measured to be almost 2 cm t h i c k ! ) . These s u b s t r a t a f o r the a s s o c i a t i o n occurred under shaded and s h e l t e r e d c o n d i t i o n s as we l l as i n open and exposed s i t e s adjacent to sreams. The character species f o r the Hypnetum d i e c k i i i s Hypnum d i e c k i i , a western North American - Japan d i s j u n c t bryophyte taxon. Most commonly 131 i t was accompanied by Marsupella emarginata, a species which occurs considerably l e s s f r e q u e n t l y i n the recognized 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 are shared by both syntaxa 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 the a l l i a n c e Racomitrion a c i c u l a r i s are Dichodontium pellucidum, Pel 1ia neesiana, Rhizomniurn glabrescens and Brachytheciurn piumosum. However, bryophyte species which d i f f e r e n t i a t e the Hypnetum d i e c k i i - "typicum" (1 i n Table XXIII) from the Scapanio - Jungermannie-tosum obovatae, and, thus, c h a r a c t e r i z e t h i s syntaxon are Scapania  s u b a l p i n a , BIepharostoma t r i c h o p h y l l u m , Pogonatum urnigerum, S c l e r o - podium o b t u s i f o l i u m , Diplophyllum t a x i f o l i urn and Oligotrichum p a r a l l e l urn. Scapanio - Jungermannietosum obovatae subass. nov. Occurring at 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 reported and documented from streamside rocks i n Lynn Canyon, at Shannon F a l l s and at Brandywine F a l l s . C h a r a c t e r i s t i c a l l y , the Scapanio - Jungermannietosum obovatae (2 i n Table XXIII) grows on gently s l o p i n g rock surfaces under shaded and s h e l t e r e d c o n d i t i o n s , only a f f e c t e d by d i r e c t m o i s ture-input, through spray, at times of highest water l e v e l s . Only a t h i n l a y e r of s o i l or sometimes no accumulated material at a l l was found on the rock s u r f a c e . The abundance and constancy of Hypnum d i e c k i i i n d i c a t e the present syntaxon's a f f i n i t y to the Hypnetum d i e c k i i - typicum. The major 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 s u b a s s o c i a t i o n Table XXIII. Hypnetum dieckii ass, nov. Number of releves 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 Rel eve-number 52 53 56 130 85 190 250 251 186 189 108 184 109 110 209 183 185 187 188 Elevation(m) 61 61 61 457 457 30 1487 1487 30 30 30 30 61 61 457 30 30 457 457 Direction ofQexposure Inclination( ) N N E SW W NW E E-SE NW W W-NW NW W-SW S-SW E-SE W N-NW W NW 22 24 30 24 20 78 40 26 76 82 48 62 32 36 66 70 34 58 48 Total cover(X) 100 100 90 90 100 100 100 100 90 80 100 100 95 100 100 100 100 80 100 Hypnum dieckii 4.5 3.4 2.4 2.4 4.5 2.4 4.5 5.5 2.4 4.5 4.5 3.4 3.4 2.4 2.4 2.4 2.4 Dichodontium pelluddum + .1 2.2 2.2 1.2 2.2 + .1 1.2 2.3 2.3 1.2 3.3 Pellia neesiana 1.4 2.4 2.4 + .1 2.4 1.4 2.4 2.4 2.4 Brachythecium plumosum 1.4 1.4 1.4 1.4 1.4 2.4 3.5 1.4 Rhizomnium glabrescens 2.2 1.2 1.2 1.2 1.2 1.2 2.2 1.2 Cephalozia bicuspidata + .4 1.4 1.4 1.4 1.4 Diplophyllum albicans 2.4 2.4 2.4 2.4 Drepanocladus uncinatus 2.4 1.4 1.4 Blindia acuta 1.2 1.2 1.2 Philonotis fontana 1.2 2.2 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 blytt i i Platydictya jungermmanioides Bartramia ithyphylla Tritomaria quinquedentata Jungermannia obovata Scapania undulata Riccardla multifida Stokesiella praelonga var. praelonga Hypnum subimponens Porotrichum blgelovii Pleuroclada albescens Hygroblella laxifolla Scouleria aquatica Geocalyx graveolens 3.4 + .1 3.4 4.5 + .4 + .1 3.4 3.4 2.4 + .4 2.4 1.4 +.1 1.2 2.4 2.4 1.4 1.1 1.4 2.4 1.4 + .4 1.4 2.2 1.2 2.4 1.4 2.4 2.4 4.5 2.2 2.4 1.4 + .1 1.2 1.4 1.2 + .1 1.2 1.4 1.2 2.4 2.4 2.4 2.4 1.1 2.4 2.4 1.4 2.4 1.4 +.4 C O ro Table XXIII (continued). Found only once: 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, Atrichum s e l w y n i i , 01 igotrichum aligerum, Racomitrium aquaticum, Polytrichum alpinum var. macounii, Racomitrium f a s c i c u l a r e , Scapania scandica, Isopterygium elegans, Lophocolea h e t e r o p h y l l a , Diplophyllum plicatum, 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 , Plagiothecium roeseanum, P o r e l l a cordaeana, D i c r a n e l l a heteromalla, 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 , Plagiothecium laetum, Rhytidiadelphus squarrosus. R e l e v e - l o c a t i o n s : 52,53,56,109,110 Shannon F a l l s 85,130,187,188,209 Brandywine F a l l s 108,183,184,185,186,189,190 Lynn Canyon 250,251 Black Tusk Meadows 134 are Jungermannia obovata and Scapania undulata. Additonal d i f f e r e n t i a -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 var. praelonga, Porotrichum b i g e l o v i i , Pleuroclada albescens and Hypnum  subimponens. 8.3.3.2. Order PLATYHYPNIDIETALIA RUSCIFORMIS ( P h i l i p p i 1956) prov. 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 Hygrohypnetum s m i t h i i ass. nov. (Table XXIV). An e v a l u a t i o n of the synsystematics of the "aquatic bryophyte communities" i n d i c a t e s P h i l i p p i ' s (1956) c l a s s i f i c a t i o n to be the best 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 of stream adjacent bryophyte vegetation i n the study area. This system emphasizes a d i v i s i o n on b a s i s of the h a b i t a t , rather than the f l o r i s t i c composition which shows considerable overlap between bryophyte assemblages w i t h species o f wide e c o l o g i c a l amplitude ^ . The newly described a s s o c i a t i o n Hygrohypnetum s m i t h i i i s reported and described from several a l p i n e l o c a l i t i e s w i t h i n Garbaldi P r o v i n c i a l Park. I t occurred on s o i l - c o v e r e d rocks d i r e c t l y along f a s t f l o w i n g streams t h a t descend from l a t e snow areas. The s u b s t r a t e i s c o n s t a n t l y under i n f l u e n c e of spray or i s o c c a s i o n a l l y inundated. A s s o c i a t i o n s from t h i s type of environment are c l a s s i f i e d by P h i l i p p i (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 contained i n the order 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 Hertel (1974), d e s c r i b i n g and proposing a new a s s o c i a t i o n , the Hygrohypnetum o c h r a c e i , and P h i l i p p i (1965), r e p o r t i n g on a Hygrohypnum ochraceum -F i s s i d e n s c r assipes var. r u f i p e s - community, c l a s s i f y these syntaxa i n g the a l l i a n c e P latyhypnidion r u s c i f o r m i s P h i l i p p i 1956 . However, G e i s s l e r (1976), employing K r a j i n a ' s (1933) c l a s s i f i -c a t i o n , c l a s s i f i e s a newly proposed a s s o c i a t i o n , the Solenostomo -Hygrohypnetum, i n t o the a l l i a n c e Hygrohypniori d i l a t a t i K r a j i n a 1933, which i s contained i n the order Hygrohypnetalia d i l a t a t i K r a j i n a 1933. I t i s t h i s a s s o c i a t i o n which shows considerable 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 to the a s s o c i a t i o n Hygrohypnetum s m i t h i i , as described i n t h i s study. The character species of G e i s s l e r ' s (1976) Solenostomo -Hygrohypnetum are Solenostoma c o r d i f o l i u m (=Jungermannia c o r d i f o l i a ) , Scapania undulata and Hygrohypnum s m i t h i i . The a s s o c i a t i o n character species of the Hygrohypnetum s m i t h i i i n the present study are 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 Chiloscyphus polyanthos and Pel 1i a neesiana. The f o l l o w i n g bryophyte taxa i n d i c a t e additonal a f f i n i t i e s , w i t h respect to f l o r i s t i c composition, s p e c i e s ' abundance and frequency, between the present a s s o c i a t i o n Hygrohypnetum s m i t h i i and the Solenostomo -Hygrohypnetum, as described 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 fontana, Jungermannia obovata, Dicho- dontium pel 1ucidum and Bryum pseudotriquetrum. Further research, 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 at subalpine Table XXIV. Hygrohypnetum s m i t h i i ass. nov. Number of rel e v e s Releve-number Elevation(m) D i r e c t i o n ofgexposure I n c l i n a t i o n ( ) Total cover(%) 1 2 3 4 5 6 232 260 234 327 328 233 1487 1492 1487 2286 2286 1487 SW E NW N-NE NE NW 18 56 38 68 84 22 100 100 100 100 100 100 Hygrohypnum s m i t h i i Chiloscyphus polyanthos Jungermannia e x s e r t i f o l i a Pel 1ia neesiana Scapania subalpina Hygrohypnum ochraceum Blepharostoma t r i c h o p h y l l u m S c o u l e r i a aquatica Scapania undulata Bryum pal l e s c e n s P h i l o n o t i s fontana Dichodontium pellucidum Jungermannia obovata Cephalozia b i c u s p i d a t a Brachythecium plumosum H y g r o b i e l l a l a x i f o l i a Jungermannia pumila Bryum pseudotriquetrum Conocephalum conicum Drepanocladus uncinatus 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 4.5 4.5 2.4 4.5 3.4 2.4 1.4 1.4 1.4 2.4 2.4 2.4 + .1 2.4 1.4 1.4 1.4 1 .4 1.4 + .1 1.4 + .4 1 .4 + .4 + .4 3.4 2.4 2.4 3.4 2.4 1.4 2.4 1.4 1.4 2.4 1.4 + .1 1.4 1 .4 1 .4 1 .4 1.4 3.3 2.2 2.2 1.2, 1.2 2.2 1 .2 1.2 2.2 1.4 1.4 1.4 1.4 1 .4 + .1 + .4 1.4 1.4 + .4 + .1 1.2 1.1 2.4 4.4 1.2 2.2 Re l e v e - l o c a t i o n : 232, 233, 234 Black Tusk Meadows 260 Panorama Ridge 327, 328 W h i s t l e r Mountain 137 and a l p i n e vegetations might shed more l i g h t on the i d e n t i t y of t h i s vegetation type and i t s r e l a t i o n s h i p to 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 to European syntaxa i n the same environment. 1. i . e . w i t h i n the Racomitrion a c i c u l a r i s : Dichodontietum 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), Brachythecietum plumosi (Krusenstjerna 1945, Norr 1970, Hertel 1974, M a r s t a l l e r 1973, 1980), Racomitrietum 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 reported from Vancouver Island by Hubschmann (1978). Within the Scapanion undulatae: Scapanietum undulatae (Schwickenrath 1944, P h i l i p p i 1956, Norr 1969, Lecointe and Provost 1970, Dunk 1971, M a r s t a l l e r 1980). 2. Hubschmann (1978), studying the bryophyte vegetation on Vancouver I s l a n d , reports and describes the Scoulerietum aquaticae i n which S c o u l e r i a aquatica i s the a s s o c i a t i o n character species and S c l e r o - podium o b t u s i f o l i u m i s designated as the a l l i a n c e character species f o r the ( p r o v i s i o n a l ) 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 ) , although standing as an apparent u n i t i n the c l u s t e r a n a l y s i s (Appendix V ) , i s considered as a "fragment" of the Racomitrio - Scoulerietum aquaticae. This i s because o f the t o t a l absence of S c o u l e r i a aquatica and the lower frequency and constancy of Brachytheciurn plumosum. Further research 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 the a s s o c i a t i o n Racomitrietum 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) report on an a s s o c i a t i o n , P e l l i o -Conocephaletum conicae, synonymous to the Fegatelletum conicae Schade 1934. In 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 fabbroniana or Conocephalurn  conicum are the dominant species. The companion species are charac-t e r i s t i c f o r both syntaxa: Mnium punctatum, F i s s i d e n s adianthoides 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. Also Hertel (1974) discusses the s t a t u s of t h i s syntaxon and s t a t e s that the "Fegatelletum" does not deserve the rank o f a s s o c i a t i o n . 138 Conocephalum conicum occurs a f t e r the d e p o s i t i o n of some mineral s o i l as a r e s u l t of submergence. Because of i t s r a p i d and vigorous growth the species can outcompete pleurocarpous mosses (Brachy- thecium spp. and S t o k e s i e l l a spp.). Eventually i t can become dominant and develop i n t o a "one-species-community" (Neumayr 1971). Hertel (1974) f u r t h e r s t a t e s that Conocephalum conicum can become ' f a c i e s ' - b u i l d i n g w i t h i n several communities which proove the species n e i t h e r to be f a i t h f u l nor to be constant. Another b a s i c requirement to a character species i s i t s confinement to a narrow e c o l o g i c a l amplitude. In obvious c o n t r a s t Hertel (1974) reports Conocephalum  conicum dominating on humid to wet s o i l , rock bases adjacent and d i r e c t above the water surface. The species i s , according to Boros (1968) i n d i f f e r e n t ; "Apinis (1939) reports on a pH-range between s i t e s of t h i s s p e c i e s ' occurrence from 5.0 to 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 ubiquitous s p e c i e s , should be designated as ' f a c i e s ' . 6. Under extensive spray-zone c o n d i t i o n s M u l l e r (1938) described an a s s o c i a t i o n Scapanietum paludosae i n which, besides vascular p l a n t s p e c i e s , the f o l l o w i n g mosses occurred: P h i l o n o t i s s e r i a t a , Drepano-cladus exanulatus and D i c r a n e l l a squarrosa. 7. K r a j i n a (1933) erected the order Hygrohypnetalia which was to contain bryophyte 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 of the streams. However, as Hertel (1974) r i g h t l y s t a t e s , t h i s order includes groups of 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 inundated 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 waters and streams over s i l i c e o u s rocks. This c l a s s i f i c a t i o n was a l s o included i n Hubschmann's (1957) o v e r a l l synsystematic c l a s s i f i c a t i o n o f the "aquatic bryophyte 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 long-l a s t i n g emerged and almost c o n s t a n t l y submerged bryophyte 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 obvious 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 include only o c c a s i o n a l l y emerged bryophyte a s s o c i a t i o n s , yet 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. Further reports on Hygrohypnum-dominated a s s o c i a t i o n s and subassocia-t i o n s are provided by Herzog (1943), P o e l t (1954), Hagel (1966), Neumayr (1971), P h i l i p p i (1973), Hubschmann (1973, 1974), G e i s s l e r (1976), Mamczarz (1978) and M a r s t a l l e r (1980). 139 8.4 BRYOPHYTE ASSOCIATIONS FROM CALCAREOUS ROCKS. Table XXV. Bryophyte a s s o c i a t i o n s from calcareous rocks. A l l i a n c e ENCALYPTION PROCERAE a l l . nov. C l u s t e r 1. A s s o c i a t i o n Lophozio - Bryoerythrophylletum r e c u r v i r o s t r i ass. nov. C l u s t e r 2. A s s o c i a t i o n Metaneckeretum menziesii ass. nov. Barbula v i n e a l i s - P I a t y d i c t y a jungermannioides - Jungermannia a t r o v i r e n s - community C l u s t e r 3A. " i n s i d e - g r o t t o - t y p e " with D i s t i c h i u m capillaceum and Fissidens bryoides C l u s t e r 3B. " o u t s i d e - g r o t t o - t y p e " with S c h i s t i d i u m r i v u l a r e and Bryum pseudotriquetrum 141 8.4. Bryophyte a s s o c i a t i o n s from calcareous rocks (Table XXV). 8.4.1. Synsystematics. The substrate 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;" calcareous rocks ^. The d e c i s i v e d i s t i n c t i o n between calcareous and s i l i c e o u s rock bryophyte communities focuses on the chemical composition o f the rock surface and/or the t h i n l a y e r of d i s i n t e g r a t e d rock m a t e r i a l accumulated om that surface (Wirth 1972, Hertel 1974). Concerning the nature o f bryophyte communities on calcareous rocks i t i s e s s e n t i a l to r e a l i z e t h a t these are r e f e r r e d to as combi-nations of species which occur predominantly on limestone and show t h e i r optimal development on t h a t substratum. When bryophyte assemblages are described at the border area or even outside t h e i r e c o l o g i c a l range, fragments or l o c a l r epresentations can r e c e i v e a s s o c i a t i o n rank. These "communities" are then not recognized according to the Braun-Blanquet 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 of separate species. The taxa become a s s o c i a t i o n character species although most of 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 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 attempts and proposals by various b r y o - s o c i o l o g i s t s , Hertel (1974) provides a s a t i s f y i n g synsystematic h i e r a r c h y f o r the bryophyte communities on 142 calcareous rock s u b s t r a t a . Based on the f l o r i s t i c composition of the 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 the author d i s -t i n g u i s h e s (1) an order from exposed, dry rocks [ S c h i t i d i e t a l i a apocarpi (Jezek and Vondracek 1962) Hertel 1974], (2) an order from s h e l t e r e d , humid rocks ( N e c k e r e t a l i a complanatae Jezek and Vondracek 1962 and (3) an order from more o r l e s s s h e l t e r e d rocks and/or s o i l ( C t e n i d i e t a l i a m o l l u s c i Hadac and Smarda 1944). Subsequently, the a s s o c i a t i o n s w i t h i n these orders are c l a s s i f i e d i n t o the a l l i a n c e s S c h i s t i d i o n apocarpi Jezek and Vondracek 1962, Neckerion complanatae Hadac and Smarda 1944 2 and C t e n i d i o n m o l l u s c i Stefureac 1941, r e s p e c t i v e l y . The present data show 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 to a s s o c i a t i o n s described 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 treatments by Hertel (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 at the l e v e l o f a l l i a n c e and order were not d e t e c t a b l e . In a d d i t i o n , the 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 America, and the r e l a t i v e l y low number of releves per recognized a s s o c i a t i o n , have con t r i b u t e d to the present treatment of the data. Where p o s s i b l y suppor-ted with character s p e c i e s , new syntaxa are proposed and a f f i n i t i e s with vegetations i n other regions i n the northern Hemisphere are i n d i c a t e d . A new a l l i a n c e i s proposed, the Encalyption procerae, best c h a r a c t e r i z e d by Encalypta procera, Homalothecium fulgescens and P o r e l l a  cordaeana. Within 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 described: (A) a Lophozio - Bryoerythrophylletum r e c u r v i r o s t r i with the character 143 species Bryoerythrophyl1um r e c u r v i r o s t r u m , Lophozia heterocolpos and Gymnostomum rec u r v i r o s t r u m and (B) a Metaneckeretum m e n z i e s i i with the character species Metaneckera m e n z i e s i i . F i n a l l y , a t h i r d "vegetation type" i s des c r i b e d , which i s c h a r a c t e r i z e d by species such as Barbula v i n e a l i s , P l a t y d i c t y a junger- mannioides and Jungermannia a t r o v i r e n s . However, the rel e v e s o f t h i s 'community' l a c k constant (and abundant) and f a i t h f u l s p e c i e s , hence, cannot be given any syntaxonomic status and be f u r t h e r c l a s s i f i e d . 8.4.2. Phytogeographical s e t t i n g . The c o n t r i b u t i o n o f the circumboreal element to the f l o r i s t i c composition of the limestone releves i s expressed by a s i g n i f i c a n t number of species. Among the hepatics are Conocephalum conicum, Jungermannia a t r o v i r e n s , Lophozia heterocolpos and P l a g i o c h i l a a s p l e n i o i - des. The mosses Bryoerythrophyl!um r e c u r v i r o s t r u m , Plagiopus oederi and to 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 the circumboreal aspect of the bryophyte vegetation on calcareous rocks i n the study area. A predominantly' 1imestone h e p a t i c , circumboreal i n i t s d i s t r i -b u t i o n , but missing i n eastern A s i a , i s Lophozia g i l l m a n i i . Except f o r Homalothecium fu l g e s c e n s , which i s designated as one of the character species f o r the a l l i a n c e Encalyption procerae, the c o n t r i b u t i o n of western North American endemics to the o v e r a l l lime-stone bryophyte vegetation i s 'expressed by l e s s frequent companions. 144 Among these species are P o r e l l a r o e l l i i , Brachythecium f r i q i d u m , Isothecium s t o l o n i f e r u m and to a l e s s e r extent Leucolepis menziesi i , Plagiomnium venustum, Claopodium bolanderi and Porotrichum b i g e l o v i i . The d i s j u n c t phytogeographical elements play only 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 to the f l o r a and vegetation of the calcareous rocks i n the study area. The western North American - western 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 the western North American -Japan d i s j u n c t moss, Hypnum subimponens, are frequent companion species i n the a l l i a n c e Encalyption procerae. B i p o l a r d i s j u n c t bryophytes among the limestone f l o r a here i n c l u d e species such as Ditrichum  f l e x i c a u l e and Brachythecium - veltitinum. 8.4.3. A l l i a n c e Encalyption procerae a l l . nov. Lophozio - Bryoerythrophylletum r e c u r v i r o s t r i ass. nov. (Table XXVI). This a s s o c i a t i o n i s described from exposed calcareous r o c k 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 l a y e r . Depending on the s/lope o f the rock surface where the sampling was done, t h i s s o i l accumulation was measured from 0.5 to 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 character species are Bryoerythrophyl1 urn  r e c u r v i r o s t r u m and Lophozia heterocolpos. A l l i a n c e character species which are 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 with these two taxa are EncaTypta procera and Homalothecium fulgescens. A d d i t i o n a l s i g n i f i c a n t a ssociates are Plagiopus oederi and Conocephalurn Table XXVI. Lophozio-Bryoerythrophylletum r e c u r v i r o s t r i ass. nov. Number of rel e v e s Releve-number Elevation(m) D i r e c t i o n ofgexposure I n c l i n a t i o n ( ) Total cover(%) 1 2 3 4 36 163 164 165 266 266 258 240 W W-NW W-NW W-NW 58 38 38 36 100 100 100 100 5 6 7 8 153 166 167 168 266 266 258 258 W N NE NE 39 74 60 68 100 60 90 75 Bryoerythrophyllum r e c u r v i r o s t r u m 2, .3 4 .3 3.3 2, .3 2, .3 2.3 2.3 2.3 Encalypta procera 2, .2 3, .2 3.2 4, .3 2, .2 1.2 3.2 Plagiopus oederi 2. .2 2.2 2, .2 2, .2 2.2 1 .2 1 .2 Homalothecium fulgescens 1, .4 2.4 2. .4 2, .4 2.4 2.4 1.4 Conocephalurn conicum 2, .4 3.4 1, .4 1.4 1.4 Lophozia heterocolpos 1, .4 2, .4 2.4 3. .4 2.4 Racomitrium heterostichum var. heterostichum 1 , .2 1.2 1, .2 3. .3 Hypnum subimponens 1 , .4 1.4 1, .4 1. .4 P o r e l l a cordaeana 2. .4 2.4 3, .4 Ditrichum f l e x i c a u l e 2.2 3.3 2.2 Gymnostomum r e c u r v i rostrum 3.2 4.3 3.2 Brachythecium f r i g i d u m 2.4 2.4 1 .4 Mm* urn thorns on i i + , .1 2.2 1 .2 Geocalyx g r a v e d ens 2.4 1.4 Barbula r e f l e x a 2.2 1.2 S e l i g e r i a donniana 1.1 2.1 Jungermannia a t r o v i r e n s 3. ,4 1.4 Found only 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 pellucidum, 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 var. falcatum, Tritomaria e x s e c t i f o r m i s , Thamnobryum neckeroides, Lophozia g i l l m a n i i , Cephalozia 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 Sl e s s e Creek; others: C h i l l i w a c k River V a l l e y . 146 conicum. O c c a s i o n a l l y , the predominantly "limestone bryophytes" S e l i g e r i a donniana and Jungermannia a t r o v i r e n s , occur w i t h i n the present a s s o c i a t i o n . The Lophozio - Bryoerythrophyl!etum r e c u r v i r o s t r i occurs w i t h i n the present data set with two "types", of which one occurs on rocks, hardly 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 species such as Racomitrium heterostichum var. a f f i n e and Hypnum subimponens. The second "type" i s a group of species found on rocks with extensive s o i l accumulation. This combination o f species contains 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 , Mnium  thomsonii, Geocalyx graveolens and Barbula r e f l e x a . However, the present data f a i l to i n d i c a t e n e i t h e r the b r y o s o c i o l o g i c a l s i g n i f i c a n c e nor the syntaxonomic r e l a t i o n of the vegetation "types" to the synsystematic treatments of calcareous rock bryophyte communities of Hertel (1974) and M a r s t a l l e r (1979, 1980) 3. Metaneckeretum me n z i e s i i ass. nov. (Table XXVII). The present syntaxon has been described from s h e l t e r e d and shaded limestone boulders i n s i d e f o r e s t s on the slopes of the C h i l l i -wack River V a l l e y near SI esse Creek adn Chipmunk Creek. Except f o r a few occasions, most o f the rock surfaces d i d not have much s o i l mate-r i a l accumulated on them. T h i s , o b v i o u s l y , depends on the i n c l i n a t i o n of these 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 phytosocio-l o g i c a l t a b l e . Table, XXVII. Metaneckeretum menziesii ass. nov. Number of re l e v e s Releve-number Elevation(m) D i r e c t i o n ofgexposure I n c l i n a t i o n ( ) Total cover(%) 1 2 3 4 5 6 7 8 9 10 11 12 143 144 149 145 148 150 152 151 • 321 324 322 323 266 266 258 230 258 258 230 230 266 266 268 268 NW NW W-NW NW NW SW SW SW W W NW W-NW 64 62 82 58 58 84 64 64 82 60 78 76 100 100 100 100 100 100 100 90 100 100 100 100 Encalypta procera 2.2 2.2 1.2 1.2 2.2 2.2 2.2 Metaneckera m e n z i e s i i 3.4 3.4 3.4 3.4 Plagiopus oederi 2.2 2.2 2.2 2.2 Homalothecium fulgescens 4.4 1.4 1.4 1.4 1.4 Eurhynchium pulchellum 2.4 2.4 +.4 Hypnum subimponens 2.4 2.4 1.4 Isothecium s t o l o n i f e r u m 2.4 2.4 1.4 Dichodontium pellucidum 2.2 3.2 Claopodium bolanderi 2.4 2.4 Porotrichum b i g e l o v i i 2.4 +.4 Gymnostomum aeruginosum 1.2 3.2 2.2 1.2 Barbula v i n e a l i s 4.3 +.2 2.2 P l a t y d i c t y a jungermmanioides 2.4 2.4 1.4 Po h l i a cruda 2.2 4.3 Blepharostoma trichophyl1um 2.4 4.4 Plagiomnium venustum 4.3 1.2 P o r e l l a r o e l l i i 3.4 Claopodium c r i s p i l u m 2.4 Bryoerythrophyllum r e c u r v i r o s t r u m Mnium ambiguum Apometzgeria pubescens Barbula r e f l e x a Plagiomnium rostratum 1.2 1 .2 + .2 1.2 1.4 3.4 1.4 1.4 3.2 1.2 1.2 2.2 1.4 1.4 1.4 + .4 1.4 1.4 2.4 + .4 + .4 2.4 2,. 4 1.4 2.4 1.2 2.2 3.2 3.3 + .2 1.2 1 .2 1 .2 1.4 2.4 1.2 1 .2 1.2 1.2 Table XXVII (continued). Found only once: 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 heterostichum var. a f f i n e , Bryum c a p i l l a r e , Conocephalum conicum, Mnium thomsonii, P o r e l l a cordaeana, Isopterygium elegans, 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 praelonga var. praelonga, 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 : 143,144,145,148,149 150,151,152 321,322,323,324 C h i l l i w a c k River Chipmunk Creek Slesse Creek V a l l e y 149 The a s s o c i a t i o n ' s character species 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 associated with the a l l i a n c e c h a r a c t e r species Encalypta procera, and Homalothecium fulgescens. Other s i g n i f i c a n t accompanying species are Plagiopus o e d e r i , Eurhynchium pulchel 1 urn and Hypnum subimponens. Within the Metaneckeretum m e n z i e s i i two subassociations have been d i s t i n g u i s h e d , based on species composition and h a b i t a t d i f f e r e n c e s . The subassociations Gymnostometosum aeruginosum occurs on s t e e p l y s l o p i n g , dry rocks surfaces ( 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, Barbula v i n e a l i s and P l a t y d i c t y a jungermannioides. A second s u b a s s o c i a t i o n , the Porelletosum r o e l l i i grows on gently as w e l l as s t e e p l y s l o p i n g , humid rock surfaces where i t s lush vegetation 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 species 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 present a s s o c i a t i o n Metaneckeretum m e n z i e s i i and European bryophyte syntaxa, the present data do not allow i n t e g r a t i o n i n the synsystematic h i e r a r c h y of bryophyte communities on calcareous rocks as c l a s s i f i e d by Hertel. (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 type o f bryophyte assemblage has been de s c r i b e d : the Barbula 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 XXVIII). This c h a r a c t e r i s t i c combination of species (extended with the t h a l l o s e hepatic Conocephalurn conicum) Table XXVIII Barbula 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. Number of releves 1 2 3 4 5 6 7 8 9 10 11 Rel eve-number 146 147 280 281 282 277 278 279 283 284 285 El evation(m) 266 266 258 258 258 240 246 246 262 268 250 D i r e c t i o n ofgexposure I n c l i n a t i o n ( ) NW NW NW N-NW N-NW NE E NW M-NE NE N n o 96 60 48 52 58 66 54 54 42 18 Total cover(%) 40 60 50 60 50 90 100 90 95 100 100 Conocephalum conicum 1 .4 1 .4 1.4 1.4 2.4 2.4 3.4 3.4 1.4 1.4 Barbula v i n e a l i s 3.2 1.2 1 .2 1.2 + .2 1.2 1 .2 1 .2 + .2 Leucolepis m e n z i e s i i 1 .2 1 .2 + .2 1.4 1.4 3.3 2.2 P l a t y d i c t y a jungermannioides 2.4 1.4 2.4 1.4 Porel1 a r o e l I i i 1 .4 1 .4 2.4 1.4 1 .4 S c h i s t i d i u m r i v u l a r e 2.2 1 .2 1 .2 1 .2 3.2 Bryum pseudotriquetrum 1.2 1 .2 + .2 + .2 4.3 P l a g i o c h i l a a s p l e n i o i d e s 2.4 1.4 1.4 1.4 Racomitrium canescens 3.3 1 .2 1 .2 + .2 Bryoerythrophyl 1um r e c u r v i rostrum 2 2.2 1 .2 1.2 2.2 Plagiopus oederi 2.2 1 .2 1 .2 1.2 1 .2 Plagiomnium rostratum 1 .2 1 .2 1 .2 1.2 Ditrichum f l e x i c a u l e + .2 Gymnostomum aeruginosum 1 .2 + .2 1 .2 1.4 Brachythecium r i v u l a r e 1 .4 + .4 Homalothecium fulgescens . 1 .4 1 .4 Brachythecium velutinum Jungermannia a t r o v i r e n s 1.4 1.4 1 .4 1.4 1.4 1 .4 1.4 + .4 D i s t i c h i u m capillaceum 1 .2 2.2 + .2 + .2 F i s s i d e n s bryoides 2.2 1.2 4.2 Lophozia g i l l m a n i i 1 1 .4 2.4 S e l i g e r i a donniana 2.2 1.2 Timmia a u s t r i a c a 1.2 1.2 T o r t u l a princeps 1.2 1.2 Apometzgeria pubescens 2.4 1.4 Gymnostomum re c u r v i r o s t r u m 1 .2 1.2 Table XXVIII (continued). Only found once: Dichodontium pellucidum, Cephalozia b i c u s p i d a t a , Scapania undulata, Mnium thomsonii. R e l e v e - l o c a t i o n s : 146,147,283,284,285 Slesse Creek 280,281,282,277,278,279 Chipmunk Creek 152 predominantly occurs under the very shaded and s h e l t e r e d c o n d i t i o n s of "r o c k - g r o t t o s " and the immediately adjacent rock bases o u t s i d e these g r o t t o s . Confined to the dry i n t e r i o r of the rock grottos was the f o l l o -wing combination of species: D i s t i c h i u m capillaceum, F i s s i d e n s b r y o i d e s , 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 Gymno-stomum re c u r v i r o s t r u m (1 i n Table XXVIII). The humid to moist rock bases d i r e c t l y o u t s ide these grottos showed a more diverse and 1uxuriant vegetation (2 i n Table XXVIII). Species confined ( w i t h i n the limestone releves) to t h i s type of 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 canescens, Brachytheciurn r i v u l a r e , B_. velutinum 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 the group of species outside the rock grottos) and the a l l i a n c e Encalyption procerae are expressed by species such as Bryoerythrophyllum r e c u r v i - 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 P l a g i o c h i l a a s p l e n i o i d e s . Further research i s needed i n order to d i s t i n g u i s h and recognize the nature of the bryophyte vegetation on calcareous rock s u b s t r a t a i n western North America, i n general and i n B r i t i s h Columbia, 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 the d e s i g n a t i o n of 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 character species and w i l l allow a more r e l i a b l e comparison with data from limestone bryophyte vegetations as described and c l a s s i f i e d i n Europe. 153 8.4.4. Evaluation of the o r d i n a t i o n techniques. In order to d i s p l a y and v i s u a l i z e r e l a t i o n s h i p s between the releves from limestone rocks, contained i n major group number 4, o r d i -nation techniques were a p p l i e d to the data s e t . The use of the o r d i -n a t i o n , namely, provides a means whereby a l l releves can be compared with one another. The r e s u l t s of the centered P r i n c i p a l Components A n a l y s i s are shown i n Figure 8. The 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 percentage d i s s i m i l a r i t y between the releves which are arranged along the X-axis and Y-axis according to t h e i r sample scores ( l i s t e d i n Appendix IV). A f i r s t group of releves (the a s s o c i a t i o n Lophozio - Bryoery-throphylletum r e c u r v i r o s t r i ) , predominantly occurs i n the bottom r i g h t corner of the o r d i n a t i o n diagram (Figure 8). A second group of releves (the 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 top l e f t corner and a t h i r d group of r e l e v e s , the Barbula 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, appears confined to the bottom l e f t corner of 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 Figure 9, i n which the centroids and the standard d e v i a t i o n s place the three groups of releves i n the r e s p e c t i v e quadrants. A comparison of the r e s u l t of t h i s centered P r i n c i p a l Components An a l y s i s with 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 c l u s t e r a n a l y s i s , shows a r e c o g n i t i o n and a separation of the same groups. However, rele v e number 11 was part of 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 (Figure 11), but i n the centered P.C.A. (Figure 8) i s grouped with releves 2, 3, 4, 154 1 1 1 1 2 4 6 8 0 2 4 6 1 2 2 2 2 2 8 0 2 4 6 8 3 3 3 3 3 4 4 4 4 4 5 5 5 . 5 5 6 6 6 6 6 7 7 7 7 7 8 8 8 8 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 1 9 9 9 9 9 0 0 2 4 6 8 0 % d i s s i m i l a r i t y Figure 8. P r i n c i p a l components a n a l y s i s of the 27 limestone r e l e v e s . (A.=Metaneckeretum m e n z i e s i i ; • =Lophozio-Bryoerythrophylletum 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 jungermannioides-Jungermannia atrovirens^commumtyif = m s i d e grotto type; O=outside gr o t t o type.) 155 2 4 6 8 0 1 ( 2 2 2 2 2 3 3 6 8 0 2 4 6 8 0 2 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 % d i s s i m i l a r i t y Figure 9. Scatter-diagram of the 27 limestone r e l e v e s from the PCA-cent o r d i n a t i o n . l=Lophozio-Bryoerythrophylletum 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 are i n d i c a t e d by l i n e s . 156 7, 8 and 10 (=cluster 2). A manual t a b u l a t i o n p r i o r to the c l u s t e r ana-l y s i s had included r e l e v e number 11 i n the Metaneckeretum m e n z i e s i i , which, a l s o , i s represented by c l u s t e r 2. The r e l a t i v e l y high cover value of Racomitrium heterostichum var. a f f i n e and the presence of Bryum c a p i l l a r e are shared with releves numbers 12 and 1, r e s p e c t i v e l y . However, the occurrences of Isothecium s t o l o n i f e r u m , P l a t y d i c t y a  jungermannioides and Gymnostomum aeruginosum i n d i c a t e i t s " a f f i n i t y " to the Metaneckeretum me n z i e s i i r a t h e r than the Lophozio - Bryoerythro-phylletum. The presence of Encalypta procera excludes re l e v e number 11 from the t h i r d group of r e l e v e s , the Barbula v i n e a l i s - P l a t y d i c t y a 5 jungermannioides - Jungermannia a t r o v i r e n s - community . The P r i n c i p a l Components A n a l y s i s shows t h i s t h i r d group of releves as a more or l e s s " i s o l a t e d " c o l l e c t i o n of limestone samples a l l r e s t r i c t e d to the l e f t lower quadrant of the o r d i n a t i o n diagram (Figure 9 ) . The c l u s t e r a n a l y s i s (Figure 11) separated t h i s c l u s t e r 3 i n t o two smaller c l u s t e r 3A and 3B. The f l o r i s t i c and e c o l o g i c a l r e l a t i o n s h i p s between these " i n s i d e - g r o t t o " and " o u t s i d e - g r o t t o " types are described on page 152. This separation i n t o two more or l e s s autonomous groups i s s u b s t a n t i a t e d by a subsequent p o l a r o r d i n a t i o n (Figure 10). Here, the d i s t i n c t i o n between 3A ( r e l e v e s 5, 6, 22, 23 and 24) and 3B (releves 19, 20, 21, 25, 26 and 27) becomes apparent i n t h e i r d i s t r i b u t i o n along the Y-axis. With respect to the r e l a t i o n s h i p s between the releves and t h e i r environments, the X-axis i n the p o l a r o r d i n a t i o n 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 of the 157 1 0 0 9 8 9 E 9 4 9 2 9 0 8 8 8 6 8 4 8 2 8 0 78 7 6 74 7 2 7 0 6 8 6 6 6 4 6 2 6 0 >> 5 8 4-> 5 6 t l 5 4 (0 5 2 5 0 »r- 4 8 E 4 6 • r - 4 4 i / i 4 2 to 4 0 • r - 38 3 6 34 32 3 0 2 8 2 6 24 2 2 2 0 18 16 14 12 10 26 10 O 27 S 2?, 20 °19 7 3 23. • •22 •, 24 12 13 15 16 14 IB 17 5 1 1 2 4 2 2 2 2 2 0 2 4 6 8 3 3 3 3 2 4 6 8 4 4 4 0 2 4 4 4 5 5 6 8 0 2 5 5 5 6 6 4 6 8 0 2 6 6 6 8 7 7 0 2 7 7 7 4 6 8 8 8 8 0 2 4 1 8 8 9 9 9 9 9 0 6 8 0 2 4 6 8 0 Figure 10, % d i s s i m i l a r i t y P o l a r o r d i n a t i o n o f the 27 limestone r e l e v e s (^=Meta-neckeretum m e n z i e s i i ;•= Lophozio-Bryoerythrophyl letum 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 jungermanni-oides-Jungermannia atrovires-community:a =inside g r o t t o type;0=outside g r o t t o type.) s T E P 2 2 2 2 2 2 2 1 1 1 1 1 1 1 1 6 5 4 3 2 1 0 9 8 7 G 5 4 3 2 1 0 9 --I I - - I -8 7 6 5 4 3 2 1 I-I--I I-I I----I + -I + - I -+ + -I i - B -co D 7 7 8 8 8 1 1 1 2 2 3 3 4 4 5 5 G 7 7 7 7 8 1 1 1 1 I 4 6 9 3 7 5 7 4 7 3 9 1 1 2 3 7 5 1 3 5 8 S 2 5 0 0 8 2 4 0 8 T 5 0 0 0 3 7 0 5 0 6 7 2 5 0 0 1 5 5 1 4 2 8 . . . . A 0 0 0 0 3 5 0 4 0 6 5 5 0 0 0 2 4 0 6 2 5 3 2 2 3 4 N 0 0 0 0 3 0 0 2 0 7 0 0 0 0 0 5 2 0 7 5 0 3 4 8 0 9 C E S Figure 11. C lu s ter ana lys i s of the 27 limestone re leves . Symbols and numerical and a lphabet ic characters are discussed in the text . Distance is euc l idean. 159 releves along t h i s a x i s . In a d d i t i o n , a l s o the measured thickness o f s o i l accumulation on the s u b s t r a t a reveals a gradient along the X-axis. The grotto r e l e v e s , on the l e f t p o r t i o n of the a x i s , d i d not show any s o i l accumulation, whereas the exposed bryophyte assemblages of 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 of the X - a x i s , were growing on s o i l d e positions on rocks up to 2(3) cm t h i c k . The d i s t r i b u t i o n of the releves along the Y-axis suggests a gradient from dry to humid. The i n s i d e - g r o t t o bryophytes, c h a r a c t e r i s -t i c a l l y , occurred on dry, c r u s t y limestone, whereas the Lophozio -Bryoerythrophylletum r e c u r v i r o s t r i grew on exposed, dry s o i l - c o v e r e d rocks. Both groups of releves are p o s i t i o n e d along the lower p o r t i o n of the Y-axis. The o u t s i d e - g r o t t o releves and the releves 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 , occurred on s h e l t e r e d rock bases and shaded boulders i n s i d e f o r e s t vegetations. T h e i r subsequent p o s i t i o n i n g along the Y-axis i s more or l e s s concentrated i n the upper p o r t i o n of the o r d i n a t i o n diagram (Figure 10). To 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 was performed on the r e l e v e s 1 scores used i n the centered P r i n c i p a l Compo-nents A n a l y s i s . The r e s u l t of t h i s c l u s t e r a n a l y s i s i s presented i n Figure 12. The f i r s t c l u s t e r a n a l y s i s (Figure 11) placed r e l e v e number 11 i n the Lophozio -Bryoerythrophylletum 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 i s placed i n c l u s t e r 2, recognized and designated before as the a s s o c i a t i o n Metaneckeretum m e n z i e s i i . O N oo r o t o oo i — . — i — CSJ Distance 0 0 o o o o - - - -• • • • • • o o o o 0 0 o o o o •si" CO LO 00 i — (O M J \ 1 — 1 — OO o r -00 LO OO I — LO O LO O O O 1 — • — 1 — 1 — CM 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 ^j- r\ r-~ 00 r o o Figure 12. C l u s t e r a n a l y s i s o f P.C.A.-scores (7 axes) of the 27 limestone r e l e v e s . Distance i s Euclidean. 160 1. Since limestone s u b s t r a t a are very 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 area, sampling of the bryophyte vegetation on calcareous rocks i s performed only i n the C h i l l i w a c k R iver V a l l e y . 2. Synsystematic treatment of the order 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 the a l l i a n c e S c h i s t i d i o n apocarpi Jezek and Vondracek 1962 are given by M a r s t a l l e r (1979) and (1980), respec-t i v e l y 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 detected between the p r e s e n t l y recognized Lophozio - Bryoerythrophyl1etum 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 provided by P o e l t (1954), Haybach (1956), Hb'fler (1959), P h i l i p p i (1965), Hagel (1966), Nagano (1969), Norr (1970), Neumayr (1971), S t r a s s e r (1972), He r t e l (1974), Stuchly (1976), Hebrard (1978), 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 the Metaneckeretum me n z i e s i i and European limestone bryophyte vegetations are found i n the f o l l o w i n g references: Demaret (1944), P o e l t (1954), Koppe (1955), H o f l e r (1959), Breuer (1962), P h i l i p p i (1965), Hagel (1966), Hubschmann (1967), Norr (1970), Neumayr (1971), S t r a s s e r (1972), He r t e l (1974), Wilczynska (1974), Mamczarz (1978), M a r s t a l l e r (1979, 1980). 5. Also the type of h a b i t a t , i n t h i s case shaded, humid boulders i n s i d e f o r e s t s , i n d i c a t e the placement of r e l e v e number 11 i n the Meta-neckeretum m e n z i e s i i . The Lophozio - Bryoerythrophyl1etum recur-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 rocks. 161 9. SUMMARY. In a study concerned with the e p i l i t h i c bryophyte vegetation i t i s important to r e a l i z e that s u b s t r a t a such as boulders and rock-w a l l s u s u a l l y are "discontinuous" with the surrounding (vascular) vegetation. Rocky slopes i n subalpine and a l p i n e areas u s u a l l y contain a pioneer stage i n which bryophytes (and l i c h e n s ) are the s o l e i n h a b i t a n t s , and v a s c u l a r plants can e s t a b l i s h only i n l a t e r stages of succession. 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 trees and shrubs may become anchored, remain i n d e f i n i t e l y i n a semi-barren s t a t e conditioned by the rat e and amount of s o i l accumulation. In most in s t a n c e s , exposed and s h e l t e r e d boulders and rocks are u s u a l l y covered by a vegetation which i s , f l o r i s t i c a l l y , considerably u n l i k e the p l a n t cover o f otherwise wooded t e r r i t o r y . The vegetation growing on a bare rock surface i s r e l a t e d p r i -m a r ily to the kind of rock, i t s chemical composition (Shacklette 1961, P h i l i p p i 1961, Nagano 1972), the s t a t e of weathering and surface micro-s t r u c t u r e (Sjogren 1964). I t i s also i n f l u e n c e d by macroclimate 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 (Krusenstjerna 1965). Hence, many e c o l o g i c a l f a c t o r s such as moisture, l i g h t , temperature and s o i l accumulation determine the d i s t r i -b ution of e p i l i t h i c bryophytes and t h e i r communities. (Oosting and Anderson 1947, Keever, Oosting and Anderson 1951). Also the relevance of bryophyte r h i z o f d s i n a t t a c h i n g the plants to the rock substrate 162 i s o f major importance i n the establishment and development of the bryophyte vegetation (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 f u r t h e r r e f l e c t e d i n the v a r i a t i o n i n f l o r i s t i c composition o f bryophyte assemblages from d i f f e r e n t environments. The bryophyte communities from exposed, dry ro c k s , from shaded, humid boulders and from streamside rocks a l l e x h i b i t "discontinuous" 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 composition as compared to the (vascular) vegetation of t h e i r immediate environment. This observation i m p l i e s some conside-r a b l e r e l a t i o n s h i p between the species of a community which, i n growing together, s u f f i c i e n t l y modify the environment to form a recognizable and r e p e t i t i v e v egetational grouping (Anderson 1965). Any procedure o f vegetation c l a s s i f i c a t i o n i n v o l v e s an arrange-ment o f communities i n t o c l a s s e s : the members of 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 serve to set them apart from members of another c l a s s . I m p l i c i t i n t h i s approach i s the suggestion that there i s some d i s c o n t i n u i t y i n species ( a t t r i b u t e ) composition between both concrete samples of vegetation ( r e l e v e s ) i n the f i e l d and t h e o r e t i c a l u n i t s (syntaxa) ab s t r a c t e d from such f i e l d data. In the present attempt to c l a s s i f y the e p i l i t h i c bryophyte communities i n southwestern B r i t i s h Columbia, the Braun-Blanquet method of vegetation study has been a p p l i e d . This approach i s based on the concept o f e x i s t i n g discontinuous v a r i a t i o n w i t h i n the p l a n t cover o f a given area and r e s u l t s i n the r e c o g n i t i o n o f more or l e s s r e a d i l y 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 are 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 of syntaxa. In a d d i t i o n to the t a b u l a t i o n technique of o r d e r i n g species and r e l e v e s , c l u s t e r analyses have been used to recognize patterns i n the data matrix of 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 ) . This search f o r groups of s i m i l a r releves has l e d to the i d e n t i f i c a t i o n o f separate a s s o c i a t i o n s , 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 , each c h a r a c t e r i z e d by t h e i r character s p e c i e s . In order to recognize these u n i t s 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 the preparation of raw t a b l e s and presence t a b l e s by the computerized t a b u l a r s o r t i n g program VEGTAB, has been found s u c c e s s f u l . I t reveals d i s t i n c t i v e patterns of species d i s t r i b u t i o n throughout the data set and provides an adequate basis f o r the c l a s s i f i c a t i o n of e p i l i t h i c bryophyte a s s o c i a t i o n s and lower syntaxonomical u n i t s . However, above 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 provide a means to combine a s s o c i a t i o n s i n t o the higher syntaxa of a l l i a n c e s and orders. The c l u s t e r a n a l y s i s algorithm detects d i f f e r e n c e s i n species composition and 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 vegetation type. The c l a s s i f i c a t i o n o f a s s o c i a t i o n s i n t o higher syntaxa, on the other hand, i s based on t h e i r occurrences 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 . Although 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 species composition and q u a n t i t i e s are detected 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 , t h i s 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 of the major groups (Tables I I I , XI, XV and XXV) The l e v e l of 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 of the syntaxa i s , thus, somewhat a r b i t r a r y . 164 However, attempts have been made to define bryophyte communities to correspond to the e p i l i t h i c a s s o c i a t i o n s described by various European b r y o - s o c i o l o g i s t s . These workers mostly a p p l i e d t a b u l a t i o n techniques, standard w i t h i n the Braun-Blanquet method, to t h e i r vegetation data. 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 analyses to a r r i v e at the separate vegetation u n i t s . I t i s t h i s comparison between data from European vegetations and the present study's data m a t r i c e s , which revealed some remarkable s i m i l a r i t i e s i n species c o n t r i b u t i o n to the separate a s s o c i a t i o n s and i n the m i c r o - s i t e s p e c i f i c i t y of the bryophyte communities i n general. Hence, where p o s s i b l e , the pr 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 units have been placed 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 f o r bryophyte communities o c c u r r i n g i n d i f f e r e n t types of 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 the present study, has been welcomed as an a d d i t i o n a l means to d i s p l a y and i n t e r p r e t patterns i n the data matrix. The r e c o g n i t i o n o f these patterns 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-axis of the o r d i n a t i o n diagrams (Figures 8 and 10). D i f f e -rences i n the p o s i t i o n s of the vegetation samples have c o n t r i b u t e d to the i d e n t i f i c a t i o n of "groups of s i m i l a r r e l e v e s " , and a comparison with the c l u s t e r a n a l y s i s ' arrangement f o l l o w e d . The P r i n c i p a l Components A n a l y s i s , which ordinates releves w i t h respect to a l l others simultaneously, s u b s t a n t i a t e d and supported the recog-165 n i t i o n of three limestone bryophyte vegetation 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 species. A subsequent p o l a r o r d i n a t i o n , which ordinates releves on the ba s i s of comparison with two endpoint r e l e v e s , not only revealed d i f f e r e n c e s w i t h i n one of the u n i t s (3A and 3B; Figure 10), but provided a means to c o r r e l a t e the p o s i t i o n s of the groups of s i m i l a r releves along the axes with e c o l o g i c a l f a c t o r s such as exposure, moisture 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 , already showed some r e l a t i o n s h i p s of releves to one another and to the environment. This provided a basis 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 expres-sed and c l a r i f i e d those r e l a t i o n s h i p s . Further research on the e p i l i t h i c bryophyte vegetation i s needed. E s p e c i a l l y i n subalpine 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 supply, the v a r i a t i o n i n species composition and the m i c r o - s i t e s p e c i f i c i t y of bryophyte assemblages need a d d i t i o n a l work. More releves may s u b s t a n t i a t e f u r t h e r the 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 the e p i l i t h i c bryophyte vegetation o f south-western B r i t i s h Columbia and other areas i n the P a c i f i c North West, i n g e n e r a l , and regions elsewhere i n North America and Europe. The knowledge on these vegetations w i l l gain i n understanding and meaning when these methods are e x t e n s i v e l y a p p l i e d over wider areas i n the temperate regions of the Northern Hemisphere. On a more l o c a l s c a l e research may be d i r e c t e d on the nature 166 of the bryophyte assemblages. F i r s t , more d e t a i l e d measurements o f environmental f a c t o r s are required to c h a r a c t e r i z e the e c o l o g i c a l d i f f e r e n c e s on and between the various rock s u b s t r a t a . Second, aut-e c o l o g i c a l ( 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 ) information i s needed f o r the various species i n order to properly examine t h e i r a b i l i t y and success to occupy the e p i l i t h i c h a b i t a t , and to more a c c u r a t e l y describe niche 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 quadrats and using information from the previous two suggestions f o r f u r t h e r research, the dynamic nature of the e p i l i t h i c vegetation could be s t u d i e d and contrasted between e c o l o g i c a l l y d i f f e r e n t environments. REFERENCES. 167 Anderson, 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 vegetation science: controversy over a non-existent problem? J . 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Moosflora und Bryophytengesellschaften der Gory Kaczawskie (nordwestliche Sudeten). Monographiae Botanicae 44: 1-111. Wirth, V. 1972. Die S i l i k a t f l e c h t e n g e m e i n s c h a f t e n im ausseralpinen Zentraleuropa. D i s s e r t a c i a e Botanicae, Bd. 17. Lehre. Worley. I. W. 1972. The bryo-geography of Southeastern A l a s k a . Part I & I I . Ph. D. T h e s i s , Univ. of B r i t i s h Columbia. Yarranton, G. A. 1965. 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 ecology of some sax 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. of Exeter. 62 p. . 1967a. P r i n c i p a l components a n a l y s i s of data from s a x i c o -lous bryophyte vegetation at Steps Bridge, Devon. I. A q u a n t i -t a t i v e assessment of 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. I b i d . I I . An experiment with heterogeneity. Can. J . Bot. 45: 229-247. . 1967c. I b i d . I I I . C o r r e l a t i o n of v a r i a t i o n i n the vege-t a t i o n with 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 of the bryophyte and macro-l i c h e n vegetation of 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%) of 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%) of the p l o t + covering <1/100 ( <1% ) of the p l o t 181 SOCIABILITY SCALE 5 compact mat - 20 cm 4 loose mat or loose t u r f - 8-20 cm 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 p l ants - t u r f - between 1 cm i n acrocarpous mosses and 3-4 (5) cm i n pleurocarpous mosses 182 APPENDIX I I CHARACTER TAXA. F i d e l i t y degree 5: Ex c l u s i v e taxa: taxa completely or almost completely confined to one phytocoenon (vegetation u n i t ) ; F i d e l i t y degree 4: S e l e c t i v e taxa: taxa o c c u r r i n g w i t h c l e a r preference f o r one phytocoenon but a l s o , though with a conside r a b l y lower presence degree, i n other coena: F i d e l i t y degree 3: P r e f e r e n t i a l taxa: taxa present i n several coena, perhaps with about equal presence degree, but with a higher combined est i m a t i o n value and (or) with a higher v i t a l i t y degree i n one p a r t i c u l a r coenon; COMPANIONS. F i d e l i t y degree 2: I n d i f f e r e n t taxa: taxa without pronounced preference f o r any coenon; ACCIDENTALS. F i d e l i t y degree 1: Strange taxa: taxa having a d e f i n i t e presence degree optimum and mostly a l s o a cover-abundance optimum outside the considered coenon. These are 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-bouring coena or r e l i c s from a coenon t h a t preceded i n succession. 183 APPENDIX I I I Bryophyte species l i s t 184 CLASS HEPATICAE Family Pseudolepicoleaceae BIepharostoma tric h o p h y l l u r n (L.) Dum. Family Herbertaceae Herbertus aduncus (Dicks.) S. Gray Family P t i l i d i a c e a e 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.) Vain. Family Lepidoziaceae Bazzania denudata (Torr. ex Gott.) Trev. B_. t r i c r e n a t a (Wahlenb.) Lindb. i n Broth L e p i d o z i a reptans (L.) Dum. Family Calypogeiaceae Calypogeia i n t e g r i s t i p u l a Steph. C_. muelleriana ( S c h i f f n . ) K. M u e l l . Family Antheliaceae A n t h e l i a j u r a t z k a n a (Limpr.) Trev. Family Cephaloziaceae Cephalozia b i c u s p i d a t a (L.) Dum. 185 Family Cephaloziaceae (continued) H y g r o b i e l l a l a x i f o l i a (Hook.) Spruce Pleuroclada albescens (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. var. byssacea C;. byssacea var. a s p e r i f o l i a (Tayl.) Macv. C. phyllacantha (Mass. & Carest.) K. M u e l l . CL t u r n e r i (Hook.) K. M u e l l . Family Geocalycaceae Geocalyx graveolens (Schrad.) Nees Family Lophocoleaceae Chiloscyphus polyanthos (L.) Corda Lophocolea cuspidata (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 Gyrothyra underwoodiana M.A. Howe Family Jungermanniaceae Anastrophyllum a s s i m i l e ( M i t t . ) Steph. A. minutum (Schreb.) Schust. B a r b i l o p h o z i a f l o e r k e i (Web. et Mohr.) Loeske 186 Family Jungermanniaceae (continued) B a r b i l o p h o z i a hatcheri.(Evans) Loeske B_. lycopodioides (Wallr.) Loeske Chandonanthus f i 1 i formi s Steph. C. s e t i f o r m i s ( E h r h J Lindb. Jamesoniella 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. Lophozia elongata Steph. L_. e x c i s a (Dicks.) Dum. I. heterocolpos (Thed. ex Hartm.) M.A. Howe IL. i n c i s a (Schrad.) Dum. L_. sudetica (Nees ex Hueb.) G r o l l e L_. v e n t r i c o s a (Dicks.) Dum. var. v e n t r i c o s a L_. wenzelii~("Nees) Steph. M y l i a t a y l o r i (Hook.) S. Gray Nardia b r e i d l e r i (Limpr.) Lindb. N_. geoscyphus (DeNot) Lindb. N_. japonica 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) Joerg. J_. quinquedentata (Huds.) Buch Family Gymnomitriaceae Gymnomitrion concinnatum ( L i g h t f . ) Corda i n Opiz. G. obtusum ( l i n d b . ) Pears. Marsupella breyissima (Dum.) G r o l l e M. condensata (Angstr.) S c h i f f n . M. emarginata (Ehrb.) Dum. M. s p a r s i f o l i a (Lindb.) Dum. M. sphacelata (Gieseke ex Lindenb.) Dum. M. s t a b l e r i Spruce 187 Family Scapaniaceae Douinia ovata (Dicks.) Buch Diplophyllum 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 Lindb. D. t a x i f o l i u r n (Wahlenb.) Dum. Scapania mucronata Buch S. paludosa (K. Muell.) K. M u e l l . S_. scandica (H. Arn. & Buch) Macv. S_. subalpina (Nees ex Lindenb.) Dum. S. u l i g i n o s a (Sw. ex Lindenb.) Dum. undulata~~(L.) Dum. Family Radulaceae Radula bolanderi Gott. 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.) Lindb. 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 (Gott.) Limpr. Family Metzgeriaceae Apometzgeria pubescens (Schrank) Kuwah. Metzgeria conjugata Lindb. 188 Family Blasiaceae B l a s i a p u s i l l a L. Family Aneuraceae Aneura pinguis (L.) Dum. R i c c a r d i a m u l t i f i d a (L.) S. Gray Family Aytoniaceae Reboulia hemisphaerica (L.) Raddi Family Conocephalaceae Conocephalurn conicum (L.) Dum. ex Lindb. Family Marchantiaceae P r e i s s i a quadrata (Scop.) Nees 189 CLASS MUSCI SUBCLASS SPHAGNIDAE Family Sphagniceae Sphagnum gir 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 Tetraphis p e l l u c i d a Hedw. SUBCLASS POLYTRICHIDAE Family P o l y t r i c h a c e a e Atrichum s e l w y n i i Aust. A. undulatum (Hedw.) P. Beauv. Bartramiopsis Tescur'i'i (James) Ki 190 Family P o l y t r i c h a c e a e (continued) 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 . ) Kindb. Pogonatum contortum ( B r i d . ) Lesq. _P. urnigerum (Hedw.) P. Beauv. Polytrichum alpinum Hedw. var. alpinum _P. alpinum var. 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. sexangulare Floerke 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 capillaceum (Hedw.) B.S.G. Ditrichum 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 acuta (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 (Dicks.) 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 Family Dicranaceae (continued) Dichodontium pellucidum Ren. & Card. D. olympicum (Hedw.) Schimp. D i c r a n e l l a heteromalla (Hedw.) Schimp. Dicranoweisia c i r r a t a (Hedw.) Lindb. i n Milde D. c r i s p u l a (Hedw.) Lindb. i_n Mi 1 de Dicranum fuscescens Turn JJ. scoparium Hedw. K i a e r i a b l y t t i i (B.S.G.) Broth. 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 Fissidentaceae F i s s i d e n s bryoides Hedw. F. osmundoides Hedw. Family Encalyptaceae Encalypta c i l i a t a Hedw. E. procera Bruch Family Pottiaceae Anoectangium aestivum (Hedw.) M i t t . Barbula r e f l e x a (.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. & Tayl.) A.J.E. Smith T o r t e l l a f r a g i l i s (Hook f. & Wils.) Limpr. T o r t u l a princeps De Not 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 var. h o l z i n g e r i (Card. & 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. torquata 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. "brevipes" 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 . var. heterostichum R. heterostichum var. a f f i n e (C. Jensen) Kindb. R. sudeticum (Funck) B.S.G. R. variurn ( M i t t . ) Jaeg. & Sauerb. S c h i s t i d i u m apocarpum (Hedw.) B. & S. var. apocar.pum S_. apocarpum var. 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 aquatica Hook, i j i Drumm. Family Schistostegaceae Schi s t o s t e g a pennata (Hedw.) Web. & Mohr. Family Bryaceae Bryum c a e s p i t i c i u m Hedw. B_. mini at urn Lesq. B_. pallescens S c h l e i c h . ex Schwaegr. B_. pseudotriquetrum (Hedw.) Schwaegr. P o h l i a c a r d o t i i (Ren.,ex_Re. & Card.) Broth. P_. cruda (Hedw.) Lindb. P_. l o n g i b r a c t e a t a Broth, vn R o e l l . P_. l u d w i g i i (Schwaegr.) Broth. P_. nutans (Hedw.) Lindb. IP. p r o l i g e r a (Kindb. e_x Limpr.) Broth. 193 Family Mniaceae Leucolepis m e n z i e s i i (Hook.) Steere i j i Koch Mnium ambiguum H. M u e l l . MT~5Tyttii B.S.G. M. spinulosum B.S.G. M. thomsoniT~Schimp. Plagiomnium rostratum (Schrad.) Kop. P_. venustum ( M i t t . ) Kop. Rhizomnium glabrescens (Kindb.) Kop. R. nudum~TBritt. & Williams) Kop. Family Aulacomniaceae Aulacomnium androgynum (Hedw.) Schwaegr. Family Bartramiaceae A n a c o l i a m e n z i e s i i (Turn.) Par. Bartramia i t h y p h y l 1 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_. fontana (Hedw.) B r i d . var. fontana Plagiopus oederi ( B r i d . ) Limpr. Family 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. Muell.) Broth 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.) Broth. Family Neckeraceae Homalia trichomanoides (Hedw.) B.S.G. Metaneckera m e n z i e s i i (Hook ijn Drumm.) Steere P o r o t r i c h u i r n b i g e l o v i i ( S u l l . ) Kindb. Thamnobryum neckeroides (Hook.) Lawt. Family Leskeaceae Pseudoleskea a t r i c h a (Kindb. j_n Macoun & Kindb.) Kindb. P_. b a i l e y i Best & Grout j_n Grout P_. in c u r v a t a (Hedw.) Loeske JP. patens (Lindb.) Kindb. P_. radicosa ( M i t t . ) Macoun & Kindb. P_. s t e n o p h y l l a Ren. & Card, ex Roe! 1. Pseudoleskeella tectorum (Funck ex Br i d . ) Kindb. e_x Broth. Pterigynandrum f i l i f o r m e Hedw. Family Thuidiaceae Claopodium bolanderi Best C_. c r i s p i f o l i u m (Hook.) Ren. & Card. C_. whippleanum ( S u l l . ) Ren. & Card. Heterocladium dimorphum ( B r i d . ) B.S.G. H_. macouni i Best H_. procurrens ( M i t t . ) Rau & Herv. Family Amblystegiaceae Cratoneuron commutatum (Hedw.) Roth. var. falcatum ( B r i d . ) Moenk. 195 Family Amblystegiaceae (continued) Drepanocladus uncinatus (Hedw.) Warnst. Hygrohypnum b e s t i i (Ren. & Bryhn.) Holz. H. 1uridurir(Hedw.) Jenn. H. ochraceum (Turn. j_n Wils.) Loeske TT. s m i t h i i ("Sw. vn L i l j . ) Broth. P l a t y d i c t y a jungermannioides ( 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. Muell.) Besch. B_. pi umosum (Hedw.) B.S.G. B_. reflexum (Starke 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_. velutinum (Hedw.) B.S.G. Eurhynchium pulchellum (Hedw.) Jenn. Homalothecium fulgescens ( M i t t , ex C. Muel1.) Jaeg. H. nevadense (Lesq.) Ren. & Card. H_. n u t t a l l i i (Wils.) Jaeg. & Sauerb. Isothecium s t o l o n i f e r u m B r i d . Scleropodium cespitans (C. Muell.) 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 Plagiothecium 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 11. \\_. subimponens Lesq. Isopterygium elegans ( B r i d . ) Lindb. J_. pulchellum (Hedw.) Jaeg. & Sauerb. Family Rhytidiaceae Pleurozium schreberi ( B r i d . ) M i t t . Rhytidiadelphus loreus (Hedw.) Warnst. R. squarrosus (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 the bryophyte vegetation from calcareous rocks. ORDINATION: ^CA-CENT AXIS EIGENVALUE %EV SUM %EV SORT EV SCALE 1 0. . 1 170347772E+03 16 .20772 16 .20772 0. . 1081826E+02 99 .99998 2 0. .8845344522E+02 12 .24960 28 .45731 0. 940496BE+01 86 .93604 3 0. 6636100796E+02 9 .19010 37 .64740 0, 8146227E+01 75 .30069 4 0. 6478132645E+02 8 .97134 46 .61873 0. 8048685E+01- 74 .39905 5 0, 5392733340E+02 7 .46820. 54 .08693 0, 7343522E+01 67 .88078 6 0. 4053179768E+02 5 .61310 59 .70003 0. 6366458E+01 58 .84917 7 0. . 3963538360E+02 5 .48896 65 . 18898 0. 6295663E+01 58 . 19476 8 0 . 3443028234E+02 4 .76812 69 .95709 0. 5867731E+01 54 .23912 9 0 .3260502171E+02 4 .51535 74 .47244 0. 5710080E+01 52 .78186 10 0. 3161708391E+02 4 . 37854 78 .85097 0. 5622907E+01 51 .97606 1 1 0. 2669218426E+02 3 .69650 82 .54747 0. 5166447E+01 47 .75671 12 0. . 2080173402E+02 2 .88076 85 .42822 0. 4560891E+01 42 . 15918 13 0. 1665683272E+02 2 .30674 87 .73495 0. 4081278E+01 37 .72581 14 0. 1584265435E+02 2 .19399 89 .92894 0. 3980283E+01 36 .79225 15 0. 1501971595E+02 2 , .08003 92 .00896 0. 3875527E+01 35 82393 16 0. 1271493798E+02 1 . .76085 93 .76979 0. 3565801E+01 32 .96094 17 0. 8595393369E+01 1 . .19034 94 .96013 0. 2931790E+01 27 .10037 18 0. 8348086760E+01 1 . . 15610 96 . , 1 1621 0. 2889305E+01 26. ,70766 19 0. 6880249695E+01 0. 95282 97 . 06903 0. 2623023E+01 24 . 24625 20 0. 5611985637E+01 0. 77718 97 . 84621 0. 2368963E+01 21 . 89781 21 0. 5560991204E+01 0. 77012 98 . 61632 0. 2358175E+01 21 . 79810 22 0. 3052311112E+01 0. 42270 99. 03902 0. 1747087E+01 16 . 14941 23 0. 2238079501E+01 0. 30994 99. 34895 0. 1496021E+01 13. 82867 24 0. 1854'169694E+01 0. 25678 99 . 60573 0. 1361679E+01 12. 58686 25 0. 1659231780E+01 0. 22978 99. 83549 0. 1288112E+01 11 . 90683 26 0. 1 186387531E+01 O. 16430 99. 99979 0. 1089214E+01 10. 06829 27 0. 3036854776E-11 0. 00000 99. 99979 0. 1742657E-05 0. 00002 TRACE OF XX' = 0.7220924050278433E+03 AND SUM OF EV = O.7220924050278437E+03 ORDINATION: PCA-CENT SAMPLES SCORES N NAME AXIS 1 2 3 4 5 6 7 RANKED 1 RANKED 2 1 B36 66 0030 35 6590 59 8346 26 91 16 1 8629 100 OOOO 18 8461 26 B284 0 0 18 B168 0 0 2 B143 26 2603 59 1064 100 OOOO 87 8431 46 5316 0 0 27 9370 10 B151 3 5636 17 B167 9 7192 3 B144 3€ 4258 71 7089 47 6094 53 5039 52 9131 53 5233 58 2319 27 B285 8 9835 19 B277 9 7351 4 B145 50 4282 80 4470 46 9423 94 2649 69 5486 55 8857 100 OOOO 22 B280 9 1590 16 B166 1 1 5155 5 B146 21 3299 31 3806 46 2422 54 5822 9 4780 62 8040 72 2147 24 B282 14 6789 26 B284 17 1301 6 B147 24 4550 26 9038 50 2728 54 8831 7 7983 56 9247 59 1054 23 B281 19 1 176 21 B279 18 4446 7 B146 44 7655 100 OOOO 67 2378 100 OOOO 69 8990 52 0397 51 1418 5 B146 21 3299 20 B278 18 7065 8 B149 61 2145 71 4598 64 0598 65 3617 54 6609 67 0627 5 5325 6 B147 24 4550 25 B283 18 7716 9 B150 36 4539 81 5256 17 5231 38 5738 0 4417 4 0432 0 0 2 B143 26 2603 24 B282 25 6669 10 B151 3 5636 63 9928 0 0 14 3559 87 8954 43 6296 19 1456 21 B279 26 3815 22 B280 26 8562 1 1 B152 37 6696 61 7573 37 1 138 7 4050 8 8278 26 3210 78 6231 20 B278 31 9602 6 B147 26 9038 12 B153 80 8430 58 7905 53 3360 0 0 36 1447 43 1 148 67 1834 19 B277 33 3277 23 B281 27 3289 13 B163 93 3895 36 7245 69 6514 3 1 193 43 3161 26 5457 47 3408 25 B283 33 8506 5 B146 31 3806 14 B164 98 9754 38 1574 70 3888 1 1 7809 57 0537 31 7446 55 7872 3 B144 36 4258 27 B285 35 5084 15 B165 94 8663 52 8111 62 8172 20 5123 41 2860 32 2026 64 7493 9 B150 36 4539 1 B36 35 6590 16 B166 83 5083 1 1 5155 31 0888 89 0513 53 8592 38 4869 40 8244 1 1 B152 37 6696 13 B163 36 7245 17 B167 100 OOOO 9 7192 23 6481 97 0443 42 9194 1 1 73 16 49 5281 7 B148 44 7655 14 B164 38 1574 18 B168 68 6938 0 0 28 0138 95 5070 30 1735 50 0632 49 2352 4 B145 50 4282 15 B165 52 8111 19 B277 33 3277 9 7351 59 7458 32 4831 100 OOOO 54 3182 27 4161 8 B149 61 2145 12 B153 58 7905 20 B278 31 9602 18 7065 53 4055 40 6673 76 1382 41 3269 48 0960 1 B36 66 0030 2 B143 59 1064 21 B279 26 3815 18 4446 62 9720 32 0479 75 8416 36 3524 27 4426 18 B168 68 6938 1 1 B152 61 7573 22 B280 9 1590 26 8562 73 6195 60 8522 7 2431 29 0414 40 8096 12 B153 80 8430 10 B151 63 9928 23 B281 19 1 176 27 3289 60 8359 53 4068 13 2324 40 6389 43 1 162 16 B166 83 5083 8 B149 71 4598 24 B282 14 6789 25 6669 64 3800 53 1944 0 0 38 •1 169 41 9849 13 B163 93 3895 3 B144 71 7089 25 B283 33 8506 18 7716 60 6374 35 5916 79 0765 39 1343 31 3225 15 B165 94 8663 4 B145 80 4470 26 B284 0 0 17 1301 60 0098 36 4922 62 5152 33 2157 83 4620 14 B164 98 9754 9 B150 81 5256 27 B285 8 9835 35 5084 39 5620 31 7584 57 1403 30 9834 86 1852 17 B167 100 OOOO 7 B148 100 OOOO MIN MAX - 0 . 2 8 2 2 8 3 8 E + 0 0 0 .3495352E+00 - 0 . 3 0 2 0 5 2 7 E + 0 0 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 . 3 8 6 8 8 4 4 E + 0 0 0 .4195840E+00 ZERO 0 .4467795E+02 0 .4036324E+02 0.5225732E+02 0.4782201E+02 0.4391841E+02 0.4071303E+02 0 .4797266E+02 ORDINATION: POLAR OF SAMPLES USING PD ENDPOINTS 26 AND 17 AUTOMATIC FIRST AXIS (BASED ON ORDINATION NUMBER 7) BASELINE* 0.94737E+02 AND PD =100.00000 - PS N NAME XREL X E 1 B36 73 .37032 0 6950874E+02 0 7189250E+02 2 B143 53 .14667 0 5034949E+02 0 7156238E+02 3 B144 53 .15382 0 5035628E+02 0 7106398E+02 4 B145 54 .85742 0 5197020E+02 0 6790128E+02 5 B146 33 .59898 0 3183063E+02 0 7773550E+02 € 8147 48 .3994 1 0 4585210E+02 0 7975952E+02 7 B148 63 .43445 0 6009581E+02 0 7265181E+02 8 B 149 69 20155 0 6555939E+02 0 7551135E+02 9 B150 65 .82291 0 6235857E+02 0 7817549E+02 10 B151 49 .25737 0 4666489E+02 0 8252521E+02 1 1 B152 54 40120 0 5153799E+02 0 7819402E+02 12 B153 77 01830 0 7296472E+02 0 6838237E+02 13 B163 76 74149 O 7270248E+02 0 5918958E+02 14 B164 81 50082 0 7721133E+02 0 5438899E+02 15 B165 83 53162 0 7913524E+02 0 5026601E+02 16 B166 93 92409 0 8898074E+02 0 3118045E+02 17 B167 99 99994 0 9473682E+02 0 3053246E-04 18 B168 91 14244 0 8634549E+02 0 3863406E+02 19 B277 40 51532 0 3838295E+02 0 5169130E+02 20 B278 38 83049 0 3678679E+02 0 4310268E+02 21 B279 32 98726 0 3125110E+02 0 5285600E+02 22 B280 34 79553 0 3296420E+02 0 7089256E+02 23 B281 32 80728 0 3108060E+02 0 6825682E+02 24 B282 36 38588 0 3447084E+02 0 7126794E+02 25 B283 38 87265 0 3682674E+02 0 4819656E+02 26 B284 0 0 0 1105946E- 10 0 4577635E-04 27 B285 22 13565 0 2097063E+02 0 4986845E+02 DI D2 RANKED XREL 0 10O000OE+O3 0 7619048E+02 26 B284 0 0 0 8749998E+02 0 8421051E+02 27 B285 22 13565 0 8709676E+02 0 8378378E+02 23 B281 32 80728 0 8550723E+02 0 8024692E+02 21 B279 32 98726 0 8400000E+02 0 1000000E+03 5 B146 33 59898 0 9199998E+02 0 9354837E+02 22 B280 34 79553 0 9428571E+02 0 8048781E+02 24 B282 36 38588 0 1000000E+03 0 8095238E+02 20 B278 38 83049 0 1000000E+03 0 8461537E+02 25 B283 38 87265 0 9480519E+02 0 9550562E+02 19 B277 40 51532 0 9365079E+02 0 8933333E+02 6 B147 48 39941 0 1000000E+03 0 7176471E+02 10 B151 49 25737 0 9374998E+02 0 6315788E+02 2 B143 53 14667 0 9444444E+02 0 5714285E+02 3 B144 53 15382 0 9374998E+02 0 5263156E+02 1 1 B152 54 40120 0 9428571E+02 0 3170731E+02 4 B145 54 85742 0 9473683E+02 - 0 3051758E-04 7 B148 63 43445 0 9459459E+02 0 3953487E+02 9 B150 65 82291 0 6438356E+02 0 7647058E+02 8 B149 69 20155 0 5666666E+02 0 7222221E+02 1 B36 73 37032 0 6140349E+02 0 8260869E+02 13 B163 76 74149 0 7818181E+02 0 9402985E+02 12 B153 77 01830 0 7499998E+02 0 9333333E+02 14 B164 81 50082 0 7916666E+02 . 0 9333333E+02 15 B165 83 53162 0 6065573E+02 0 7534247E+02 18 B168 91 14244 - 0 4577637E-04 0 9473683E+02 16 B166 93 92409 0 5409834E+02 0 8904109E+02 17 B167 99 99994 r o O O 201 X IS £ 2 m T 5 2™ i n i n i n c o p ~ T r o 3 a i i n o i u J O O T P j p - r - - o i c o 2 < or 10 CM co o — rt Pl PI IX) 1 10 p~ in r- pi co 0) 01 co in Ol CO o •— co CP oo 03 L0 CO CP CO 00 LO CO p » 03 P~ in co in in in t * CM OJ *~ co Ol »- Ol Ol Ol Ol Ol CO 00 00 CQ 00 oo 00 00 00 00 00 m 00 00 m CO m m 03 00 CO 00 00 00 co m in •<T CO OJ ro Ol PI in CO LO r~ 0) in o _ 00 Ol P) p- Cl o Ol CN CN *- OJ *- Ol Ol OJ OJ a. in o ~ Q x 5 < o o o 6 z o o O 11 a UJ CL v> a a. 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UJ O Q i n cc z < o U J aa CMCOOIOIPIPICMCMCMT OOOOOOOOOO + + + + + + + + + i L U U I U J U J U J U J U J U J U J U J i n o o ^ O O c o c o i n o o O Q O c o Q Q c o o j o i i n p - O Q i n o O c M c o O P ~ t O O U O O C l B N -K O O n o O M n i s i n i ^ O T i n O O O O O l O Ol — C O C O ' - ' - C O O l L D P l Ol CM 0 O + + UJ UJ in o> 01 *r co io r- TT in o — to 00 00 Ol Ol 0 O + + LU UJ 01 — in T 0 oi CO OJ •s- in 01 01 oi oi 0 O + + LU UJ 01 — in O 0 oo co ••-•<r in 01 01 oi CM oi O O O + + + UJ U J UJ oi O 01 LP CO *T in CM ID 0 O <T in o in in LO 01 o CO OJ CM O O + + UJ UJ in m LO — r-LO in in co CO OJ 03 00 CO CO CO O O O O O O O O O O O O O O O O O O O O O OJ oi OJ O O O + + + LU UJ U J CO 01 00 p~ — 01 co in CM co o t~ r- co oi co v CM co o p*-U J U J U J U J U J U J U J U J I T c o O c o c o i n o O i T C i O c n o i o i Q O i C O O I O T P O C O O O J O o i O i n r o o i O O i - o i O ^ n c i O O i ' o1 o o o O O o o O O O O o O O o o o CO CO CO CM OJ CM Ol OJ OJ OJ CM OJ OJ CM CM CM CM CM O O o o O O o O O O O O o o O O O O + + + + + + + + + + + + + + + + + + UJ UJ UJ UJ UJ UJ UJ UJ UJ UJ Ul UJ UJ U J UJ UJ UJ U J O O O CO CO 00 CO p- L0 01 L0 co in co un O Q o 01 oo 01 —— CO L0 •*r CO CO P- in OJ CO Ol P l O O o 01 CO 01 p~ co CO O T p~ Ol O oo 01 01 Ol Q o 01 O 01 in L0 CO O 01 o r- co CO oi oo o o o 01 CM 01 CO in LO co CO ID CP in T 01 *T o o o L0 -- CM CO LO 0) T— o in L0 O *- oi oo Ol 01 O) L0 CO 01 01 p- p- 01 Ol Ol o o o O O O o O O O O o O O O o o o CM OJ 0 o + + UJ UJ I P- CM in in i 1 oo co CO — UJ Ol CM o o CM TT o o — CO 0 -01 in co 1 ot CM cc 01 CO GO r* LP in co o oi r- co L0 01 CO CM OJ LP co p-OJ OJ OJ O O O + + + UJ UJ UJ — LP CO r~ O O) in co in in LP LO CO P- CM co O in p- co LO 1 CM O O ID CO T 01 CM CO CO 03 in i O -=r CO f -Ol CM O O + + UJ UJ CM LP r- in in — p» ID CO 03 P» CD I- p-OJ OJ O O + + U l UJ Pl LO p- in O -OJ UJ rr CO in oo r- i -CM CM 0 O + + Ul UJ n co co oi 01 in O) O in oi oi O p» co CM CM Ol O O O + + + U J UJ UJ CM Ol 1 CM CO O O oo O oo co — O OJ in — CM *-ID P- P-CM CM 0 O + + UJ UJ 01 -CO CP co O 01 t- Ol o oi P- LP CM CM o o + + UJ UJ ID ro O co O in LP T tD O O UJ r- L0 Ol OJ Ol O O O + + + UJ UJ UJ CO LP O co in co 01 — ^ in UJ in p- co *-*- 03 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 OIOJCMOI — O C M C M C M O I O I C M O I C M O I 0 0 0 0 0 * - 0 0 0 0 0 0 0 0 0 + + + + + I + + + + + + + + + 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 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 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 cpo io i in incoo ico ino iTroo icoo i u j ^ ' - p ~ i n O ' - u 5 L o o i c M c o p - i n p -U J C 0 P - O i n ' - r T 0 1 P ~ 0 1 P ~ O l C 0 P - C 0 i n o i T j o i n o i p - o o i n o i L P o i p - ^ p * T r n i D i n i n o i u j i n r - - o i u > u 3 ^ ^ , * r Ol Oi O O + + UJ UJ 03 t -CO CO in oi ^ , 03 *-p~ oo CM Ol I O O ' + + U J UJ I CM CO I CM in i •fl- p~ I * - CM I o co i LP CO I CM in i O O O O O O O O O O O O O i Ol CM CM CM CM CM CM O O O O O O O + + + + + + + UJ UJ U J UJ UJ UJ U J Ol in !- co in co p~ p- T 03 CO Ol Pl oo Pl CM 01 UJ O CO CO in P- CM CO in Ol O) p- CO CM LP co Ol •sr UJ p- in Ul CM CM CM in *r LP O o O O O O O p - L o c n c o t ^ o i n c o i n c o r - c M o o i n o i i n T r p - o o i n p j O C 0 O 0 1 C 0 Q 0 0 C M C 0 r > 0 0 C M i n L 0 P ~ — P - t c O O O C M o i o B i n o i T r o o i c o o i i n o i ' j - p - o i c o ^ c o p i c M o i L O — c p i p o i o i o o p - i n p - — v - c i i t O i o n ^ u i S ( M oi oo oi T in oi oi P~ in UJ oi oi in in LP — ' - p ~ i n ^ . .^ i P C ^ O i n O T O i p - o i P - o i c o r - P i c o — p - r o r - e o i n c M o i p - c o o i £ S J ^ 9 m o ^ ® m r o w M ^ ^ ^ ^ 0 3 < c n c p c o » Q ^ ' u ^ ^ l ^ ^ n UJ in io in p. oi CP co * ^ ^ ^ ^ CM in in in CM CM OI in ? 3J < z ao p- 03 01 O CM CO f in UJ p~ r- r- CO oo CO 00 CO CO Ol CM Ol CM OJ CM CM OJ CM oo 00 00 00 00 00 00 00 m OQ CO 01 O 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 of the bryophyte syntaxa. 203 Racomitrio - 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 Tabular s o r t i n g t a b l e s and c l u s t e r analyses f o r the four major groups of bryophyte vegetation r e l e v e s . 226 BRYOPHYTE ASSOCIATIONS FROM EXPOSED, DRY ROCKS. Releve-numbers " t r a n s l a t i o n " l i s t . ( in the l e f t column are the numbers as given by the c l u s t e r ana l y s i s ; in the r i gh t column are the f i e l d releve-numbers as l i s t e d in the separate a s soc ia t ion t ab le s . ) 1 = 9 68 = 249 39 = 155 3 = 11 71 = 254 23 = 116 6 — 15 72 = 255 24 = 117 52 — 179 74 = 257 42 = 169 26 = 119 76 = 262 43 = 170 53 = 181 4 = 12 29 = 122 10 26 9 = 22 30 = 123 62 = 243 11 = 27 47 = 174 64 = 245 12 = 31 48 = 175 63 — 244 13 = 32 46 = 173 115 = 331 40 = 157 65 = 246 117 = 333 41 = 158 66 = 247 83 — 288 81 = 286 59 = 240 84 = 289 82 = 287 60 = 241 85 — 290 89 = 294 61 = 242 21 = 65 91 = 296 49 = 176 8 zz 17 90 = 295 50 = 177 116 zz 332 92 = 297 118 = 334 51 = 178 93 = 300 119 = 335 36 = 134 94 = 301 122 = 338 37 — 135 123 = 341 120 = 336 54 = 182 125 = 343 121 = 337 55 — 222 124 = 342 104 = 311 56 — 223 98 = 305 105 = 312 20 = 46 100 = 307 108 = 315 128 zz 347 99 = 306 106 = 313 126 - 345 101 = 308 107 = 314 129 = 348 7 = 16 25 = 118 127 = 346 17 = 38 31 = 124 2 10 16 = 37 102 = 309 69 — 252 19 = 44 114 = 330 58 — 236 18 = 43 103 = 310 5 = 14 86 = 291 32 = 125 75 258 88 = 293 80 = 266 78 = 264 87 = 292 70 = 253 79 265 109 = 316 95 = 302 22 = 68 110 = 317 96 = 303 33 — 131 111 = 318 97 = 304 35 — 133 112 = 319 27 = 120 34 zz 132 113 = 320 28 = 121 57 — 235 14 = 33 73 = 256 77 — 263 15 = 35 44 = 171 67 = 248 38 = 154 45 = 172 227 BRYOPHYTE ASSOCIATIONS FROM SHADED, HUMID ROCKS. Rel eve-numbers " t r a n s l a t i o n " l i s t . ( 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 t h column are the f i e l d releve-numbers as l i s t e d i n the separate a s s o c i a t i o n t a b l e s . ) 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 = 231 82 = 339 83 = 340 54 = 102 67 = 225 68 = 226 18 = 30 24 = 47 42 = 72 40 = 70 41 = 71 61 = 138 11 = 20 21 = 41 22 = 42 43 = 73 58 = 114 47 = 82 16 = 28 25 = 49 27 = 51 30 = 57 76 = 298 77 = 299 66 = 214 37 = 64 59 = 136 60 = 137 19 = 39 20 = 40 56 = 112 57 = 113 78 = 321 81 = 324 79 = 322 80 = 323 12 = 21 15 = 25 71 = 237 72 = 38 73 = 39 228 BRYOPHYTE ASSOCIATIONS FROM SILICEOUS ROCKS ALONG STREAMS. Releve-numbers " t r a n s l a t i o n " l i s t . ( 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 separate a s s o c i a t i o n t a b l e s . ) 1 = 6 53 = 197 84 = 250 86 = 259 54 = 198 85 = 251 88 = 261 55 = 199 42 = 186 24 = 104 63 = 207 45 = 189 25 = 105 89 = 267 28 = 108 26 = 106 90 = 268 40 = 184 27 = 107 70 = 215 29 = 109 80 = 229 95 = 275 30 = n o 6 = 66 71 = 216 65 = 209 36 = 129 74 = 219 39 = 183 94 = 274 73 = 218 41 185 38 = 159 75 = 220 43 = 187 97 = 235 72 = 217 44 = 188 98 = 326 76 = 221 77 224 33 = 126 2 = 48 34 = 127 32 = 115 35 = 128 11 = 81 66 = 210 59 = 203 67 = 211 12 = 83 78 = 227 102 = 344 79 = 228 47 = 191 68 = 212 48 = 192 69 = 213 50 = 194 15 = 86 58 = 202 19 = 90 61 = 205 17 = 88 7 = 76 16 = 87 8 = 77 18 = 89 31 = 111 96 = 276 10 = 79 101 = 329 13 = 84 62 = 206 9 = 78 64 = 208 21 = 92 91 = 269 20 = 91 93 = 271 49 = 193 92 = 270 22 = 93 81 = 232 23 = 94 87 = 260 57 = 201 83 = 234 60 = 204 99 = 327 3 = 52 100 = 328 4 = 53 82 = 233 5 = 56 51 = 195 37 = 130 52 = 196 14 = 85 56 = 200 46 = 190 229 BRYOPHYTE ASSOCIATIONS FROM CALCAREOUS ROCKS. Rel eve-numbers " t r a n s l a t i o n " l i s t . ( 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 separate a s s o c i a t i o n t a b l e s . ) 1 = 36 13 = 163 14 = 164 15 = 165 11 = 152 12 = 153 16 = 166 17 = 167 18 = 168 2 = 143 3 = 144 8 = 149 4 = 145 7 = 148 9 150 10 = 151 5 = 146 6 = 147 22 = 280 23 = 281 24 = 283 19 = 277 20 = 278 21 = 279 25 = 283 26 = 284 27 = 285 2 3 0 1 1111111111 12345678901234567890 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 PHILFON 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 5-4--4--1 + 1 1 1 " 1 3 3-"- 1--- 43222- -323 2--S+ 2-3 + --3 + ---1 2 — 21-42-1--3 4 1 + + -1 -51 -21 — -. _-, - + -- + -— 1--- + -1- + -++ — - 1 -22-1 21-1 1 32443 _ _ 3 3 S--323 4--134 1-111 — 322111--_ --23-2 1211 43233321 1 1—2+1 1 1 + 1---2322 __ +31--- 1212 _ 2 2 1  """-—14 3-121 1-1-+1111 2 _ — 3 - + 1 + -1 11-— 11 — 4+13-2 +1 --++++-12++ + 31-2-2 +1 ---32-2-51344321 1 1 2 1 131 1 22 + 1-2- 11----+ --1 1 1 1 32 1 + ---+1135212-71 1 11-122 11 — ----412+ 211--2 + 2 1 — 1-1-+1 31-2-2 42 — 3-1 ---2-32222 22 — 322 31211 111 21 1 1 1 11 1-+ 1_+1++__11 t 2+ ++ 1-2221 + 21-111 2 1-+1 112 ---32 --- 11-1---1231 11 11---222114-112 1 +++ 1 + 1 13 1 + 1 +1 _323 12331 — 1 21 + -—1 + 211 121 1 — ---21-121 1 121 1 2122++21 + 2232 ' + -1++++--2 -4 1  H 32--2+21-3 1-+-1 14 + -1 13-3---2 1---3 _ 4 * _ + 2 5 _ . 1 - • + + 2 - . 2 + 1 124 + 3  — - 1 2  1  4  - -1---1 1 —1-3 — ---1-+1-1312+-•--31 •1 .1--112—1 +1+11-1--+1--+11 — 411 •1-21-11 11 — 12+11 1211 1 — - -+ 1131 + -21122+12 223--2+3 12-32-2 11--21 1 ' 43-1 . 1 - - 1  •1 ++21+ --1-2-12 +12 111 " 1 — 1--•132 — . + 1 +-—1-111 — 1 + 1 — 1 1--+2-3  - + ---+2---+1+ 4324 1--1 •13342--11-1 -++-+ -2-22 -32- •12211--3 -11111- •12--43333-21-41 12552233-422431-2-232-11311--23121 + 21 11-3+1 1-•11-2221+-1 1 -•1+2-43__ 1_2+- + ---24223333-3 — -2423323--44-2144--423 — -4411 —111--2111--31 1 21 1 1222-•11 1+1 + -++-1--1+1411-+-13313 •1-. 23 .1 -123231-1—22121-31321-44 11- 1--22-24— 233-31 21233 3223-311 11 — •12-3 23 1-21 — 1 4 — 1 1 4 11--111- •1-1-1221 112131 11-41 1122+--2 — --13 — -1++-•11 — 1--++1-•11 + 1-•11- •11-•11-•1--+-•1-1 — -31-•1-• 1--•11- •12-•1-1222-+ 2--+-++1- •141 — •1- •12--22--3 — -31-•113--3-121111 — 1 11111 —-221-._ 1 1 1 1-221 12111 1---11 -31 — 1--1 — -1 + --21-•111-•1 — -3-1--2-•111-•11- -211- 1+--1-•1 1 - 2 : --2-2-12-321 • 1 134-•1 + 11 •1--221- •1-1-1 22++-1+++-12 — -31----4333--J -11+3-1-11- -2 — -1 + -• — 2211-..-—1++ — — — - 1 - 1 --31--1 — -1---33- •11-•11--1--•13--22+ -111 ---1 1 1 313 -11-1-2 — — --11 1 --11 :>.12-11 132-122—1-211--+1--11-1211-1--+13 1311---2-4 1 1 + 2 — -•12-11-•1+->0 •1-+-•11+----+11--+1+-0 ° p -m \ 231 2 11111111111111111111111 n 11111 1 1 1 1 1 1 1 1 112222222222333333333344444444445555555555G666666GG677777777778888888888999999999900000000001 111 11 1 1 1 12222P22222 12345G7890123456789012345678901234567890123456789012345678901234567890123456789012345G7890123456789012345678901 234567890123456789 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 31-2-2 42 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 1 2 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 1 1 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 __ 3 4 3333 41 12552233-422431- 2 r-' 1 1-2221+ 1--3 4 1 + 33 -3--323 221 3221 1 1 • + 31 1212 32-2-51344321 1 1 2- 1 131 1 11 1 1 1 13 --+ 3 4+13-2 +1 --++++- 12++ + -- - 1 + •-- + + --++ + . u — ! : 11 + ---22+- •1- 2-11-•12 — -- 1---2+3--24223333-3-v 44 -51-12-32-2 11-44-214 ! • 33 -21 — ,_. 4. -11 -2111 •--3-121 1 1 1--1--14 3-121--1--3-1-1-•1-1222- •11-. 1 + 2  -11111---221 1-1-+1111-1 12--1 -, _ 1 1  1+++1 + -++21+ -1+1411-+ -3+- -2-3--+1+11-- + -++1 - •141 — •1-•1-+2-2- -++1-•1 13-•1-•1111-221-• 132--+1 11 1 + 1--1 1 +1 + -•11 — -32--2+21--23-3-1--+11--1 + 1-•1- -411 — •122++-- + 2-121111 11-223 -4333--+ +-•1 + -•124 + - •11--32- 1221 1--2 2--—112 -221 1 - in 11+3-1 — -111 1 1---3 13 •111 3-1--2--211--2-2-12-22 1 1 •11-1211-1-•1-1-111--11+1--321- •134--23 1 11-•11 212-•111-1++ 1--4324 -11-1 •1 + 11 -+1+++-•11-1-2 1 1 132-• 1 1 + •112 -+12--31--2+--2-22 -111-•1++--33- • 11 •122--1-•1311-•11+2-1--1 - + 2-•22+-•111--21 1 --+13-+ -1--21--+1 1 -+1 + -+1-12--23--1 — - - - - - — ' i T --11--13--+1---2-4-•12-•11-• 1 + -K J ( !* c<-i'( >•• o4"4 Do ID u P\V>7 v >AVI <CLUSTEI? OPTION=MINVAR FUNCTION" 2EUCLIDEAN R E L * * C A S E S = 1 - 1 2 9 VAR=1-152 MAX=*> C L U S T E R A N A L Y S I S CASES=CASE/f : 1 - 129 C A S E W I S E N=129 ALGORITHM-MINVAR DI STANCE = E U C L I D E A N U S I N G : 1.V1 . 2.V2, 3 . V3, 4.V4, 5.V5, 6.V6, 7.V7, 8.V8. 9.V9, 1 0 . V 1 0 . 1 1 . V 1 1 , 12.V12. 1 3 . V 1 3 . 14.V14. 15.V15, 1 6 . V 1 6 , 1 7 . V 1 7 , 1 8 . V 1 8 . 1 9 . V 1 9 , 2 0 . V 2 0 . 2 1 . V 2 1 , 2 2 . V 2 2 , 2 3 . V 2 3 . 2 4 . V 2 4 , 2 5 . V 2 5 , 2 6 . V 2 6 , 2 7 . V 2 7 , 2 8 . V 2 8 , 2 9 . V 2 9 , 3 0 . V 3 0 , 3 1 . V 3 1 . 3 2 . V 3 2 . 3 3 . V 3 3 , 3 4 . V 3 4 . 3 5 . V 3 5 , 3 6 . V 3 6 . 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 , 4 2 . V 4 2 , 4 3 . V 4 3 , 4 4 . V 4 4 , 4 5 . V 4 5 . 4 6 . V 4 6 . 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 , 5 5 . V 5 5 , 5 6 . V 5 6 , 5 7 . V 5 7 , 5 8 . V 5 8 , 5 9 . V 5 9 , 6 0 . V 6 0 , 6 1 . V 6 1 . 6 2 . V 6 2 . 6 3 . V 6 3 , 6 4 . V 6 4 , 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 . 71 .V7 1 , 7 2 . V 7 2 , 7 3 . V 7 3 , 7 4 . V 7 4 , 7 5 . V 7 5 , 7 6 . V 7 6 , 7 7 . V 7 7 , 7 8 . V 7 8 , 7 9 . V 7 9 , 8 0 . V 8 0 , 8 1 .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 , 9 4 . V 9 4 , 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 , 1 0 0 . V 1 0 0 , 1 0 1 . V 1 0 1 . 1 0 2 . V 1 0 2 , 1 0 3 . V 1 0 3 . 1 0 4 . V 1 0 4 , 1 0 5 . V 1 0 5 . 1 0 6 . V 1 0 6 , 1 0 7 . V 1 0 7 . 1 0 8 . V 1 0 8 . 1 0 9 . V 1 0 9 . 1 1 0 . V 1 1 0 , 1 1 1 . V 1 1 1 , 1 1 2 . V 1 1 2 , 113.V1 1 3, 1 14.V 1 14, 115.V1 15. 1 1 6 . V 1 1 6 , 1 1 7 . V 1 1 7 . 1 1 8 . V 1 1 8 . 1 1 9 . V 1 1 9 . 1 2 0 . V 1 2 0 , 1 2 1 . V 1 2 1 . 1 2 2 . V 1 2 2 , 1 2 3 . V 1 2 3 . 1 2 4 . V 1 2 4 . 1 2 5 . V 1 2 5 , 1 2 6 . V 1 2 6 , 1 2 7 . V 1 2 7 , 1 2 8 . V 1 2 8 . 1 2 9 . V 1 2 9 , 1 3 0 . V 1 3 0 , 131 .V131, 1 3 2 . V 1 3 2 , 1 3 3 . V 1 3 3 , 1 3 4 . V 1 3 4 , 1 3 5 . V 1 3 5 . 1 3 6 . V 1 3 6 , 1 3 7 . V 1 3 7 . 1 3 B . V 1 3 8 . 1 3 9 . V 1 3 9 . 1 4 0 . V 1 4 0 , 14< .V141, 1 4 2 . V 1 4 2 , 1 4 3 . V 1 4 3 , 144.V144, 1 4 5 . V 1 4 5 , 1 4 6 . V 1 4 6 , 1 4 7 . V 1 4 7 , 1 4 8 . V 1 4 8 , 1 4 9 . V 1 4 9 , 1 5 0 . V 1 5 0 . 1 5 1 . V 1 5 1 , 1 5 2 . V 1 5 2 CASE/*1 1 3 6 52 26 53 10 62 64 63 1 15 t 17 83 84 85 21 8 1 16 31 36 37 54 55 56 2 0 128 126 129 127 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 4 0 41 81 «!2 89 91 9 0 92 9 3 ' 34 123 12T. 124 98 100 £ 9 101 7 17 16 19 18 8 6 88 87 109 1 10 1 1 1 i 1 2 1 13 14 — T 5 -38 39 23 2 4 42 43 29 3 0 47 48 46 6 5 6S 59 6 0 61 49 5 0 1.13 1 19 122 "1.20 121 "104 105 108 106 107 25 31 102 1 14 103 32 8 0 7 0 95 96 97 27 28 73 44 4 5 1 1 2 2 8 4 9 7 9 1 1 1 0 9 8 8 8 8 7 4 2 0 7 7 6 7 0 4 6 6 2 1 5 4 4 1 9 8 3 3 3 3 9 7 6 5 2 1 1 0 9 8 1 1 4 3 1 1 2 1 9 8 7 6 5 4 3 2 4 - ~ + - • + - • + - • + - • 4 I • I - -+ + + + + I + • I-+ I + I " " -+ - I + - I + + I I-I - I -I + J + I-+ + + + I + j -+ 1 + 1-+ -I • I - I I-- - - I ••-I I-- T 4> - - I - - I -+ + + 1-+ -+ -+ -+ -1 1 -Cn - - i i -i - - i +--+--CD D I S T A N C E S 1 4 5 9 1 1 1 1 1 1 2 2 2 2 2 3 3 3 3 3 4 4 4 4 5 5 5 5 6 6 6 6 7 7 7 8 8 8 9 9 2 4 6 7 8 9 2 5 7 8 9 1 3 4 6 8 1 2 5 9 0 2 8 9 0 2 7 8 0 4 6 2 5 8 2 7 O O O O 0 0 O. 0 0 0 0 O 2 4 5 2 5 1 8 0 0 7 3 0 0 4 4 2 7 1 5 5 5 7 0 0 0 0 5 8 4 0 4 0 0 0 0 4 2 0 0 0 5 2 8 8 8 2 9 5 6 0 2 7 6 5 6 O 3 6 5 8 1 7 6 6 4 7 7 4 7 7 0 8 0 0 8 7 1 5 0 7 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 8 5 3 9 5 7 4 7 7 1 3 4 5 7 6 1 3 4 7 4 0 7 3 9 7 3 5 7 1 0 < F I N I S H > COMMAND USES LOADER STORAGE MONITOR V M ( P G S ) C P U ( S E C ) E L A P S ( M I N ) COST READ 1 .016 .084 135 6 4 9 .0 $ 30 C L U S T E R 1 .020 .013 189 2. 306 . 1 $1 32 T O T A L S .036 .097 177 2. 9 5 5 . 1 $1 62 E x e c u t 1 o n T e r m 1 n a t e d 0 8 : 5 5 : 2 5 T = 2 . 9 9 3 RC=0 $1 .65 $ S I G 233 19 ANDRRUP 62 RACOSUD 26 LOPHVENV 53 DIPLTAX 3 MARSEMA 17 LOPHSUD 55 GYMNCON 2 0 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 DICRSCD 15 CYNOdEN 23 CLAOBOL 85 GRIMTOR 107 ANTHdUR 30 CEPHBYSA 34 CERAPUR 11 MARSSPA 44 ANASMIN 49 ANDRROT 7 0 BARTITH 28 HEDWCIL 36 LOPHINC 51 DICRFUS 104 KIAEBLY 109 NARDdAP 125 HOMANEV 89 SCHISTR 94 AMPHMOU 98 DICRHET 112 PHILFON 133 PSEUATR 39 BARBHAT G9 PSEUBAI 9 9 P T I L C A L r"lT6 TRITQUI 128 PLAGP IL 21 HYPNCIR 33 POLYuu^ 45 LOPHCUS 46 CAMPATR 48 CHANFIL 6 5 ISOPELE 101 DIPLALB 108 MARSCON 126 PSEUCAL 129 PTERFIL 148 PLAGASP 8 DICHPEL 38 BRYUCAP 68 POHLLUD 4 0 RACOLAN 4 1 PLEUSCH 5 0 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 dUNGEXS 150 POREROE 151 HETEPRO 5 CEPHTUR 7 SCLEOBT 18 CAMPPAR 97 AMPHCAL 27 CHANSET• 31 AULAAND 35 LOPHEXC 58 OLIGHER 59 BLINACU 6 0 RACOACI 64 LOPHWEN 105 BRACALB 106 PSEUPAT 71 dUNGOBO 78 DITRHET 79 OLIGALI 84 PORENAV 88 HOMANUT 91 RACOVAR 9 3 GRIMMON 1 15 KIAEFAL 12 1 RADUBOL 123 DOUIOVA 132 PORECOR 136 SCAPPAL 141 GRIMALPA 142 CAMP FRA 143 ARCTFUL-145 RHYTTRI 146 TIMMAUS 147 PTILCIL 9 POROBIG 29 DICRCRI 37 PREIOUA 43 BAZZDEN 57 MARSSPH 67 PSEUSTE 73 PELLNEE 77 CHILPOL 8 0 PLAGOED 82 SCHIPEN 86 DISTCAP 9 0 ATRIUND 95 BRACSTA 96 PSEUINC 139 NARDBRE 140 HYPNDIC 100 RACOFAS 102 BARBLYC 103 BAZZTRI 144 dUNGCON 114 MARSBRE 116 POLYSEX 117 POHLCAR 1 18 ANEUPIN 149 HETEMAC 119 BARBVIN 122 HETEDIM 152 ISOTSTO 11 • 1 11111 5 2 5 1 6 6 6 1 18882 1533555222221 1 3 6 2 6 3 0 2 4 3 5 7 3 4 5 1 8 6 1 6 7 45608697E2985589 - - 1 - - - 2 3 2 1 121 1 12 2 2 2 2 2 3 2 2 4 3 2 2 2 | 13 1 21 1 --4 1+221 3 1 1 1 1 3 ---1 11-2+11+ 1 - 2 1 3 1 1 — 1 — 3 - 2 + - 1 - 1-2 44322223| — 1-1 — 1 1 111 + 2 -I — + 2 + -1-1 + 1+ +-1 + 2--221 1 1 21 --2 2+1 — 1-11 — -5 4 4 4 3 3 + 3 3 2 3 2 1 3 4 3 - 2221 1 1 12| 1 - - - 2 1 2 + 1 3 2 2 + 4 3 3 3 3 3 2 3 1 11221 •1--+- -++--1-1 -+1 +-++) -21--1 + --12--+ — •-11 3-•1 1 — •1-1--3 — -+11 1 — --223+--1--3--+1-•111--2 12-•1-1 -32443] -+-11 • 1 + -111 •1-1--2222 -+--1-•13--+-11 + 1--1-•12-• 1 + -65 7 7 7 2 3 3 3 5 7 6 6 7 7 7 7 11 2 3 5 4 7 7 7 8 1 2 4 6 4 9 1 2 3 0 1 -1-321 1-322-1 1 1--13 -1-12 --1--11-+-112 --1 1 1 + -----4412 —-2-1--221 11 1 1 1 222 1 1 - 2 1 2 - 1 — 3 2 3 --3---1 1 11 H 2 1 - 2 3 1 3 4 3 3 4 4 5 5 3 3 4 3 •12---2-11-- - +++ •-- -23+1-11--21 3 - - 1 - 1 1 121 644-32.1 1-1-•1 + -3 4 - — , - - 1 31-111-1 1 1 1 1| 1 1 1 1 , 1 1 1 1 1 1 1 1 1 1| 1 1 1 14488899999222PO90I 1 1 1 1 8 8 3 0 1 111 1 1 3 3 2 2 4 4 2 3 4 4 4 6 6 5 6 6 4 5 1 1 2 1 3 4 5 8 9 b 4 2 3 9 0 7 8 6 5 6 9 0 1 9 0 3 9 2 0 1 4 5 8 G 7 | 5 1 2 4 3 2 0 0 5 6 7 7 8 3 4 129 1 0 2 3 4 3 5 4 B 0 9 1 | 7 7 6 9 8 6 8 7 9 0 - 1 1 1 2 2 1 1 1 -1+-1+ — + 1 - - -1111-1-C 5 3 4 4 3 1 1 1 1 3 1 -P--2-1 1 + -- 12 -111-- 3 - 3 — 1 2 1 -•1 14 1323--3-11--32- -+1 -1 + •11-1 •1+211-1--41 1433 34+---1 1331 -31 131 — - 2 -•11 12121 12l| 111-- - -212- -•11-132-•1+1 •121 -33 1 - 1 -122--22--211--+11 -++1 -31--21---2-1 11 12+1 111 1 + -13 131 1 + 12 •11-•13-b4-L _ _ 3 1 _ . „ 1 i 3 1 + _ - _ - 2 - - 1 1 1 3 2 11-11 1 + 2 + --12 1 2 5 2 1 3 3 2 1 2 3 3 — 1 3 2 2 | 14-22 21 111 12 1 1 3-321 |33 12 ++-+4+12| 1 1 •12 + -•-11---+1 +1 1 1 1+-++ 22| •1-2 11 - 1 - 1 2 C — 1 + 1 + 2 1 2 - - -1-- + --4443--31--4-+2-21+4---1-1-•1-21 11---11--13-- + 2- 12^ •111-12-12f -23 1 2 2 2 0 0 0 0 C I 2 3 0 1 0 3 8 7 9 9 3 2 2 7 4 1 i 2++ + 2121 - - 2 1 - 3 1 - 2 1 1 1 211 1 1-222+1 + -1 1- - 1 2 - 2 2 2 1 2 3 1 3 2 1++-+1 1 1 4 2 2 1 2 2 — 1 1 1 1 1+43323J 21-2+1 — 1 1 1 2 3 — 1 1+121 | 4 3 2 4 2 + 2 ? 3 4 4 4 2 3 — 1 3 3 3 1 1 1 2 2 2 ] ! 1--++2 1222- 1 — 4 3 3 32121 121 1 — -1 + 2 2 ( 3 4 4 4 1 4 4 4 2 1 3 3 2 1 3 1 1 1--1 1-+--1-12--+1-•11-2-1-1244433441--32+++21-•1 + 1 14--+2--1--++1-•11311--23-•122-•13234 -3212121-•112-•1 + 1--+1 + 1 1--32-•1 1231--+1 + -•111-23-•11-• 1 + -2 - 1 + - - 3 1 1 1 - 1 2 -312-12-1-3 3 3 1 -• 4 4 3 3 221 — 1--121 1-1 1-3-11-1 221-•-- 1 -211 -1 1 1 --2-1 1-• -a • 1-•1--+13 1-2-RC=0 $5.12 1 1 1 1 1 1 1 1 1 12222222222333333333344444444445555555555666668666677777777778 888 12345678901234567890123456739012345678901234567890123456789012345678901234567890 123 1 DIPLALB 2 RACOHETH 3 ISOPELE 4,.RHI ZGLA 5 POGOCON 6' DICRHET •T, BARTPOM . 8 . LEPIREP 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 PLAGLAE 22 DICRFUS 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 BLEPTRI 46 DIPLTAX 47 BAZZDEN 48 POHLCRU 49 CEPHBIC 50 ANASMIN 51 HERBADU 52 SCAPUND 53 GYMNOBT 54 ANACMEN 55 DOUIOVA 56 GRIMTOR 57 TRITQUI 58 PLAGPIL 59 FRULTAMN 60 EURHPUL 61 ANDRRUP 62 CHANFIL 63 MARSSPA 64 UAMEAUT 65 MYLITAY 66 BAZZTRI 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 CALYTRI 80 GYROUND 81 OLIGALI 82 POLYLON 83 RHYTLOR 84 POLYALPM 85 LOPHGIL 86 RACOACI 87 BRACPLU 88 ATRISEL 89 PHILFON 90 PELLNEE 91 DICHPEL 92 BRACFRI 93 JUNGOBO 94 DIPLPLI 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 PSEUTEC 109 DREPUNC 1 10 BARBHAT 1 1 1 RACOSUD 1 12 K I A E B L Y 113 DICRCRI 1 14 PTILCIL 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 ENCACIL 128 BRYUCAP 5432 1 2- + --+5S--2 332-12-2 4-+1 - - 1 ++ 2++22-333-42-2—1-1-243--1 + 21 + + -- 1 ++--2++ + + + - + + 2 +2 + -+ -•--44-22+433+ 3--+--1-214 32+24 1+1-2+2+11 1--1--12---•-- + +1-342 2++ + --11---11--2--- -441++--+41 1 + -2-1--341 -— 3 113-1142+311---- 1--3--2 ---1 1 + 2 1 + 2  53 3+-•11- 1 + 3 1--1 25 -243412333 31++4 +13---23 14 +1-3+31 1 + -2532 + 32 43132+1 1 323 34 2+ -32-2--1++1 1 122---33 1 121 43--2++2--4423--1--1- -31 -32--2+13+---12---1 + ----1--+--S+-2 - -++1+++12 - + 3+ 1-22-+ 3-+++-1-2--•1--- - + 3+ -++2++- 132 -3---3---21- -241 1 1-21 + --4--1-11--1+---1 1-2 1 11 11 ' 3412 1-21 1 12 -5-- 111----1 1 + 1-+11-- 1 4-15--22--21--++3-++-•1 143-122--43-U • — .++--+ + + 11--11 ++ ; -11 + 1 + +-3 + --43+-1 + 1 121 14221+ -+13131-4-43 214 + i -_3__ + 1 2+ 1--33 111 1 ++- + 1 + ++1 + 21 ' 1 ---1 + 2+-- 2 +1 --31 1 2-++ 1 — 3 2 : — 4 2 121-++ 23 1 1 1 + 1 R-.__ + :._ 1 . 4 : 1-12-1 22 1 -3-++ 22-332 324 1 _ 2 4 + -1 1 2 + + +1 111 1 , + ______ -3 2 — 3 1 : . 2 + + + ' -3 3 + +-+ _ 3  331 2 + 1 2 11 + 2  3 , 1 i 3 _ _ ++ 4-1 1 , i 2  2 ' 5-13 2 1 . 1 : . 2  1 21-1- + 3 1 : — 4--1 + 12 -- + 22-1-22 12-1 22 :::::::.;:::::::::::±:: -1-21 ! + _ 1 3 ,n L__ •1 32------1 1+11--1--1--+11,1--1332--•1 1.- — • -12 •1-1--1-I 11 SCAPAME 10 MARSEMA 16 PLAGASP 2 RACOHETH 3 ISOPELE 12 CLAOBOL 18 ISOTSTO 46 DIPLTAX 19 HETEMAC 1 DIPLALB 4 RHIZGLA 61 ANDRRUP 27 LOPHVENV 41 POREROE 47 BAZZDEN 59 FRULTAMN 15 HYPNSUB 26 LOPHCUS 45 BLEPTRI 48 POHLCRU 49 CEPHBIC 55 DOUIOVA 7 BARTPOM 9 NARDSCA 13 POLYALPA 29 PLAGUND 31 HYPNCIR 35 PLAGDEN 37 PLAGOED 17 STOKPRAP 50 ANASMIN 58 PLAGPIL 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 BARTITH 104 PSEUINC 8 LEPIREP 22 DICRFUS 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 DIPLOBT 52 SCAPUND 54 ANACMEN 68 RACOBRE 80 GYROUND 69 RACOHETA 70 HYPNREV 83 RHYTLOR 87 BRACPLU 88 ATRISEL 89 PHILFON 106 BARBLYC 112 KIAEBLY 116 HOMATRI 119 BARBREF 124 PLAGROS 126 SCHISTR 93 JUNGOBO 94 DIPLPLI 95 RICCMUL 96 GRIMHAR 97 RADUCOM 30 SCAPMUC 99 DRYPPAT 100 GYMNCON 36 PORECOR 102 CLAOWHI 38 PLAGINS 43 BARBFLO 53 GYMNOBT 57 TRITQUI 64 JAMEAUT 108 PSEUTEC 73 HOMANUT 110 BARBHAT 7 7 POHLPRO 7 8 CEPHPHY 113 DICRCRI 114 PTILCIL 115 STOKPRAS 79 CALYTRI 1 17 PSEURAD 81 OLIGALI 82 POLYLON 85 LOPHGIL 86 RACOACI 122 THAMNEC 90 PELLNEE 91 DICHPEL 125 TIMMAUS 92 BRACFRI 127 ENCACIL 128 BRYUCAP 1 1 1 1 1 1 1 1 1 12222222222 '33333333344444444445555555555666666666677777777778 1234567890123456789012345678901234567 9901234567890123456789012345678901234567890 888 123 --31 13- 1 142+31 1-3 1 1- 1 + 3-2434 12333-2532 + 32 43132+1 1 33 1 121 43 1 -+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 1 1 1--3--2 3 •- 1 323 34 2 + 2 + + 2--4423-- 1 - - 1 j-31 12 1-22- + 3 1 — 21 241 1 1 34 12 1-21 j--1 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 + + + • 1 2+++1 1 11 1 4_ 1 5.. + + + 1 2 1 , 1 4 3  4 2 3 12 2 2 111 + +:2- + +13131-4-43 214 + j 1-3-++ 22-332 324 1 i 53 1 3 4--1-1 1--1 + -|--1 11 + 1-+11 22 --• 2-2 ++-- + + + 1 1_ - ! 1 ++ i:  :++- + 1 + ++1 + 21 121- + + 23 1 1 1 1 + + __2 441++ 3--21 _1 1 31++4 +13 : 1 1+2 1 + 2- + 2- + :  ++3_ + + _ — 43 . + 2-- + 3 2--1--3 32 21-1 4 + 21 + --2-1 1 • +1 2+ 312 1--+ + -- + 3+ , + 2-j 1+2+ 2 +1 ' 4 1-12-1 -22 2+ 1 211 + • 1-- + 2 21 + -2 1 + 1-----1 5--1 1 1 1 2 + + _ + 1 2 2 . .4_-4 + + _+ 2---1 + 212' _2 21 1-31 1--- 2-++ 2---++ + 2 : 21-__ 2 + 1 t , -3 1 1 l l 3 - -•1 21-1- + -22-1-22-+ + 1 ++ 1--++ + 2 :--2 1--1 1--2 4 '• + -1> -34 14 -2--2 1 1 + --.-1+++ -1 + 1 1--+11 1--133 2-1_. 1 J ---1 — --32 -1 _ 2--++---12-1 1-21---•13 -2-11-.4 2-2 1--3 --•13-•12--22-•11--32 1-12 1 . 2 1  2 : .__1 .__2 -• 2 " — I : : - : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : u> __„,____!„_ 2 - \ + . _ + •_ 5 + 2  :-- 1--1 _ •!_ 1-RC=0 $1. <CLUSTER OPTION-MINVAR FUNCTION = EUCLIDEAN REL = MAX=*> * CASESM-83 VAR=1-128 CLUSTER ANALYSIS CASES = CASE/>': 1-83 CASEWISE N = 83 ALGORITHM=MINVAR DI ST ANCE = EUCL. I DE AN USING: 18.V18 34.V34 50.V50 66.V66 82 . V82 98.V98 1 1 I . V 1 1 1 , 124.V124, 1 . V 1 . 2 19.V19, 35.V35, 51 . V51 , 67.V67. 83.V83, 99.V99, V2, 3.V3, 4.V4 20.V20, 21.V21 36.V36 52.V52 68.V68 84.V84 100.V100, 112.V1 12, 125.V125, 5.V5, 6 22.V22, 38.V38, 54.V54 , 70.V70, 86 . V86, 101.V101, 102 .V6. 7.V7, 8.V8, 23.V23, 24.V24, 37.V37 53.V53 69.V69 85.V85 113.V1 13, 126.V126, 114.V1 14, 127.V127, 39.V39, 55.V55, 71.V71, 87.V87, V102. 115.V115 , 128.V128 LO 9.V9, 10.V10, 11.V11, 12.V12, 13.V13, 14.V14. 15.V15, 16.V16 17 V17 25.V25, 26.V26. 27.V2"? 40.V40, 41.V41, 42.V42, 43.V43 56.V56, 57.V57, 58.V58. 59.V59 72.V72, 73.V73, 74.V74, 75.V75 88.V88, 89.V89, 90.V90, 91.V91 28.V28 44 . V44 60.V60 76 . V76 92 . V92 29.V29, 30.V30, 31.V31 45.V45, 46.V46. 47.V47 61.V61, 62.V62 77.V77, 78.V78 93.V93, 94.V94, 63.V63, 79.V79, 95.V95, 32.V32. 33.V33, 48.V48, 49.V49, 64.V64, 80.V80, 65.V65, 81.V81 , 96.V96, 97.V97, 103.V103, 104.V104, 105.V105, 106.V106, 107.V107, 108.V108 109V109 1 1 0 V 1 1 0 116.V116, 117.V117, 118.V118, 119.V119, 120.V120, 121 .V121 , 122.V122, 123.V123, 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 8 8 7 7 7 7 2 0 9 7 6 2 7 6 0 8 6 6 7 5 5 5 4 2 0 9 4 4 3 3 3 0 9 6 9 8 7 6 5 4 3 2 1 + -+ -+ -+ -+ -+ + + 1 + 1 + + + -I + -+ -+ -+ -+ -+ -+ -+ -+ -— I - -1 0 I s T A N C E S 3 6 8 9 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 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 5 5 O 0 O O 0 2 0 5 O O 0 7 8 5 3 O 5 5 0 0 0 O 0 2 0 5 O O 5 7 0 5 O 0 1 8 9 0 6 0 1 1 1 1 1 1 1 1 1 2 2 4 0 0 1 3 4 5 6 8 8 3 3 0 4 6 3 6 1 1 1 0 9 6 9 9 5 3 8 O 6 4 7 6 5 3 1 6 O O 3 0 9 9 <FINISH> COMMAND USES LOADER STORAGE MONITOR VM(PGS) CPU(SEC) ELAPS(MIN) COST READ 1 .016 .055 1 10 .376 .0 $ . 10 CLUSTER 1 .020 .012 135 .867 .0 $.30 TOTALS .036 .067 128 1 . 243 . 1 $.40 11 SCAPAME 10 MARSEMA 16 PLAGASP 2 RACOHETH 3 ISOPELE 12 CLAOBOL 18 ISOTSTO 46 DIPLTAX 19 HETEMAC 1 DIPLALB 4 RHIZGLA 61 ANDRRUP 27 LOPHVENV 41 POREROE 47 BAZZDEN 59 FRULTAMN 15 HYPNSUB 26 LOPHCUS 45 BLEPTRI 48 POHLCRU 49 CEPHBIC 55 DOUIOVA 1 BARTPOM 9 NARDSCA 13 POLYALPA 29 PLAGUND 31 HYPNCIR 35 PLAGDEN 37 PLAGOED 17 STOKPRAP 50 ANASMIN 58 PLAGPIL 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 ITH 104 PSEUINC 8 LEPIREP 22 DICRFUS 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 71 AMPHCAL . 84. POLYALPM 105 PLAGROE 109 DREPUNC 111 RACOSUD 123 HOMAFUL 5 POGOCON 23 RACOCAN 24 LOPHHET 44 POLYFOR 76 DIPLOBT 52 SCAPUND 54 ANACMEN 68 RACOBRE 80 GYROUND 69 RACOHETA 70 HYPNREV 83 RHYTLOR 87 BRACPLU 8 LEPIREP 8 LEPIREP 89 PHILFON 106 BARBLYC 1 12 KIAEBLY 1 16 HOMATRI 1 19 BARBREF 124 PLAGROS 126 SCHISTR 93 JUNGOBO 94 DIPLPLI 95 RICCMUL 96 GRIMHAR 97 RADUCOM 30 SCAPMUC 99 DRYPPAT 100 GYMNCON 36 PORECOR 102 CLAOWHI 38 PLAGINS 43 BARBFLO 53 GYMNOBT 57 TRITOUI 64 JAMEAUT 108 PSEUTEC 73 HOMANUT 110 BARBHAT 77 POHLPRO 78 CEPHPHY 1 13 DICRCRI 114 PTILCIL 115 STOKPRAS 79 CAL Y TRI 117 PSEURAD 81 OLIGALI 82 POLYLON 85 LOPHGIL 86 RACOACI 122 THAMNEC 90 PELLNEE 91 DICHPEL 125 TIMMAUS 92 BRACFRI 127 ENCACIL 4463452232231 355 1177 1436633351 6 1645 124565282363894P61383745441329855B6990490702347884201 1| -- 1 142 .-+1 --2- + + + -+ + 2 33 2 + - + -2- 1 1 + 1 ---- 1 32243333+ 2+323122-1--- + - - + H 1 y 542443523-23+H + - 1 + + + + +111 1 13343124335231 |45332332244 123322 +2+-2+11+-1-2 -- 1 1 + -2422434|12 1-2 1-1 1+++1--: + + +i 1 1 1 — 3 if I + 1 + 1 1 _ _ -1-2-++ •I3243 + --2--2 + - + - + 1 1 + - 1 1 1 1 -34243 1| + 1- + + --1+2--3-2-+ 2--2 + -•1344 -323--2++-•1--3-- 1 1 --2--+ 2--21--2 + --2+-13214-31-1--1 11-1 + 32 f 1 1 — 1 -3232^ + + + 1 1-- + + + 331 1--1 + t 1-4312+ -2-22C4333 124431332 2|-1 1-- 22- 1234- + -- 1 1| 1-1-2-1+ •1-2-• 1 1 1 - -- - 1 + 1 -P34--3-- 1 + 2----+-3-• 1 1 --+11-- 1---. + -51-11-11-7885661244461122454 -|2 1 13 + -3 1--2+-+ + + 34-- + + ---422312J -2334 •11- - + -2 •1- 333-| - + + - 1 1 + 2-2 21 1 1 + 4421---334 +1-1 + - -+--2-r - 2 - - -1 21-21-D43 P 332--2 + 122-12 . 1 1 + + -2 h 1 + 2 1-4  1 — 11- M + 2-1 •1--1- --2--22-• 1 + -•1 1-133 •1 12--+13--42--3+--22-•12-12237763561255787811 777 11238765706767909067819025 123 1-+ ! 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SCAPUND R/ICOACX UL NGPUM PELLNEE NARDSCA 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 BLEPTRI 21 BARTPOM 22 RACOHETH 23 POLYALPA 21 CERAPUR 23 DIPLALB 23 HYGRLAX 27 ANASMIN 2B RACOBRE 29 ATRISEL 30 UUNGOBO 31 DITRHFT 32 DIPLPLI 33 0LIGA1 I 34 RACOHETA 35 RACOFAS 36 CONOCON 37 RICCMUL 38 P0R08IG 39 STOKPRAP 40 GEOCGRA 4 1 POHLCRU 42 AMPHCAL 43 METZCON 44 BRACFRI 45 CYNOJEN 46 HETEMAC 47 POHLLON 48 POHLNUT 49 ISOPELE 50 JAMfcAUT* 5 1 RACOAOU 52 SCLEOBT 53 HYPNSUB 54 DIPLTAX 55 PHILFON 56 CHILPOL 57 BART ITH 58 POHLPRO 59 SCAPSCA GO RACOVAR 61 PLEUALB 62 OLIGPAR 63 BAZZTRI 64 CLAOBOL 65 ANEUPIN 66 BARTLES 67 DICRHET 68 SPHAGIR G9 OXYSTEN 70 PHILARN 71 JUNGATR 72 ATRIUND 73 RADUBOL 74 ISOPPUL 75 TRITQUI 7G MYLITAY 77 PLAGLAE• 78 DREPUNC 79 RHYTSOA 80 BRYUPAL 8 1 RACOLAN 82 SCHISTR 83 SCHIRIV 84 BRACVEL 85 GRIMTOR 86 PREIQUA 87 SCAPPAL 88 FISSOSM 89 JUNGEXS 90 DRYPPAT 91 ANOEAES 92 PSEUPAT 93 BRYUPSE 94 JUNGCON 95 ISOTSTO 96 FISSBRY 97 POREROE 98 PLAGROS 99 HYGRSMI 100 RHIZNUD 101 PORECOR 102 MNIUBLY 103 PLATJUN 104 LOPHHET 105 BARBLYC 10G TORTFRA 107 HYGRLUR 108 BARBREF 10? BRYUCAP 110 SCAPMUC 111 BRYUMIN 112 STOKPRAS 113 DICRCRI 114 TRITPOL 115 PLAGPIL 116 BLASPUS 117 LOPHSUD 1 238 111 11 1 1 1111 1 122222222223333333333444444.44445555555_-556G6666G66677777777778 8888888889999999999000 1234.678301234567890123456789012345678^01234567890123456789012345G789012345G7890 1234567890123456789012 4 ; 3344 245 2-1 — - 2+ '• 3223 21 312 + - -3 2 : 223 2 + -42--+ 31 124 + - - + 2 2121 132 23122 + 3+21 122-2 2232222 12 1 - - 1 221 1+1-243-3 12 1-1 1 1 1 + 2 1 1-- + -22223 122 — - 2 3 - 3 3 + 4433343-2-24 143- - 1 1 1 + + - - 2 2 2 - - 1 1 13 1 2- + 2 1 +++ + - - 2 - 3 2 2 3 - 2 22 321 1 2221 + 3-133 -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 2222 -2 - -1 1 1 +++ + -1432 + 31 1 + 23- 4 + 2+--2 1 124332-22332124-222-22424 1 12 1 + 1+42-1 212+1 4 5 3-12 + - - 1 3 - 3 - - 31 322122 1-2-3343 . - -1 -144+-31 -1 1+1 12+J--1 _1 2 1 +1 121 1 222 14 1 1-4 1-44444 34-1 1 11 44 14111--+1 -1 123-+--2-2- - 123+1 121 1 1-1--1 12223 121-1 2 11- 1 -21-1 •1-1 -33424524213+--111 -3344 12 + 121+3 23 2 22223-4331231+-232- -2 -21121-111-1- • 1+1 + - -•1- -1-1--21 22+1 1 1 + 111 + 21+-111- -2121--2 + -111 •11 -1 - - -•-2 1-• 1 + + - -2+1- .-2 — 2 - 2 1 2 + 32121-- 1 - 2 2 1334442-1 -2 - - •1 1-1 1+4- -+1-1 -13 - - 12-2- 2 + + -31 + 1 1 1- - + 32-32 — 22 — 22- -2 -22 -2 21234- 1-1- - -2 -4 1-- 2 - -- - 1-11--43335-- 2 - - - 1 --+1++---+4-1 1--211-+-•12 -212113--4 1 2 — 1 --211-11--1 - 1 -•1221112- -21 1-- - - 2 --232-• 1-•11-•12- -__2 — - 2 - - 2 -13 31 •1 1 1--2++-- - 12 - -4 — 21- - 2 --23- -21--22--2+- . + 21-1 2 -434- 1 1 1-•111 1 4 1 2 - - 1 3 1 — 3 + - 1 - - - 1 1 1-1 -1 - - -•11- 1--21--2-1112 2- -1131+-2- -2 2 - -22 -+1+14-1-112++3--22 1-- 2 2 - - 2 1 2 - - 1-121-1-1--2 -21 --21 1 + 1 322-1 - •1-1-121212--22- •1-1--322 1 12 + -- 13244 12---131 1+1-2-1 + -•212 -1222 - - - •+1---+2-1 -+1-1 1-•22-• 3 2 2 - - - 4 - -44-•1--•11-11-- 2 - 3 -•11--11-- + + -— 2-i i -li § I i Do I f ^ " r- ci,' ft-x3 1 1 1 239 111 1 1 1 1 1 1 1 1 1 122222222223333333333444444444455555555556666G666G6777777777788888888889999999999000 1234567890123456789012345G789012345G78901234567890123456789012345678901234567890 123456789012345G789012 10 HYPNDIC 3 DICHPEL 4 SCAPUND 7 PELLNEE 12 BLINACU 15 MARSEMA 30 UUNG0B0 9 CEPHBIC 5 RACOACI 13 RHIZGLA 11 SCAPSUB 8 NARDSCA 16 BRACPLU 20 BLEPTRI 2G HYGRLAX 1 SCOUAQU 37 RICCMUL 25 DIPLALB 56 CHILPOL 18 PLAGASP 6 JUNGPUM 55 PHILFON 17 POGOURN 80 BRYUPAL 89 JUNGEXS 39 STOKPRAP 62 OLIGPAR 3E POROBIG 44 BRACFRI 65 ANEUPIN 71 UUNGATR 83 SCHIRIV 87 SCAPPAL 14 SCAPAME 36 CONOCON 52 SCLEOBT 2 HYGROCH 40 GEOCGRA 78 DREPUNC 86 PREIOUA 24 CERAPUR 75 TRITQUI 88 FISSOSM 99 HYGRSMI 6 1 PLEUALB 29 ATRISEL 33 OLIGALI 48 POHLNUT 51 RACOAQU 53 HYPNSUB 54 DIPLTAX 57 BARTITH 84 BRACVEL 93 BRYUPSE 94 JUNGCON 23 POLYALPA 27 ANASMIN 35 RACOFAS 59 SCAPSCA GO RACOVAR 4 1 POHLCRU 43 METZCON 49 ISOPELE 64 CLAOBOL 68 SPHAGIR 69 OXYSTEN 82 SCHISTR 104 LOPHHET 109 BRYUCAP 32 D IPLPLI 34 RACOHETA 46 HETEMAC 58 POHLPRO 74 ISOPPUL 63 BAZZTRI 66 BARTLES 81 RACOLAN 90 DRYPPAT 95 ISOTSTO 102 MNIUBLY 103 PLATJUN 107 HYGRLUR 108 BARBREF 110 SCAPMUC 114 TRITPOL 19 PLAGROE 2 1 BARTPOM' 22 RACOHETH 28 RACOBRE 31 DITRHET 91 ANOEAES 92 PSEUPAT 42 AMPHCAL 45 CYNOJEN 47 POHLLON 9G FISSBRY 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 PLAGLAE 79 RHYTSQA 111 BRYUMIN 1 12 STOKPRAS 113 DICRCRI 85 GRIMTOR 115 PLAGPIL 116 BLASPUS 117 LOPHSUD -1432+31 1+23+4 + 2 + - - 2 1 124332-22332 1 24-222-22424 1 1 2 1 + 1 + 4 2 - 1 - - - 2 1 2+ 1 - - -45 3-12+--13 4 2 - - + 31 124 + - - t-2 2 12 1 132 23122 + 3 + 2 1 122-2 2232222 12 1 - - 1 221 1+1-243-3 121-1- 1 11+2 1 1--+-22223 122 23-33 + 4433343-2-24 143--1 1 1-- - + + - - 2 2 2 - - 1 1 13 + 2 + - - 1 + - + +12- + 22 -2 -22 -1 + +1 1 1+1 3++ 1- 1 1-111-111 + 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 - -334245242 13+ 3344 12+121+3 23 2 - - - 2 2 2 2 3 21 12 1 2 12-2 2 + + -31+1 1 1 + 3 2 - 3 2 - - 2 2 - - 2 2 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 2 2 32 11 2221 + 3 - 1 3 3 - - 2 2 -1 123- + - -2 -2 - -123+1 12 1 1 1-1 — 1 12223 121-1 2 1 1-1 - 3 - - 31 322122 1-2-3343 1- 144 +-31-1 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 2 1 2 1 - -1 ,-1 1-22 1334 442 -1 -2 1 1-1 1+4 +1-1 4 3 3 4 4 245 2-1 2 + 3223 21 312 »---2 1 2121 13 1221 1 12 21 1 232 + 111 1 + + 2+1 2 - - 2 -212 + 32121 •! 1 n t 4 1 2 - - 13 1 3 + - 1 11 1-•1- 1-1- -22- 1--222-1- -2 1- 1 2 - - 2 - 1 - 1--2 + - -21-1 2-434- 1 1 1-•1- -1--2 11+1-2 --322-1 1--212-1222 +1 + 2---+4-11 -+1++-4-12--21 1-11 2 - - 1 131 + -412- -1 -+1+14-' 2- -2 - 2 1-1 12 ++3-- 2 2 - - 2 1 2 - - 1 1 - 1 2 1 1324442 13 31 1-121212- ... 1-• 1 + --21-1- •1 + - • 1--43335-12- -2 4 - - 2 1 - 1--223 2-- 2 -- + --21 1- + - 1-• ! 2 - 2 - 2 1 - - 1 -•-322 1 12 + -2+-• - 1 - 1 - - -1311+1--4-+1--322- - - 4 4 --2-1 112-•13-•111--2++--21-- 2 - - 2 2 -•1-1-•11-11--21--11-- 2 - - 2 2 --221-•11--41-- - 1 -1 i -- 2 - - 1-- - 2 2 - -•11-11-- 2 - 3 -•11-•11--3 -+-- 1 -0 ° C) ^  ^  IT) 1 1 •' v j y t ,c1 <CLUSTER OPTION-MINVAR FUNCTION'EUCLIDEAN REL = " CASES=1-MAX=*> 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. 20.V20. 21.V21, 22.V22, 23.V23, 24.V24, 25.V25, 26.V26, 27.V27, 28.V28. 29.V29, 30.V30, 31.V31, 32.V32, 33.V33. 34.V34. 35.V35, 36.V36, 37.V37. 38.V38, 39.V39. 40.V40, 41.V41. 42.V42, 43.V43, 44.V44, 45.V45. 46.V46, 47.V47, 48.V48, 49.V49, 50.V50, 51.V51. 52.V52, 53.V53. 54.V54, 55.V55, 56.V56. 57.V57, 58.V58. 59.V59, 60.V60. 61.V61. 62.V62, 63.V63. 64.V64, 65.V65, 66.V66, 67.V67. 68.V68, 69.V69. 70.V70, 71.V71, 72 . V72. 73 . V73, 74.V74, 75.V75, 76.V76. 77.V77, 78.V78, 79.V79, 80.V80, 81 . V8 1 , 82.V82, 83.V83, 84.V84. 85.V85. 86.V86, 87.V87, 88.V88, 89.V89, 90.V90, 91.V91, 92.V92, 93.V93, 94.V94, 95.V95. 96.V96, 97.V97, 98.V98. 99.V99, 100.V100, 101.V101. 102.V102. 103.V103. 104.V104, 105.V105, 106.V106, 107.V107, 108.V108, 109.V109, 110.V110, 111.V111. 112.V112. 113.V113, 114.V114, 115.V115, 116.V116, 117.V117 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 S T E P 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 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 2 2 1 1 1 1 1 0 9 8 7 6 1 1 3 1 9 8 7 6 5 4 3 2 1 + 1 + ---•£• + - - I I T-I-I + -I + - I -+ + -I + + I + I - I - I + 1 I-+ j + + + + + 1 + 1 + + _. + -. ro T 1 1 I I ; 1 j -p. •to-co: o D I S T A N C 2 5 0 7 0 5 8 1 . 0 2 . 5 1 1 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 5 6 8 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 6 2 0 5 6 7 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 1 8 0 0 0 6 5 0 0 6 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 0 0 0 7 0 0 0 7 0 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 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 3 5 6 1 0 4 7 1 7 8 . 2 8 7 8 7 3 0 3 5 5 7 6 8 241 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 93 BRYUPSE 94 d l jmCON 23 POLYALPA 27 ANASMIN 35 RACOFAS 551 SCAPSCA 60 RACOVAR 4• POHLCRU 4C. METZCON 4<ISOPELE 64 CLAOBOL GE SPHAGIR 6£ OXYSTEN SCHISTR 104 LOPHHET 109 BRYUCAP 32 DIPLPLI 34 RACOHETA 46 HETEMAC 58 POHLPRO 74 ISOPPUL 63 BAZZTRI 66 BARTLES 8 1 RACOLAN 90 DRYPPAT 95 ISOTSTO 102 MNIUBLY 103 FLATdUN 107 HYGRLUR 108 BARBREF 110 SCAPMUC 1 14 TRITPOL 19 PLAGROE 2 1 BARTPOM 22 RACOHETH 28 RACOBRE 31 DITRHET 91 ANOtAES 92 P S E S J P A T 42 AMPHCAL 45 CYNO.JEN 47 POHLLCN 9S FISSBRY 97 POREROE 98 PLAGROS. bO dAMEAUT 100 RHIZNIJD 101 PORECOR 67 DICRHET 70 PHILARN 72 ATRIUND 105 BARBLYC 106 TORTFRA 73 RA3UB0L 76 MYLITAY 77 PLAGLAE 79 RHYTSQA 111 BRYUMIN 1 12 STOKPRAS 113 DICRCRI 85 GRIMT! 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