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UBC Theses and Dissertations

The bryophytes of the semi-arid steppe of South-Central British Columbia McIntosh, Terry T. 1986

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THE BRYOPHYTES OF THE SEMI-ARID STEPPE OF SOUTH-CENTRAL BRITISH COLUMBIA by TERRY T. MCINTOSH B.A., W i l f r i d L a u r i e r U n i v e r s i t y , 1975 M.Sc, U n i v e r s i t y Of Canterbury, 1978 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in THE FACULTY OF GRADUATE STUDIES Botany Department We accept t h i s t h e s i s as conforming to the r e q u i r e d standard THE UNIVERSITY OF BRITISH COLUMBIA J u l y 1986 © T e r r y T. Mcintosh, 1986 g 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 available for 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 publication 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 The University of B r i t i s h Columbia 1956 Main Mall Vancouver, Canada V6T 1Y3 DE-6 (3/81) i i A b s t r a c t T h i s d i s s e r t a t i o n p r o v i d e s keys and c o n c i s e d e s c r i p t i o n s of t h e g e n e r a , s p e c i e s and v a r i e t i e s of b r y o p h y t e s known from t h e s e m i - a r i d s t e p p e r e g i o n o f s o u t h - c e n t r a l B r i t i s h C o l u m b i a . T h i r t y - s e v e n g e n e r a and s e v e n t y - s e v e n s p e c i e s a r e i n c l u d e d . D e t a i l e d i l l u s t r a t i o n s a r e p r o v i d e d f o r s e l e c t e d s p e c i e s . L o c a l d i s t r i b u t i o n maps a r e i n c l u d e d f o r a l l t h e t a x a . F o u r of t h e s p e c i e s i n c l u d e d h e r e a r e new t o N o r t h A m e r i c a : C r o s s i d i u m r o s e i , Phaseurn v l a s s o v i i , P o t t i a c f . w i l s o n i i and P t e r y g o n e r u m k o z l o v i i . I n a d d i t i o n , t h e g e n e r a C r o s s i d i u m and T r i c h o s t o m o p s i s a s w e l l a s e i g h t s p e c i e s and two v a r i e t i e s a r e new t o Canada. F i v e a d d i t i o n a l s p e c i e s a r e r e p o r t e d f o r t h e f i r s t t i m e f r om B r i t i s h C o l u m b i a . D i s t r i b u t i o n s o f t h e t a x a i n t h e s t u d y a r e a a r e d i s c u s s e d a t l o c a l and w o r l d l e v e l s . L o c a l d i s t r i b u t i o n s a r e i n f l u e n c e d most by s u b s t r a t u m t y p e , a s p e c t and e l e v a t i o n a l change, and e f f e c t s of d i s t u r b a n c e . Most o f t h e t a x a a r e t e r r i c o l o u s and r e s t r i c t e d t o o r w i t h a p p a r e n t p r e f e r e n c e t o one o f f i v e s o i l t y p e s . Many t a x a a r e s a x i c o l o u s , b u t d i s t r i b u t i o n s w i t h r e s p e c t t o r o c k t y p e c o u l d n o t be d e t e r m i n e d . The r e g i o n a l b r y o f l o r a i s compared a l s o t o t e m p e r a t e d r y l a n d b r y o p h y t e f l o r a s g l o b a l l y . S u r p r i s i n g l y , t h e r e g i o n a l f l o r a has v e r y s t r o n g a f f i n i t i e s , a t b o t h t h e s p e c i e s and genus l e v e l s , n o t o n l y t o d r y l a n d f l o r a s i n t h e N o r t h A m e r i c a n s o u t h w e s t b ut a l s o w i t h t h e a r i d r e g i o n s o f E u r a s i a and N o r t h A f r i c a . A l t h o u g h some o f t h e t a x a i n t h e s e a r e a s a r e w i d e s p r e a d t h r o u g h o u t t h e N o r t h e r n h e m i s p h e r e , many s p e c i e s e x p r e s s d i s j u n c t i o n s between t h e a r i d r e g i o n s . Two c o n t r a s t i n g c a u s e s , l o n g - d i s t a n c e d i s p e r s a l and v i c a r i a n c e , a r e d i s c u s s e d w i t h r e s p e c t t o t h e o r i g i n of t h e d i s j u n c t i v e p a t t e r n s . I n most i n s t a n c e s v i c a r i a n c e a p p e a r s a more l i k e l y c a u s e o f t h e d i s j u n c t i o n , a l t h o u g h t h i s c o n c l u s i o n i s h i g h l y s p e c u l a t i v e . I f v i c a r i a n c e i s a c a u s e , t h e n t h e d i s j u n c t t a x a a r e p r o b a b l y a n c i e n t and c o n s e r v a t i v e . B i p o l a r d i s j u n c t i o n s a r e a l s o d i s c u s s e d . Long d i s t a n c e d i s p e r s a l as w e l l as hop d i s p e r s a l a p p e a r t o be more l i k e l y c a u s e s o f t h i s t y p e o f p a t t e r n t h a n v i c a r i a n c e . Many of t h e t a x a r e q u i r e c r i t i c a l t a x o n o m i c r e v i s i o n . T h i s r e f l e c t s t h e l a c k of r e s e a r c h i n a r i d r e g i o n s . S u i t e s o f g e n e r a t h a t a r e p r o b l e m a t i c i n c l u d e Grimmia, S c h i s t i d i u m a n d C o s e i n o d o n , and Phaseurn, P o t t i a , Desmatodon and T o r t u l a . A l s o , a t l e a s t 41% o f t h e s p e c i e s p o s e p a r t i c u l a r t a x o n o m i c p r o b l e m s . Some p o s s i b l e s o l u t i o n s f o r t h e s e p r o b l e m s a r e s u g g e s t e d h e r e . T a b l e of C o n t e n t s A b s t r a c t i i L i s t of F i g u r e s v i Acknowledgement . : v i i C h a p t e r I INTRODUCTION .1 1 . P u r p o s e .1 2. F l o r i s t i c R e s e a r c h i n A r i d and S e m i - A r i d R e g i o n s .2 3. L o c a t i o n and G e n e r a l D e s c r i p t i o n o f t h e S t u d y A r e a ......9 C h a p t e r II METHODS 21 1. F i e l d Methods 21 2. L a b o r a t o r y Methods 22 C h a p t e r I I I THE BRYOPHYTES OF THE SEMI-ARID STEPPE OF SOUTH CENTRAL BRITISH COLUMBIA 24 1. I n t r o d u c t i o n 24 2. Key t o t h e G e n e r a ...27 3. L i s t o f B r y o p h y t e s Found i n t h e S t u d y A r e a 38 4. D i s c u s s i o n s and D e s c r i p t i o n s of Taxa 41 THE BRYOPHYTE GENERA C e r a t o d o n 41 D i t r i c h u m 43 E n c a l y p t a 44 A l o i n a 47 B a r b u l a 50 B r y o e r y - t h r o p h y l l u m 53 C r o s s i d i u m 56 Desmatodon 61 Didymodon 67 Phaseurn 78 P o t t i a 82 P s e u d o c r o s s i d i u m 88 P t e r y g o n e u r u m 90 S t e g o n i a 97 T o r t u l a 98 T r i c h o s t o m o p s i s 105 W e i s s i a 106 C o s c i n o d o n 108 Grimmia 110 S c h i s t i d i u m 121 E n t o s t h o d o n 124 F u n a r i a 125 O r t h o t r i c h u m 127 Bryum 135 L e p t o b r y u m 137 P o h l i a 138 P s e u d o l e s k e e l l a 138 C o n a r d i a 1 39 B r a c h y t h e c i u m 140 V E u r h y n c h i u m 143 H o m a l o t h e c i u m 144 Hypnum 145 P o l y t r i c h u m 147 C e p h a l o z i e l l a 150 M a n n i a 151 A t h a l a m i a 153 R i c c i a 154 C h a p t e r IV TAXONOMIC CONSIDERATIONS 1 57 1. I n t r o d u c t i o n 157 2. Taxonomic C h a r a c t e r s and C l a s s i f i c a t i o n 157 3. Taxonomic C o n s i d e r a t i o n s a t t h e Genus L e v e l 162 4. Taxonomic C o n s i d e r a t i o n s a t t h e S p e c i e s and V a r i e t a l L e v e l s 179 C h a p t e r V BIOGEOGRAPHICAL CONSIDERATIONS 199 1. I n t r o d u c t i o n 199 2. New and N o t e w o r t h y D i s t r i b u t i o n a l R e c o r d s 201 3. R e g i o n a l D i s t r i b u t i o n a l P a t t e r n s 204 4. W o r l d D i s t r i b u t i o n P a t t e r n s 216 C h a p t e r VI GENERAL DISCUSSION AND CONCLUSIONS 247 LITERATURE CITED 256 FIGURES 284 APPENDIX A C l i m a t e D a t a From S e l e c t e d S t a t i o n s i n B r i t i s h C o l u m b i a , A l b e r t a and W a s h i n g t o n S t a t e 339 APPENDIX B L o c a t i o n s of C o l l e c t i o n S i t e s i n S o u t h - C e n t r a l B r i t i s h C o l u m b i a 340 APPENDIX C S e c o n d a r y S i t e s and B r y o p h y t e s C o l l e c t e d i n A l b e r t a , Oregon and W a s h i n g t o n S t a t e s 343 v i L i s t o f F i g u r e s 1. G e n e r a l R e f e r e n c e Map o f t h e S t u d y A r e a 284 2. G e n e r a l i z e d S u r f i c i a l G e o l o g y o f t h e S t u d y A r e a 285 3. S t e p p e V e g e t a t i o n a t S i t e 23 n e a r S p e n c e s B r i d g e 286 4. S t e p p e V e g e t a t i o n a t S i t e 35 n e a r Cache C r e e k .286 5. S p o t t e d L a k e ; S i t e 95 n o r t h - w e s t o f Osoyoos .....287 6. L a c u s t r i n e T e r r a c e s a t S i t e 68 n e a r Summerland 287 7. Rock O u t c r o p a t S i t e 88 n o r t h o f Osoyoos 288 8. B r y o e r y t h r o p h y l l u m columbianum ...289 9. Desmatodon c o n v o l u t u s 290 10. C r o s s i d i u m a b e r r a n s 291 11. C r o s s i d i u m r o s e i 292 12. C r o s s i d i u m sp 292 13. P t e r y q o n e u r u m ovatum 293 14. Didymodon r i g i d u l u s v a r . g r a c i l i s 294 15. Didymodon v i n e a l i s v a r . v i n e a l i s 294 16. Phascum v l a s s o v i i 295 17. P s e u d o c r o s s i d i u m r e v o l u t u m 295 18. S i m i l a r i t i e s a t t h e S p e c i e s L e v e l Between S i x D r y l a n d B r y o p h y t e F l o r a s U s i n g S o r e n s e n s Index 296 19. S i m i l a r i t i e s a t t h e Genus L e v e l Between S i x D r y l a n d B r y o p h y t e F l o r a s U s i n g S o r e n s e n s Index ....297 20. t o 98. Maps o f t h e L o c a l D i s t r i b u t i o n of t h e B r y o p h y t e s From t h e S t u d y A r e a 299-338 v i i A cknowledgement I would l i k e t o thank my s u p e r v i s o r , W.B. S c h o f i e l d , f o r h i s s u p p o r t and p a t i e n c e o v e r t h e y e a r s . A l s o , t h a n k s t o t h e B.C. M i n i s t r y of F o r e s t s , t h e Government of B r i t i s h C o l u m b i a (G.R.E.A.T. S c h o l a r s h i p ) , t h e U n i v e r s i t y o f B r i t i s h C o l u m b i a and Dr . S c h o f i e l d f o r f i n a n c i a l s u p p o r t . Thankyou t o my c o m m i t t e e , D r s . Hughes, B r a d f i e l d and S t r a l e y , f o r t h e i r t i m e and e n e r g y . And t o F r e d G a n d e r s who k i n d l y e d i t e d t h e m a n u s c r i p t . To my many f r i e n d s , who p r o v i d e d h e l p and e n c o u r a g e m e n t , t h a n k y o u . S p e c i a l t h a n k s t o John C h r i s t y , T r e v o r Goward, John A. Spence, A r t h u r Leo K r u c k e b e r g , K a l i D. Robson, C y n t h i a D u r a n c e , Rene B e l l a n d , O l i v i a L e e , D. Watson, Mike Hawkes, S.P. K i n g , K e v i n G i l e s and many o t h e r s who must f o r g i v e me f o r n o t i n c l u d i n g them by name. A l s o , t o D e b b i e and A l e c Roos and C h r i s t i n e and Normand B a j a r d , f o r p u t t i n g up w i t h me d u r i n g t h e l a s t d a y s . I e s p e c i a l l y t h a n k my f a m i l y , Mary, P h a e d r a and J e s s i c a , who have w e a t h e r e d t h e whole p r o c e s s . 1 I . INTRODUCTION 1. PURPOSE A r i d e n v i r o n m e n t s t h r o u g h o u t t h e w o r l d have y i e l d e d a v a r i e t y o f r a r e and m o r p h o l o g i c a l l y u n u s u a l b r y o p h y t e s (Agnew and V o n d r e c e k 1975, M a g i l l 1981), many o f w h i c h a r e r e s t r i c t e d t o t h i s t y p e of h a b i t a t . Y e t f l o r i s t i c s t u d y i n t h e s e r e g i o n s i s , a t b e s t , s u p e r f i c i a l a nd f r a g m e n t a r y . T h i s s i t u a t i o n i s m i r r o r e d i n t h e s t e p p e and d e s e r t r e g i o n s of t h e w e s t e r n N o r t h A m e r i c a n C o r d i l l e r a , where no s i n g l e c o m p r e h e n s i v e s u r v e y of t h e b r y o p h y t e s has been c o m p l e t e d . Lawton (1971) and F l o w e r s (1973) have p r o v i d e d f l o r i s t i c t r e a t m e n t s of t h i s a r e a , b u t t h e y a r e i n c o m p l e t e w i t h r e s p e c t t o t h e r e g i o n a l a r i d l a n d b r y o p h y t e f l o r a . The p r i m a r y p u r p o s e o f t h e p r e s e n t s t u d y i s t o i n i t i a t e d e t a i l e d f l o r i s t i c b r y o l o g i c a l r e s e a r c h i n t h e C o r d i l l e r a n r e g i o n and t o p r o v i d e a d e s c r i p t i v e manual o f t h e b r y o p h y t e s of t h e s e m i - a r i d s t e p p e of s o u t h - c e n t r a l B r i t i s h C o l u m b i a . A l t h o u g h t h i s i s but a s m a l l p o r t i o n o f t h e e x t e n s i v e C o r d i l l e r a n d r y l a n d r e g i o n t h a t s t r e t c h e s from B r i t i s h C o l u m b i a i n t o M e x i c o , i t s g e o m o r p h o l o g i c a l and c l i m a t i c d i v e r s i t y p r o v i d e s a wide s p e c t r u m o f r e p r e s e n t a t i v e a r i d h a b i t a t s . A d d i t i o n a l aims of t h i s s t u d y a r e : " i ) t o p r e p a r e k e y s f o r t h e i d e n t i f i c a t i o n o f t h e b r y o p h y t e s t h a t a r e f o u n d i n t h e s t u d y a r e a i i ) t o p r o v i d e d i s t r i b u t i o n maps and, i n some c a s e s , i l l u s t r a t i o n s o f t h e r e s p e c t i v e t a x a 2 i i i ) t o d i s c u s s t a x o n o m i c p r o b l e m s r e l a t e d t o t h e s e t a x a i v ) t o d e s c r i b e and d i s c u s s t h e l o c a l and g l o b a l d i s t r i b u t i o n s and b i o g e o g r a p h i c a f f i n i t i e s o f t h e t a x a 2. FLORISTIC RESEARCH IN ARID AND SEMI-ARID REGIONS Few a r i d r e g i o n s have been t h o r o u g h l y i n v e s t i g a t e d (Koponen and H a t t o r i 1982; M a g i l l 1982a; S c o t t 1982a, 1982b), y e t numerous p u b l i c a t i o n s a r e a v a i l a b l e , r a n g i n g f r o m s m a l l a r t i c l e s d e s c r i b i n g n o t e w o r t h y l o c a l c o l l e c t i o n s o r p r e s e n t i n g c h e c k l i s t s , t o an o c c a s i o n a l i l l u s t r a t e d f l o r a . T h e s e works must be s y n t h e s i z e d i n o r d e r t o d e v e l o p c o m p l e t e , a c c u r a t e l i s t s o f b r y o p h y t e s f r o m s t e p p e and d e s e r t h a b i t a t s . The f o l l o w i n g r e v i e w r e p r e s e n t s an i n i t i a l s y n t h e s i s . F i r s t , t h e h i s t o r y o f f l o r i s t i c r e s e a r c h i n t h e s t u d y a r e a i s p r e s e n t e d , f o l l o w e d by a r e v i e w o f r e s e a r c h done i n o t h e r a r e a s . The more u s e f u l p u b l i c a t i o n s from e a c h r e s p e c t i v e a r e a a r e g i v e n . T h e s e were c o m p i l e d from t h e r e v i e w s o f H e b r a r d ( 1 9 8 2 ) , Lane ( 1 9 7 8 ) , S c h o f i e l d and M i l l e r (1982) and Vana ( 1 9 8 2 ) , as w e l l a s f r o m o t h e r s o u r c e s . i ) REGIONAL AND NORTH AMERICAN RESEARCH The f i r s t b r y o p h y t e c o l l e c t i o n s f r o m t h e s t u d y a r e a a p p e a r t o have been made by J ohn Macoun from s t e p p e s i t e s a l o n g t h e F r a s e r and Thompson R i v e r s between L y t t o n and Kamloops ( K i n d b e r g , 1893, 1907; Macoun and K i n d b e r g 1892; Macoun 1902). A few o t h e r e a r l y c o l l e c t i o n s made i n t h e s t u d y a r e a a r e i n t h e U n i v e r s i t y o f B r i t i s h C o l u m b i a h e r b a r i u m . I n c l u d e d a r e 3 c o l l e c t i o n s by E. W i l s o n i n 1904, A. H. Brinkman i n 1910 ( s e e Brinkman 1923 and 1933 f o r e a r l y l i s t s o f B r i t i s h C o l u m b i a h e p a t i c s ; see S c h o f i e l d , 1968a, f o r a r e c e n t l i s t i n g ) and F. MacFadden i n 1928 ( s e e MacFadden 1929). S i n c e t h e n t h e r e g i o n has been more o r l e s s n e g l e c t e d u n t i l r e l a t i v e l y r e c e n t l y when F.M. Boas, R . J . B a n d o n i , F. D i l l , T. T a y l o r and W.B. S c h o f i e l d p r o v i d e d a s e r i e s of c o l l e c t i o n s . However, few a d d i t i o n a l b r y o p h y t e s have been added t o Macoun an d K i n d b e r g ' s o r i g i n a l l i s t s . S c h o f i e l d (1968b, 1976) commented b r i e f l y on t h e h a b i t a t and d i s t r i b u t i o n a l p a t t e r n s of some of t h e s e t a x a . Somewhat more f l o r i s t i c r e s e a r c h has been u n d e r t a k e n i n t h e a d j a c e n t a r i d r e g i o n s o f W a s h i n g t o n (Cooke 1955, Daubenmire 1970, J o n e s 1929, Lawton and I r e l a n d 1963, P i c k e t t and J o n e s 1928), Idaho (Hong 1976, J o n e s 1929, P i c k e t t and J o n e s 1928, M c C l e a r y and G r e e n 1971) and Oregon ( C h r i s t y 1980, 1983; C h r i s t y e t a l . 1982; D u V a l l 1938; Hong 1978; J o n e s 1936; Rossman 1977, 1978) t h a n i n t h e B r i t i s h C o l u m b i a s t e p p e , but c o m p r e h e n s i v e a n d s y s t e m a t i c r e s e a r c h i s l a c k i n g i n t h e s e a r e a s as w e l l . The f l o r a o f Lawton (1971) c o m p e n s a t e s somewhat f o r t h i s , i l l u s t r a t i n g and d e s c r i b i n g t h e m a j o r i t y o f t h e r e g i o n a l a r i d l a n d t a x a i n c l u d i n g t h o s e known from B r i t i s h C o l u m b i a . J o n e s (1929) and Lawton (1971) p r o v i d e b r i e f h i s t o r i c a l a c c o u n t s of b r y o l o g i c a l c o l l e c t i n g i n t h e r e g i o n . The e x t e n s i v e a r i d r e g i o n s o f s o u t h w e s t N o r t h A m e r i c a , e x c l u d i n g M e x i c o , have been r e a s o n a b l y w e l l s u r v e y e d b r y o l o g i c a l l y . Some o f t h e more r e l e v a n t p u b l i c a t i o n s i n c l u d e , f o r A r i z o n a : E v a n s ( 1 9 1 7 ) , H a r i n g ( 1 9 6 1 ) , L i t t l e ( 1 9 3 9 ) , 4 M c C l e a r y (1953, 1959, 1961); f o r C a l i f o r n i a : H a r t h i l l e t a l . ( 1 9 7 9 ) , Howe ( 1 8 9 9 ) , Koch (1950, 1954), S t e e r e ( 1 9 5 4 ) ; f o r C o l o r a d o : E v a n s ( 1 9 1 5 ) , Weber (1963, 1973); f o r Nevada: L a v i n (1981, 1982), Lawton ( 1 9 5 8 ) ; f o r New M e x i c o : I r e l a n d e t a l . (1981, 1984), M a h l e r ( 1 9 7 8 ) , S t a r k and C a s t e t t e r ( 1 9 8 2 ) ; f o r T e x a s : B a r t r a m ( 1 9 2 9 ) , E l l i s o n ( 1 9 6 4 ) , M a g i l l (1974; 1976, an e x c e l l e n t r e v i e w i n c l u d i n g k e y s ) , M a h l e r ( 1 9 7 5 ) , W h i t e h o u s e and M c A l l i s t e r ( 1 9 5 4 ) ; and f o r U t a h : F l o w e r s (1933, 1961, 1973) and Graham ( 1 9 3 7 ) . M a g i l l (1982b) p u b l i s h e d a s h o r t b u t u s e f u l r e v i e w o f b r y o p h y t e d i s t r i b u t i o n and s y n e c o l o g y i n t h e s o u t h w e s t r e g i o n . Nash e t aj^. (1977) p r o v i d e g e n e r a l e c o l o g i c a l n o t e s f o r t h r e e of t h e s o u t h w e s t e r n d e s e r t s . D e l g a d i l l o (1982) p r e s e n t e d an o v e r v i e w o f M e x i c a n b r y o p h y t e f l o r i s t i c s and n o t e d t h a t many p o r t i o n s of t h e c o u n t r y , i n c l u d i n g t h e d e s e r t and s t e p p e r e g i o n s , r e m a i n p o o r l y c o l l e c t e d . However, t h e few s i g n i f i c a n t g e n e r a l c o n t r i b u t i o n s i n c l u d e : B a r t r a m and R i c h a r d s ( 1 9 4 1 ) , C a r d o t ( 1 9 1 1 ) , Crum (1951) and D e l g a d i l l o and Zander ( 1 9 8 4 ) . Some p a r t s o f B a j a C a l i f o r n i a a r e b e t t e r c o l l e c t e d . P u b l i c a t i o n s by Bowers e t a_l. ( 1 9 7 6 ) , p r o v i d i n g a b r i e f h i s t o r y of b o t a n i c a l e x p l o r a t i o n i n t h e a r e a , Crum and M i l l e r ( 1 9 5 6 ) , Crum and S t e e r e (1958) and Koch and Crum (1950) a r e a v a i l a b l e . A manual of M e x i c a n mosses i s p r e s e n t l y b e i n g p r o d u c e d ( s e e D e l g a d i l l o 1982) and s h o u l d p r o v e i n v a l u a b l e f o r f l o r i s t i c r e s e a r c h of a l l s e m i - a r i d r e g i o n s . The A m e r i c a n p r a i r i e s have been c o l l e c t e d s p o r a d i c a l l y , a l t h o u g h t h e r e c e n t p u b l i c a t i o n s o f C h u r c h i l l (1977a, 1977b, 1979a, 1979b, 1980, 1983) as w e l l as h i s c o m p r e h e n s i v e l i s t 5 c o v e r i n g t h e whole r e g i o n ( C h u r c h i l l 1976) have i m p r o v e d t h e s i t u a t i o n s i g n i f i c a n t l y . O t h e r p e r t i n e n t a r t i c l e s on t h e A m e r i c a n p r a i r i e s i n c l u d e : C h u r c h i l l and R e d f e a r n ( 1 9 7 7 ) , C h u r c h i l l e t a l . ( 1 9 8 1 ) , E v a n s ( 1 9 2 4 ) , Eversman and S h a r p ( 1 9 8 1 ) , Hong (1977a, 1977b), I k e n b e r r y ( 1 9 3 2 ) , Koch ( 1 9 7 1 ) , Lawton ( 1 9 5 3 ) , L i t t l e . ( 1 9 3 6 ) , M cGregor ( 1 9 5 5 ) , P o r t e r (1933, 1935), S h a r p (1930) and W i l l i a m s ( 1 9 0 2 ) . I m p o r t a n t a r t i c l e s on C a n a d i a n p r a i r i e b r y o p h y t e s i n c l u d e B i r d (1960, 1962, 1967, 1973), C o n a r d ( 1 9 5 7 ) , I r e l a n d e t a l . (1980) and Macoun and K i n d b e r g ( 1 8 9 2 ) . M i t t e n (1864) r e p o r t e d on t h e p r a i r i e c o l l e c t i o n s o f D. L y a l l and J . B u t t l e . B i r d (1962) d i s c u s s e d t h e b r y o p h y t e c o l l e c t i o n s of T. Drummond from 1825-1827. A number o f i l l u s t r a t e d f l o r a s a r e a l s o u s e f u l f o r t h e a r e a . I n c l u d e d a r e F l o w e r s ' h e p a t i c f l o r a (1961) and moss f l o r a ( 1 9 7 3 ) , t h a t , a l t h o u g h w r i t t e n p r i m a r i l y f o r use i n U t a h , a r e t h e most v a l u a b l e o f t h e i r k i n d f o r N o r t h A m e r i c a n a r i d l a n d b r y o p h y t e r e s e a r c h , p r o v i d i n g b o t h e x c e l l e n t keys and i l l u s t r a t i o n s as w e l l as e c o l o g i c a l i n s i g h t . Lawton (1971) i s u s e f u l i n t h e n o r t h e r n p o r t i o n s o f t h e r e g i o n . Crum and A n d e r s o n (1981) i s h e l p f u l f o r most o f t h e common t a x a , and a l s o f o r some l e s s common p r a i r i e t a x a . In a d d i t i o n t o F l o w e r s ( 1 9 6 1 ) , S c h u s t e r (1953) and G o d f r e y (1977) a r e u s e f u l f o r t h e s t u d y of t h e h e p a t i c s . O l d e r f l o r a s , i n p a r t i c u l a r t h o s e o f G r o u t (1928-1941 f o r t h e mosses) and F r y e and C l a r k e (1937-1947 f o r h e p a t i c s ) a r e a l s o v a l u a b l e r e f e r e n c e s . 6 i i ) EURASIAN AND NORTH AFRICAN RESEARCH The w e s t e r n E u r a s i a n - n o r t h A f r i c a n r e g i o n has been e x t e n s i v e l y c o l l e c t e d by b r y o l o g i s t s . As a r e s u l t , p u b l i c a t i o n s a r e numerous. Some n o t a b l e c o n t r i b u t i o n s i n c l u d e t h o s e by Agnew and V o n d r e c e k ( 1 9 7 5 ) , B i l e w s k e y (1965, 1970, 1974), B i z o t ( 1 9 6 2 ) , Brown ( 1 9 3 7 ) , E l - S a a d a w i ( 1 9 7 6 ) , Frahm ( 1 9 7 8 ) , F r e y (1969, 1972, 1974), F r o e l i c h (1949, 1964), G r a d s t e i n ( 1 9 7 0 ) , H e n d e r s o n (1957, 1958, 1961), H e n d e r s o n and M u i r h e a d ( 1 9 5 5 ) , H e n d e r s o n and P r e n t i c e ( 1 9 6 9 ) , Imam and Ghabbour ( 1 9 7 2 ) , J o v e t -A s t and B i s c h l e r (1966, 1970, 1971, 1972), Long ( 1 9 7 9 ) , N o g u c h i ( 1 9 5 6 ) , P e t t e t ( 1 9 6 6 ) , R o b i n s o n and G o d f r e y ( 1 9 6 0 ) , Ros and L l i m o n a ( 1 9 8 4 ) , Rungby ( 1 9 5 9 ) , S c h i f f n e r ( 1 9 1 3 ) , Townsend (1966a, 1966b, 1969, 1984), T r a b u t (1927) and V o n d r e c e k ( 1 9 6 5 ) . However, d e s p i t e t h e l a r g e number o f p u b l i c a t i o n s , o n l y Agnew and V o n d r e c e k (1975) and B i l e w s k y (1965) have p r o d u c e d i l l u s t r a t e d moss m a n u a l s . A l t h o u g h i n t e n d e d f o r I r a q and I s r a e l , r e s p e c t i v e l y , t h e s e two p u b l i c a t i o n s a r e a p p l i c a b l e t h r o u g h o u t t h e r e g i o n . A l s o u s e f u l a r e t h e g e n e r a l E u r o p e a n f l o r a s o f B r o t h e r u s ( 1924-1925), Monkemeyer ( 1 9 2 7 ) , M u l l e r (1951-1958), and S a v i c z - L j u b i t z k a j a and S m i r n o v a ( 1 9 7 0 ) , w h i c h p r o v i d e keys and some i l l u s t r a t i o n s f o r many o f t h e d r y l a n d t a x a . No f l o r a s have been c o m p l e t e d f o r t h e h e p a t i c s , a l t h o u g h t h e work of J o v e t - A s t and B i s c h l e r n o t e d above i s f a i r l y c o m p r e h e n s i v e , d e t a i l i n g l o c a l d i s t r i b u t i o n s and e c o l o g y , a l t h o u g h u n f o r t u n a t e l y l a c k i n g k e y s and i l l u s t r a t i o n s . B o r o s (1968) and M a r s t a l l e r (1980a, 1980b) p r e s e n t e d u s e f u l e c o l o g i c a l and d i s t r i b u t i o n a l i n f o r m a t i o n w i t h r e s p e c t t o many 7 a r i d l a n d b r y o p h y t e s . C o r l e y et. a l . (1981) p r o v i d e d a c o m p r e h e n s i v e l i s t f o r E u r o p e and t h e A z o r e s , n o t i n g p r o b l e m a t i c and d o u b t f u l s p e c i e s . T h i s l i s t i s e s s e n t i a l f o r f l o r i s t i c r e s e a r c h i n t h e i r r e g i o n . The n o r t h - c e n t r a l and e a s t e r n s t e p p e s and d e s e r t r e g i o n s o f A s i a ( f r o m t h e C a s p i a n sea t o M o n g o l i a and C h i n a ) a r e p o o r l y i n v e s t i g a t e d (Koponen and H a t t o r i 1982, Vana 1982). J a r v i n e n and Koponen (1975) d i s c u s s e d t h e e x c u r s i o n of V. B r o t h e r u s i n t h i s r e g i o n i n 1896, l i s t i n g h i s b r y o p h y t e c o l l e c t i o n s and p o i n t i n g o u t t h e more i n t e r e s t i n g d i s c o v e r i e s . S a v i c z - L j u b i t z k a j a and S m i r n o v a (1970) p r o v i d e a manual t h a t i s u s e f u l i n t h i s a r e a . The M o n g o l i a n r e g i o n i s b e t t e r known, w i t h a v e r y u s e f u l f l o r a by Abramova and Abramov (1983) a v a i l a b l e . O t h e r s t u d i e s i n M o n g o l i a i n c l u d e t h o s e o f D i x o n ( 1 9 2 8 ) , Duda ( 1 9 6 8 ) , K a r c z m a r z (1981) and S c h u b e r t e t a l . ( 1 9 7 7 ) . i i i ) SOUTHERN HEMISPHERE RESEARCH A l t h o u g h t h e s o u t h e r n h e m i s p h e r e b r y o f l o r a has been c o l l e c t e d f r o m a s e a r l y a s t h e l a t e e i g h t e e n t h and e a r l y n i n e t e e n t h c e n t u r i e s ( M a g i l l 1981, S c o t t and S t o n e 1976), most o f t h e a r i d r e g i o n s were n e g l e c t e d i n f a v o r o f t h e more b r y o l o g i c a l l y d i v e r s e w e t t e r a r e a s . P o r t i o n s o f S o u t h A f r i c a , however, s u c h a s t h e Cape r e g i o n , have been w e l l c o l l e c t e d . The i l l u s t r a t e d and d e s c r i p t i v e f l o r a o f M a g i l l (1981) i s e s s e n t i a l f o r t h i s r e g i o n , even t h o u g h i t i n c l u d e s o n l y a c r o c a r p o u s t a x a up t o t h e G r i m m i a c e a e . I t i n c l u d e s a l s o a h i s t o r y o f b r y o l o g i c a l r e s e a r c h i n S o u t h A f r i c a . 8 O t h e r p u b l i c a t i o n s of n o t e i n c l u d e A r n e l l ( 1 9 6 3 ) , D i x o n ( 1 9 2 0 ) , D i x o n and Gepp ( 1 9 2 3 ) , D i x o n and Wager ( 1 9 3 0 ) , M a g i l l ( 1 9 8 0 ) , M a g i l l and S c h e l p e ( 1 9 7 9 ) , S c h e l p e (1970, 1979) and Sim ( 1 9 2 6 ) . Two r e c e n t A u s t r a l i a n f l o r a s ( C a t c h e s i d e 1980, S c o t t and S t o n e 1976) i n c l u d e many of t h e r e g i o n a l a r i d - l a n d b r y o p h y t e t a x a . The numerous d i s c o v e r i e s and d i s c u s s i o n s o f S t o n e f o r t h e a r i d r e g i o n s (1973, 1976, 1980, f o r e x a m p l e ) , s t r e s s t h a t a g r e a t d e a l o f work r e m a i n s t o be c o m p l e t e d i n A u s t r a l i a . S a i n s b u r y ' s New Z e a l a n d f l o r a ( S a i n s b u r y 1955) i s a l s o h e l p f u l . No m a j o r f l o r a s e x i s t t o a i d i n f l o r i s t i c r e s e a r c h i n a r i d v e g e t a t i o n o f S o u t h A m e r i c a , a l t h o u g h t h o s e o f S c o t t and S t o n e (1976) and M a g i l l (1981) a r e a p p l i c a b l e i n p a r t s i n c e t h e y i n c l u d e many S o u t h A m e r i c a n t a x a . Of t h e few a v a i l a b l e p u b l i c a t i o n s , t h e most u s e f u l a r e B a r t r a m (1943, 1952), D e g u c h i ( 1 9 8 4 ) , H a s s e l de Menendez ( 1 9 6 2 ) , H a s s e l de Menendez e t a l . ( 1 9 8 4 ) , Hermann ( 1 9 7 6 ) , H e r z o g (1954, 1960), Kuehnemann (1938, 1949), R o b i n s o n ( 1 9 7 5 ) , S e k i (1974) and W i l l i a m s ( 1 9 1 5 ) . M i t t e n ( 1 8 6 9 ) , t h o u g h d e a l i n g p r i m a r i l y w i t h t h e Amazon r e g i o n , l i s t e d many b r y o p h y t e t a x a i n and west o f t h e Andes, and i s t h e r e f o r e u s e f u l . In a d d i t i o n , t h e work of B r o t h e r u s (1924-1925) r e m a i n s i n v a l u a b l e t h r o u g h o u t t h e S o u t h e r n H e m i s p h e r e , as i t documented and o c c a s i o n a l l y i l l u s t r a t e d many o f t h e e a r l y c o l l e c t i o n s f r o m t h e r e g i o n . 9 3. LOCATION AND GENERAL DESCRIPTION OF THE STUDY AREA In B r i t i s h C o l u m b i a , s e m i - a r i d s t e p p e i s r e s t r i c t e d p r i m a r i l y t o t h e narrow b a s a l p o r t i o n s of two n o n - c o n t i g u o u s v a l l e y s y s t e m s i n t h e s o u t h - c e n t r a l p a r t o f t h e p r o v i n c e : t h e F r a s e r - T h o m p s o n - N i c o l a s y s t e m and t h e O k a n a g a n - S i m i l k a m e e n s y s t e m ( F i g . 1 ) . However, s m a l l e r , more i s o l a t e d a r e a s of s t e p p e a r e f o u n d i n r i v e r v a l l e y s t o t h e e a s t of t h e s e s y s t e m s and i n l o w l a n d p o c k e t s i n t h e n o r t h o f t h e p r o v i n c e , but t h e s e were n o t i n c l u d e d i n t h e p r e s e n t s u r v e y . The s a m p l i n g l o c a t i o n s v i s i t e d d u r i n g t h e c o u r s e of t h e r e s e a r c h a r e shown on F i g . 1 ( s e e A p p e n d i x B f o r s p e c i f i c l o c a t i o n s o f t h e s t u d y s i t e s ) . The f o l l o w i n g s e c t i o n p r o v i d e s a b r i e f o v e r v i e w o f t h e g e o l o g y , c l i m a t e , v e g e t a t i o n and s o i l s o f t h e s t u d y a r e a . i ) GEOLOGICAL AND GLACIAL HISTORY The s t u d y a r e a l i e s w i t h i n t h e c o n f i n e s o f t h e W e s t e r n C o r d i l l e r a n P h y s i o g r a p h i c R e g i o n , an e x t e n s i v e o r o g e n i c s y s t e m s t r e t c h i n g f r o m A l a s k a s o u t h i n t o M e x i c o . T h r e e m a j o r s u b d i v i s i o n s c o m p r i s e t h e C o r d i l l e r a n R e g i o n i n B r i t i s h C o l u m b i a ( H o l l a n d 1976). Of t h e s e , t h e w e s t e r n and e a s t e r n s u b d i v i s i o n s a r e e x t e n s i v e l y m o u n t a i n o u s , whereas t h e I n t e r i o r s u b d i v i s i o n i s p r i m a r i l y an a r e a o f low l y i n g p l a t e a u s . Most o f t h e s t u d y r e g i o n i s l o c a t e d on t h e s o u t h e r n p o r t i o n o f t h e I n t e r i o r s u b d i v i s i o n , a l t h o u g h s m a l l s e c t i o n s a r e l o c a t e d on t h e b o r d e r s of t h e e a s t e r n m o u n t a i n o u s s y s t e m ( s o u t h of Keremeos and f r o m j u s t e a s t o f L y t t o n t o L i l l o o e t ) . The W e s t e r n C o r d i l l e r a has had a complex g e o l o g i c a l 10 h i s t o r y , a l t e r n a t i n g between p e r i o d s o f m a s s i v e u p l i f t i n g and s u b s e q u e n t e r o s i o n a l d o w n c u t t i n g ( H o l l a n d 1976, K i n g 1977, McKee 1972). One o f t h e most w i d e s p r e a d p e r i o d s o f u p l i f t o c c u r r e d d u r i n g t h e e a r l y T e r t i a r y , e l e v a t i n g much of t h e a r e a o u t of t h e s h a l l o w s of t h e P a c i f i c Ocean (McKee 1972). F o l l o w i n g t h i s a c t i v i t y , l a r g e a r e a s of t h e i n t e r i o r p l a t e a u l a n d s c a p e were c o v e r e d by b a s a l t l a v a from two p e r i o d s of v u l c a n i s m , one d u r i n g t h e E o c e n e and a n o t h e r i n t h e M i o c e n e . A n o t h e r p e r i o d o f u p l i f t f o l l o w e d i n t h e P l i o c e n e , and e r o s i o n o f t h i s s u r f a c e i n i t i a t e d t h e r e l i e f and d r a i n a g e p a t t e r n s e x p r e s s e d i n t h e modern l a n d s c a p e ( D o u g l a s 1970). The complex g e o l o g i c a l h i s t o r y of t h e r e g i o n i s r e f l e c t e d i n t h e h e t e r o g e n e o u s n a t u r e of t h e s u r f i c i a l b e d r o c k , g e n e r a l l y r e p r e s e n t e d i n F i g u r e 2. The a d v e n t o f t h e P l e i s t o c e n e marked t h e b e g i n n i n g o f a s e r i e s o f e x t e n s i v e g l a c i a t i o n s . D u r i n g t h e most r e c e n t g l a c i a l maximum, B r i t i s h C o l u m b i a was c o v e r e d by a t h i c k m a n t l e o f i c e t h a t s t r e t c h e d i n t o n o r t h e r n W a s h i n g t o n S t a t e . At t h e i c e -b e d r o c k i n t e r f a c e , s t r o n g e r o s i o n a l f o r c e s m o d i f i e d t h e l a t e T e r t i a r y l a n d s c a p e . The e f f e c t s i n c l u d e d t h e d e e p e n i n g and w i d e n i n g o f v a l l e y s ( H o l l a n d 1976), e s p e c i a l l y t h o s e v a l l e y s r u n n i n g p a r a l l e l t o t h e f l o w o f i c e , i n c l u d i n g t h e Okanagan and F r a s e r s y s t e m s ( F u l t o n 1969). A f t e r t h e r e c e s s i o n o f t h e g l a c i e r s , g r e a t q u a n t i t i e s o f e r o s i o n a l d e b r i s r e m a i n e d i n t h e v a l l e y b o t t o m s (summarized by Ryder 1978, 1982). To a l a r g e e x t e n t , t h e s e d e p o s i t s have p e r s i s t e d , p a r t i c u l a r l y a l o n g t h e s i d e s o f t h e v a l l e y s where t h e y have e s c a p e d t h e d o w n c u t t i n g e f f e c t s o f t h e major r i v e r s . The d e p o s i t s mask much o f t h e 11 s u p e r f i c i a l b e d r o c k . The most w i d e s p r e a d and l o c a l l y d o minant t y p e s of d e p o s i t a r e m o r a i n a l and f l u v i o g l a c i a l t e r r a c e s , p r i n c i p a l l y composed o f c o a r s e m a t e r i a l , and g l a c i o l a c u s t r i n e t e r r a c e s ( F i g . 6 ) , composed o f f i n e c l a y s and s i l t s and formed i n i c e - o r debris-dammed p o s t - g l a c i a l l a k e s ( F u l t o n 1969). In some a r e a s , s u c h as a r o u n d Kamloops, S p e n c e s B r i d g e and s o u t h o f P e n t i c t o n , l a c u s t r i n e t e r r a c e s d o m i n a t e t h e lo w e r v a l l e y l a n d s c a p e . F o l l o w i n g i t s d e p o s i t i o n , g l a c i a l d e b r i s has been r e w o r k e d by v a r i o u s f orms o f e r o s i o n . V a l l e y d r i f t has been c u t i n t o by r i v e r s and d e p o s i t e d downstream as modern a l l u v i u m . L a t e r a l headward e r o s i o n has been r a p i d a t t i m e s , p r o d u c i n g e x t e n s i v e g u l l e y s y s t e m s i n t h e v a r i o u s t e r r a c e s (Ryder 1978). T r i b u t a r y s t r e a m s , when t h e y l o s t t h e i r c a r r y i n g c a p a c i t y , d e p o s i t e d d e b r i s a l o n g s i d e t h e main v a l l e y , f o r m i n g a l l u v i a l f a n s (Ryder 1978). Most of t h e f a n s d e v e l o p e d i m m e d i a t e l y f o l l o w i n g d e g l a c i a t i o n when m e l t w a t e r s were most e x t e n s i v e . O t h e r t y p e s o f p o s t - g l a c i a l d e p o s i t s , i n c l u d i n g c o l l u v i a l d e b r i s and a e o l i a n d e p o s i t s s u c h as dunes, a r e s c a t t e r e d t h r o u g h o u t . As a r e s u l t o f o r o g e n i c , g l a c i a l and s u b s e q u e n t e r o s i o n a l e v e n t s , t h e c o n t e m p o r a r y l a n d s c a p e i n t h e r i v e r b o t t o m s c o m p r i s i n g t h e s t u d y a r e a i s e x t r e m e l y d i v e r s e . A l t h o u g h some a r e a s e x p r e s s a r e l a t i v e l y u n i f o r m t o p o g r a p h y , l o c a l v a r i a t i o n i n a s p e c t , s l o p e and s u p e r f i c i a l m a t e r i a l s t e n d s t o be h i g h . 12 i i ) CLIMATE The c l i m a t e of t h e s t u d y r e g i o n i s t h e d r i e s t and h o t t e s t i n C anada. T h i s i s p r i n c i p a l l y t h e r e s u l t o f a r a i n s h a d o w e f f e c t c a u s e d by t h e e x t e n s i v e m o u n t a i n s y s t e m t o t h e w e s t . A p p e n d i x A l i s t s d a t a from c l i m a t e s t a t i o n s i n o r v e r y n e a r t h e s t u d y a r e a , as w e l l a s , f o r c o m p a r a t i v e p u r p o s e s , d a t a from W a s h i n g t o n S t a t e , s o u t h e r n A l b e r t a and V a n c o u v e r . W i t h r e s p e c t t o t h e s t u d y a r e a , a number o f t r e n d s a r e e x p r e s s e d . Summers a r e h o t and d r y e x c e p t f o r an e a r l y r a i n f a l l peak, u s u a l l y i n J u n e . A c c o r d i n g t o S c h a e f e r ( 1 9 7 8 ) , t h i s peak r e f l e c t s an i n t e r p l a y o f l a t e s u r g e s of u n s t a b l e c o l d a i r comb i n e d w i t h a h i g h amount o f i n s o l a t i o n . C o n v e c t i v e a c t i v i t y i s s t r e n g t h e n e d and t h u n d e r s t o r m s o f t e n r e s u l t . W i n t e r s , on t h e o t h e r hand, a r e c o l d and w e t t e r , w i t h most of t h e a n n u a l p r e c i p i t a t i o n o c c u r r i n g t h e n . S n o w f a l l i s s p o r a d i c , u s u a l l y r e m a i n i n g on t h e g r o u n d f o r o n l y s h o r t p e r i o d s . In a d d i t i o n , t h e i n t e r i o r v a l l e y s have h i g h amounts o f s u n s h i n e , o v e r 1800 sun h o u r s p e r y e a r on a v e r a g e ( S c h a e f e r 1978), a d i r e c t r e s u l t o f t h e low d e g r e e o f c l o u d c o v e r , e s p e c i a l l y i n summer. M a i n l y b e c a u s e o f t h e s t r o n g i n s o l a t i o n v a l u e s , d i f f e r e n c e s i n m i c r o c l i m a t e c a u s e d by s l o p e and a s p e c t a r e e x t r e m e , p a r t i c u l a r l y e v i d e n t i n t h e h i g h l y d i v e r s e t o p o g r a p h i c a l s e t t i n g of t h e s t u d y r e g i o n . T h u s , o v e r s h o r t d i s t a n c e s t h e r e a r e a v a r i e t y o f v e r y d i f f e r e n t h a b i t a t s a v a i l a b l e f o r b r y o p h y t e s t o e x p l o i t . The s t e p p e r e g i o n of W a s h i n g t o n S t a t e i s g e n e r a l l y h o t t e r and d r i e r t h a n t h e s t e p p e i n B r i t i s h C o l u m b i a , and a summer 13 r a i n f a l l peak i s l e s s a c u t e . T h i s warming and d r y i n g t r e n d c o n t i n u e s i n t o Oregon and s o u t h w a r d s i n t o t h e s o u t h w e s t d e s e r t r e g i o n . N o n e t h e l e s s , t h e c l i m a t e i n t h e s t u d y a r e a i s more s i m i l a r t o t h e C o r d i l l e r a n s t e p p e t o t h e s o u t h t h a n i t i s t o t h e c l i m a t e o f t h e p r a i r i e r e g i o n . The A l b e r t a n c l i m a t e i s c o m p a r a t i v e l y w e t t e r and c o l d e r t h a n t h a t of t h e s t u d y r e g i o n , w i t h most o f t h e p r e c i p i t a t i o n o c c u r r i n g d u r i n g t h e summer s e a s o n . F u r t h e r m o r e , t h e p e r i o d o f d r o u g h t i s s i g n i f i c a n t l y l o n g e r i n t h e s t u d y a r e a t h a n i n t h e p r a i r i e r e g i o n s (Weaver 1980). The c l i m a t e o f t h e s t u d y r e g i o n i s c o n s i d e r e d by K r a j i n a ( 1 9 6 5 ) a s r e p r e s e n t a t i v e o f t h e m i d d l e l a t i t u d e s e m i - a r i d s t e p p e c l i m a t e . C h a r a c t e r i s t i c of t h i s c l i m a t e i s a p r e v a i l i n g w a t e r d e f i c i e n c y , w i t h e v a p o r a t i o n f r o m s o i l and v e g e t a t i o n b e i n g g r e a t e r t h a n th e p r e c i p i t a t i o n . In a d d i t i o n , w i n t e r s a r e c o l d and summers a r e warm t o h o t . The p r e d o m i n a n t v e g e t a t i o n t y p e s a r e g r a s s e s and, i n some r e g i o n s , s h r u b s . A s e m i - a r i d s t e p p e c l i m a t e o c c u p i e s an i n t e r m e d i a t e p o s i t i o n between d e s e r t c l i m a t e s and more humid g r a s s l a n d o r d r y f o r e s t c l i m a t e s , a l t h o u g h , c h a r a c t e r i s t i c a l l y , t h e b o u n d a r i e s between t h e s e t h r e e t y p e s a r e e x t r e m e l y d i f f u s e . i i i ) VEGETATION The v e g e t a t i o n o f t h e s t u d y r e g i o n r e p r e s e n t s t h e n o r t h e r n -most e x t e n s i o n o f t h e C o r d i l l e r a n s t e p p e or s h r u b - s t e p p e . T h i s v e g e t a t i o n t y p e o c c u p i e s t h e d r i e s t p a r t s o f t h e w e s t e r n i n t e r m o u n t a i n r e g i o n from B r i t i s h C o l u m b i a and W a s h i n g t o n s o u t h 14 t o n o r t h e r n C a l i f o r n i a and A r i z o n a and e a s t w a r d t o s o u t h e r n Wyoming and C o l o r a d o (Daubenmire 1978). I t has been v a r i o u s l y c l a s s i f i e d , e i t h e r a s s e m i - d e s e r t ( D a n s e r e a u 1957, E y r e 1968, W a l t e r 1983), s a g e b r u s h g r a s s l a n d ( T i s d a l e 1982), b l u e b u n c h -w h e a t g r a s s g r a s s l a n d s o r p a l o u s e p r a i r i e ( C o u p l a n d 1979) o r , as n o t e d e a r l i e r , s e m i - a r i d s t e p p e ( K r a j i n a 1965). A l t h o u g h a number o f v a s c u l a r p l a n t t a x a a r e d i a g n o s t i c , two s p e c i e s a r e most c h a r a c t e r i s t i c : bunch g r a s s , A q r o p y r o n  s p i c a t u m ( P u r s h ) S c r i b . and S m i t h , o r b l u e b u n c h - w h e a t g r a s s , and s a g e b r u s h or b i g s a g e , A r t e m i s i a t r i d e n t a t a N u t t . ( F i g . 3 and 4 ) . Gro w i n g among t h e s e d o m i n a n t s a r e o t h e r common but l e s s c o n s p i c u o u s t a x a . I n c l u d e d a r e t h e g r a s s e s , Poa s a n d b e r q i i V a s e y , K o e l e r i a c r i s t a t a P e r s . , S t i p a comata T r i n . and Ru p r . and S p o r o b o l u s c r y p t a n d r u s ( T o r r . ) G r a y ( t h e l a t t e r two g r a s s e s o f t e n r e p l a c e A g r o p y r o n s p i c a t u m on sandy s o i l s ) and t h e s h r u b s P u r s h i a t r i d e n t a t a ( P u r s h ) D.C. ( i n t h e s o u t h e r n h a l f of t h e Okanagan and K o o t e n a y V a l l e y s o n l y ) and Ch r y s o t h a m n u s n a u s e o s u s ( P a l l . ) B r i t t . Common f o r b s i n c l u d e A n t e n n a r i a d i m o r p h a ( N u t t . ) T.and G., B a l s a m o r h i z a s a g i t a t t a ( P u r s h ) N u t t . , Lomatium  macrocarpum ( N u t t . ) C o u l t . and Rose, O p u n t i a f r a g i l i s ( N u t t . ) Haw. and P h l o x l o n g i f o l i a N u t t . The v a s c u l a r p l a n t l a y e r i s d i s c o n t i n u o u s i n t h e s t e p p e and i s i n t e r s p e r s e d e i t h e r w i t h b a r e s o i l o r , i n a r e a s o f l i t t l e d i s t u r b a n c e , w i t h a t h i n , f r a g i l e l a y e r composed o f b l u e - g r e e n " a l g a e " ( C y a n o b a c t e r i a ) , a l g a e , l i c h e n s and mosses ( s e e A n d e r s o n and R u s h f o r t h 1977, f o r one l i s t i n g o f r e p r e s e n t a t i v e c r u s t t a x a ) . I t has been d e m o n s t r a t e d t h a t t h i s c r u s t i s b e n e f i c i a l i n 15 t h e m a i n t e n a n c e of a h e a l t h y and v i g o r o u s s t e p p e community ( A n d e r s o n e t a l . 1982a, 1982b, R o g e r s 1977). I t p r e v e n t s o r r e d u c e s b o t h r a i n and wind i n d u c e d s o i l e r o s i o n ( A n d e r s o n e t a l . 1982b, B o o t h 1941) and r e t a r d s e v a p o r a t i o n o f s o i l m o i s t u r e f o l l o w i n g p e r i o d s of p r e c i p i t a t i o n ( B r o t h e r s o n and R u s h f o r t h 1983). The f r e e l i v i n g c y a n o b a c t e r i a , a s w e l l as t h e s y m b i o t i c c y a n o b a c t e r i a i n some l i c h e n s p e c i e s , f i x a t m o s p h e r i c n i t r o g e n , t h u s making i t a v a i l a b l e f o r use by o t h e r p l a n t s (Cameron and F u l l e r 1960, S h i e l d s e t a l . 1957). In a d d i t i o n , t h e c r u s t i s s u s p e c t e d t o r e t a r d i n v a s i o n s by a l i e n p l a n t t a x a (Mack and Thompson 1982). A u n i q u e a s s o c i a t i o n of p l a n t t a x a i s f o u n d a d j a c e n t t o a l k a l i n e d e p r e s s i o n s ( F i g . 5 ) , a r e a s r i c h i n s a l t s ( s e e t h e d i s c u s s i o n r e g a r d i n g s o l o n e t z i c s o i l s ) . T y p i c a l s t e p p e t a x a a r e a b s e n t o r r a r e i n t h e s e s i t e s . The t h r e e most c h a r a c t e r i s t i c t a x a a r e D i s t i c h l i s s t r i c t a ( T o r r . ) Rydb., P o t e n t i 1 1 a a n s e r i n a L. and S a l i c o r n i a r u b r a N e l s . A l t h o u g h u s u a l l y a b s e n t from t h e S a l i c o r n i a z o n e , a number of r a t h e r u n u s u a l and r a r e b r y o p h y t e s have been f o u n d i n t h e l e s s s a l i n e D i s t i c h l i s z o n e . In t h e s t u d y r e g i o n , a l k a l i n e d e p r e s s i o n s a r e r e l a t i v e l y few i n number and s m a l l i n a r e a , whereas t h e y a r e common and v e r y e x t e n s i v e n o r t h o f t h e r e g i o n , p r i m a r i l y between C l i n t o n and W i l l i a m s L a k e , a s w e l l a s i n t h e G r e a t B a s i n r e g i o n t o t h e s o u t h . V a r i o u s f a c t o r s , i n p a r t i c u l a r t h e d i v e r s e and o f t e n u n s t a b l e t o p o g r a p h y , i n t e r a c t t o p r o d u c e a m o s a i c o f p a t t e r n s w i t h i n t h e v e g e t a t i o n c o v e r ( F i g . 3, 4, 5 and 6 i l l u s t r a t e t h i s t o some d e g r e e ) . S u p e r i m p o s e d on t h e s e p a t t e r n s a r e t h e n e a r l y 16 u b i q u i t o u s e f f e c t s of d o m e s t i c h e r d s o f c a t t l e and, i n some a r e a s , h o r s e s . As a d i r e c t r e s u l t o f b o t h t r a m p l i n g and g r a z i n g p r e s s u r e t h e r e i s u s u a l l y a d r a m a t i c d e c r e a s e i n p r e f e r r e d f o r a g e s p e c i e s , i n p a r t i c u l a r b l u e b u n c h - w h e a t g r a s s , a c c o m p a n i e d by t h e i n c r e a s e and l o c a l dominance of a v a r i e t y o f e x o t i c weedy p l a n t s p e c i e s , most i m p o r t a n t of w h i c h a r e Bromus t e c t o r u m L. ( c h e a t g r a s s ) and C e n t a u r e a s p p . ( k n a p w e e d s ) . Weakly o r non-p a l a t a b l e n a t i v e t a x a , i n c l u d i n g b i g sage and A r t e m i s i a f r i g i d a W i l l d . , a l s o i n c r e a s e i n some s i t e s . The c r y p t o g a m i c c r u s t i s a l s o e a s i l y damaged by t h e t r a m p l i n g o f l i v e s t o c k , w i t h b o t h s p e c i e s r i c h n e s s and t o t a l p l a n t c o v e r d e c r e a s i n g ( A n d e r s o n e_t a l . 1982b, K l e i n e r and H a r p e r 1972). T h i s a f f e c t s n u t r i e n t and m o i s t u r e l e v e l s , as w e l l as i n c r e a s i n g t h e p o t e n t i a l f o r s o i l e r o s i o n . The s t e p p e i s b o r d e r e d by a v a r i e t y o f o t h e r v e g e t a t i o n t y p e s . A l o n g r i v e r s , s t r e a m s and l a k e s a t low e l e v a t i o n s a r e f o u n d r i p a r i a n c o m m u n i t i e s d o m i n a t e d by s h r u b s , i n c l u d i n g Rosa spp. and S a l i x s p p . , and t r e e s , i n c l u d i n g P o p u l u s t r i c h o c a r p a T. and G. ( B l a c k C o t t o n w o o d ) , P i n u s p o n d e r o s a D o u g l . ( P o n d e r o s a P i n e ) and P s e u d o t s u g a m e n z i e s i i ( M i r b . ) F r a n c o ( D o u g l a s F i r ) . T h i s t y p e o f a s s o c i a t i o n a l s o a p p e a r s a l o n g some t r i b u t a r y s t r e a m s and g u l l i e s t h a t l e a d i n t o t h e main v a l l e y s . W i t h i n c r e a s e d e l e v a t i o n , t h e s t e p p e u s u a l l y i n t e r g r a d e s i n t o f o r e s t . A t l o w e r e l e v a t i o n s (below a b o u t 600m, d e p e n d i n g on a s p e c t ) t h e f o r e s t i s p r i m a r i l y a s a v a n n a h - l i k e o r , o c c a s i o n a l l y , a c l o s e d f o r e s t l a n d s c a p e , composed o f P o n d e r o s a P i n e and, l e s s commonly, D o u g l a s F i r . The u n d e r s t o r y v e g e t a t i o n 17 i s , i n many r e s p e c t s , s i m i l a r t o t h e s t e p p e , w i t h v a s c u l a r t a x a , i n c l u d i n g s a g e b r u s h and b l u e b u n c h w h e a t g r a s s , l o c a l l y f r e q u e n t , a l t h o u g h i t i s a l s o c h a r a c t e r i z e d by t h e p r e s e n c e o f t h e s h r u b s A r t e m i s i a f r i g i d a , A r c t o s t a p h y l o s u v a - u r s i ( L . ) S p r e n g . , C h r y s o t h a m n u s n a u s e o s u s ( P a l l . ) B r i t t . and A m e l a n c h i e r a l n i f o l i a N u t t . The t r a n s i t i o n t o f o r e s t i s r a r e l y smooth, r e s u l t i n g from t o p o g r a p h i c i n f l u e n c e s , as w e l l as f r o m t h e e f f e c t s o f d i s t u r b a n c e and s o i l t y p e (McLean 1970). F o r example, on c o o l e r , n o r t h - f a c i n g s l o p e s o r i n shaded draws, f o r e s t may o c c u r a t e l e v a t i o n s n o r m a l l y d o m i n a t e d by s t e p p e . T r e e s o c c u r on some s t e e p , r o c k y s l o p e s and on c o a r s e t e x t u r e d s o i l s i n t h e s t e p p e p r o p e r . D o u g l a s F i r f o r e s t r e p l a c e s t h e P o n d e r o s a P i n e b e l t a t h i g h e r e l e v a t i o n s ( a b o u t 850m; M i t c h e l l and G r e e n 1981). In a few a r e a s , most n o t a b l y n o r t h w e s t of Kamloops, e a s t o f M e r r i t t and west o f O s o y o o s , t h e i n c r e a s e i n e l e v a t i o n does n o t c o i n c i d e w i t h a s t e p p e t o f o r e s t t r a n s i t i o n a l s e q u e n c e . I n s t e a d t h e s e m i - a r i d s t e p p e c h a n g e s i n t o a more m e s i c s t e p p e , ( t h e m i d d l e and upper g r a s s l a n d z o n e s o f S p i l s b u r y and T i s d a l e , 1944, and T i s d a l e , 1947) c h a r a c t e r i z e d by a d e c r e a s e i n t h e i m p o r t a n c e o f t h e low e l e v a t i o n d o m i n a n t s , b i g sage and b l u e b u n c h -w h e a t g r a s s , and a c o n t r a s t i n g i n c r e a s e i n o t h e r t a x a , most i m p o r t a n t l y t h e g r a s s F e s t u c a s c a b r e l l a T o r r . , and t h e s h r u b s , C h r y s o t h a m n u s n a u s e o s u s and A r t e m i s i a f r i g i d a . In s u c h a r e a s t h e v e g e t a t i o n i s more c o n t i n u o u s t h a n i n t h e d r i e r s t e p p e , l i t t e r i s more common and a c r y p t o g a m i c c r u s t l a y e r i s e i t h e r w e a k l y d e v e l o p e d o r a b s e n t . T h i s g r a s s l a n d i n t e r g r a d e s i n t o D o u g l a s F i r f o r e s t . 18 D e t a i l e d d i s c u s s i o n and d e s c r i p t i o n s o f s p e c i f i c t r e n d s i n t h e v e g e t a t i o n o f t h e s t u d y r e g i o n a r e g i v e n by Brayshaw ( 1 9 6 5 ) , K r a j i n a ( 1 9 6 5 ) , McLean ( 1 9 7 0 ) , McLean and Ma r c h a n d ( 1 9 6 8 ) , McLean and T i s d a l e ( 1 9 7 2 ) , M i t c h e l l and G r e e n ( 1 9 8 1 ) , T i s d a l e (1947) and van Ryswyk e t a l . ( 1 9 6 6 ) . Many o f t h e s e a u t h o r s a t t e m p t e d t o c l a s s i f y t h e v e g e t a t i o n a t a l o c a l and p r o v i n c i a l l e v e l , and t h e i r works a r e u s e f u l i n t h i s r e g a r d . A l t h o u g h t r e a t e d a s a s u b s e t o f t h e c e n t r a l N o r t h A m e r i c a n p r a i r i e v e g e t a t i o n t y p e by some w o r k e r s (Knapp 1979), t h e v e g e t a t i o n o f t h e s t u d y r e g i o n i s d i s t i n c t c o m p o s i t i o n a l l y and p h y s i o g n o m i c a l l y . P r a i r i e v e g e t a t i o n i s d o m i n a t e d p r i m a r i l y by r h i z o m a t o u s , s o d - f o r m i n g g r a s s e s , B o u t e l o u a g r a c i l i s (H.B.K.) L a g . i n p a r t i c u l a r , but a l s o i n c l u d i n g A g r o p y r o n s m i t h i i Rydb. S h r u b b y p l a n t s a r e r a r e o r a b s e n t . The g r a s s e s form a dense c o v e r o v e r t h e s o i l and, i n c o n t r a s t t o t h e d r i e r p a r t s o f t h e C o r d i l l e r a n s t e p p e , b a r e s o i l i s n o t common e x c e p t i n u n s t a b l e and d i s t u r b e d s i t e s . A l t h o u g h i n B r i t i s h C o l u m b i a t h e C o r d i l l e r a n s t e p p e i s r e s t r i c t e d i n i t s d i s t r i b u t i o n , s o u t h w a r d s from c e n t r a l W a s h i n g t o n t o C a l i f o r n i a and A r i z o n a i t i s more e x t e n s i v e , d o m i n a t i n g much o f t h e C o r d i l l e r a n l a n d s c a p e . The s t e p p e s o u t h o f t h e s t u d y a r e a a l s o t e n d s t o be much r i c h e r i n p l a n t s p e c i e s a n d community t y p e s , r e f l e c t i n g n o t o n l y t h e i n c r e a s e d a r i d i t y and i n c r e a s e d t e m p e r a t u r e s w i t h p r o g r e s s i o n s o u t h w a r d s b u t a l s o t h e l a c k o f g l a c i a t i o n s i n t h e a r e a ( S t e b b i n s and M a j o r 1965). However, t h e g e n e r a l p hysiognomy i s much t h e same. T h a t i s , s h r u b s (and i n some a r e a s t r e e s ) and h e r b a c e o u s p l a n t s , 19 i n t e r s p a c e d w i t h c r y p t o g a m c r u s t s , a r e c h a r a c t e r i s t i c ( s e e Daubenmire 1970 and 1978 f o r f u r t h e r comment). i v ) SOILS B r y o p h y t e s i n h a b i t o n l y t h e t o p few c e n t i m e t e r s o f t h e s o i l s u r f a c e . No s t a n d a r d c l a s s i f i c a t i o n i s a v a i l a b l e t o d e s c r i b e t h i s h i g h l y v a r i a b l e and h e t e r o g e n o u s v e n e e r . Numerous and u s u a l l y i n t e r r e l a t e d s o i l c h a r a c t e r i s t i c s , m a i n l y r e l a t i v e s t a b i l i t y , t e x t u r e , o r g a n i c c o n t e n t , c h e m i c a l c o m p o s i t i o n and d e p o s i t i o n of wind blown m a t e r i a l ( l o e s s ) , upon w h i c h t h e h i g h d e g r e e o f d i s t u r b a n c e i s s u p e r i m p o s e d , v a r y not o n l y f r o m s i t e t o s i t e , b ut w i t h i n s i t e s as w e l l . F i v e s u p e r f i c i a l s o i l t y p e s were r e c o g n i z e d d u r i n g t h e s u r v e y ( i n p a r t b a s e d on V a l e n t i n e and L a v k u l i c h 1978): a) loamy s o i l s - composed of a h e t e r o g e n o u s m i x t u r e o f medium ( s a n d s ) and f i n e ( c l a y s and s i l t s ) p a r t i c l e s ; o r g a n i c component h i g h l y v a r i a b l e , o f t e n a b s e n t i n open a r e a s but a b undant n e a r v a s c u l a r p l a n t s ; c a l c i u m l e v e l u s u a l l y h i g h , a l t h o u g h sometimes low; m i c r o t o p o g r a p h i c a l l y d i v e r s e ; medium r e s i s t a n c e t o d i s t u r b a n c e . b) c l a y and s i l t r i c h s o i l s - v e r y f i n e l y t e x t u r e d ; v e r y r i c h i n c a l c i u m and u s u a l l y low i n o r g a n i c s ; h i g h l y s t a b l e , v e r y h a r d when d r y ; i n most a r e a s n o t a s common as t h e f i r s t s o i l t y p e ; common n e a r a n i m a l e x c a v a t i o n s o r r o a d c u t s , l o c a l l y d o m i n a n t , however, on or n e a r l a c u s t r i n e b a n k s . c) sandy s o i l s - m o s t l y composed o f sand, o n l y s m a l l amounts of f i n e s ; v e r y u n s t a b l e and p o r o u s , r e a d i l y d i s t u r b e d 20 when d r y ; c a l c i u m c o n t e n t h i g h l y v a r i a b l e , o f t e n a b s e n t , o r g a n i c s v a r i a b l e , o f t e n low; uncommon, u s u a l l y i n m i c r o s i t e s i n a r e a s d o m i n a t e d by t h e loamy s o i l s . d) r o c k y o r s t o n y s o i l s - s i m i l a r t o t h e loamy s o i l t y p e , e x c e p t much r o c k i e r and sometimes s a n d i e r ; u s u a l l y on s l o p e s a d j a c e n t t o o u t c r o p s . e) a l k a l i n e s o i l s - a d j a c e n t t o d e p r e s s i o n s b u t a l s o f l a n k i n g some i n t e r m i t t e n t s t r e a m g u l l i e s ; v e r y h a r d when d r y , wet d u r i n g p a r t of t h e y e a r , u s u a l l y t h r o u g h w i n t e r i n t o e a r l y s p r i n g ; a l t h o u g h t h i s s o i l t y p e i s e x t r e m e l y h i g h i n s a l t c o n t e n t , t h e v e r y t o p l a y e r may be l e a c h e d o f s a l t s ( F l o w e r s 1973), an i m p o r t a n t c o n s i d e r a t i o n w i t h r e s p e c t t o b r y o p h y t e d i s t r i b u t i o n s . 21 I I . METHODS 1. F I E L D METHODS S a m p l i n g l o c a t i o n s were s e l e c t e d i n i t i a l l y by a u t o m o b i l e r e c o n n a i s a n c e , t h e n were s u r v e y e d on f o o t . P r e f e r r e d s i t e s were t h o s e w i t h a h i g h d e g r e e of h a b i t a t d i v e r s i t y and m i n i m a l d i s t u r b a n c e . S t r e a m and l a k e m a r g i n s , wooded g u l l e y s and c l o s e d -c a n o p i e d p i n e f o r e s t s were e x c l u d e d s i n c e t h e i r b r y o f l o r a s , as w e l l as t h e i r l i c h e n and v a s c u l a r f l o r a s , were n o t i n d i c a t i v e of s t e p p e c l i m a t e s . Open f o r e s t s i t e s a t lower e l e v a t i o n s ( u s u a l l y below 600m) were i n c l u d e d , however. A l s o , b e c a u s e o f a f f i n i t i e s t o t h e f o r e s t z o n e , h i g h e l e v a t i o n s t e p p e ( e q u i v a l e n t t o t h e u p p e r and m i d d l e g r a s s l a n d z o n e s o f S p i l s b u r y and T i s d a l e 1944) was e x c l u d e d . The m a r g i n s o f a l k a l i n e d e p r e s s i o n s were i n c l u d e d i n t h e s u r v e y even t h o u g h t h e y a r e a t t i m e s w e t t e r t h a n t h e s u r r o u n d i n g s t e p p e . They a r e u s u a l l y c o n s i d e r e d t o be c h a r a c t e r i s t i c of s e m i - a r i d c l i m a t e s (Daubenmire 1978, W a l t e r 1983) and t h e i r f l o r a i s not r e p r e s e n t a t i v e o f v e g e t a t i o n of h i g h e r e l e v a t i o n s . The b e s t t i m e s t o c o l l e c t were i n p e r i o d s when t h e s o i l was damp, s u c h as a f t e r r a i n s o r , sometimes, i n t h e e a r l y m o r n i n g . A t s u c h t i m e s some of t h e b r y o p h y t e t a x a were r e v i v e d and b r i g h t g r e e n , c o n t r a s t i n g w i t h t h e s u b s t r a t u m and making them more v i s i b l e . S p r i n g and autumn s e a s o n s t e n d t o be w e t t e r , t h u s t h e y a r e t h e p r e f e r r e d s e a s o n s t o c o l l e c t . In a d d i t i o n , most s p o r o p h y t e s , i f p r e s e n t a t a l l , t e n d t o mature d u r i n g t h e s p r i n g p e r i o d . N e v e r t h e l e s s , s a t i s f a c t o r y c o l l e c t i o n s c a n be made 22 t h r o u g h o u t t h e y e a r . A few of t h e t a x a a r e e p h e m e r a l s and, a l t h o u g h t h e y may not a l w a y s be i n i d e a l c o n d i t i o n f o r d e t e r m i n a t i o n , most t a x a a r e i d e n t i f i a b l e a t a l l t i m e s . W i t h t h e e x c e p t i o n of a few autumn and w i n t e r c o l l e c t i n g t r i p s , most o f t h e f i e l d w o r k was done d u r i n g t h e s p r i n g and summer months from 1979 t o 1982. A t o t a l o f one h u n d r e d and t h i r t y - e i g h t d r y l a n d s i t e s were sampled, one h u n d r e d and t e n o f w h i c h a r e i n s o u t h - c e n t r a l B r i t i s h C o l u m b i a ( F i g . 1 ) . Twenty-e i g h t a d d i t i o n a l s i t e s were sampled i n Oregon, W a s h i n g t o n and A l b e r t a . L o c a t i o n s o f t h e v a r i o u s s i t e s a r e f o u n d i n A p p e n d i c e s B and C. 2. LABORATORY METHODS C o l l e c t i o n s were u s u a l l y e x amined w i t h b o t h d i s s e c t i n g and compound m i c r o s c o p e s . A number o f l e a v e s were s t r i p p e d f r o m t h e stem o f i n d i v i d u a l p l a n t s . B o t h d o r s a l and v e n t r a l s u r f a c e s o f l e a v e s were e x p o s e d . In many c a s e s i t was n e c e s s a r y t o make a s e r i e s o f t r a n s v e r s e s e c t i o n s o f t h e u p p e r h a l v e s o f one or more l e a v e s . A l t h o u g h t i m e c o n s u m i n g , t h e p r o d u c t i o n o f c r o s s -s e c t i o n s i s c r i t i c a l t o t h e i d e n t i f i c a t i o n o f many t a x a . Permanent s l i d e s r e p r e s e n t i n g most o f t h e t a x a were p r e p a r e d u s i n g H o y e r ' s s o l u t i o n . T h e s e s l i d e s were o f t e n u s e d f o r r e f e r e n c e b o t h f o r i d e n t i f i c a t i o n p u r p o s e s and f o r t h e p r e p a r a t i o n o f f i g u r e s . Most i d e n t i f i c a t i o n s were made u s i n g t h e f o l l o w i n g s e r i e s o f f l o r a s : F l o w e r s ( 1 9 7 3 ) , G r o u t ( 1 9 2 8 - 1 9 4 1 ) , Lawton ( 1 9 7 1 ) , 23 S m i t h ( 1 9 7 8 a ) , and, l e s s f r e q u e n t l y , Crum and A n d e r s o n ( 1 9 8 1 ) , B i l e w s k y ( 1 9 6 5 ) , Agnew and V o n d r e c e k (1975) and Nyholm (1954-1969). Many f a m i l i a l and g e n e r i c monographs were a l s o c o n s u l t e d ( e . g . , Kramer 1980, S a i t o 1975 and V i t t 1973). P u z z l i n g s p e c i m e n s were s e n t t o v a r i o u s a u t h o r i t i e s f o r i d e n t i f i c a t i o n o r c o n f i r m a t i o n . T h e s e a u t h o r i t i e s a r e c i t e d when a p p r o p r i a t e . C o l l e c t i o n s a r e d e p o s i t e d i n t h e U n i v e r s i t y o f B r i t i s h C o l u m b i a (UBC, see A p p e n d i x D ) , t h e N a t i o n a l Museum of Canada i n Ottawa (CANM) and t h e B u f f a l o Museum o f N a t u r a l H i s t o r y ( BUF). C o l l e c t i o n s made by o t h e r r e s e a r c h e r s were a l s o examined. Most of t h e s e a r e a t t h e U n i v e r s i t y o f B r i t i s h C o l u m b i a but some were f o u n d a t t h e U n i v e r s i t y o f W a s h i n g t o n i n S e a t t l e and a t W a s h i n g t o n S t a t e U n i v e r s i t y i n P u l l m a n . In t o t a l o v e r 6500 c o l l e c t i o n s were i d e n t i f i e d , most o f w h i c h had more t h a n one t a x o n p r e s e n t . T h u s , w e l l o v e r 10,000 i d e n t i f i c a t i o n s were made d u r i n g t h e c o u r s e of t h e s t u d y . 24 I I I . THE BRYOPHYTES OF THE SEMI-ARID STEPPE OF SOUTH-CENTRAL BRITISH COLUMBIA 1. INTRODUCTION S e v e n t y - s e v e n s p e c i e s o f mosses and f i v e s p e c i e s o f l i v e r w o r t s , r e p r e s e n t i n g 16 f a m i l i e s and 37 g e n e r a , were c o l l e c t e d d u r i n g t h e s u r v e y . The s y s t e m a t i c p r e s e n t a t i o n o f t h e mosses u s e d h e r e f o l l o w s t h e s y s t e m d e v e l o p e d by F l e i s c h e r (1900-1923) and s u b s e q u e n t l y employed by B r o t h e r u s ( 1 9 2 4 - 1 9 2 5 ) , Crum and A n d e r s o n ( 1 9 8 1 ) , G r o u t (1928-1941), Lawton (1971) and S m i t h ( 1 9 7 8 a ) . The c l a s s i f i c a t i o n of t h e h e p a t i c s i s b a s e d on t h e f o r m a t of S t o t l e r and C r a n d a l l - S t o t l e r ( 1 9 7 7 ) . The n o m e n c l a t u r e , as a r u l e , f o l l o w s Crum e t a l . (1973) and W i j k e t a l . (1959-1969) f o r t h e mosses, and, f o r t h e h e p a t i c s , S t o t l e r and C r a n d a l l - S t o t l e r ( 1 9 7 7 ) . E x c e p t i o n s a r e n o t e d i n t h e t e x t . D e s c r i p t i o n s t h a t accompany e a c h s p e c i e s and v a r i e t y f o l l o w a p a r t i c u l a r s e q u e n c e . F i r s t , commonly u s e d o r h i s t o r i c a l l y i m p o r t a n t synonyms a r e l i s t e d . T h e s e a r e a c c o m p a n i e d by a b r i e f d e s c r i p t i o n o f t h e t a x o n . The p u r p o s e of t h i s d e s c r i p t i o n i s t w o - f o l d . In most c a s e s , i t p r e s e n t s a c o n c i s e summary of t h e more r e l e v a n t m a c r o s c o p i c c h a r a c t e r s u s e d i n i d e n t i f y i n g t h e t a x o n . In a few c a s e s , i t p r o v i d e s a more c o m p l e t e d e s c r i p t i o n o f a t a x o n t h a t i s n o t f u l l y d e s c r i b e d i n r e a d i l y a v a i l a b l e f l o r a s (examples i n c l u d e B r y o e r y t h r o p h y l l u m columbianum, Phaseurn  v l a s s o v i i and P t e r y q o n e u r u m k o z l o v i i ) . T e r m i n o l o g y i n t h e d e s c r i p t i o n s f o l l o w s Crum and A n d e r s o n ( 1 9 8 1 ) , I r e l a n d (1982) 25 and S m i t h ( 1 9 7 8 a ) . R e p r e s e n t a t i v e s p e c i m e n s a r e l i s t e d . I l l u s t r a t i o n s a r e o f t e n c r i t i c a l f o r t h e p o s i t i v e i d e n t i f i c a t i o n of some t a x a . I l l u s t r a t i o n s a v a i l a b l e e l s e w h e r e t h a t a r e c o n s i d e r e d t o be u s e f u l a r e c i t e d f o l l o w i n g t h e d e s c r i p t i o n s o f t h e v a r i o u s t a x a . I l l u s t r a t i o n s p r e s e n t e d i n t h i s d i s s e r t a t i o n a r e a l s o l i s t e d h e r e . The g e o g r a p h i c a l d i s t r i b u t i o n o f e a c h t a x o n i s d i v i d e d i n t o t h r e e c a t e g o r i e s ; A: r e g i o n a l d i s t r i b u t i o n ( w i t h i n t h e s t u d y a r e a ) B: N o r t h A m e r i c a n d i s t r i b u t i o n C: w o r l d d i s t r i b u t i o n . M a j o r r a n g e e x t e n s i o n s a r e n o t e d i n t h e r e s p e c t i v e c a t e g o r i e s . S u m m a r i z i n g r e m a r k s f o l l o w t h e d i s t r i b u t i o n s e c t i o n . T h e s e remarks i n c l u d e r e l e v a n t d i s t i n g u i s h i n g c h a r a c t e r s and comments on m o r p h o l o g i c a l l y s i m i l a r t a x a . Comments r e g a r d i n g t a x o n o m i c and n o m e n c l a t u r a l d e c i s i o n s a r e i n t r o d u c e d h e r e o r i n t h e f a m i l y and g e n e r i c i n t r o d u c t i o n s ( t h e s e a r e c o n s i d e r e d f u r t h e r i n C h a p t e r V I ) . O t h e r r e l e v a n t t o p i c s , i n c l u d i n g p h y t o g e o g r a p h y and e c o l o g y , a r e d i s c u s s e d i n t h e remarks s e c t i o n . The h e p a t i c s a r e t r e a t e d f o l l o w i n g t h e mosses. A c o m p l e t e l i s t o f t h e s p e c i e s from t h e s t u d y a r e a f o l l o w s t h e g e n e r i c key. The keys t h a t have been d e v e l o p e d h e r e a r e b a s e d p r i m a r i l y on v e g e t a t i v e c h a r a c t e r s . S p o r o p h y t i c c h a r a c t e r s a r e u s e d s p a r i n g l y and u s u a l l y o n l y s e c o n d a r i l y . I n some c a s e s , v e g e t a t i v e c h a r a c t e r s a r e i n s u f f i c i e n t t o d i s t i n g u i s h between t a x a , and s p o r o p h y t i c c h a r a c t e r s must be u s e d . F o r t u n a t e l y , 26 s p o r o p h y t e s a r e common i n t h e s e i n s t a n c e s . A r e l i a n c e on v e g e t a t i v e c h a r a c t e r s i s p r a c t i c a l , s i n c e most d r y l a n d b r y o p h y t e c o l l e c t i o n s a r e s t e r i l e . F u r t h e r m o r e , even when s p o r o p h y t e s a r e f o u n d , o f t e n t h e y a r e immature, p a r t i a l l y decomposed o r d e f o r m e d , and t h u s a r e n o t v e r y u s e f u l f o r i d e n t i f i c a t i o n . When u s i n g some a v a i l a b l e f l o r a s t o i d e n t i f y d r y l a n d b r y o p h y t e s , a m a j o r drawback i s t h e i r r e l i a n c e on s p o r o p h y t i c c h a r a c t e r s . T h i s i s u n f o r t u n a t e , s i n c e i t makes t h e keys u s a b l e f o r o n l y f e r t i l e c o l l e c t i o n s . N e v e r t h e l e s s , a number of keys have p r o v e n t o be h e l p f u l , a t l e a s t i n p a r t , f o r t h e i d e n t i f i c a t i o n of s t e p p e b r y o p h y t e s . T h e s e i n c l u d e F l o w e r s (1961, 1973), G r o u t ( 1 9 2 8 - 1 9 4 1 ) , M a g i l l (1976) and, t o a l e s s e r d e g r e e , Agnew and V o n d r e c e k (1975) and B i l e w s k y ( 1 9 6 5 ) . U n l e s s o t h e r w i s e n o t e d , t h e l e a f c e l l s r e f e r r e d t o i n t h e keys a r e t h o s e o f t h e u p p e r and m e d i a l p o r t i o n and not of t h e b a s a l s e c t i o n o f t h e l e a f , w h i c h u s u a l l y e x h i b i t s a v e r y d i f f e r e n t a r e o l a t i o n . B a s a l c e l l s a r e u s u a l l y l o n g e r , l e s s d i s t i n c t l y c o l o u r e d and t h i n n e r - w a l l e d t h a n t h e c e l l s a b o v e . F u r t h e r m o r e , c e n t r a l l a m i n a l c e l l s s h o u l d be u s e d b e c a u s e t h e m a r g i n a l and p a r a c o s t a l c e l l s sometimes d i f f e r i n s i z e , shape and p a p i l l o s i t y f r o m t h e o t h e r s . C e l l s o f a v e r a g e s i z e s h o u l d be u s e d f o r c a l c u l a t i n g measurements, and a b e r r a n t c e l l s s h o u l d be a v o i d e d . S p e c i e s and g e n e r a t h a t have n o t y e t been f o u n d i n t h e s t u d y a r e a , b u t a t some p o i n t m i g h t be c o l l e c t e d b e c a u s e t h e y a r e i n a d j a c e n t a r e a s , a r e i n c l u d e d i n t h e keys and, o c c a s i o n a l l y , i n t h e d i s c u s s i o n s . 27 2. KEY TO THE GENERA 1 mosses; p l a n t s c o n s p i c u o u s l y l e a f y , t h e l e a v e s u n l o b e d ....2 1 1 h e p a t i c s ; p l a n t s t h a l l o s e , o r , i f l e a f y , t h e n l e a v e s i n c o n s p i c u o u s , l o b e d and p u r p l e t o b l a c k 49 2 p l a n t s e r e c t o r a s c e n d i n g , a n c h o r e d a t t h e b a s e , sometimes a p p e a r i n g b u d - l i k e ( a c r o c a r p o u s m o s s e s ) ; l e a v e s o f t e n t w i s t e d a r o u n d t h e stem; upper and m e d i a l l e a f c e l l s u s u a l l y i s o d i a m e t r i c , o c c a s i o n a l l y h e x a g o n a l - r h o m b o i d a l or r e c t a n g u l a r , r a r e l y l i n e a r , o f t e n p a p i l l o s e ; c o s t a u s u a l l y e x c u r r e n t o r p e r c u r r e n t ...3 2 1 p l a n t s p r o s t r a t e o r c r e e p i n g , u s u a l l y a n c h o r e d by r h i z o i d s on u n d e r s i d e o f t h e stem, not a p p e a r i n g b u d - l i k e ( p l e u r o c a r p o u s m o s s e s ) ; l e a v e s n e v e r t w i s t e d a r o u n d t h e stem; upper and m e d i a l l e a f c e l l s l i n e a r t o r h o m b o i d a l o r o v a l , n e v e r q u a d r a t e , a l w a y s smooth; c o s t a u s u a l l y n o t r e a c h i n g apex 45 3 l e a v e s w i t h l a m e l l a e , f i l a m e n t s o r c o n s p i c u o u s l y b u l g i n g , p a p i l l o s e c e l l s on t h e v e n t r a l s u r f a c e of t h e c o s t a ( F i g . 10-13, 16), sometimes c o s t a u n e v e n l y b u l g i n g w i t h w e a k l y d e v e l o p e d l a m e l l a e , o f t e n c r e s t e d w i t h i n d i v i d u a l c e l l s , r e a d i l y s een from above w i t h d i s s e c t i n g m i c r o s c o p e ( F i g . 12) 4 3 1 l e a v e s w i t h o u t l a m e l l a e o r f i l a m e n t s on t h e v e n t r a l s u r f a c e o f t h e c o s t a ; some t a x a have a w e l l - d e v e l o p e d , e v e n l y b u l g i n g c o s t a ( F i g . 8 and 9 ) , but i n d e p e n d e n t , b u l g i n g c e l l s a r e a b s e n t 9 4 l e a v e s w i t h l a m e l l a e t h a t a r e o c c a s i o n a l l y i n d i s t i n c t 5 4 1 l e a v e s w i t h f i l a m e n t s or b u l g i n g c e l l s ( F i g . 10-12, 16) 7 5 c o s t a w i t h numerous l a m e l l a e (>20) e x t e n d i n g most of t h e l e n g t h of t h e l e a f ; l e a f m a r g i n s i n f o l d e d o v e r t h e l a m e l l a e ; l e a v e s l a n c e o l a t e from a b r o a d s h e a t h i n g b a s e P o l y t r i c h u m 5 1 c o s t a w i t h 2 t o 4 l a m e l l a e on t h e upper h a l f o f t h e l e a f o n l y ; l e a f m a r g i n s p l a n e t o r e c u r v e d ; l e a v e s o v a t e t o s p a t u l a t e 6 6 l a m e l l a e i n c o n s p i c u o u s , p a p i l l o s e , o f t e n f u s e d a t t h e base t o f o r m a c u s h i o n ( F i g . 12); l a m i n a l c e l l s p a p i l l o s e ; s p o r o p h y t e s r a r e ; c a p s u l e s a l w a y s o p e r c u l a t e and s h o r t e x s e r t e d C r o s s i d i u m 28 6 1 l a m e l l a e w e l l d e v e l o p e d , smooth t o r a r e l y w e a k l y p a p i l l o s e , n e v e r f u s e d a t t h e base ( F i g . 13); l a m i n a l c e l l s smooth t o o c c a s i o n a l l y weakly p a p i l l o s e ; s p o r o p h y t e s common; c a p s u l e s c l e i s t o c a r p o u s o r o p e r c u l a t e , immersed o r s h o r t - e x s e r t e d P t e r y q o n e u r u m 7 f i l a m e n t s 15-30 and 4-7 c e l l s h i g h , c o v e r i n g most of t h e upper h a l f o f t h e l e a f s u r f a c e ; l e a f m a r g i n s s t r o n g l y i n f o l d e d ; l e a v e s a p p e a r i n g f l e s h y A l o i n a 7 1 f i l a m e n t s 3-5, o f t e n p o o r l y d e v e l o p e d (1-4 c e l l s h i g h ) , r e s t r i c t e d t o c e n t r a l p a r t of t h e a d a x i a l l e a f s u r f a c e , sometimes t o t h r e e o r f o u r c e l l s h i g h b u t u s u a l l y r e p r e s e n t e d o n l y by b u l g i n g , b a r r e l - o r b o t t l e - s h a p e d c e l l s , w i t h p r o j e c t i n g upper ends ( F i g . 16); l e a f m a r g i n s r e c u r v e d t o p l a n e , n e v e r i n f o l d e d ; l e a v e s n e v e r a p p e a r i n g f l e s h y 8 8 c a p s u l e s s p h e r i c a l , c l e i s t o c a r p o u s and immersed; l e a v e s o v a t e - l a n c e o l a t e t o l a n c e o l a t e , u s u a l l y t a p e r e d t o t h e apex, r a r e l y r o u n d e d above; l a m i n a l p a p i l l a e low and s c a t t e r e d o r on t a l l , h o l l o w p r o j e c t i o n s ( F i g . 16); b u l g i n g c e l l s on c o s t a b a r r e l - o r b o t t l e - s h a p e d ; awns o f t e n l o n g and o r a n g e i n c o l o u r Phaseurn 8 1 c a p s u l e s c y l i n d r i c a l , o p e r c u l a t e and e x s e r t e d ; l e a v e s o v a t e -l a n c e o l a t e t o o v a t e or s p a t u l a t e , o f t e n r o u n d e d above; l a m i n a l p a p i l l a e low, e i t h e r w e l l d e v e l o p e d o r i n d i s t i n c t a s weak l y f o r m e d mammillae i n t h e c e n t e r o f t h e c e l l ; c o s t a l f i l a m e n t s u s u a l l y 2-4 c e l l s h i g h , sometimes t a l l e r , o c c a s i o n a l l y a s b u l g i n g c e l l s and , i f s o , t h e n b a r r e l -s h a ped, n e v e r b o t t l e - s h a p e d ; awns l o n g t o o f t e n s h o r t o r a b s e n t , u s u a l l y r e d o r o c c a s i o n a l l y o r a n g e i n c o l o u r C r o s s i d i u m 9 lumens o f upper o r m e d i a l l a m i n a l c e l l s p a p i l l o s e 10 9 1 lumens o f upper o f m e d i a l l a m i n a l c e l l s smooth 29 10 awns p r e s e n t on some l e a v e s 11 10 1 awns a b s e n t , but c o s t a may be e x c u r r e n t a s an a p i c u l u s o r c u s p i d a t e p o i n t 16 11 p l a n t s y e l l o w - g r e e n o r p a l e g r e e n ; l e a v e s u s u a l l y <1.5mm l o n g , o v a t e t o l a n c e o l a t e , u s u a l l y t a p e r e d t o t h e apex; b a s a l p o r t i o n o f t h e l e a f n o t d i f f e r i n g d i s t i n c t l y i n c o l o u r from u p p e r p a r t ; c a p s u l e s c l e i s t o c a r p o u s (and e i t h e r immersed a n d s p h e r i c a l , o r s h o r t - e x s e r t e d and o v o i d t o s h o r t - c y l i n d r i c ) o r s t e g o c a r p o u s ; i f p e r i s t o m e p r e s e n t t h e n b a s a l t u b e s h o r t o r a b s e n t 12 29 11 1 p l a n t s g r e e n t o d a r k g r e e n , o f t e n b l a c k i s h ; l e a v e s g e n e r a l l y >1.5mm, o f t e n >2mm l o n g , a t l e a s t some l e a v e s l i g u l a t e o r o b l o n g , u s u a l l y w i t h an o b t u s e t o e m a r g i n a t e apex, o c c a s i o n a l l y t a p e r e d ; b a s a l p a r t of t h e l e a f u s u a l l y l i g h t e r i n c o l o u r t h a n t h e upper p o r t i o n of t h e l e a f ; c a p s u l e s s t e g o c a r p o u s c y l i n d r i c a l , e x s e r t e d ( i f s h o r t - e x s e r t e d t h e n c o v e r e d by l o n g - c a m p a n u l a t e c a l y p t r a ) ; i f l o n g - e x s e r t e d t h e n b a s a l t u b e o f p e r i s t o m e w e l l d e v e l o p e d ) 14 12 c a p s u l e s immersed, s p h e r i c a l and c l e i s t o c a r p o u s , sometimes i n c o n s p i c u o u s and h i d d e n i n t h e l e a v e s Phaseurn 12 1 c a p s u l e s e x s e r t e d , o v o i d t o s h o r t c y l i n d r i c a l , c l e i s t o c a r p o u s t o s t e g o c a r p o u s 13 13 upper m a r g i n s o f l e a f o f t e n composed o f l o n g e r and smoother c e l l s t h a n a d j a c e n t l a m i n a l c e l l s , sometimes f o r m i n g t e e t h ; c a p s u l e s s h o r t - c y l i n d r i c a l t o o v o i d , s h o r t - e x s e r t e d ; s e t a e u s u a l l y <1 cm l o n g , e i t h e r c l e i s t o c a r p o u s o r o p e r c u l a t e , sometimes s y s t y l i o u s ( o p e r c u l u m r e t a i n e d on c a p s u l e by c o l u m e l l a ) ; p e r i s t o m e a b s e n t o r v e r y r e d u c e d P o t t i a 13 1 upper m a r g i n s o f l e a f n o t o r w e a k l y m o d i f i e d ( o c c a s i o n a l l y composed o f l o n g e r and smoother c e l l s t h a n a d j a c e n t l a m i n a l c e l l s , b u t i n d i s t i n c t l y s o , n e v e r f o r m i n g t e e t h ) ; c a p s u l e s s h o r t - c y l i n d r i c and l o n g - e x s e r t e d ; s e t a e u s u a l l y >1 cm, a l w a y s o p e r c u l a t e , n e v e r s y s t y l i o u s ; p e r i s t o m e w e l l d e v e l o p e d Desmatodon ( g u e p i n i i ) 14 b a s a l c e l l s w i t h t r a n s v e r s e w a l l s u s u a l l y o r a n g e and t h i c k e r t h a n l o n g i t u d i n a l w a l l s ; s p o r o p h y t e s common; c a l y p t r a l o n g and c a m p a n u l a t e , c o m p l e t e l y c o v e r i n g t h e c a p s u l e ; c a p s u l e s s h o r t - e x s e r t e d E n c a l y p t a 14 1 b a s a l c e l l s w i t h t r a n s v e r s e w a l l s u s u a l l y o f t h e same t h i c k n e s s a s l o n g i t u d i n a l w a l l s , n o t o r a n g e c o l o u r e d ; s p o r o p h y t e s i n f r e q u e n t t o common i n a few t a x a ; c a l y p t r a c u c u l l a t e , n o t c o v e r i n g a l l o f t h e c a p s u l e ; c a p s u l e s l o n g -e x s e r t e d 15 15 awns smooth, u s u a l l y t a p e r e d ( w i d e r a t t h e b a s e t h a n a b o v e ) ; l e a v e s u s u a l l y t a p e r e d t o apex; upper m a r g i n s w e a k l y r e c u r v e d , many l e a v e s l y i n g on t h e i r s i d e on a s l i d e ; b a s a l p o r t i o n o f t h e l e a f n o t a l w a y s c o n s p i c u o u s l y l i g h t e r i n c o l o u r t h a n u p p e r p a r t o f l e a f ; p e r i s t o m e s h o r t , l a c k i n g b a s a l t u b e o r b a s a l t u b e i n c o n s p i c u o u s Desmatodon 15 1 awns r o u g h o r smooth, i f smooth t h e n n o t d i s t i n c t l y t a p e r e d and l e a v e s o b o v a t e t o o v a t e w i t h w e a k l y r e c u r v e d t o r e v o l u t e 30 u pper m a r g i n s , u s u a l l y l y i n g f l a t on a s l i d e ; b a s a l p o r t i o n . of t h e l e a f u s u a l l y c o n s p i c u o u s l y l i g h t e r i n c o l o u r t h a n u p p e r p a r t of l e a f ; p e r i s t o m e l o n g , w i t h t a l l b a s a l t u b e T o r t u l a 16 p l a n t s g r o w i n g on r o c k , r a r e l y on wood, o f t e n g r e e n above and b l a c k i s h below, sometimes b l a c k i s h t h r o u g h o u t ; l e a v e s u s u a l l y e r e c t t o o c c a s i o n a l l y s l i g h t l y t w i s t e d ; c o s t a e n d i n g b e f o r e apex; c a p s u l e s common, immersed t o s l i g h t l y e x s e r t e d , c o v e r e d by c o m p a n u l a t e , h a i r y c a l y p t r a e O r t h o t r i c h u m 16 1 p l a n t s u s u a l l y on s o i l , o c c a s i o n a l l y a s red-brown c u s h i o n s on r o c k f a c e s (Didymodon o c c i d e n t a l i s ) , o t h e r w i s e b l a c k i s h or brown t o o c c a s i o n a l l y g r e e n t h r o u g h o u t ; l e a v e s e r e c t t o c r i s p e d ; c o s t a e x c u r r e n t a s mucro o r p e r c u r r e n t ; c a p s u l e s uncommon, l o n g - e x s e r t e d ( s h o r t - e x s e r t e d i n W e i s s i a o n l y ) , c o v e r e d by s m a l l , smooth and c u c u l l a t e c a l y p t r a e 17 17 upper m a r g i n s of l e a f s t r o n g l y i n v o l u t e ; l e a v e s g r e e n , c r i s p e d when d r y ; u s u a l l y n e a r b o u l d e r s on s l o p e s i n open p i n e s t a n d s We i s s i a 17 1 upper m a r g i n s of l e a f r e c u r v e d , p l a n e o r w e a k l y ^ i n v o l u t e ; l e a v e s o f v a r i o u s d i s p o s i t i o n when d r y , o n l y a few t a x a w i t h c r i s p e d l e a v e s and i f s o , u s u a l l y b l a c k i s h o r brown i n c o l o u r ; h a b i t a t v a r i o u s 18 18 upper h a l f o f l e a f l a m i n a m o s t l y b i s t r a t o s e , o c c a s i o n a l l y t r i - s t r a t o s e i n p a t c h e s ; u p p e r l a m i n a l c e l l s s t r o n g l y p a p i l l o s e ; l e a v e s o v a t e t o s p a t u l a t e , o f t e n b r o a d l y c u c u l l a t e T o r t u l a ( c a n i n e r v i s ) 18 1 l e a v e s u s u a l l y u n i s t r a t o s e , r a r e l y b i s t r a t o s e a t apex o r a l o n g m a r g i n s , n e v e r t r i s t r a t o s e ; upper l a m i n a l c e l l s o b s c u r e l y t o s t r o n g l y p a p i l l o s e ; l e a v e s l a n c e o l a t e t o s p a t u l a t e , r a r e l y o v a t e , o c c a s i o n a l l y c u c u l l a t e 19 19 l e a v e s u s u a l l y l a n c e o l a t e t o s h o r t - t r i a n g u l a r l a n c e o l a t e ; c o s t a n e v e r b r o a d l y c o n v e x on upper a d a x i a l s u r f a c e ; m i d d l e s u p e r f i c i a l c e l l s o f t h e c o s t a on t h e a b a x i a l s u r f a c e s i m i l a r t o c e l l s of t h e a d j a c e n t l a m i n a ( q u a d r a t e t o v e r y s h o r t - r e c t a n g u l a r ) ; e l o n g a t e c e l l s i n f r e q u e n t ; a r e o l a t i o n o f c o s t a l s u r f a c e n ot o b s c u r e 20 19 1 l e a v e s s p a t u l a t e t o o v a t e - l a n c e o l a t e , o c c a s i o n a l l y t r i a n g u l a r - l a n c e o l a t e ; c o s t a sometimes b r o a d l y c o n v e x on t h e upper a d a x i a l s u r f a c e ( F i g . 8 and 9 ) ; most s u p e r f i c i a l c e l l s of t h e c o s t a on t h e a b a x i a l s u r f a c e e l o n g a t e o r l i n e a r , c o n s p i c u o u s l y d i f f e r e n t t h a n t h e l a m i n a l c e l l s (examine 31 c e l l s f r o m m i d - l e a f o r j u s t a b ove, some t a x a have s h o r t e r , n e a r l y i s o d i a m e t r i c c e l l s n e a r apex of c o s t a ) ; a r e o l a t i o n o f c o s t a l s u r f a c e o f t e n o b s c u r e 21 20 l e a v e s g r e e n t o r e d d i s h - b r o w n and wea k l y c r i s p e d t o e r e c t ; m a r g i n s u s u a l l y u n i s t r a t o s e a bove, u s u a l l y r e c u r v e d ; c o s t a w e a k l y b u l g i n g t o f l a t i n c r o s s - s e c t i o n , b a s a l c e l l s n o t t h i n - w a l l e d or b u l g i n g Didymodon 2 0 1 l e a v e s u s u a l l y b l a c k i s h and s t r o n g l y c r i s p e d ; m a r g i n s b i s t r a t o s e a b ove, w e a k l y r e c u r v e d t o p l a n e ; c o s t a u s u a l l y b u l g i n g - c o n v e x i n c r o s s - s e c t i o n ; b a s a l c e l l s o f t e n t h i n -w a l l e d a n d o c c a s i o n a l l y a p p e a r i n g i n f l a t e d , e s p e c i a l l y i n l a r g e r l e a v e s T r i c h o s t o m o p s i s 21 c o s t a b r o a d ( o f t e n >8 c e l l s wide) and s t r o n g l y c o n v e x on v e n t r a l s u r f a c e , composed of v e r t i c a l l y e l o n g a t e d c e l l s ( F i g . 8 and 9) 22 2 1 1 c o s t a n a r r o w e r ( u s u a l l y <5 c e l l s wide; e . g . , F i g . 17) and o n l y o c c a s i o n a l l y c o n v e x ( i f so t h e n l e a v e s l a n c e o l a t e t o l i g u l a t e a n d composed o f v e r t i c a l l y more o r l e s s i s o d i a m e t r i c c e l l s ) 23 22 l e a v e s , when d r y , u s u a l l y s p i r a l l y t w i s t e d a r o u n d stem, r e d g r e e n t o d a r k o l i v a c e o u s ; apex d e e p l y c o n c a v e t o c u c u l l a t e , u s u a l l y b l u n t and p a r t l y composed o f s u b - q u a d r a t e c e l l s , sometimes a c u t e ( F i g . 9 ) ; p l a n t s f o u n d a s i n d i v i d u a l s o r s m a l l c o l o n i e s on s o i l Desmatodon ( c o n v o l u t u s ) 2 2 1 l e a v e s , when d r y , u s u a l l y e r e c t o r s l i g h t l y t w i s t e d a bove, d a r k r e d - b r o w n ; apex a c u t e and composed o f s u b - l i n e a r c e l l s , o f t e n s l i g h t l y c o n c a v e , n e v e r c u c u l l a t e , a l t h o u g h r a r e l y s u b - c u c u l l a t e ( F i g . 8 ) ; p l a n t s u s u a l l y f o u n d a s compact t u r f s on s o i l n e a r o u t c r o p s B r y o e r y t h r o p h y l l u m (columbianum) 23 u p p e r m a r g i n s s t r o n g l y r e v o l u t e ( F i g . 17); upper l a m i n a b r o a d l y c h a n n e l l e d ( F i g . 17) P s e u d o c r o s s i d i u m 2 3 1 m a r g i n s r e c u r v e d t o p l a n e , n o t s t r o n g l y r e v o l u t e ; upper l a m i n a n o t b r o a d l y c h a n n e l l e d 24 24 b a s a l l e a f c e l l s t h i n - w a l l e d and a p p e a r i n g w e a k l y i n f l a t e d ; c o s t a b r o a d ( u s u a l l y >5 c e l l s wide) and c o n v e x i n c r o s s s e c t i o n ; apex of l e a f u s u a l l y m u c r o n a t e ; p l a n t s red-brown t o d a r k g r e e n ; l e a v e s u s u a l l y s t r o n g l y c r i s p e d B r y o e r y t h r o p h y l l u m ( r e c u r v i r o s t r u m ) 24 1 b a s a l l e a f c e l l s t h i n - o r t h i c k - w a l l e d , n e v e r a p p e a r i n g 32 i n f l a t e d ; c o s t a n a r r o w e r and e i t h e r w e a k l y c o n v e x or i n a l a m i n a l c h a n n e l ; apex m u c r o n a t e o r r o u n d e d ; p l a n t s u s u a l l y g r e e n t o b l a c k i s h ; l e a v e s r a r e l y c r i s p e d 25 25 p l a n t s of s e a s o n a l l y wet s i t e s ; l e a v e s t a p e r e d t o apex; m a r g i n s u s u a l l y t o o t h e d and b o r d e r e d by smooth, t h i c k - w a l l e d and r e c t a n g u l a r c e l l s ; s p o r o p h y t e s common; o p e r c u l u m u s u a l l y r e t a i n e d on c a p s u l e by c o l u m e l l a ( s y s t y l i o u s ) P o t t i a ( h e i m i i ) 2 5 1 p l a n t s of d r i e r s t e p p e ; l e a v e s u s u a l l y b r o a d l y r o u n d e d above, r a r e l y t a p e r i n g t o apex; m a r g i n s n o t t o o t h e d , o c c a s i o n a l l y w e a k l y b o r d e r e d by smooth, t h i c k - w a l l e d and r e c t a n g u l a r c e l l s ; s p o r o p h y t e s o c c a s i o n a l ; c a p s u l e s n ot s y s t y l i o u s 26 26 b a s a l c e l l s w i t h t r a n s v e r s e w a l l s o r a n g e and t h i c k e r t h a n l o n g i t u d i n a l w a l l s ; s p o r o p h y t e s o f t e n p r e s e n t ; c a l y p t r a l o n g and c a m p a n u l a t e , c o m p l e t e l y c o v e r i n g t h e c a p s u l e ; c a p s u l e s s h o r t - e x s e r t e d E n c a l y p t a 2 6 1 b a s a l c e l l s w i t h t r a n s v e r s e w a l l s u s u a l l y o f t h e same t h i c k n e s s a s l o n g i t u d i n a l w a l l s , n o t o r a n g e c o l o u r e d ; s p o r o p h y t e s i n f r e q u e n t ; c a l y p t r a c u c u l l a t e , c o v e r i n g o n l y t h e t o p p a r t of t h e c a p s u l e ; c a p s u l e s l o n g - e x s e r t e d 27 27 upper a d a x i a l c o s t a i n narrow c h a n n e l B a r b u l a 2 7 1 u p p e r a d a x i a l c o s t a f l a t o r c o n v e x , n o t i n a narrow c h a n n e l 28 28 l e a v e s u s u a l l y y e l l o w - g r e e n t o p a l e g r e e n ; b a s a l p o r t i o n o f t h e l e a f u s u a l l y t h e same c o l o u r a s t h e upper p a r t o r o n l y s l i g h t l y l i g h t e r ; a d a x i a l s u r f a c e o f t h e c o s t a e v e n l y c o n v e x ; l e a f l a m i n a u n i s t r a t o s e ; m a r g i n a l c e l l s sometimes l o n g e r and smoother t h a n a d j a c e n t l a m i n a l c e l l s Desmatodon 28 1 l e a v e s d a r k g r e e n t o b l a c k i s h , o c c a s i o n a l l y brown; b a s a l p o r t i o n o f t h e l e a f u s u a l l y d i s t i n c t l y l i g h t e r t h a n upper p a r t ; a d a x i a l s u r f a c e of t h e c o s t a f l a t t o , r a r e l y , w e a k l y c o n v e x ; l a m i n a u n i - t o t r i s t r a t o s e ; m a r g i n a l c e l l s t h e same s i z e a s t h e l a m i n a l c e l l s T o r t u l a 29 l e a v e s l i n e a r t o l o n g - l a n c e o l a t e ; m a r g i n s p l a n e t o i n c u r v e d , i n w h i c h c a s e t h e l e a v e s a p p e a r t u b u l a r , e s p e c i a l l y i n t h e l o w e r h a l v e s 30 2 9 1 l e a v e s o v a t e - l a n c e o l a t e o r l a n c e o l a t e t o o b l o n g o r 33 s p a t h u l a t e , not l i n e a r o r l o n g - l a n c e o l a t e ; m a r g i n s o f t e n r e c u r v e d , o c c a s i o n a l l y p l a n e ; l e a v e s n e v e r t u b u l o s e below, sometimes s u b - t u b u l o s e i n u p p e r t h i r d 32 30 l e a v e s s t r o n g l y c r i s p e d t o e r e c t ; a l a r c e l l s y e l l o w i s h and u s u a l l y i n f l a t e d ; p l a n t s common i n a d j a c e n t a r e a s , not y e t i n s t u d y a r e a ( Dicranum) 3 0 1 l e a v e s c o n t o r t e d o r e r e c t , n e v e r c r i s p e d ; a l a r c e l l s not i n f l a t e d o r c o l o u r e d . . . 31 31 p l a n t s f o r m i n g d ense t u r f s ; l e a v e s e v e n l y d i s t r i b u t e d a l o n g stems, n e v e r as a r o s e t t e a t apex of stem, l e a v e s e r e c t , u s u a l l y d a r k g r e e n ; s p o r o p h y t e s n o t f o u n d ; common on s h a d e d , d r y s l o p e s o f l a c u s t r i n e banks o r n e a r r o c k o u t c r o p s D i t r i c h u m 3 1 1 p l a n t s u s u a l l y as s c a t t e r e d i n d i v i d u a l s , n e v e r a s d e n s e t u r f s , u s u a l l y w i t h a r o s e t t e o f l e a v e s a t t h e apex o f t h e stem; l e a v e s wide s p r e a d i n g and c o n t o r t e d , g r e e n t o y e l l o w -g r e e n ; s p o r o p h y t e s common; c a p s u l e s p y r i f o r m ; u s u a l l y i n waste p l a c e s or s e a s o n a l l y wet a r e a s L e p t o b r y u m 32 p l a n t s d a r k g r e e n , b l a c k i s h o r d a r k brown, r a r e l y l i g h t g r e e n , o f t e n f o r m i n g c u s h i o n s on r o c k or t u r f s on s o i l ; u p p e r l e a f c e l l s w i t h t h i c k w a l l s , q u a d r a t e and a b o u t 10 m i c r o m e t e r s i n l o n g e s t d i m e n s i o n , t o , r a r e l y , o v a t e -r e c t a n g u l a r and a b o u t 15 m i c r o m e t e r s i n l o n g e s t d i m e n s i o n ; n e v e r i n s e a s o n a l l y wet s i t e s 33 3 2 1 p l a n t s y e l l o w - g r e e n t o l i g h t g r e e n , r a r e l y b r o w n i s h o r d a r k g r e e n , u s u a l l y on s o i l , r a r e l y on r o c k ; upper l e a f c e l l s w i t h t h i c k o r t h i n w a l l s , u s u a l l y i r r e g u l a r l y i s o d i a m e t r i c t o r h o m b o i d a l o r e l o n g a t e , and u s u a l l y much l o n g e r t h a n 15 m i c r o m e t e r s i n l o n g e s t d i m e n s i o n , o c c a s i o n a l l y >50 m i c r o m e t e r s ; some of t h e s e t a x a f o u n d o n l y i n o r a d j a c e n t t o s e a s o n a l l y wet s i t e s 39 33 p l a n t s d a r k g r e e n t o b l a c k , o c c a s i o n a l l y g r e e n o r r e d - b r o w n , f o r m i n g d e n s e c u s h i o n s o r mats on r o c k o r on v e r y s t o n y s o i l s u r f a c e s , r a r e l y on f i n e g r a i n e d s o i l ; a t l e a s t some l e a v e s awned (awns o f t e n v e r y s h o r t ) , r a r e l y o t h e r w i s e 34 3 3 1 p l a n t s g r e e n t o red-brown, o c c a s i o n a l l y b l a c k i s h and i f s o , t h e n n o t f o r m i n g c u s h i o n s , u s u a l l y a s l o o s e t o open c o l o n i e s o r a s i s o l a t e d i n d i v i d u a l s on s o i l o r r o c k , o c c a s i o n a l l y f o r m i n g c u s h i o n s on r o c k (Didymodon o c c i d e n t a l i s ) ; l e a v e s n e v e r awned 36 34 l e a v e s o c c a s i o n a l l y p l i c a t e a b o v e ; awns v e r y l o n g (1/2 t o 1X l e n g t h o f l a m i n a ) ; l e a f m a r g i n s w e a k l y r e c u r v e d t o , r a r e l y , 34 p l a n e ; u p p e r l e a f l a m i n a i r r e g u l a r l y b i s t r a t o s e t o u n i s t r a t o s e ; b a s a l c e l l s u s u a l l y l o n g r e c t a n g u l a r ; , c a p s u l e s common, s h o r t - e x s e r t e d o r n e a r l y immersed, c o v e r e d by l a r g e c a m p a n u l a t e and p l i c a t e c a l y p t r a e Cose i n o d o n 34 1 l e a v e s n e v e r p l i c a t e ; awns s h o r t e r ( u s u a l l y much s h o r t e r t h a n 1/2 l e n g t h o f l e a f l a m i n a ) , sometimes i n c o n s p i c u o u s ; l e a f m a r g i n s u s u a l l y p l a n e , sometimes i n c u r v e d , r a r e l y r e c u r v e d ; u p p e r l e a f l a m i n a e v e n l y b i s t r a t o s e ( o c c a s i o n a l l y p a t c h y - t r i s t r a t o s e ) t o u n i s t r a t o s e (sometimes w i t h o n l y m a r g i n s b i s t r a t o s e ) , r a r e l y u p p e r l a m i n a i r r e g u l a r l y b i s t r a t o s e ; b a s a l c e l l s u s u a l l y s h o r t - r e c t a n g u l a r , i f l o n g -r e c t a n g u l a r t h e n a t l e a s t some c e l l s , e i t h e r a t l e a f b a s e o r i n m i d - l e a f , somewhat s i n u o u s ; c a p s u l e s common i n some t a x a but g e n e r a l l y r a r e , e x s e r t e d o r immersed; c a l y p t r a e s m a l l and smooth, c u c u l l a t e or m i t r a t e , n e v e r c a m p a n u l a t e o r c o v e r i n g a l l of t h e c a p s u l e 35 35 p l a n t s f o r m i n g c u s h i o n s o r , o f t e n , v e r y b r o a d t u r f s ; a t l e a s t some l e a v e s n e a r apex long-awned ( t o 1/2 t h e l e n g t h of t h e l e a f o r l o n g e r ) ; c a p s u l e s e x s e r t e d o r immersed, i f immersed t h e n t h e c a p s u l e i s g l o b o s e and a s y m m e t r i c a l l y v e n t r i c o s e a t b a s e , t h e s e t a c u r v e d and t h e c o l u m e l l a n o t f a l l i n g w i t h o p e r c u l u m Grimmia 3 5 1 p l a n t s f o r m i n g s m a l l t o , r a r e l y , e x t e n s i v e t u r f s , r a r e l y f o r m i n g c u s h i o n s ; l e a v e s u s u a l l y s h o r t - a w n e d ( u s u a l l y much s h o r t e r t h a n 1/2 o f t h e l e a f , o f t e n i n c o n s p i c u o u s or a b s e n t ) ; c a p s u l e s immersed, c y l i n d r i c a l a n d n o t v e n t r i c o s e ; s e t a s t r a i g h t and t h e c o l u m e l l a f a l l i n g w i t h o p e r c u l u m S c h i s t i d i u m 36 p l a n t s g r o w i n g on r o c k , u s u a l l y b l a c k i s h t h r o u g h o u t , sometimes g r e e n above a n d b l a c k i s h below; l e a v e s u s u a l l y e r e c t t o s l i g h t l y t w i s t e d ; c o s t a e n d i n g b e f o r e apex; c a p s u l e s common, immersed t o e m e r g e n t , c o v e r e d by c a m p a n u l a t e , w e a k l y h a i r y c a l y p t r a O r t h o t r i c h u m 3 6 1 p l a n t s u s u a l l y g r o w i n g on s o i l , r a r e l y on r o c k , b l a c k i s h t o e v e n l y brown o r g r e e n ; l e a v e s e r e c t t o c r i s p e d ; c o s t a o f t e n e x c u r r e n t a s mucro o r p e r c u r r e n t ; c a p s u l e s r a r e , e x s e r t e d ; c a l y p t r a c u c u l l a t e , smooth, n e v e r c a m p a n u l a t e or h a i r y 37 37 l e a f m a r g i n s r e c u r v e d , u s u a l l y w e a k l y t o o t h e d above; a r e o l a t i o n o f t h e a d a x i a l s u r f a c e o f upper c o s t a o b s c u r e , composed o f n a r r o w l y e l o n g a t e c e l l s ; e x t r e m e l y common, f o u n d a t e v e r y s i t e C e r a t o d o n 3 7 1 l e a f m a r g i n s p l a n e t o r e c u r v e d , n ot t o o t h e d above; 35 a r e o l a t i o n of t h e a d a x i a l s u r f a c e o f upper c o s t a e a s i l y s e e n , composed of q u a d r a t e o r s h o r t - r e c t a n g u l a r c e l l s ; common t o o c c a s i o n a l 38 38 p l a n t s g r e e n t o red-brown; l e a v e s weakly c r i s p e d t o e r e c t ; m a r g i n s u s u a l l y u n i s t r a t o s e a b o v e , u s u a l l y r e c u r v e d ; c o s t a w e a k l y b u l g i n g t o f l a t i n c r o s s - s e c t i o n , b a s a l c e l l s n o t t h i n - w a l l e d o r b u l g i n g Didymodon 38 1 p l a n t s b l a c k i s h , o c c a s i o n a l l y d a r k b r o w n i s h - g r e e n o r g r e e n ; l e a v e s u s u a l l y c r i s p e d ; m a r g i n s b i s t r a t o s e a b o v e , w e a k l y r e c u r v e d t o p l a n e ; c o s t a u s u a l l y b u l g i n g - c o n v e x i n c r o s s -s e c t i o n ; b a s a l c e l l s o f t e n t h i n - w a l l e d and o c c a s i o n a l l y a p p e a r i n g i n f l a t e d , e s p e c i a l l y i n l a r g e r l e a v e s T r i c h o s t o m o p s i s 39 l e a v e s l a n c e o l a t e t o o v a t e - l a n c e o l a t e ; u p p e r and m i d d l e c e l l s l i n e a r t o e l o n g a t e - r h o m b o i d a l , u s u a l l y narrow (commonly <15 m i c r o m e t e r s w i d e ) ; s p o r o p h y t e s uncommon 40 3 9 1 l e a v e s b r o a d l y o v a t e o r o v a t e - l a n c e o l a t e t o o b l o n g o r s p a t u l a t e ; upper and m i d d l e c e l l s i r r e g u l a r - i s o d i a m e t r i c t o r h o m b o i d a l ( u s u a l l y >15 m i c r o m e t e r s w i d e ) ; s p o r o p h y t e s common 41 40 c o s t a e x c u r r e n t as awn o r e n d i n g b e f o r e apex; upper and m e d i a l l e a f c e l l o b l o n g - h e x a g o n a l t o r h o m b o i d a l ; l e a v e s b o r d e r e d o r not Bryum 40 1 c o s t a p e r c u r r e n t ( n e v e r a s an awn) o r e n d i n g w e l l below apex; u p p e r c e l l s l i n e a r t o l i n e a r - r h o m b o i d a l ; l e a v e s n o t c l e a r l y b o r d e r e d P o h l i a 41 l e a v e s o v a t e , w i t h t a p e r i n g a p i c e s and l o n g awns; upper l e a f c e l l s e l o n g a t e and t h i c k - w a l l e d , f o r m i n g a d i s t i n c t h y a l i n e p a t c h ; m a r g i n s r e v o l u t e ; l a m i n a l c e l l s , a s v i e w e d i n c r o s s -s e c t i o n , w i t h d o r s a l w a l l s much t h i c k e r t h a n v e n t r a l w a l l s ; p l a n t s i n s t e p p e , r e s e m b l i n g t i n y c a b b a g e s S t e g o n i a 4 1 1 l e a v e s o v a t e - l a n c e o l a t e t o o b l o n g o r s p a t u l a t e , w i t h o r w i t h o u t awns; m a r g i n s p l a n e t o w e a k l y r e c u r v e d , n e v e r r e v o l u t e ; upper l e a f c e l l s o n l y s l i g h t l y d i f f e r e n t f r o m m e d i a l c e l l s , n o t f o r m i n g a h y a l i n e p a t c h ; l a m i n a l c e l l s more o r l e s s e q u a l l y t h i c k e n e d d o r s a l l y and v e n t r a l l y ; p l a n t s u s u a l l y f o u n d i n s e a s o n a l l y wet s i t e s , o c c a s i o n a l l y i n s t e p p e , v a r i o u s l y s h a p e d , but n e v e r r e s e m b l i n g t i n y c a b b a g e s 42 42 l e a v e s u s u a l l y b o r d e r e d by a band o f narrow, t h i c k - w a l l e d 36 and s m a l l - l u m e n e d c e l l s ( c h e c k a t t h e m i d d l e o f t h e l e a f ) ; m e d i a l l a m i n a l c e l l s u s u a l l y <20x1fJ m i c r o m e t e r s , some c e l l s t o 30x20 m i c r o m e t e r s , r a r e l y l a r g e r ; c a p s u l e s c e r n u u s and not p y r i f o r m , or c a p s u l e s e r e c t , c y l i n d r i c a l a nd o p e r c u l u m r e m a i n i n g a t t a c h e d t o c a p s u l e by c o l u m e l l a . .. 43 42 1 l e a v e s n o t b o r d e r e d , o r , r a r e l y , i n d i s t i n c t l y b o r d e r e d by a band of somewhat n a r r o w e r c e l l s ; m e d i a l l a m i n a l c e l l s o f t e n l a r g e , o f t e n >50x20 m i c r o m e t e r s ; c a p s u l e s p y r i f o r m and c u r v e d , o r e r e c t and c y l i n d r i c a l t o s u b - p y r i f o r m , o p e r c u l u m d e c i d u o u s ...44 43 c a p s u l e s c e r n u o u s , u s u a l l y w i t h a w e l l d e v e l o p e d p e r i s t o m e , sometimes l a c k i n g , n e v e r s y s t y l i o u s ; l e a v e s , e s p e c i a l l y below t h e m i d d l e , b o r d e r e d by t h i c k - w a l l e d , narrow c e l l s ; n ot y e t f o u n d i n s t u d y a r e a , b u t v e r y common i n a l k a l i n e d e p r e s s i o n s n o r t h - w e s t o f t h e a r e a ( i . e . , between C l i n t o n and W i l l i a m s L a k e ) (Desmatodon c e r n u u s ) 43 1 c a p s u l e s e r e c t , p e r i s t o m e a b s e n t and s y s t y l i o u s ; l e a v e s w e a k l y b o r d e r e d by somewhat t h i c k - w a l l e d , narrow c e l l s P o t t i a ( n e v a d e n s i s ) 44 c a p s u l e s i n c l i n e d , p y r i f o r m a n d a s y m m e t r i c F u n a r i a 44 1 c a p s u l e s e r e c t , s h o r t - c y l i n d r i c t o s u b - p y r i f o r m and s y m m e t r i c E n t o s t h o d o n 45 c o s t a d o u b l e o r s i n g l e , s h o r t , r a r e l y r e a c h i n g m i d l e a f ; l e a v e s b r o a d l y o v a t e - l a n c e o l a t e 46 45 1 c o s t a s i n g l e , e x t e n d i n g beyond m i d l e a f , o f t e n t o apex; l e a v e s l a n c e o l a t e t o o v a t e - l a n c e o l a t e 47 46 p l a n t s d a r k g r e e n t o g r e e n , b r a n c h e s j u l a c e o u s ; l e a v e s t i n y , u s u a l l y a b o u t 0.5 mm l o n g , s h o r t a c u m i n a t e and w e a k l y f a l c a t e ; c o s t a d o u b l e o r s i n g l e ; f o r m i n g compact p a t c h e s on r o c k P s e u d o l e s k e e l l a 4 6 1 p l a n t s g r e e n o r y e l l o w g r e e n , b r a n c h e s not j u l a c e o u s ; l e a v e s l a r g e r ( g e n e r a l l y > 1mm), u s u a l l y l o n g a c u m i n a t e and f a l c a t e ; c o s t a d o u b l e ; f o r m i n g l o o s e mats on r o c k , o r s o i l a d j a c e n t t o r o c k Hypnum 47 p l a n t s g r e e n t o y e l l o w - g r e e n , u s u a l l y w i t h m e t a l l i c l u s t r e ; l e a v e s d e e p l y p l i c a t e and s t r a i g h t ; many b r a n c h e s h o o ked a t t i p s when d r y ; on r o c k , o c c a s i o n a l l y on s o i l H o m a l o t h e c i u m 37 4 7 1 p l a n t s y e l l o w - g r e e n t o p a l e g r e e n , w i t h o u t m e t a l l i c l u s t r e ; l e a v e s p l a n e t o w e a k l y p l i c a t e , o f t e n somewhat f a l c a t e ; b r a n c h e s n o t hooked a t t i p s ; on s o i l , r a r e l y on r o c k . . . .48 48 l e a v e s d e l t o i d t o o v a t e - l a n c e o l a t e ; l e a f apex a c u t e t o o b t u s e , o f t e n w e a k l y t o o t h e d , t h e a p i c a l c e l l s much s h o r t e r t h a n t h e l i n e a r m e d i a l c e l l s E u r h y n c h i u m 48 1 l e a v e s o v a t e - ^ l a n c e o l a t e t o l a n c e o l a t e ; l e a f apex u s u a l l y a c u m i n a t e , n o t t o o t h e d , t h e a p i c a l c e l l s n o t much d i f f e r e n t t h a n l i n e a r m e d i a l c e l l s B r a c h y t h e c ium 49 p l a n t s l e a f y and c o m p r i s e d of f i l a m e n t o u s b r a n c h e s , o f t e n i n t e r t w i n e d w i t h mosses; u s u a l l y i n r o c k y s i t e s C e p h a l o z i e l l a 4 9 1 p l a n t s t h a l l o s e ; h a b i t a t v a r i o u s 50 50 s m a l l p l a n t s , u s u a l l y < 1cm i n d i a m e t e r , f o r m i n g c i r c u l a r p a t c h e s ; s c a l e s i n c o n s p i c u o u s on u n d e r s u r f a c e o f t h a l l u s R i c e i a 5 0 1 l a r g e r p l a n t s , u s u a l l y much l a r g e r t h a n 1cm wide, composed of s t r a p - l i k e t h a l l i f o r m i n g i r r e g u l a r p a t c h e s ; l a r g e s c a l e s p r e s e n t on u n d e r s u r f a c e o f t h a l l u s .....51 51 v e n t r a l s c a l e s d a r k p u r p l e ; p l a n t s f o r m i n g d a r k , o f t e n f r o s t e d , w o r m l i k e s t r u c t u r e s on s o i l s u r f a c e when d r y , a r e s u l t o f t h e i n c u r v i n g o f t h e t h a l l u s m a r g i n s ; c e l l s o f upper t h a l l u s s u r f a c e more o r l e s s i s o d i a m e t r i c , o f t e n w i t h c o r n e r t h i c k e n i n g s ; a i r p o r e s n o t s t e l l a t e - t h i c k e n e d a s v i e w e d f r o m a b o v e ; c a r p o c e p h a l a p r o d u c e d a t apex o f t h a l l u s M a n n i a 5 H v e n t r a l s c a l e s h y a l i n e t o p a l e p u r p l e ; p l a n t s n o t f o r m i n g w o r m l i k e s t r u c t u r e s on s o i l when d r y ; c e l l s on upper s u r f a c e o f t h e t h a l l u s r e c t a n g u l a r t o e l o n g a t e - r e c t a n g u l a r and l a c k i n g c o r n e r t h i c k e n i n g s ; a i r p o r e s s t e l l a t e - t h i c k e n e d a s v i e w e d from a b o v e ; c a r p o c e p h a l a p r o d u c e d i n t h e c e n t e r o f t h e t h a l l u s A t h a l a m i a 38 3. L I S T OF BRYOPHYTES FOUND IN THE STUDY AREA A l o i n a b i f r o n s (De Not.) D e l g a d i l l o A l o i n a b r e v i r o s t r i s (Hook, e t G r e v . ) K i n d b e r g A l o i n a r i g i d a (Hedw.) L i m p r . A t h a l a m i a h y a l i n a (Sommerf.) H a t t . B a r b u l a c o n v o l u t a Hedw. B a r b u l a u n g u i c u l a t a Hedw. B r a c h y t h e c i u m a l b i c a n s (Hedw.) B.S.G. B r a c h y t h e c i u m c o l l i n u m (C. M u e l l . ) B.S.G. B r y o e r y t h r o p h y l l u m columbianum (Herm. e t Lawt.) Zander B r y o e r y t h r o p h y l l u m r e c u r v i r o s t r u m (Hedw.) Chen Bryum a r g e n t e u m Hedw. Bryum c a e s p i t i c i u m Hedw. C e p h a l o z i e l l a d i v a r i c a t a (Sm.) S c h i f f . C e r a t o d o n p u r p u r e u s (Hedw.) B r i d . C o n a r d i a compacta (C. M u e l l . ) H. R o b i n s . C o s c i n o d o n c a l y p t r a t u s (Hook.) C. J e n s . C o s c i n o d o n c r i b r o s u s (Hook.) C. J e n s . C r o s s i d i u m a b e r r r a n s H o l z . e t B a r t r . C r o s s i d i u m r o s e i W i l l i a m s C r o s s i d i u m s p . Desmatodon c o n v o l u t u s ( B r i d . ) G r o u t Desmatodon q u e p i n i i B.S.G. Desmatodon o b t u s i f o l i u s (Schwaegr.) Schimp. Didymodon f a l l a x (Hedw.) Zander Didymodon o c c i d e n t a l i s Zander Didymodon r i q i d u l u s v a r . g r a c i l i s ( S c h l e i c h . ex H o o k . e t G r e v . ) 39 Zander Didymodon v i n e a l i s v a r . b r a c h y p h y l l u s ( S u l l . i n W h i p p l . ) Zander Didymodon v i n e a l i s v a r . l u r i d u s ( H o r n s c h . i n S p r e n g . ) Zander Didymodon v i n e a l i s v a r . v i n e a l i s Z ander D i t r i c h u m f l e x i c a u l e v a r . densum (B.S.G.) B r a i t h w . E n c a l y p t a r h a p t o c a r p a Schwaegr. E n c a l y p t a v u l g a r i s Hedw. E n t o s t h o d o n r u b i g i n o s u s ( W i l l i a m s ) G r o u t E u r h y n c h i u m p u l c h e l l u m (Hedw.) J e n n . F u n a r i a h y g r o m e t r i c a Hedw. F u n a r i a m u h l e n b e r g i i T u r n . Grimmia a l p e s t r i s (Web. e t Mohr) S c h l e i c h . ex. Nees. Grimmia anodon B.S.G. Grimmia l a e v i g a t a ( B r i d . ) B r i d . Grimmia o v a l i s (Hedw.) L i n d b . Grimmia p l a g i o p o d i a Hedw. Grimmia p u l v i n a t a (Hedw.) Sm. Grimmia t r i c h o p h y l l a G r e v . H o m a l o t h e c i u m aenium ( M i t t . ) Lawt. Hypnum r e v o l u t u m ( M i t t . ) L i n d b . Hypnum v a u c h e r i L e s q . L e p t o b r y u m p y r i forme (Hedw.) W i l s . Mannia f r a g r a n s ( B a l b i s ) F r y e e t C l a r k O r t h o t r i c h u m anomalum Hedw. O r t h o t r i c h u m c u p u l a t u m B r i d . O r t h o t r i c h u m h a l l i i S u l l . e t L e s q . O r t h o t r i c h u m l a e v i g a t u m Z e t t . 40 O r t h o t r i c h u m r u p e s t r e S c h l e i c h . e t Schwaegr. Phascum c u s p i d a t u m Hedw. Phascum v l a s s o v i i L a z a r . P o h l i a c r u d a (Hedw.) L i n d b . P o l y t r i c h u m j u n i p e r i n u m Hedw. P o l y t r i c h u m p i l i f e r u m Hedw. P o t t i a b r y o i d e s ( D i c k s . ) M i t t . P o t t i a h e i m i i (Hedw.) F u r n . ex. Hampe P o t t i a n e v a d e n s i s C a r d , e t T h e r . P o t t i a c f . w i l s o n i i (Hook.) B.S.G. P s e u d o c r o s s i d i u m r e v o l u t u m ( B r i d . i n S c h r a d . ) Zander P s e u d o l e s k e e l l a t e c t o r u m ( B r i d . ) B r o t h . P t e r y q o n e u r u m k o z l o v i i L a z a r . P t e r y q o n e u r u m l a m e l l a t u m ( L i n d b . ) J u r . P t e r y q o n e u r u m ovatum (Hedw.) D i x . P t e r y g o n e u r u m s u b s e s s i l e ( B r i d . ) J u r . R i c c i a c r y s t a l l i n a L . R i c c i a s o r o c a r p a B i s c h . S c h i s t i d i u m apocarpum (Hedw.) B.S.G. S c h i s t i d i u m h e t e r o p h y l l u m ( K i n d b . ) M c i n t o s h comb. nov. i n e d . S t e q o n i a l a t i f o l i a v a r . p i l i f e r a ( B r i d . ) B r o t h . T o r t u l a b r e v i p e s ( L e s q . ) B r o t h . T o r t u l a c a n i n e r v i s ( M i t t . ) B r o t h . T o r t u l a i n t e r m e d i a ( B r i d . ) De N o t . T o r t u l a r u r a l i s (Hedw.) G a e r t n . T r i c h o s t o m o p s i s a u s t r a l a s i a e (Hook, e t G r e v . ) H. R o b i n s . W e i s s i a b r a c h y c a r p a C M . 41 4. DESCRIPTIONS AND DISCUSSIONS OF TAXA DITRICHACEAE A l t h o u g h o n l y t h e D i t r i c h a c e a e i s t r e a t e d h e r e , t h r e e s p e c i e s o f D i c r a n u m i n t h e c l o s e l y r e l a t e d f a m i l y D i c r a n a c e a e , D. b r e v i f o l i u m , D. s c o p a r i u m and D. t a u r i c u m , a r e f o u n d i n m e s i c s i t e s a d j a c e n t t o t h e s t u d y a r e a . Two g e n e r a i n t h e D i t r i c h a c e a e , C e r a t o d o n and D i t r i c h u m , e a c h r e p r e s e n t e d by a s i n g l e s p e c i e s , a r e p r e s e n t i n t h e s t u d y a r e a . CERATODON B r i d . CERATODON PURPUREUS (Hedw.) B r i d . C. c o n i c u s (Hamp.) L i n d b . S m a l l p l a n t s ; g a m e t o p h y t e s u s u a l l y <0.5cm t a l l , g r e e n t o y e l l o w - g r e e n , sometimes r e d - b r o w n ; s c a t t e r e d o r i n l o o s e t u f t s on s o i l o r , r a r e l y , l o g s , u s u a l l y m i x e d w i t h o t h e r m o s s e s . S p o r o p h y t e s o c c a s i o n a l and p r i m a r i l y i n m e s i c s i t e s , w i t h d i s t i n c t i v e r e d d i s h , b a n a n a - s h a p e d and r i b b e d c a p s u l e s . R e p r e s e n t a t i v e s p e c i m e n s : B88993, B88994, B88995 I l l u s t r a t i o n s : Crum and A n d e r s o n 1981, F l o w e r s 1973 D i s t r i b u t i o n : A: t h r o u g h o u t t h e s t u d y a r e a ( F i g . 33) B: w i d e s p r e a d i n N o r t h A m e r i c a C: c o s m o p o l i t a n , more common i n t e m p e r a t e l a t i t u d e s Remarks : C e r a t o d o n p u r p u r e u s i s t a x o n o m i c a l l y n o t o r i o u s n o t o n l y 42 b e c a u s e i t i s m o r p h o l o g i c a l l y v a r i a b l e but a l s o b e c a u s e i t i s o f t e n s t e r i l e and m i x e d w i t h o t h e r s u p e r f i c i a l l y s i m i l a r s p e c i e s . I f t h e key c h a r a c t e r s a r e f o l l o w e d c a r e f u l l y , however, i t s h o u l d n o t be m i s t a k e n f o r o t h e r t a x a . I t s most d i s t i n c t i v e f e a t u r e s a r e t h e o b s c u r e a r e o l a t i o n o f t h e a d a x i a l c o s t a l s u r f a c e , t h e r e c u r v e d l e a f m a r g i n s and t h e i r r e g u l a r l y d e n t a t e , u p p e r b o r d e r o f t h e l e a f . S p e c i e s o f Didymodon w i t h r e c u r v e d m a r g i n s l a c k t e e t h and t h e c o s t a l c e l l s a r e q u a d r a t e and e a s i l y n o t e d . P s e u d o c r o s s i d i u m r e v o l u t u m may be c o n f u s e d w i t h C. p u r p u r e u s b e c a u s e o f i t s s i m i l a r s i z e , growth h a b i t and c o l o u r but d i f f e r s m i c r o s c o p i c a l l y i n i t s d e n s e p a p i l l o s i t y and s t r o n g l y r e v o l u t e m a r g i n s . Bryum c a e s p i t i c i u m , w i t h w h i c h C. p u r p u r e u s o f t e n grows, has l o n g awns and l e a v e s t h a t a r e e r e c t t o o n l y s l i g h t l y t w i s t e d , whereas C. p u r p u r e u s has s h o r t awns o r none a t a l l , and t h e upper l e a v e s a r e u s u a l l y a l w a y s t w i s t e d a b o v e . M i c r o s c o p i c a l l y , t h e q u a d r a t e c e l l s o f C. p u r p u r e u s r e a d i l y s e p a r a t e i t f r o m B. c a e s p i t i c i u m , w h i c h has e l o n g a t e c e l l s . C e r a t o d o n c o n i c u s , a c c o r d i n g t o Lawton ( 1 9 7 1 ) , d o e s n o t d e s e r v e s p e c i e s s t a t u s . I a g r e e . The main c h a r a c t e r u s e d t o d e f i n e t h e t a x o n , t h e p r e s e n c e o f y e l l o w i s h awns, i s t o o v a r i a b l e w i t h i n p o p u l a t i o n s t o be of t a x o n o m i c s i g n i f i c a n c e . 43 DITRlCHUM Hampe DITRlCHUM FLEXICAULE (Schwaegr.) Hampe v a r . DENSUM (B.S.G.) B r a i t h w . P l a n t s f o r m i n g s h i n y - g r e e n t o y e l l o w - g r e e n t u r f s , u s u a l l y i n s h a d e d s i t e s i n open s i t u a t i o n s on l a c u s t r i n e b anks o r r o c k o u t c r o p s . S p o r o p h y t e s not c o l l e c t e d . R e p r e s e n t a t i v e s p e c i m e n s : B888996, B88997 I l l u s t r a t i o n s : Crum and A n d e r s o n 1981, I r e l a n d 1982 D i s t r i b u t i o n : A: f a i r l y common but s c a t t e r e d ( F i g . 49) B: p r o b a b l y w i d e s p r e a d a c r o s s t h e c o n t i n e n t C: c i r c u m p o l a r Remarks : The d i s t i n c t i v e a r e o l a t i o n , t h e somewhat c l a s p i n g , s u b -t u b u l a r l e a v e s and t h e s h i n y , y e l l o w g r e e n t u r f s a r e a l l c h a r a c t e r i s t i c o f D. f l e x i c a u l e . A l s o , t h e s e c h a r a c t e r s s h o u l d s u f f i c e i n s e p a r a t i n g i t f r o m m o r p h o l o g i c a l l y s i m i l a r t a x a , i n c l u d i n g C e r a t o d o n p u r p u r e u s and Didymodon s p p . ENCALYPTACEAE The c h a r a c t e r i s t i c l a r g e and l o n g - c a m p a n u l a t e c a l y p t r a t h a t c o v e r s a l l o f t h e c y l i n d r i c a l c a p s u l e , combined w i t h a l a r g e r a v e r a g e l e a f c e l l s i z e , a p r e v a i l i n g s y n o i c o u s c o n d i t i o n , and, i n most t a x a , t h i c k e n e d and c o l o u r e d t r a n s v e r s e b a s a l l e a f c e l l w a l l s , s e p a r a t e t h e E n c a l y p t a c e a e f r o m t h e P o t t i a c e a e (Agnew and V o n d r e c e k 1975, B i l e w s k y 1965, H o r t o n 1981). 44 ENCALYPTA Hedw. E n c a l y p t a r h a p t o c a r p a and, r a r e l y , E . v u l g a r i s , were f o u n d i n t h e s t u d y a r e a . A t h i r d s p e c i e s , E . i n t e r m e d i a , was f o u n d i n a d j a c e n t s i t e s . D. H o r t o n d e t e r m i n e d my c o l l e c t i o n s . T h e s e t h r e e t a x a may be t r e a t e d more r e a s o n a b l y a s v a r i e t i e s o f a s i n g l e h i g h l y v a r i a b l e t a x o n ( E . r h a p t o c a r p a ) , but a r e k e p t s e p a r a t e h e r e , f o l l o w i n g H o r t o n ( 1 9 8 1 ) . Key t o t h e S p e c i e s o f ENCALYPTA I most l e a v e s w i t h awns; s e t a l o n g and base of mature c a l y p t r a u s u a l l y p o s i t i o n e d w e l l above upper l e a v e s ; p e r i s t o m e p r e s e n t and u s u a l l y w e l l d e v e l o p e d ; c a p s u l e s u s u a l l y f u r r o w e d , r i b s p r o m i n e n t E. r h a p t o c a r p a I I l e a v e s u s u a l l y m u t i c o u s ; s e t a l o n g o r s h o r t , i n w h i c h c a s e t h e c a l y p t r a r e s t s on o r w i t h i n l e a v e s ; p e r i s t o m e a l w a y s a b s e n t t o v e s t i g i a l , n e v e r w e l l d e v e l o p e d ; c a p s u l e s more o r l e s s smooth t o l i g h t l y f u r r o w e d , r a r e l y w i t h p r o m i n e n t r i b s 2 b a s e o f c a l y p t r a p o s i t i o n e d w e l l above l e a v e s ; c a p s u l e more or l e s s smooth E . v u l g a r i s 2 1 b a s e o f c a l y p t r a immersed w i t h i n l e a v e s ; c a p s u l e u s u a l l y r i b b e d ; n o t y e t i n s t u d y a r e a ( E . i n t e r m e d i a ) ENCALYPTA RHAPTOCARPA Schwaegr. In open, b r o w n i s h - g r e e n t u r f s on s o i l , o f t e n n e a r o u t c r o p s o r on m o d e r a t e l y d i s t u r b e d s o i l i n s t e p p e . S p o r o p h y t e s common; c a p s u l e o f t e n r i b b e d ; p e r i s t o m e w e l l d e v e l o p e d t o , o f t e n , e r o d e d away o r p o o r l y d e v e l o p e d . R e p r e s e n t a t i v e s p e c i m e n s : B88998, B88999 I l l u s t r a t i o n s : F l o w e r s 1973, H o r t o n 1981 45 D i s t r i b u t i o n : A: w i d e s p r e a d ( F i g . 50) B: m a i n l y w e s t e r n N o r t h A m e r i c a , s c a t t e r e d p o p u l a t i o n s t h o u g h o u t h i g h a r c t i c and b o r e a l a r e a s ( s e e H o r t o n 1981 f o r maps) C: more o r l e s s c i r c u m p o l a r Remarks : E n c a l y p t a r h a p t o c a r p a i s an e x t r e m e l y common s t e p p e moss, e s p e c i a l l y i n a r e a s o f moderate d i s t u r b a n c e . S t e r i l e c o l l e c t i o n s c an be c o n f u s e d r e a d i l y w i t h s m a l l p l a n t s of T o r t u l a r u r a l i s ( s p a t u l a t e t o b r o a d - l a n c e o l a t e and s t r o n g l y p a p i l l o s e l e a v e s t h a t a r e t w i s t e d when d r y ) . I t d i f f e r s , however, i n i t s o r a n g e -c o l o u r e d and t h i c k - w a l l e d t r a n s v e r s e b a s a l c e l l w a l l s , f e a t u r e s l a c k i n g i n T o r t u l a . F o r t u n a t e l y , t h e s h o r t - e x s e r t e d c a p s u l e s and a c c o m p a n y i n g l o n g - c a m p a n u l a t e c a l y p t r a a r e u s u a l l y p r e s e n t , and r e a d i l y d i s t i n g u i s h p o p u l a t i o n s o f E n c a l y p t a . S p e c i e s o f T o r t u l a have l o n g - e x s e r t e d c a p s u l e s and s m a l l e r , c u c u l l a t e c a l y p t r a e , w h i c h c o v e r o n l y t h e t o p p o r t i o n s o f t h e c a p s u l e s . ENCALYPTA VULGARIS Hedw. In open, g r e e n t o b l u e - g r e e n t u f t s on s o i l . S p o r o p h y t e s common; c a p s u l e s l o n g - c y l i n d r i c a l , s h o r t - e x s e r t e d . R e p r e s e n t a t i v e s p e c i m e n : B89000 I l l u s t r a t i o n s : F l o w e r s 1973, H o r t o n 1981 D i s t r i b u t i o n : A: r a r e , s i t e 1 a t t h e base o f B o t a n i e M o u n t a i n n e a r L y t t o n ( F i g . 51) 46 B: m a i n l y s o u t h - w e s t e r n N o r t h A m e r i c a ; C a l i f o r n i a and New M e x i c o t o s o u t h e r n B r i t i s h C o l u m b i a ( s e e H o r t o n 1981 f o r maps) C: w e s t e r n N o r t h A m e r i c a / E u r a s i a ; b i p o l a r ( A u s t r a l i a , New Z e a l a n d ) Remarks : E n c a l y p t a v u l g a r i s i s a r a r e t a x o n i n t h e s t e p p e , f o u n d o n l y o n c e . See t h e r e m a r k s w i t h E . r h a p t o c a r p a . POTTIACEAE T h i s l a r g e and d i v e r s e f a m i l y i s r i c h l y r e p r e s e n t e d i n a r i d r e g i o n s . F o r example, i n t h e moss f l o r a s o f t h e p r i n c i p a l l y a r i d c o u n t r i e s o f I r a q and I s r a e l t h e P o t t i a c e a e make up about 35% o f t h e g e n e r a and 45% o f t h e s p e c i e s (Agnew and V o n d r e c e k 1975, B i l e w s k y 1965). In t h e p r e s e n t s t u d y t h e f a m i l y i n c l u d e s 24% o f t h e moss g e n e r a and 46% o f t h e s p e c i e s i n t h e a r e a . T h i s p r o m i n e n c e of t h e P o t t i a c e a e i n d r y r e g i o n s p r o b a b l y r e s u l t s , i n p a r t , f r o m a h i g h t o l e r a n c e t o d e s s i c a t i o n e x h i b i t e d by many members o f t h e f a m i l y . The P o t t i a c e a e i s a t a x o n o m i c a l l y t r o u b l e s o m e f a m i l y . In s p i t e o f v a l u a b l e and i n t e r e s t i n g r e g i o n a l s t u d i e s ( e . g . , S a i t o 1975, Zander 1981b) and t a x o n o m i c r e v i s i o n s (Kramer 1980, D e l g a d i l l o 1975, Zander 1978b) c o n s i d e r a b l e r e s e a r c h r e m a i n s t o be done. Zander (1982) r e v i e w e d t a x o n o m i c p r o b l e m s i n t h e f a m i l y . 47 ALOINA S p e c i e s o f A l o i n a a r e d i s t i n c t i v e i n t h e f i e l d , c h a r a c t e r i z e d by t h e s m a l l , b u d - l i k e and g r e g a r i o u s p l a n t s , e a c h composed o f o n l y a few l e a v e s , and by h a b i t a t , t e n d i n g t o be most common on c l a y - r i c h s o i l s u r f a c e s . M i c r o s c o p i c a l l y , t h e c o n c a v e l e a v e s , e a c h b e a r i n g a b u n d a n t , w e l l d e v e l o p e d f i l a m e n t s on t h e upper s u r f a c e s , a r e c h a r a c t e r i s t i c . T h r e e s p e c i e s o c c u r i n o u r a r e a . Of t h e t h r e e , A. b i f r o n s i s most d i s t i n c t i v e , w i t h i n d i v i d u a l l e a v e s b e a r i n g l o n g , h y a l i n e awns. The o t h e r two, A. b r e v i r o s t r i s and A. r i q i d a , r e q u i r e c a r e f u l s t u d y i n o r d e r t o d i s t i n g u i s h between them. D e l g a d i l l o (1975) s u p p l i e d d i s t r i b u t i o n a l maps f o r t h e t h r e e t a x a . Key t o t h e S p e c i e s o f ALOINA 1 l e a v e s p i l i f e r o u s ; p l a n t s u s u a l l y d a r k p u r p l i s h - g r e e n A. b i f r o n s 1 1 l e a v e s n o t p i l i f e r o u s , p l a n t s g r e e n o r g r e e n i s h - b r o w n 2 2 h y a l i n e l e a f b a s e u s u a l l y a s l o n g o r l o n g e r t h a n t h e b l a d e a bove; l e a v e s o f t e n a p p e a r i n g s h o r t and s t o c k y ; s t e r e i d l a y e r s a t m i d l e a f u s u a l l y one o r two; l a r g e c l e a r c e l l s between s t e r e i d s and f i l a m e n t s u s u a l l y >2 i n number; c a p s u l e g e n e r a l l y c y l i n d r i c a l w i t h c o n i c a l t o s h o r t o p e r c u l u m ; u s u a l l y s y n o i c o u s A. b r e v i r o s t r i s 2 1 h y a l i n e l e a f b a s e u s u a l l y s h o r t e r t h a n b l a d e a b o v e ; l e a v e s u s u a l l y l o n g - l i g u l a t e , r a r e l y s h o r t and s t o c k y ; s t e r e i d l a y e r s a t m i d l e a f u s u a l l y two o r t h r e e ; l a r g e , c l e a r c e l l s between s t e r e i d s and f i l a m e n t s one o r two i n number; c a p s u l e o v o i d - c y l i n d r i c a l w i t h e l o n g a t e c o n i c a l t o s u b u l a t e o p e r c u l u m ; d i o i c o u s t o i n c o n s p i c u o u s l y s y n o i c o u s A. r i g i d a 48 ALOINA BIFRONS (De Not.) D e l g a d i l l o A. p i l i f e r a (B.S.G.) Crum e t S t e e r e A. r i g i d a v a r . p i l i f e r a (B.S.G.) L i m p r . Common on sunny l a c u s t r i n e b anks i n e x t e n s i v e , u s u a l l y open c o l o n i e s , sometimes p a r t i a l l y b u r i e d , r a r e l y i n s t e p p e ; o f t e n g r o w i n g w i t h numerous t e r r i c o l o u s c r u s t o s e l i c h e n s p e c i e s . S p o r o p h y t e s uncommon and u s u a l l y s h r i v e l l e d b e f o r e m a t u r i t y . R e p r e s e n t a t i v e s p e c i m e n s : B89001, B89002 I l l u s t r a t i o n s : F l o w e r s 1973, D e l g a d i l l o 1975 D i s t r i b u t i o n : A: s c a t t e r e d , b u t common on l a c u s t r i n e d e p o s i t s ( F i g . 20) B: w e s t e r n N o r t h A m e r i c a ; s o u t h e r n B r i t i s h C o l u m b i a , Idaho s o u t h t o M e x i c o ; new t o B r i t i s h C o l u m b i a and Canada C: w e s t e r n N o r t h A m e r i c a / w e s t e r n E u r o p e - N . A f r i c a d i s j u n c t ; b i p o l a r ( S o u t h A f r i c a , New Z e a l a n d , A u s t r a l i a , C h i l e ) Remarks : A d d i t i o n a l f e a t u r e s s e p a r a t i n g A. b i f r o n s f r o m t h e o t h e r two A l o i n a s p e c i e s i n c l u d e a l a c k o f s t e r e i d s i n t h e l e a f and t h e l e s s d i f f e r e n t i a t e d b a s a l l e a f m a r g i n . In b o t h A. r i g i d a and A. b r e v i r o s t r i s t h e l e a f b a s e i s b o r d e r e d by t h i n - w a l l e d , e l o n g a t e c e l l s , a b s e n t o r w e a k l y d e v e l o p e d i n A. b i f r o n s ( D e l g a d i l l o 1975). In s o u t h e r n W a s h i n g t o n and n o r t h e r n O r e g o n , A. b i f r o n s i s an i m p o r t a n t component o f t h e s t e p p e c r y p t o g a m f l o r a . However, i t i s f o u n d o n l y i n p r o t e c t e d a r e a s i n t h e B r i t i s h C o l u m b i a s t e p p e . 49 ALOINA BREVIROSTRIS (Hook, e t G r e v . ) K i n d b . B a r b u l a r i g i d a v a r . b r e v i r o s t r i s (Hook, e t G r e v . ) B r i d . S m a l l , d a r k g r e e n p l a n t s , u s u a l l y i n open c o l o n i e s i n t h e s t e p p e , most common on l a c u s t r i n e banks, but u s u a l l y i n more sha d e d s i t u a t i o n s t h a n A. b i f r o n s . S p o r o p h y t e s common, a l t h o u g h o c c a s i o n a l l y s h r i v e l l e d b e f o r e m a t u r i t y . R e p r e s e n t a t i v e s p e c i m e n s : B89003, B89004 I l l u s t r a t i o n s : C r a i g 1939, Crum and A n d e r s o n 1981, D e l g a d i l l o 1975, S a v i c z - L j u b i t z k a j a and S m i r n o v a 1970 D i s t r i b u t i o n : A: s c a t t e r e d and uncommon ( F i g . 21) B: s c a t t e r e d a c r o s s n o r t h e r n h a l f of t h e c o n t i n e n t , a l t h o u g h m a i n l y w e s t e r n ; s o u t h t o Montana i n t h e west and t o O n t a r i o and Nova S c o t i a i n t h e e a s t ; S t e e r e (1950) p r o v i d e d a N o r t h A m e r i c a n d i s t r i b u t i o n map C: c i r c u m b o r e a l Remarks : A l o i n a b r e v i r o s t r i s i s v e r y d i f f i c u l t t o d i s t i n g u i s h f r o m A. r i g i d a . They may be t r e a t e d b e s t a s v a r i e t i e s o f one s p e c i e s . Crum and A n d e r s o n (1981) s t r e s s e d t h e shape and s i z e o f t h e upper l e a f c e l l s and t h e s i z e o f t h e i n d i v i d u a l l e a v e s as i m p o r t a n t c h a r a c t e r s i n d i s t i n g u i s h i n g t h e two t a x a . B o t h c h a r a c t e r s a r e t o o v a r i a b l e i n t h e l o c a l m a t e r i a l t o be o f a s s i s t a n c e . 50 ALOINA RIGIDA (Hedw.) L i m p r . S i m i l a r t o A. b r e v i r o s t r i s i n h a b i t and h a b i t a t . R e p r e s e n t a t i v e s p e c i m e n s : B89005, B89006 I l l u s t r a t i o n s : C r a i g 1939, Crum and A n d e r s o n 1981, D e l g a d i l l o 1975 D i s t r i b u t i o n : A: r e a s o n a b l y common, but s c a t t e r e d ( F i g . 22) B: s c a t t e r e d a c r o s s s o u t h e r n and w e s t e r n p o r t i o n of t h e c o n t i n e n t C: more o r l e s s c i r c u m t e m p e r a t e ; b i p o l a r ( S o u t h A m e r i c a -A u s t r a l i a , where i t i s p o s s i b l y i n t r o d u c e d , see C a t c h e s i d e 1980) Remarks : See t h e d i s c u s s i o n a c c o m p a n y i n g A. b r e v i r o s t r i s f o r comment. BARBULA Hedw. My t r e a t m e n t o f B a r b u l a and t h e c l o s e l y r e l a t e d genus Didymodon f o l l o w s t h e r e c e n t r e v i s i o n s of S a i t o (1975) and Zander (1978b, 1981b). T h e s e a u t h o r s s e l e c t e d a s e r i e s o f g a m e t o p h y t i c c h a r a c t e r s t h a t r e a d i l y s e p a r a t e d t h e two g e n e r a ( d i s c u s s e d i n t h e n e x t c h a p t e r ) . The t r a d i t i o n a l c o n c e p t s o f t h e two g e n e r a , b a s e d on t h e use o f s p o r o p h y t i c c h a r a c t e r s a s employed by many r e s e a r c h e r s , a r e ambiguous and d i f f i c u l t t o a p p l y . In a d d i t i o n , a t h i r d g e n u s , P s e u d o c r o s s i d i u m , i s a l s o s p l i t f r o m B a r b u l a h e r e , f o l l o w i n g Zander ( 1 9 7 9 a ) . Two s p e c i e s , B a r b u l a c o n v o l u t a and B. u n g u i c u l a t a , a r e 51 p r e s e n t i n t h e s t u d y r e g i o n . B o t h t a x a i n d i c a t e c o m p a r a t i v e l y c o o l e r and more m e s i c c l i m a t e c o n d i t i o n s and a r e more common i n open a r e a s e l s e w h e r e i n t h e p r o v i n c e . Key t o t h e S p e c i e s o f BARBULA I l e a v e s w i t h o u t e x c u r r e n t mucro (sometimes s h o r t a p i c u l u s p r e s e n t ) ; p l a n t s s m a l l , u s u a l l y < 2mm t a l l B. c o n v o l u t a I I a t l e a s t some l e a v e s w i t h e x c u r r e n t m u l t i c e l l u l a r mucro; p l a n t s u s u a l l y > 2mm t a l l B. u n g u i c u l a t a BARBULA CONVOLUTA Hedw. B. e u s t e g i a C a r d , e t T h e r . S m a l l g r e e n p l a n t s , s c a t t e r e d on s o i l i n g r a s s l a n d s a t h i g h e r e l e v a t i o n s , r a r e l y i n d r i e s t s t e p p e ; l e a v e s somewhat c o n t o r t e d t o t w i s t e d whey d r y , w e a k l y s p r e a d i n g when wet. S p o r o p h y t e s n o t c o l l e c t e d . R e p r e s e n t a t i v e s p e c i m e n s : B89007, B89008 I l l u s t r a t i o n s : Lawton 1971, F l o w e r s 1973 (as B. e u s t e g i a ) , Crum and A n d e r s o n 1981 D i s t r i b u t i o n : A: s c a t t e r e d , more common i n t h e s o u t h - e a s t e r n s e c t i o n ( F i g . 24) B: a c r o s s t h e c o n t i n e n t , a l t h o u g h more common n o r t h w a r d , s o u t h t o C a l i f o r n i a i n t h e west and t o N. C a r o l i n a i n t h e e a s t C: c i r c u m p o l a r ; b i p o l a r (New Z e a l a n d ) 52 Remarks : In t h e p r e s e n t s t u d y B a r b u l a e u s t e g i a has been i n c l u d e d w i t h i n B. c o n v o l u t a , a l t h o u g h most r e s e a r c h e r s p r e f e r t o s e p a r a t e i t ( S t e e r e 1938, 1939a; Lawton 1971; F l o w e r s 1973; Tan 1980). See t h e n e x t c h a p t e r . M a c r o s c o p i c a l l y , B. c o n v o l u t a c a n be c o n f u s e d e a s i l y w i t h Desmatodon o b t u s i f o l i u s . The a d a x i a l s u r f a c e o f t h e c o s t a of B. c o n v o l u t u s , however, i s w e a k l y d e v e l o p e d , n ot b r o a d and convex as i s t h e c a s e w i t h D. o b t u s i f o l i u s , and i s p o s i t i o n e d i n a l a m i n a l g r o o v e , a f e a t u r e a b s e n t i n D. o b t u s i f o l i u s . BARBULA UNGUICULATA Hedw. P l a n t s d a r k ( d u l l ) g r e e n t o g r e e n , o c c u r r i n g a s i s o l a t e d i n d i v i d u a l s o r open c o l o n i e s , u s u a l l y admixed w i t h o t h e r mosses on s o i l i n s h a d e d s i t u a t i o n s ; l e a v e s o f t e n t w i s t e d when d r y , s p r e a d i n g when wet. S p o r o p h y t e s o c c a s i o n a l . R e p r e s e n t a t i v e s p e c i m e n s : B89009, B89010 I l l u s t r a t i o n s : Lawton 1971, F l o w e r s 1973, Crum a n d A n d e r s o n 1981 D i s t r i b u t i o n : A: s c a t t e r e d ( F i g . 25) B: m a i n l y i n t h e s o u t h e r n h a l f o f t h e c o n t i n e n t C: c i r c u m t e m p e r a t e ; b i p o l a r (New Z e a l a n d ) Remarks B a r b u l a u n g u i c u l a t a i s u s u a l l y a d i s t i n c t i v e s p e c i e s , w i t h t h e m u l t i - c e l l u l a r mucro e a s i l y s e p a r a t i n g most c o l l e c t i o n s from B. c o n v o l u t a , a l t h o u g h some c o l l e c t i o n s of t h i s l a t t e r t a x o n a r e 53 w e a k l y m u c r o n a t e . B r y o e r y t h r o p h y l l u m r e c u r v i r o s t r u m o f t e n r e s e m b l e s B. u n g u i c u l a t a . The c a n a l i c u l a t e a d a x i a l c o s t a of B a r b u l a , however, i s d i s t i n c t f r o m t h e b r o a d , r a i s e d c o s t a of B r y o e r y t h r o p h y l l u m . Reduced p l a n t s o f B. u n g u i c u l a t a a l s o c a n be c o n f u s e d w i t h Desmatodon o b t u s i f o l i u s ( s e e t h e d i s c u s s i o n t h e r e ) . BRYOERYTHROPHYLLUM Chen B r y o e r y t h r o p h y l l u m i s b e s t c h a r a c t e r i z e d by t h e p a p i l l o s e a n d r a i s e d a d a x i a l c o s t a , e l o n g a t e , t h i n - w a l l e d a n d somewhat i n f l a t e d b a s a l c e l l s of t h e l e a f , and, i n many c a s e s , a d a r k r e d c o l o u r a t i o n (Chen 1941; S a i t o 1975; Zander 1978a, 1981a). Two s p e c i e s a r e f o u n d i n t h e s t u d y r e g i o n : B. r e c u r v i r o s t r u m o f m e s i c h a b i t a t s and B. columbianum o f x e r i c h a b i t a t s . Key t o t h e S p e c i e s of BRYOERYTHROPHYLLUM I l e a v e s l i n e a r - l a n c e o l a t e t o b r o a d l y l a n c e o l a t e , apex a c u t e t o b l u n t , a p i c u l a t e and sometimes t o o t h e d , u s u a l l y c r i s p e d when d r y ; u s u a l l y i n shaded h a b i t a t s B. r e c u r v i r o s t r u m I I l e a v e s o v a t e - l a n c e o l a t e w i t h a c u t e apex, n e v e r t o o t h e d , e r e c t t o s l i g h t l y t w i s t e d when d r y ; u s u a l l y i n sunny h a b i t a t s B. columbianum BRYOERYTHROPHYLLUM COLUMBIANUM (Herm. e t Lawt.) Zander Didymodon c o l u m b i a n u s Herm. e t Lawt. In compact r e d d i s h - b r o w n ( r a r e l y d a r k g r e e n ) t u f t s o r sods on s o i l , u s u a l l y a d j a c e n t t o o u t c r o p s , o c c a s i o n a l l y on l a c u s t r i n e s i l t s ; l e a v e s o v a t e - l a n c e o l a t e and somewhat c o n c a v e , 54 u s u a l l y < 1mm l o n g , s t i f f and e r e c t when d r y , o c c a s i o n a l l y s l i g h t l y t w i s t e d above, e r e c t - s p r e a d i n g when wet, a p i c e s of most l e a v e s a c u t e ; m a r g i n s w e a k l y r e c u r v e d i n m i d d l e p a r t o f t h e l e a f ; c o s t a p e r c u r r e n t t o s h o r t e x c u r r e n t as a c o n i c a l c e l l , h i g h l y o r n a m e n t e d and r a i s e d on a d a x i a l s u r f a c e , composed of numerous ( o f t e n > 8) p a p i l l o s e , u s u a l l y v e r t i c a l l y e l o n g a t e d c e l l s ; upper l e a f c e l l s brown, m o s t l y 7-12 m i c r o m e t e r s , q u a d r a t e , r e c t a n g u l a r t o i r r e g u l a r , u s u a l l y p a p i l l o s e e s p e c i a l l y n e a r c o s t a ( m a r g i n a l c e l l s o f t e n n e a r l y s m o o th); b a s a l c e l l s t h i n n e r w a l l e d , h y a l i n e t o p a l e g r e e n , q u a d r a t e t o s h o r t r e c t a n g u l a r , o f t e n b r o a d e r t h a n l o n g , c o r n e r s s l i g h t l y t h i c k e n e d . S p o r o p h y t e s n o t c o l l e c t e d i n s t u d y a r e a , r a r e t h r o u g h o u t r a n g e . R e p r e s e n t a t i v e s p e c i m e n s : B89011, B89012 I l l u s t r a t i o n s : F i g . 8, Lawton 1971, Hermann and Lawton 1968 D i s t r i b u t i o n : A: uncommon, m a i n l y i n s o u t h p a r t o f t h e Okanagan V a l l e y ( F i g . 28) B: N o r t h w e s t T e r r i t o r i e s , O r e g o n , W a s h i n g t o n ; new t o B r i t i s h C o l u m b i a C: w e s t e r n N o r t h A m e r i c a ; b i p o l a r ( S o u t h A m e r i c a ; B o l i v i a ) Remarks : U n t i l r e c e n t l y , B. columbianum was c o l l e c t e d o n l y a l o n g t h e C o l u m b i a R i v e r i n W a s h i n g t o n and Oregon S t a t e s where i t i s common, and i n t h e N o r t h w e s t T e r r i t o r i e s , where i t i s a p p a r e n t l y r a r e ( S t e e r e and S c o t t e r 1978). L e w i s (1981) r e p o r t e d B. c olumbianum ( v a r . a t a c a m e n s i s Zander e t L e w i s ) f r o m B o l i v i a , 55 where i t i s f o u n d a t h i g h e l e v a t i o n s (3730m) b u t i n an a r e a of s i m i l a r v e g e t a t i o n and c l i m a t e as t h e s t u d y a r e a . B r y o e r y t h r o p h y l l u m columbianum i s r e a d i l y c o n f u s e d w i t h Desmatodon c o n v o l u t u s . The l a t t e r s p e c i e s d i f f e r s i n h a v i n g , t y p i c a l l y , s p i r a l l y t w i s t e d l e a v e s (when d r y ) , a n a r r o w e r and o f t e n h i g h e r band o f o r n a m e n t e d c o s t a l c e l l s , and a b l u n t and c u c u l l a t e l e a f a pex. O c c a s i o n a l l y , i t has a s u b - a c u t e apex, but n e v e r s h a r p l y a c u t e as i s t h e c a s e w i t h B. columbianum (compare F i g . 8 and F i g . 9 ) . Some s m a l l , r e d d i s h - b r o w n p o p u l a t i o n s o f . Didymodon s p p . c a n be c o n f u s e d w i t h B. columbianum, b u t t h e y l a c k t h e wide, h i g h l y d e v e l o p e d and o r n a m e n t e d c o s t a . F u r t h e r m o r e , Didymodon spp. r a r e l y f o r m compact s o d s t y p i c a l o f most p o p u l a t i o n s o f B. columbianum. F o r f u r t h e r d i s c u s s i o n r e f e r t o Zander (1978a) and Hermann and Lawton ( 1 9 6 8 ) . BRYOERYTHROPHYLLUM RECURVIROSTRUM (Hedw.) Chen Didymodon r e c u r v i r o s t r i s (Hedw.) J e n n . D. r u b e l l u s B.S.G. In s m a l l g r e e n t o red-brown t u f t s o r c o l o n i e s on s o i l o f s h a d e d s i t e s , u s u a l l y g r o w i n g w i t h o t h e r mosses. S p o r o p h y t e s common, w i t h r e d d i s h c a p s u l e s and s h o r t , r u d i m e n t a r y p e r i s t o m e s . R e p r e s e n t a t i v e s p e c i m e n s : B89013, B89014 I l l u s t r a t i o n s : F l o w e r s 1973, I r e l a n d 1982, Crum and A n d e r s o n 1981 D i s t r i b u t i o n : A: uncommon, m a i n l y i n Okanagan V a l l e y ( F i g . 29) 56 B: a c r o s s t h e c o n t i n e n t , s o u t h w a r d s from C a l i f o r n i a t o N o r t h C a r o l i n a C: c i r c u m p o l a r ; b i p o l a r (New Z e a l a n d ) Remarks : B e c a u s e t h e l e a v e s o f B. r e c u r v i r o s t r u m a r e u s u a l l y s t r o n g l y c r i s p e d when d r y , t h i s s p e c i e s c a n be c o n f u s e d w i t h B a r b u l a u n g u i c u l a t a , some s p e c i e s o f Didymodon, T r i c h o s t o m o p s i s  a u s t r a l a s i a e , o r W e i s s i a b r a c h y c a r p a . However, t h e b r o a d and o r n a m e n t e d c o s t a , t h e l o n g r e c t a n g u l a r , t h i n - w a l l e d and somewhat i n f l a t e d b a s a l c e l l s , and t h e b l u n t , i r r e g u l a r t o t o o t h e d l e a f apex s h o u l d r e a d i l y d i s t i n g u i s h B. r e c u r v i r o s t r u m . I p r e f e r n o t t o s t r e s s t h e t r a d i t i o n a l use of t h e red-brown c o l o u r of B. r e c u r v i r o s t r u m as a d i s t i n g u i s h i n g t a x o n o m i c c h a r a c t e r b e c a u s e o t h e r m o r p h o l o g i c a l l y s i m i l a r t a x a a r e a l s o r e d - b r o w n , and p l a n t s o f B. r e c u r v i r o s t r u m c a n be g r e e n t h r o u g h o u t . F o r f u r t h e r d i s c u s s i o n see Z a n d e r ( 1 9 7 8 a ) , F l o w e r s (1973) and Crum and A n d e r s o n ( 1 9 8 1 ) . As Z a n d e r p o i n t s o u t , t h e i l l u s t r a t i o n s o f t h i s s p e c i e s by F l o w e r s (1973) a r e e x c e l l e n t . CROSSIDIUM J u r . C r o s s i d i u m i s e s s e n t i a l l y a t r o p i c a l and t e m p e r a t e d r y l a n d genus w i t h m a j o r c e n t e r s of d i s t r i b u t i o n i n s o u t h - w e s t e r n N o r t h A m e r i c a and a r o u n d t h e M e d i t e r r a n e a n Sea, p a r t i c u l a r l y i n s o u t h e r n E u r o p e . A few s p e c i e s e x h i b i t s c a t t e r e d d i s t r i b u t i o n s i n t h e s o u t h e r n h e m i s p h e r e ( P e r u , S o u t h A f r i c a , A u s t r a l i a and New Z e a l a n d ) a n d a c r o s s s o u t h e r n A s i a i n t h e n o r t h e r n h e m i s p h e r e ( s e e D e l g a d i l l o 1975 f o r d i s t r i b u t i o n maps of most o f t h e known 57 t a x a ) . The genus i s r e p o r t e d h e r e as new t o Canada. T h r e e s p e c i e s of C r o s s i d i u m have been f o u n d i n t h e s t u d y a r e a . Of t h e s e , C. a b e r r a n s was e x p e c t e d , h a v i n g been f o u n d i n t h e a d j a c e n t s t e p p e of W a s h i n g t o n S t a t e (Lawton 1971). On t h e o t h e r hand, C. r o s e i was u n e x p e c t e d , s i n c e i t had been r e p o r t e d p r e v i o u s l y o n l y from P e r u ( W i l l i a m s 1915). I s u s p e c t , however, t h a t i t may be c o n s p e c i f i c w i t h C. d a v i d a i C a t c h e s i d e o f A u s t r a l i a ( C a t c h e s i d e 1980) and C. a s i r e n s e F r e y e t K i r s c h n e r f r o m S a u d i A r a b i a ( F r e y and K i r s c h n e r 1984). The t h i r d s p e c i e s r e m a i n s unnamed f o r now. G a m e t o p h y t i c a l l y , i t s t r o n g l y r e s e m b l e s C. s e r i a t u m , a r a r e t a x o n from B a j a C a l i f o r n i a (Crum and S t e e r e 1958) and A r i z o n a ( Z a n d e r 1977). The s p o r o p h y t e s o f t h e l o c a l p o p u l a t i o n s , however, a r e q u i t e d i f f e r e n t f r o m t h o s e d e s c r i b e d f o r C. s e r i a t u m by Crum and S t e e r e ( 1 9 5 8 ) . C h a r a c t e r i s t i c of t h e genus C r o s s i d i u m i s t h e o r n a m e n t a t i o n o f t h e u p p e r a d a x i a l s u r f a c e o f t h e c o s t a . In a l l t h r e e o f o u r s p e c i e s t h e f i l a m e n t s , i n c o n t r a s t t o A l o i n a , a r e o f t e n d i f f i c u l t t o d i s c e r n . O c c a s i o n a l l y t h e y a r e p r e s e n t o n l y as a s e r i e s o f b u l g i n g c e l l s ( as a r e f o u n d on many l e a v e s o f C. a b e r r a n s ) o r , as i s t h e c a s e w i t h t h e o t h e r two t a x a , t h e y a r e f u s e d a t t h e base f o r m i n g i r r e g u l a r l y s h a p e d c u s h i o n s o v e r t h e c o s t a l s u r f a c e . I n C r o s s i d i u m s p . t h e y o f t e n f o r m l a m e l l a e - l i k e s t r u c t u r e s . D e l g a d i l l o (1975) r e v i s e d t h e genus on a w o r l d wide b a s i s , a l s o p r o v i d i n g d i s t r i b u t i o n maps. 58 Key t o t h e s p e c i e s o f CROSSIDIUM 1 upper l e a f c e l l s smooth o r w i t h a s i n g l e , n o n - o rnamented c e n t r a l p a p i l l a ; c o s t a u s u a l l y l i g h t l y o r n a m e n t e d , f i l a m e n t s 1, r a r e l y , 2 - 3 c e l l s h i g h , n e v e r f o r m i n g c u s h i o n s o r l a m e l l a e ( s e e F i g . 9) C. a b e r r a n s 1 1 u p p e r l e a f c e l l s , e s p e c i a l l y t o w a r d s t h e c o s t a , s t r o n g l y p l u r i - p a p i l l o s e ; c o s t a u s u a l l y o r n a m e n t e d w i t h a d i s t i n c t g r o u p of f i l a m e n t s t h a t a r e u s u a l l y f u s e d , f o r m i n g i r r e g u l a r l y s h a p e d c u s h i o n s o r l a m e l l a e 2 2 l e a v e s o v a t e t o o v a t e - l a n c e o l a t e , e r e c t t o s l i g h t l y t w i s t e d when d r y , u s u a l l y w i t h a l o n g , smooth and somewhat s i n u o u s awn, apex o c c a s i o n a l l y b r o a d l y c u c u l l a t e , n e v e r n a r r o w l y c u c u l l a t e C r o s s i d i u m s p . 2 1 l e a v e s o v a t e - o b l o n g t o l i g u l a t e , t w i s t e d when d r y , sometimes a p i c u l a t e o r v e r y s h o r t awned, apex n a r r o w l y c u c u l l a t e C. r o s e i CROSSIDIUM ABERRANS H o l z . e t B a r t r . P l a n t s d a r k g r e e n t o g r e e n , o c c a s i o n a l l y r e d d i s h - b r o w n ; u s u a l l y i n s m a l l c o l o n i e s on s o i l , sometimes s c a t t e r e d among o t h e r mosses, most common i n s h a d e d s t e p p e s i t e s ; l e a v e s u s u a l l y g r e a t e r t h a n 1 mm l o n g , o v a t e - l a n c e o l a t e t o l i g u l a t e ( o c c a s i o n a l l y o v a t e ) , c o n c a v e and somewhat pr o w - s h a p e d , w e a k l y t w i s t e d a r o u n d t h e stem when d r y , s p r e a d i n g when wet; l e a f apex o f t e n e m a r g i n a t e ; m a r g i n s r e c u r v e d t o n e a r apex, sometimes i r r e g u l a r l y t o o t h e d above r e c u r v e d p o r t i o n ; c o s t a u s u a l l y o r ange-brown an d e x c u r r e n t a s an awn (many p o p u l a t i o n s l a c k i n g an awn; f . e p i l o s u m , see F l o w e r s 1973). S p o r o p h y t e s common; c a p s u l e s p a l e t o d a r k ( r e d - ) brown, u s u a l l y s t r i a t e a n d w i d e r a t t h e mouth t h a n below; p e r i s t o m e w h i t e and w i t h a b a s a l t u b e . R e p r e s e n t a t i v e s p e c i m e n s : B89015, B89016 59 I l l u s t r a t i o n s : F i g . 10, F l o w e r s 1973, G r o u t 1928-1941 D i s t r i b u t i o n : A: w i d e s p r e a d but s c a t t e r e d and e l u s i v e l o c a l l y ( F i g . 37) B: w e s t e r n N o r t h A m e r i c a : B r i t i s h C o l u m b i a s o u t h t o M e x i c o C: w e s t e r n N o r t h A m e r i c a / w e s t e r n E u r a s i a , N. A f r i c a Remarks : Even t h o u g h C r o s s i d i u m a b e r r a n s i s h i g h l y v a r i a b l e m o r p h o l o g i c a l l y , i n most c a s e s i t i s d i s t i n c t i v e and e a s y t o r e c o g n i z e . I n t h e f i e l d t h e s m a l l c l u s t e r s of d a r k g r e e n g a m e t o p h y t e s crowned, u s u a l l y , w i t h numerous d a r k s p o r o p h y t e s w i t h c o n t r a s t i n g l o n g , i v o r y - w h i t e p e r i s t o m e s , a r e c h a r a c t e r i s t i c . M i c r o s c o p i c a l l y , t h e f i l a m e n t s on t h e c o s t a a r e d i s t i n c t i v e , a l t h o u g h on most l e a v e s t h e y a r e r e d u c e d t o b u l g i n g c e l l s . CROSSIDIUM ROSEI W i l l i a m s S m a l l , b r o w n i s h - g r e e n p l a n t s u s u a l l y g r o w i n g among o t h e r mosses on c l a y - r i c h s o i l ; l e a v e s u s u a l l y g r e a t e r t h a n 1mm l o n g , sometimes s h o r t e r , b r o a d - l a n c e o l a t e t o l i g u l a t e , somewhat t w i s t e d a r o u n d t h e stem when d r y , s p r e a d i n g when wet; upper c e l l s i r r e g u l a r l y q u a d r a t e t o r e c t a n g u l a r , 13-19 m i c r o m e t e r s w i d e , u s u a l l y p l u r i p a p i l l o s e ; c o s t a e x c u r r e n t as a mucro o r a v e r y s h o r t awn, sometimes p e r c u r r e n t ; c o s t a l f i l a m e n t s 1 t o 3 c e l l s h i g h , o f t e n f u s e d below i n t o i r r e g u l a r c u s h i o n s , p a p i l l o s e . S p o r o p h y t e s p r e s e n t i n o n l y c o l l e c t i o n ; p e r i s t o m e a b s e n t ; s p o r e s l a r g e , g r e a t e r t h a n 25 m i c r o m e t e r s . R e p r e s e n t a t i v e s p e c i m e n s : B89017, B89018 60 I l l u s t r a t i o n s : F i g . 11, D e l g a d i l l o 1975, W i l l i a m s 1915 D i s t r i b u t i o n : A: r a r e , 2 s i t e s : s i t e 8, Gang Ranch a r e a , n o r t h o f L i l l o o e t t ; s i t e 70, c l a y banks west o f P e n t i c t o n ( F i g . 38) B: s o u t h e r n B r i t i s h C o l u m b i a ; new t o N o r t h A m e r i c a C: w e s t e r n N o r t h A m e r i c a ; b i p o l a r ( S o u t h A m e r i c a : Peru,. C h i l e ; p o s s i b l y A u s t r a l i a and S a u d i A r a b i a ) Remarks: T h i s i n t e r e s t i n g moss was an a c c i d e n t a l d i s c o v e r y , f i r s t f o u n d g r o w i n g w i t h a number o f o t h e r mosses c o l l e c t e d on a l a c u s t r i n e bank ne a r P e n t i c t o n . The l a r g e s p o r e s ( l a r g e s t f o r t h e genus a c c o r d i n g t o D e l g a d i l l o , 1975) and t h e c u c u l l a t e and m u c r o n a t e l e a f a p i c e s a r e d i s t i n c t i v e . I t somewhat r e s e m b l e s t h e f o l l o w i n g t a x o n , but t h e key c h a r a c t e r s , i n p a r t i c u l a r t h e l i g u l a t e , t w i s t e d and a w n l e s s l e a v e s , s h o u l d e a s i l y d i s t i n g u i s h i t . CROSSIDIUM s p . M i n u t e , d a r k g r e e n t o g o l d e n - b r o w n p l a n t s g r o w i n g as s m a l l open c o l o n i e s o r s c a t t e r e d among o t h e r mosses on c l a y - r i c h s o i l ; l e a v e s u s u a l l y < 1mm l o n g , b r o a d l y o v a t e t o o v a t e - l a n c e o l a t e , c o n c a v e , i m b r i c a t e and s l i g h t l y t w i s t e d when d r y , e r e c t s p r e a d i n g when wet; m a r g i n s r e c u r v e d t o n e a r apex; u p p e r c e l l s h e x a g o n a l - i r r e g u l a r t o q u a d r a t e , 14-17 m i c r o m e t e r s w i d e , s t r o n g l y p l u r i p a p i l l o s e ; c o s t a o r n a m e n t e d a d a x i a l l y w i t h a c u s h i o n of f u s e d f i l a m e n t s , o f t e n f o r m i n g l a m e l l a e , w h i c h a r e o c c a s i o n a l l y f i l a m e n t o u s a l o n g t h e i r c r e s t s , 1-4 c e l l s h i g h , 61 p a p i l l o s e ; c o s t a e x c u r r e n t a s a smooth, h y a l i n e awn w h i c h i s o f t e n hooked o r s i n u o u s . S p o r o p h y t e s c o l l e c t e d o n c e , immature; c a p s u l e s s h o r t - c y l i n d r i c ; p e r i s t o m e a p p a r e n t l y a b s e n t ; s p o r e s f i n e l y p a p i l l o s e , 25-35 m i c r o m e t e r s i n d i a m e t e r . R e p r e s e n t a t i v e s p e c i m e n s : B89019, B89020 I l l u s t r a t i o n s : : F i g . 12, Crum and S t e e r e 1958 ( f o r C. s e r i a t u m , b u t t h e g a m e t o p h y t i c c h a r a c t e r i s t i c s a r e i d e n t i c a l ) D i s t r i b u t i o n : A: s c a t t e r e d , but f a i r l y common i n some s i t e s ( F i g . 39) B: s o u t h e r n B r i t i s h C o l u m b i a C: w e s t e r n N o r t h A m e r i c a n endemic Remarks : T h i s t a x o n i s h a r d t o f i n d i n t h e f i e l d b e c a u s e o f i t s t i n y s i z e . Many of t h e c o l l e c t i o n s were made a c c i d e n t a l l y and were d i s c o v e r e d when i d e n t i f y i n g o t h e r t a x a i n t h e l a b o r a t o r y . I t i s e s p e c i a l l y d i f f i c u l t t o f i n d when m i x e d w i t h p o p u l a t i o n s o f t h e u b i q u i t o u s P t e r y g o n e u r u m ovatum, t h a t i t s t r o n g l y r e s e m b l e s m a c r o s c o p i c a l l y . The l a t t e r t a x o n , however, i s n e v e r s t r o n g l y p a p i l l o s e and a l w a y s has much more c l e a r l y d e v e l o p e d and m o s t l y n o n - p a p i l l o s e l a m e l l a e ( r e f e r t o F i g . 13 f o r c o m p a r i s o n ) . DESMATODON B r i d . T h r e e s p e c i e s of Desmatodon, D. c o n v o l u t u s , D. g u e p i n i i and D. o b t u s i f o l i u s , were f o u n d d u r i n g t h e s t u d y . D. c o n v o l u t u s and D. g u e p i n i i a r e r e p o r t e d a s new t o Canada. Of t h e t h r e e s p e c i e s , o n l y D. c o n v o l u t u s i s common, a l t h o u g h r e s t r i c t e d p r i m a r i l y t o t h e h a r d - p a c k e d s u r f a c e s o f t h e l a c u s t r i n e t e r r a c e s . 62 Two o t h e r t a x a , D. c e r n u u s and D. r a n d i i , were c o l l e c t e d i n a l k a l i n e and s e e p a g e a r e a s a d j a c e n t t o t h e s t u d y r e g i o n . Desmatodon c e r n u u s i s p a r t i c u l a r l y common i n t h e r e g i o n n o r t h o f C l i n t o n . D. r a n d i i i s r a r e , c o l l e c t e d o n l y a l o n g a s t r e a m e a s t o f Savona ( t h i s l o c a t i o n i s t h e f i r s t r e p o r t of D. r a n d i i from B r i t i s h C o l u m b i a ) . Crum and A n d e r s o n (1981) and C o r l e y e t a_l. (1981) p o i n t o u t t h a t Desmatodon, a s i t s t a n d s , i s n o t e n t i r e l y d i s t i n c t from t h e c l o s e l y r e l a t e d g e n e r a T o r t u l a and P o t t i a . W i t h T o r t u l a ( i n p a r t i c u l a r t h e T. b r e v i p e s g r o u p ) , i t s h a r e s a number o f g a m e t o p h y t i c c h a r a c t e r s , i n c l u d i n g , i n most c a s e s , l e a f s h a p e , c e l l s i z e and p a p i l l a t i o n , p a r t l y m o d i f i e d l e a f b o r d e r s , smooth awns, convex u p p e r a d a x i a l c o s t a e a n d s m a l l s p o r e s ( i n a few c a s e s ) . The g e n e r a d i f f e r m a i n l y i n s p o r o p h y t i c c h a r a c t e r s . C h a r a c t e r i s t i c a l l y , T o r t u l a has l a r g e r s p o r o p h y t e s ( t a l l e r s e t a e and l o n g e r c a p s u l e s ) and a b e t t e r d e v e l o p e d p e r i s t o m e ( l o n g e r , w e l l d e v e l o p e d t e e t h and a t a l l b a s a l membrane) t h a n Desmatodon w h i c h has s m a l l e r s p o r o p h y t e s and s h o r t e r , l e s s w e l l d e v e l o p e d p e r i s t o m e s ( s h o r t , i r r e g u l a r l y d i v i d e d t e e t h on a v e r y s h o r t b a s a l membrane). In a d d i t i o n , Desmatodon i s c l o s e l y a l l i e d t o P o t t i a , s h a r i n g f e a t u r e s s u c h a s , i n some c a s e s , l a r g e s p o r e s , s h o r t awns, l e a f b o r d e r m o d i f i c a t i o n and l a r g e r c e l l s . P o t t i a d i f f e r s i n h a v i n g l e s s p a p i l l o s e c e l l s , l a x e r l e a v e s , w e a k l y m o d i f i e d u p p e r c o s t a e and o c c a s i o n a l t o o t h i n g o f t h e u p p e r l e a f m a r g i n s . S p o r o p h y t i c d i f f e r e n c e s , w i t h r e s p e c t t o P o t t i a , i n c l u d e g e n e r a l l y s m a l l e r s p o r o p h y t e s , s h o r t t o o v a t e c a p s u l e s and an 63 a b s e n t o r s h o r t , b u l k y p e r i s t o m e w i t h o r w i t h o u t a l o w , b a s a l membrane. Key t o t h e S p e c i e s of DESMATODON 1 l e a v e s somewhat red-brown, u s u a l l y s t r o n g l y t w i s t e d a r o u n d stem when d r y , c u c u l l a t e above; upper p o r t i o n o f c o s t a b r o a d l y c o n v e x , w i d e r t h a n l o w e r p a r t D. c o n v o l u t u s 1 1 l e a v e s u s u a l l y l i g h t g r e e n , s l i g h t l y t w i s t e d o r a l m o s t e r e c t , n e i t h e r s t r o n g l y t w i s t e d nor c u c u l l a t e a b o v e ; u p p e r p o r t i o n o f t h e c o s t a n ot b r o a d l y c o n v e x , t h e same w i d t h o r n a r r o w e r t h a n l o w e r p a r t 2 2 l e a v e s s l i g h t l y t w i s t e d t o e r e c t when d r y ; c e l l s s m a l l 1 , u s u a l l y 10-15 m i c r o m e t e r s wide; l e a f m a r g i n s u s u a l l y s l i g h t l y m o d i f i e d w i t h smoother, somewhat l o n g e r and more t h i c k - w a l l e d c e l l s t h a n a d j a c e n t l a m i n a l c e l l s 3 2 1 l e a v e s c o n t o r t e d when d r y ; c e l l s l a r g e , u s u a l l y >20 m i c r o m e t e r s wide; m a r g i n s d i s t i n c t l y b o r d e r e d , a t l e a s t a t m i d - l e a f , by l o n g , narrow c e l l s , b i s t r a t o s e below; g r o w i n g a l o n g m a r g i n s o f a l k a l i n e d e p r e s s i o n s o r n e a r s e e p s ; not y e t known from t h e s t u d y r e g i o n 4 3 l e a v e s o v a t e - l a n c e o l a t e , u s u a l l y w i t h s h o r t awns, not b o r d e r e d o r o n l y i n d i s t i n c t l y s o ; upper l e a f c e l l s u s u a l l y 12-15 m i c r o m e t e r s wide D. g u e p i n i i 3 1 l e a v e s l i g u l a t e , most l e a v e s w i t h p e r c u r r e n t o r s h o r t e x c u r r e n t c o s t a , r a r e l y a s a s h o r t awn; u s u a l l y d i s t i n c t l y b o r d e r e d w i t h one o r more rows o f somewhat e l o n g a t e , t h i c k -w a l l e d and smooth c e l l s ; u p p e r l e a f c e l l s 8-12 m i c r o m e t e r s wide D. o b t u s i f o l i u s 4 p e r i s t o m e t e e t h p r e s e n t (D. c e r n u u s ) 4 1 p e r i s t o m e t e e t h a b s e n t o r r u d i m e n t a r y and a d h e r i n g t o t h e o p e r c u l u m when i t f a l l s (D. r a n d i i ) 64 DESMATODON CONVOLUTUS ( B r i d . ) G r o u t T o r t u l a a t r o v i r e n s (Sm.) L i n d b . P l a n t s t i n y , u s u a l l y much s h o r t e r t h a n 0.5cm, g r e e n t o d a r k g r e e n , o f t e n w i t h r e d d i s h - b r o w n t i n g e , f o u n d a s i n d i v i d u a l s o r s m a l l compact c o l o n i e s on h a r d - p a c k e d , c l a y - r i c h s o i l , common on l a c u s t r i n e b a n k s . S p o r o p h y t e s n o t c o l l e c t e d . R e p r e s e n t a t i v e s p e c i m e n s : B89021, B89022 I l l u s t r a t i o n s : F i g . 9, F l o w e r s 1973, S a v i c z - L j u b i t z k a j a and S m i r n o v a 1970 ( a s T o r t u l a c o n v o l u t a ) . D i s t r i b u t i o n : A: w i d e s p r e a d and o f t e n l o c a l l y common ( F i g . 40) B: s o u t h e r n B r i t i s h C o l u m b i a t o M e x i c o ; new t o B r i t i s h C o l u m b i a and Canada C: c i r c u m t e m p e r a t e i n N o r t h e r n H e m i s p h e r e ; b i p o l a r ( A u s t r a l i a , New Z e a l a n d , C h i l e , S o u t h A f r i c a ) Remarks : T h i s t i n y s p e c i e s r e s e m b l e s a number o f o t h e r t a x a , i n c l u d i n g B r y o e r y t h r o p h y l l u m columbianum, P s e u d o c r o s s i d i u m  r e v o l u t u m , Didymodon v i n e a l i s v a r . l u r i d u s and v a r . b r a c h y p h y l l u s . I t i s d i s t i n g u i s h e d from t h e Didymodon t a x a and from P s e u d o c r o s s i d i u m r e v o l u t u m ( F i g . 17) by i t s c u c u l l a t e l e a f apex and i t s w i d e , o r n a m e n t e d c o s t a . I t i s d i s t i n c t f r o m B. columbianum ( F i g . 8) by i t s t w i s t e d g r o w t h h a b i t when d r y and a l e s s a c u t e , c u c u l l a t e l e a f a p e x . However, some p o p u l a t i o n s o f D. c o n v o l u t u s have a s u b - a c u t e l e a f apex and t h i s c a n r e s u l t i n some c o n f u s i o n . 65 DESMATODON GUEPINII B.S.G. T o r t u l a g u e p i n i i B r o t h . P l a n t s t i n y , y e l l o w - g r e e n t o g r e e n , s c a t t e r e d among o t h e r mosses on s o i l ; l e a v e s s m a l l , u s u a l l y a b o u t 1mm l o n g , somewhat t w i s t e d when d r y , e r e c t - s p r e a d i n g when wet, o v a t e - l a n c e o l a t e ; m a r g i n s w e a k l y r e c u r v e d ; c o s t a w e l l d e v e l o p e d , w i t h u p p e r a d a x i a l s u r f a c e c o n v e x and 3-4 c e l l s w i d e , e x c u r r e n t t o a s h o r t t o medium s i z e d , o r a n g e o r h y a l i n e awn, w h i c h i s d i s t i n c t l y w i d e r a t t h e b a s e t h a n above; upper l a m i n a l c e l l s 12-15(18) m i c r o m e t e r s w i d e , s u b - q u a d r a t e t o t r a n s v e r s e l y r e c t a n g u l a r , p a p i l l o s e ; b a s a l c e l l s s h o r t - r e c t a n g u l a r , smooth, o f t e n w i t h c o r n e r t h i c k e n i n g s , e s p e c i a l l y t o w a r d s t h e median c e l l s . S p o r o p h y t e s n o t c o l l e c t e d . R e p r e s e n t a t i v e s p e c i m e n s : B89023, B89024 I l l u s t r a t i o n s : G r o u t 1928-1941, S e r g i o 1978 D i s t r i b u t i o n : A: r a r e , two s i t e s : s i t e 24, 13 km n o r t h o f S p e n c e s B r i d g e ; s i t e 35, 1 km e a s t of Cache C r e e k ( F i g . 41) B: s c a t t e r e d i n t h e w e s t e r n p a r t of t h e c o n t i n e n t ; B r i t i s h C o l u m b i a , A l b e r t a , C a l i f o r n i a , New M e x i c o , M e x i c o ; new t o B r i t i s h C o l u m b i a and Canada C: w e s t e r n N o r t h A m e r i c a / w e s t e r n E u r a s i a Remarks : T h i s t a x o n i s p r o b a b l y w i d e s p r e a d i n w e s t e r n N o r t h A m e r i c a and i n t h e s t u d y r e g i o n , y e t i s so i n c o n s p i c u o u s t h a t i t has been o v e r l o o k e d . I t s t r o n g l y r e s e m b l e s T o r t u l a b r e v i p e s i n t h e f i e l d b u t i s d i s t i n c t i n c o l o u r , T. b r e v i p e s i s u s u a l l y t i n g e d 66 r e d d i s h - b r o w n , and has l e s s r e c u r v e d l e a v e s and a l e s s t w i s t e d h a b i t when d r y . M i c r o s c o p i c a l l y , D. g u e p i n i i has a s h o r t e r , o f t e n o r a n g e awn and, u s u a l l y , s m a l l e r b a s a l l e a f c e l l s w h i c h a r e o f t e n t h i c k e n e d i n t h e c o r n e r s , a t l e a s t i n t h e u p p e r p a r t of t h e l e a f b a s e . G r o u t (1928-1 941) s u g g e s t e d t h a t D. g u e p i n i i i s p o s s i b l y a r e d u c e d form of D. l a t i f o l i u s (Hedw.) B r i d . o r D. l e u c o s t o m a (R.Br.) B e r g g r . I t a l s o a p p e a r s c l o s e l y r e l a t e d t o D. p i i n t h o b i u s S u l l . e t L e s q . A l t h o u g h t h i s i s t h e f i r s t r e p o r t of D. g u e p i n i i f o r Canada, t h e r e i s an e a r l i e r u n r e p o r t e d c o l l e c t i o n f r o m M i l k R i v e r , A l b e r t a w h i c h N. J o n e s made i n 1928 ( a t NYBG). DESMATODON OBTUSIFOLIUS (Schwaegr.) Schimp. B a r b u l a o b t u s i f o l i a Schwaegr. S m a l l , l i g h t g r e e n p l a n t s i n t i n y , u s u a l l y s c a t t e r e d c o l o n i e s o r a s i n d i v i d u a l s on open, c l a y r i c h s o i l . S p o r o p h y t e s no t c o l l e c t e d . R e p r e s e n t a t i v e s p e c i m e n s : B89025, B89026 I l l u s t r a t i o n s : Crum and A n d e r s o n 1981, F l o w e r s 1973, Lawton 1971 D i s t r i b u t i o n : A: r a r e , 3 s i t e s : s i t e 7, a l o n g F r a s e r R i v e r e a s t o f Dog C r e e k ; s i t e 52, s o u t h s i d e o f Thompson R i v e r a t t h e D a l l e s ; s i t e 69, s o u t h - w e s t s i d e o f Okanagan Lake ( F i g . 42) B: w i d e s p r e a d : B r i t i s h C o l u m b i a t o C a l i f o r n i a e a s t t o New B r u n s w i c k and s o u t h t o T e n n e s s e e ; d i s j u n c t i n t h e a r c t i c 67 C: c i r c u m t e m p e r a t e Remarks : . G r o u t (1928-1941) n o t e d t h a t , i n s t e r i l e c o n d i t i o n , Desmatodon o b t u s i f o l i u s i s sometimes c o n f u s e d w i t h B a r b u l a  u n q u i c u l a t a , t h e l a t t e r d i f f e r i n g m a i n l y i n h a v i n g l e a v e s w i t h m u c r o n a t e c o s t a e . However, C o n a r d (1951) and F l o w e r s (1973) a c c u r a t e l y p o i n t e d o ut t h a t some p o p u l a t i o n s of D. o b t u s i f o l i u s have l e a v e s w i t h s h o r t mucros s i m i l a r t o t h o s e o f B. u n g u i c u l a t a . C o n a r d (1951) s t r e s s e d h a b i t a t d i f f e r e n c e s (D.,. o b t u s i f o l i u s on c a l c a r e o u s r o c k s u r f a c e s and B. u n g u i c u l a t a on g r o u n d ) and t h e c o l l e n c h y m a t o u s t h i c k e n i n g s o f t h e upper b a s a l l e a f c e l l s a s i m p o r t a n t c h a r a c t e r s s e p a r a t i n g t h e two. However, s e v e r a l o t h e r c h a r a c t e r s , i n p a r t i c u l a r t h e c a n a l i c u l a t e a nd weak l y d e v e l o p e d a d a x i a l s u r f a c e o f t h e c o s t a , t h e l a r g e , t h i n - w a l l e d b a s a l c e l l s a nd t h e u n d e f i n e d upper l e a f m a r g i n s , a r e b e t t e r a i d s i n d i s t i n g u i s h i n g B. u n g u i c u l a t a (and B. c o n v o l u t a ) f r o m D. o b t u s i f o l i u s , w h i c h has a convex c o s t a l s u r f a c e , s m a l l e r , t h i c k e r - w a l l e d b a s a l c e l l s a nd a m o d i f i e d u p p e r l e a f m a r g i n . DIDYMODON Hedw. A number o f t a x a , t r a d i t i o n a l l y i n c l u d e d i n t h e genus B a r b u l a , a r e t r e a t e d h e r e a s s p e c i e s o r v a r i e t i e s o f Didymodon, f o l l o w i n g t h e t r e a t m e n t s o f S a i t o (1975) and Zander (1978b, 1981b). F o r f u r t h e r d i s c u s s i o n s ee t h e i n t r o d u c t i o n t o B a r b u l a and t h e f o l l o w i n g c h a p t e r . Didymodon i s t a x o n o m i c a l l y c o m p l e x . As S t e e r e (1939a) n o t e d 68 " n e a r l y e v e r y d i a g n o s t i c c h a r a c t e r u s e d i n B a r b u l a ( i n our c a s e Didymodon) v a r i e s w i t h i n r a t h e r wide l i m i t s ...". Many e a r l y r e s e a r c h e r s u s e d o n l y s i n g l e , h i g h l y v a r i a b l e c h a r a c t e r s t o d i s t i n g u i s h t a x a , whereas most s p e c i e s a r e b e s t i d e n t i f i e d by a s e r i e s o f c h a r a c t e r i s t i c s . T h i s has r e s u l t e d i n c o n s i d e r a b l e c o n f u s i o n w i t h r e s p e c t t o t h e l i m i t s of t h e s e t a x a ( S t e e r e 1 9 39a). By u s i n g a m u l t i - c h a r a c t e r a p p r o a c h , a s w e l l a s by e x a m i n i n g a wide range of m a t e r i a l f r o m b o t h N o r t h A m e r i c a and E u r o p e , Zander (1978b, 1981b) has r e s o l v e d many o f t h e g r o u p ' s t a x o n o m i c p r o b l e m s . A l t h o u g h some of Z a n d e r ' s n o m e n c l a t u r a l d e c i s i o n s a r e n o t f o l l o w e d h e r e , t h i s s u r v e y o f Didymodon i s b a s e d e s s e n t i a l l y on Z a n d e r ' s t a x o n o m i c c r i t e r i a . F o u r s p e c i e s of Didymodon have been f o u n d d u r i n g t h i s s u r v e y : D. r i g i d u l u s , D. f a l l a x , D. o c c i d e n t a l i s and D. v i n e a l i s . D. v i n e a l i s i s r e p r e s e n t e d by t h r e e v a r i e t i e s , of w h i c h v a r . b r a c h y p h y l l u s and v a r . l u r i d u s a r e r e c o r d e d a s new t o B r i t i s h C o l u m b i a and Canada. Key t o t h e S p e c i e s of DIDYMODON 1 s u p e r f i c i a l a d a x i a l c e l l s o f t h e upper c o s t a e l o n g a t e -r e c t a n g u l a r t o s u b - l i n e a r ; l e a v e s o f t e n w e a k l y f l e x u o s e ; r a r e , i n s h a d e d s i t e s D. f a l l a x 1 1 s u p e r f i c i a l a d a x i a l c e l l s o f t h e upper c o s t a m o s t l y q u a d r a t e , o c c a s i o n a l l y a few e l o n g a t e - r e c t a n g u l a r c e l l s p r e s e n t ; l e a v e s n o t f l e x u o u s ; common t h r o u g h o u t 2 2 l e a v e s u s u a l l y n a r r o w l y c h a n n e l l e d a bove, t h e c h a n n e l a l m o s t r e a c h i n g t h e apex ( F i g . 1 5); c o s t a o f t e n e x c u r r e n t a s a mucro; u p p e r l a m i n a l c e l l s and c o s t a s t r o n g l y p a p i l l o s e t o r a r e l y smooth; on s o i l o r r o c k 3 2 1 l e a v e s b r o a d l y c h a n n e l l e d above w i t h c h a n n e l e n d i n g w e l l b e f o r e apex i n most l e a v e s ; c o s t a o f t e n e x c u r r e n t a s a 69 f l e s h y , o c c a s i o n a l l y s i n u o u s p r o j e c t i o n ; u p p e r l a m i n a l c e l l s and c o s t a u s u a l l y smooth t o i n d i s t i n c t l y p a p i l l o s e ; on s o i l D. r i q i d u l u s 3 p l a n t s u s u a l l y g r o w i n g as t u f t s o r c u s h i o n s on r o c k , r a r e l y on s o i l ; l e a v e s o v a t e - l a n c e o l a t e and somewhat c o n c a v e , more o r l e s s e r e c t and s l i g h t l y s p r e a d i n g when d r y ; apex o f t h e l e a f o f t e n f l e s h y , c h a n n e l o b s c u r e ; many l e a f t i p s b r o k e n o f f ; u pper l e a f c e l l s s h a r p - a n g l e d , w e a k l y p a p i l l o s e t o smooth D. o c c i d e n t a l i s 2 1 p l a n t s o c c a s i o n a l l y f o r m i n g s m a l l t u f t s , n e v e r f o r m i n g c u s h i o n s , r a r e l y on r o c k , u s u a l l y on s o i l ; l e a v e s o v a t e -l a n c e o l a t e t o l a n c e o l a t e , n e v e r c o n c a v e , u s u a l l y t w i s t e d , r a r e l y e r e c t when d r y ; apex o f t h e l e a f c h a n n e l l e d , not f l e s h y ; l e a f t i p s r a r e l y b r o k e n ; upper l e a f c e l l s q u a d r a t e t o r o u n d e d , n o t s h a r p l y a n g l e d , u s u a l l y d i s t i n c t l y p l u r i -p a p i l l o s e D. v i n e a l i s DIDYMODON FALLAX (Hedw.) Zander B a r b u l a f a l l a x Hedw. P l a n t s d a r k g r e e n t o g r e e n , o c c a s i o n a l l y b r o w n i s h , u s u a l l y a s s m a l l c o l o n i e s s c a t t e r e d on s o i l i n s h a d e d h a b i t a t s o f h i g h e r e l e v a t i o n s . S p o r o p h y t e s common. R e p r e s e n t a t i v e s p e c i m e n s : B89027, B89028 I l l u s t r a t i o n s : Crum and A n d e r s o n 1981, F l o w e r s 1973 D i s t r i b u t i o n : A: r a r e , 3 s i t e s : s i t e 20, Monck P r o v i n c i a l P a r k , n o r t h -e a s t end of N i c o l a L a k e ; s i t e 64, n o r t h of Summerland; s i t e 82, s o u t h - w e s t of Okanagan F a l l s ( F i g . 43) B: s c a t t e r e d a c r o s s t h e s o u t h e r n and c e n t r a l s e c t i o n s o f t h e c o n t i n e n t C: c i r c u m t e m p e r a t e Remarks : T h i s s p e c i e s o f Didymodon i s t h e e a s i e s t of t h e l o c a l t a x a 70 t o i d e n t i f y . In a d d i t i o n t o t h e key c h a r a c t e r s , o t h e r d i s t i n c t i v e f e a t u r e s i n c l u d e t h e i r r e g u l a r l y t h i c k e n e d u p p e r l a m i n a l c e l l w a l l s and, u s u a l l y , p l i c a t i o n s i n t h e b a s a l p a r t of t h e l e a f . The s t a t u s of t h i s t a x o n i s u n c e r t a i n (R. Z a n d e r , p e r s . comm.). Zander s u g g e s t e d t h a t p o s s i b l y D. f a l l a x , B a r b u l a  r e f l e x a ( B r i d . ) B r i d . , D. r i g i d i c a u l i s (C. M u e l l . ) K. S a i t o , D. g i g a n t e u s ( F u n c k . ) J u r . and t h e E u r o p e a n D. s p a d i c e u s ( M i t t . ) L i m p r . a l l r e p r e s e n t a s i n g l e p o l y m o r p h i c s p e c i e s . DIDYMODON OCCIDENTAL!S ( M i t t . ) Zander D. v i n e a l i s v a r . r u b i g i n o s u s ( M i t t . ) Zander B a r b u l a r u b i g i n o s a M i t t . P l a n t s r e d - b r o w n t o , o c c a s i o n a l l y , d a r k g r e e n , d a r k e r a t b a s e , u s u a l l y a s s m a l l c u s h i o n s o r t u r f s on r o c k o u t c r o p s . S p o r o p h y t e s n o t c o l l e c t e d i n t h e s t u d y a r e a . R e p r e s e n t a t i v e s p e c i m e n s : B89029, B89030 I l l u s t r a t i o n s : Lawton 1971 (as B a r b u l a r u b i g i n o s a ) , Zander 1981b D i s t r i b u t i o n : A: s c a t t e r e d t h r o u g h o u t , a l t h o u g h m a i n l y n o r t h e r n ; o c c a s i o n a l l y a b u n d a n t l o c a l l y ( F i g . 44) B: w e s t e r n N o r t h A m e r i c a ; s o u t h e r n B r i t i s h C o l u m b i a t o Montana an d s o u t h t o M e x i c o C: N o r t h A m e r i c a n endemic Remarks : Didymodon o c c i d e n t a l i s i s d i s t i n g u i s h e d i n t h e f i e l d by i t s 71 s a x i c o l o u s h a b i t a t and i t s c u s h i o n - o r t u r f - f o r m i n g h a b i t . I t d i f f e r s f r o m l o c a l members of t h e genus Grimmia, w i t h w h i c h i t s h a r e s t h e above c h a r a c t e r s , by i t s red-brown c o l o u r a t i o n , a w n l e s s l e a v e s and, u s u a l l y , p a p i l l o s e upper l e a f c e l l s . Some c o l l e c t i o n s o f t h i s t a x o n c a n be c o n f u s e d w i t h D. r i q i d u l u s , w h i c h i t r e s e m b l e s i n h a v i n g w e a k l y p a p i l l o s e l e a f c e l l s and a s m a l l , somewhat f l e s h y l e a f apex. However, t h e i r r e g u l a r l y s h a p e d and s t r o n g l y a n g l e d upper l e a f c e l l s , t h e t r a n s v e r s e l y e l o n g a t e b a s a l l e a f c e l l s and d i s t i n c t i v e h a b i t a t s h o u l d d i s t i n g u i s h i t from t h a t t a x o n . The i n t e r i o r c o l l e c t i o n s of D. o c c i d e n t a l i s d i f f e r i n a number o f ways from c o a s t a l c o l l e c t i o n s of t h e same t a x o n . L e a v e s o f c o a s t a l p l a n t s a r e u s u a l l y more p a p i l l o s e , have a more d i s t i n c t u p p e r a d a x i a l c o s t a l g r o o v e and have b a s a l c e l l s w h i c h a r e l o n g i t u d i n a l l y r e c t a n g u l a r . In t h e s e c h a r a c t e r s , t h e c o a s t a l p l a n t s a r e more s i m i l a r t o D. v i n e a l i s t h a n a r e t h e i n t e r i o r p l a n t s . I n a d d i t i o n , c o a s t a l p o p u l a t i o n s u s u a l l y have abundant s p o r o p h y t e s , w h i c h , up t o now, have n o t been c o l l e c t e d f r o m i n t e r i o r p o p u l a t i o n s . C o a s t a l p o p u l a t i o n s a r e m a i n l y on r o c k a l o n g f l o o d e d m a r g i n s o f w a t e r c o u r s e s . A l t h o u g h t h e i n t e r i o r p o p u l a t i o n s a r e o f t e n a l o n g s i d e w a t e r c o u r s e s , t h e y a r e n o t r e s t r i c t e d t h e r e . 72 DIDYMODON RIGIDULUS Hedw. v a r . GRACILIS ( S c h l i e c h . ex Hook e t G r e v . ) Zander D. a c u t u s ( B r i d . ) K. S a i t o B a r b u l a a c u t a ( B r i d . ) B r i d . B. b e s c h e r e l l i i S a u e r b . i n J a e g . P l a n t s d a r k g r e e n or g r e e n t o red-brown ( e s p e c i a l l y i n t h e n o r t h p a r t o f t h e s t u d y a r e a ) , a s s c a t t e r e d i n d i v i d u a l s o r l o o s e t u r f s on s o i l , u s u a l l y among o t h e r mosses; l e a v e s e r e c t t o somewhat t w i s t e d when d r y , r a r e l y c r i s p e d , b r o a d l y c h a n n e l l e d above but n o t t o apex ( F i g . 14); apex f l e s h y and o f t e n w e a k l y u n d u l a t e , r a r e l y e n d i n g i n c o n i c a l mucro. S p o r o p h y t e s n o t c o l l e c t e d . R e p r e s e n t a t i v e s p e c i m e n s : B89031, B89032 I l l u s t r a t i o n s : F i g . 1 4 , S t e e r e 1938 ( a s B a r b u l a b e s c h e r e l l i i ) , F l o w e r s 1973 ( a s B. a c u t a and B. b e s c h e r e l l i i ) , Zander 1981b D i s t r i b u t i o n : A: s c a t t e r e d , f a i r l y common i n t h e n o r t h e r n r e g i o n ( F i g . 45) B: a c r o s s t h e c o n t i n e n t , more common i n t h e west C: c i r c u m p o l a r Remarks : Zander (1981b) p o i n t e d o u t t h a t D. r i q i d u l u s i s d i s t i n g u i s h a b l e f r o m o t h e r Didymodon t a x a , i n p a r t i c u l a r members of t h e D. v i n e a l i s complex, o f t e n o n l y w i t h g r e a t d i f f i c u l t y . He s t r e s s e d t h a t r e l e v a n t c h a r a c t e r s must be e v a l u a t e d c a r e f u l l y , e s p e c i a l l y w i t h r e s p e c t t o t h e d r y l a n d s i t u a t i o n where most s p e c i m e n s a r e m a r k e d l y r e d u c e d i n s i z e . F e a t u r e s o f t h e l e a f 73 apex a r e most d i a g n o s t i c , but t h e d e g r e e of p a p i l l o s i t y , a r e o l a t i o n i n t h e b a s a l r e g i o n , and p l a n t c o l o u r a r e a l s o u s e f u l as a d d i t i o n a l c h a r a c t e r s . I n my r e g i o n a l s u r v e y I f o u n d some of t h e c h a r a c t e r s u s e d by Zander t o be u n r e l i a b l e , s i n c e t h e y v a r i e d c o n s i d e r a b l y . U n r e l i a b l e c h a r a c t e r s i n c l u d e d l e a f c o l o u r a t h i g h m a g n i f i c a t i o n , c o s t a l w i d t h a t m i d - l e a f , c r o s s - s e c t i o n a l f e a t u r e s of t h e c o s t a and u p p e r l e a f c e l l a r e o l a t i o n . Z ander (1981b) r e c o g n i z e d f o u r v a r i e t i e s o f D. r i g i d u l u s : v a r . r i g i d u l u s , v a r . g r a c i l i s , v a r . i c m a d o p h i l l a ( Schimp. ex C. M u e l l . ) Zander and v a r . s u b u l a t u s ( T h e r . e t B a r t r . ex B a r t r . ) Z a n d e r . Most o f o u r c o l l e c t i o n s r e a d i l y f a l l i n t o v a r . g r a c i l i s ( s e e Zander 1981b f o r k e y s ) , a l t h o u g h t h e r e i s some i n t e r g r a d a t i o n t o w a r d s t h e f l e s h y - a w n e d v a r . i c m a d o p h i l a . DIDYMODON VINEALIS ( B r i d . ) Zander The Didymodon v i n e a l i s complex i s h i g h l y v a r i a b l e , e s p e c i a l l y i n d r y l a n d s i t u a t i o n s ( Z a n d e r 1978b, 1981b), o f t e n r e p r e s e n t e d by a p e r p l e x i n g s e r i e s o f r a c e s and h a b i t a t forms ( S t e e r e 1939a). Zander i n s p e c t e d a wide r a n g e o f N o r t h A m e r i c a n c o l l e c t i o n s and r e c o g n i z e d up t o s i x t a x a , e i t h e r as s p e c i e s and v a r i e t i e s ( Z a n d e r 1978b) o r j u s t a s v a r i e t i e s o f D. v i n e a l i s ( Z a n d e r 1981b). F o l l o w i n g Z a n d e r ' s most r e c e n t t r e a t m e n t , f o r r e a s o n s d i s c u s s e d i n t h e i n t r o d u c t i o n t o Didymodon and, i n more d e t a i l , i n t h e f o l l o w i n g c h a p t e r , t h r e e v a r i e t i e s of D. v i n e a l i s a r e - r e c o g n i z e d i n t h e s t u d y r e g i o n : v a r . b r a c h y p h y l l u s , v a r . l u r i d u s and v a r . v i n e a l i s . A n o t h e r v a r i e t y ( s e n s u Zander 1981b), D. v i n e a l i s v a r . 74 n i c h o l s o n i i (= D. n i c h o l s o n i i Culm. ), h a s been f o u n d i n t h r e e s i t e s a l o n g t h e Okanagan and S i m i l k a m e e n V a l l e y s a d j a c e n t t o t h e s t u d y a r e a . F l o w e r s (1973) i n c l u d e d i l l u s t r a t i o n s of t h i s t a x o n i n h i s f l o r a , i d e n t i f y i n g . i t as Didymodon r i g i d u l u s . E x a m i n a t i o n o f F l o w e r s ' c o l l e c t i o n s a t UBC, as w e l l a s h i s d e s c r i p t i o n s and i l l u s t r a t i o n s , c l e a r l y show t h a t t h e t a x o n i s D. v i n e a l i s v a r . n i c h o l s o n i i and not D. r i g i d u l u s . The l e a v e s o f v a r . n i c h o l s o n i i a r e b r o a d e r and l e s s a c u t e t h a n i n t y p i c a l v a r . v i n e a l i s and t h e l e a f m a r g i n s a r e b i - t o ( r a r e l y ) t r i s t r a t o s e , a f e a t u r e a b s e n t i n t h e o t h e r v a r i e t y . I f a s p e c i m e n c a n n o t be d e t e r m i n e d a s one of t h e f o u r v a r i e t i e s and l i e s i n a zone of i n t e r g r a d a t i o n , t h e n i t i s b e s t t o l e a v e i t a s "Didymodon v i n e a l i s ( B r i d . ) Z a n d e r " f o r r e a s o n s n o t e d by Zander ( 1 9 8 1 b ) . Key t o t h e V a r i e t i e s o f DIDYMODON VINEALIS 1 l e a v e s o v a t e t o , r a r e l y , o v a t e - l a n c e o l a t e , r e d - b r o w n , u s u a l l y e r e c t , r a r e l y t w i s t e d , c l a s p i n g , t i n y ( u s u a l l y much s h o r t e r t h a n 1mm), n o t o r s l i g h t l y s p r e a d i n g when wet; upper l a m i n a l c e l l s r o u n d e d and e v e n l y t h i c k - w a l l e d , m o s t l y smooth e x c e p t , o c c a s i o n a l l y , f o r s u p e r f i c i a l c e l l s o f c o s t a ; l e a f apex r o u n d e d t o , r a r e l y , a c u t e , a p i c a l g r o o v e wide; l e a f m a r g i n s u s u a l l y p a l e ; p l a n t s o f t e n g r o w i n g a s s m a l l f i n g e r - l i k e p r o j e c t i o n s , r a r e l y a s t u r f s v a r . l u r i d u s 1 1 l e a v e s o v a t e - l a n c e o l a t e t o l a n c e o l a t e , red-brown t o g r e e n , u s u a l l y l o o s e l y t w i s t e d , n o t c l a s p i n g , f r o m l e s s t h a n 1mm t o g r e a t e r t h a n 2mm l o n g , u s u a l l y s p r e a d i n g when wet; upper l a m i n a l c e l l s u s u a l l y a n g l e d and o c c a s i o n a l l y u n e v e n l y t h i c k e n e d , u s u a l l y p a p i l l o s e , o r , i f smooth, l e a v e s l o n g e r t h a n 1 mm a n d n o t o v a t e ; l e a f apex b l u n t t o a c u t e , n o t r o u n d e d , a p i c a l g r o o v e narrow; l e a f m a r g i n s n o t p a l e ; p l a n t s f o r m i n g t u r f s o r as i n d i v i d u a l s , n o t a s f i n g e r - l i k e p r o j e c t i o n s 2 2 l e a v e s u s u a l l y l e s s t h a n 1mm l o n g , o v a t e - t r i a n g u l a r ; m a r g i n s 75 u s u a l l y s t r o n g l y r e c u r v e d ; many l e a f a p i c e s b l u n t v a r . b r a c h y p h y l l u s 2 1 l e a v e s u s u a l l y g r e a t e r t h a n 1mm l o n g , u s u a l l y o v a t e -l a n c e o l a t e t o l a n c e o l a t e ; m a r g i n s l e s s d i s t i n c t l y r e c u r v e d t o p l a n e ; a p i c e s u s u a l l y t a p e r e d t o a c u t e , r a r e l y b l u n t v a r . v i n e a l i s DIDYMODON VINEALIS v a r . BRACHYPHYLLUS ( S u l l . i n W h i p p l . ) Zander D. b r a c h y p h y l l u s ( S u l l . i n W h i p p l . ) Zander. B a r b u l a b r a c h y p h y l l a S u l l . i n W h i p p l . P l a n t s d a r k g r e e n t o red-brown, u s u a l l y a s s m a l l t u r f s on s o i l i n sunny c o n d i t i o n s . S p o r o p h y t e s n o t f o u n d . R e p r e s e n t a t i v e s p e c i m e n s : B89033, B89034 I l l u s t r a t i o n s : Lawton 1971, Zander 1981b D i s t r i b u t i o n : A: o c c a s i o n a l , m a i n l y i n t h e s o u t h e r n a r e a ( F i g . 46) B: w e s t e r n N o r t h A m e r i c a : s o u t h e r n B r i t i s h C o l u m b i a t o M e x i c o ; new t o B r i t i s h C o l u m b i a and Canada C: w e s t e r n N o r t h A m e r i c a n endemic ( a l t h o u g h p o s s i b l y more w i d e s p r e a d , see t h e r e m a r k s ) Remarks : D. v i n e a l i s v a r . b r a c h y p h y l l u s i s d i s t i n g u i s h e d f r o m t h e o t h e r v a r i e t i e s m a i n l y by t h e o v a t e - t r i a n g u l a r , b l u n t i s h l e a v e s , w h i c h u s u a l l y have s t r o n g l y r e c u r v e d m a r g i n s , and t h e s m a l l l e a f s i z e ( o n l y D. v i n e a l i s v a r . l u r i d u s has s m a l l e r l e a v e s ) . O t h e r d i s t i n c t i v e f e a t u r e s i n c l u d e t h e u s u a l l y b r o a d l y c u c u l l a t e l e a f a p i c e s , e l o n g a t e - r e c t a n g u l a r a d a x i a l c o s t a l c e l l s and t h e s t r o n g l y p a p i l l o s e c o n d i t i o n , a l t h o u g h s p e c i m e n s have been examined where l o w e r stem l e a v e s a r e smooth w h i l e t h o s e above 76 a r e p a p i l l o s e . In a d d i t i o n , t h e h a b i t and s t r o n g l y r e c u r v e d m a r g i n s o f D. v i n e a l i s v a r . b r a c h y p h y l l u s a r e r e m i n i s c e n t o f P s e u d o c r o s s i d i u m  r e v o l u t u m , B r y o e r y t h r o p h y l l u m columbianum and Desmatodon  c o n v o l u t u s . However, t h e l a t t e r two t a x a have d i s t i n c t , wide and convex c o s t a e w h i c h r e a d i l y s e p a r a t e them, whereas P. r e v o l u t u m has b o t h s t r o n g l y r e v o l u t e l e a f m a r g i n s and k e e l e d l e a v e s ( F i g . 1 7 ) . Zander (1981b) q u e s t i o n e d t h e endemic s t a t u s o f D. v i n e a l i s v a r . b r a c h y p h y l l u s s u g g e s t i n g t h a t i t may be synonymous w i t h D. r e e d i i R o b i n s o n of e a s t e r n N o r t h A m e r i c a and D. t e c t o r u m S a i t o f r o m C h i n a and J a p a n . DIDYMODON VINEALIS v a r . LURIDUS ( H o r n s c h . i n S p r e n g . ) Z a n d e r D. l u r i d u s H o r n s c h . i n S p r e n g . D. t r i f a r i u s v a r . l u r i d u s ( H o r n s c h . i n S p r e n g . ) Mont. B a r b u l a l u r i d a H o r n s c h . i n S p r e n g . P l a n t s r e d - b r o w n , o c c a s i o n a l l y d a r k g r e e n , u s u a l l y a s s h o r t (<1.0 mm t a l l ) i n d i v i d u a l s o r r a r e l y a s t u r f s on s o i l i n sunny c o n d i t i o n s ; l e a v e s s h o r t - l a n c e o l a t e t o o v a t e - t r i a n g u l a r ; l e a f apex u s u a l l y r o u n d e d , r a r e l y a c u t e , u s u a l l y w e a k l y r e c u r v e d ; u p p e r l e a f c e l l s smooth, r a r e l y p a p i l l o s e , u s u a l l y w i t h r o u n d e d o r o v a l ( r a r e l y a n g l e d ) lumens. S p o r o p h y t e s n o t f o u n d . R e p r e s e n t a t i v e s p e c i m e n s : B89035, B89036 I l l u s t r a t i o n s : S m i t h 1978a (as B a r b u l a t r i f a r i a ) , Zander 1981b D i s t r i b u t i o n : A: f a i r l y common, m a i n l y i n t h e s o u t h Okanagan and e a s t o f 77 Kamloops ( F i g . 47) B: w e s t e r n N o r t h A m e r i c a : M e x i c o t o s o u t h e r n B r i t i s h C o l u m b i a ; new t o Canada and B r i t i s h C o l u m b i a C: w e s t e r n N o r t h A m e r i c a n / E u r a s i a n d i s j u n c t Remarks : D. v i n e a l i s v a r . l u r i d u s i s , g e n e r a l l y , t h e most d i m i n u t i v e member o f t h e D. v i n e a l i s c o m p l e x . T h i s i s c o n t r a r y t o t h e o p i n i o n of S t e e r e (1939a) who c o n s i d e r e d v a r . b r a c h y p h y l l u s t o be t h e s m a l l e s t f o r m . C e r a t o d o n p u r p u r e u s o c c a s i o n a l l y has a s i m i l a r g r o w t h h a b i t as w e l l a s smooth, r o u n d e d l a m i n a l c e l l s , but s u c h p l a n t s s h o u l d be r e a d i l y s e p a r a t e d u s i n g o t h e r c h a r a c t e r s ( s e e t h e r e m a r k s w i t h C. p u r p u r e u s ) . I t e n t a t i v e l y a c c e p t t h e o p i n i o n of Zander (1978b, 1981b) c o n c e r n i n g t h e s t a t u s o f t h i s t a x o n . E x a m i n a t i o n o f E u r o p e a n s p e c i m e n s and i d e n t i f i c a t i o n o f my m a t e r i a l by A. C r u n d w e l l t e n d e d t o c o n f i r m t h a t t h e y a r e t h e same ( t h e E u r o p e a n m a t e r i a l u s u a l l y i s l a r g e r , w i t h a l a x e r g r o w t h h a b i t and has r e c u r v e d m a r g i n s ) , y e t i n t e r g r a d a t i o n s between t h i s and t h e l a r g e r members o f t h e D. v i n e a l i s complex a r e n o t common. C r u n d w e l l ( p e r s . comm.) c o n s i d e r e d t h i s v a r i e t y t o be a d i s t i n c t s p e c i e s . However, f o r r e a s o n s o u t l i n e d i n t h e i n t r o d u c t i o n , Z a n d e r ' s t r e a t m e n t i s f o l l o w e d h e r e . DIDYMODON VINEALIS v a r . VINEALIS Zander B a r b u l a v i n e a l i s B r i d . P l a n t s d a r k g r e e n o r g r e e n t o red-brown, a s i n d i v i d u a l s o r 78 s m a l l t u r f s , u s u a l l y on s o i l , o c c a s i o n a l l y on r o c k . S p o r o p h y t e s not c o l l e c t e d . R e p r e s e n t a t i v e s p e c i m e n s : B89037, B89038 I l l u s t r a t i o n s : F i g . 15, S a v i c z - L j u b i t z k a j a and S m i r n o v a 1970 (as B a r b u l a v i n e a l i s and B. c y l i n d r i c a ) , Lawton 1971 (as B. v i n e a l i s ) , Z ander 1981b. D i s t r i b u t i o n : A: w i d e s p r e a d and f a i r l y common ( F i g . 48) B: w e s t e r n N o r t h A m e r i c a : B r i t i s h C o l u m b i a t o Montana s o u t h t o M e x i c o C: c i r c u m t e m p e r a t e ; b i p o l a r ( S o u t h A m e r i c a ; C h i l e , P a t a g o n i a ) Remarks : T h i s t a x o n i s u s u a l l y t h e e a s i e s t v a r i e t y o f D. v i n e a l i s t o d e t e r m i n e w i t h c o n f i d e n c e . The l o n g - l a n c e o l a t e l e a v e s w i t h d i s t i n c t c h a n n e l l e d a p i c e s a r e i t s most c h a r a c t e r i s t i c f e a t u r e s . D. v i n e a l i s v a r . v i n e a l i s s t r o n g l y r e s e m b l e s D. r i g i d u l u s v a r . g r a c i l i s m a c r o s c o p i c a l l y b u t , w i t h c a r e , c a n be s e p a r a t e d by t h e key c h a r a c t e r s . PHASCUM Hedw. The genus Phascum c o n t a i n s a b o u t s i x s p e c i e s f o r w h i c h i n n u m e r a b l e v a r i e t i e s have been d e s c r i b e d ( s e e W i j k e t a l . 1959 - 1969). A l t h o u g h s c a t t e r e d , t h e genus i s d i s t r i b u t e d p r i m a r i l y a c r o s s n o r t h t e m p e r a t e l a t i t u d e s . Up t o t h e p r e s e n t , o n l y t h r e e s p e c i e s , P. c u s p i d a t u m , P. h y a l i n o t r i c h u m and P. f l o e r k e a n u m , have been r e p o r t e d f r o m N o r t h 79 A m e r i c a , w i t h o n l y P. c u s p i d a t u m known from t h e s t u d y a r e a . The d i s c o v e r y o f P. v l a s s o v i i , p r e v i o u s l y f o u n d o n l y i n s o u t h e a s t e r n E u r o p e and c e n t r a l A s i a , was e n t i r e l y u n e x p e c t e d . Phaseurn i s not e n t i r e l y d i s t i n c t f r o m P o t t i a , a c l o s e l y r e l a t e d genus w h i c h s h a r e s many g a m e t o p h y t i c and s p o r o p h y t i c c h a r a c t e r s . See t h e remarks i n t h e f o l l o w i n g c h a p t e r . Key t o t h e S p e c i e s o f PHASCUM I upper l e a f c e l l s w i t h p a p i l l a e r a i s e d a s t a l l , h o l l o w p r o j e c t i o n s ; a d a x i a l s u r f a c e o f t h e c o s t a o r n a m e n t e d w i t h b o t t l e - s h a p e d , p a p i l l o s e c e l l s s i m i l a r t o t h o s e on t h e l a m i n a l c e l l s , b u t l a r g e r ( s e e F i g . 16) P. v l a s s o v i i I I u p p e r l e a f c e l l s w i t h low p a p i l l a e ; c o s t a n o t orn a m e n t e d , sometimes c o v e r e d w i t h b u l g i n g , i s o d i a m e t r i c and p a p i l l o s e c e l l s P. c u s p i d a t u m PHASCUM CUSPIDATUM Hedw. P l a n t s y e l l o w - g r e e n t o l i g h t brown, o f t e n p a l e when o l d , u s u a l l y i n s m a l l t u f t s of a few i n d i v i d u a l s on b a r e ( o f t e n sandy) s o i l ; common i n many sunny l o c a t i o n s i n s t e p p e . S p o r o p h y t e s common; c a p s u l e s s h i n y r e d - b r o w n , g l o b o s e and immersed i n t h e l e a v e s . R e p r e s e n t a t i v e s p e c i m e n s : B89039, B89040 I l l u s t r a t i o n s : Crum and A n d e r s o n 1981, F l o w e r s 1973 D i s t r i b u t i o n : A: w i d e s p r e a d and common ( F i g . 73) B: s c a t t e r e d a c r o s s s o u t h e r n h a l f of N o r t h A m e r i c a C: c i r c u m t e m p e r a t e ; b i p o l a r (New Z e a l a n d ) 80 Remarks : Phascum c u s p i d a t u m i s w i d e s p r e a d and, i n most c a s e s , e a s i l y r e c o g n i z a b l e i n t h e f i e l d . The g l o b o s e , immersed c a p s u l e s and t h e y e l l o w - g r e e n s l i g h t l y t w i s t e d , prow-shaped l e a v e s a r e d i s t i n c t i v e . O t h e r t h a n P. v l a s s o v i i , t h e o n l y t a x a w h i c h c o u l d be m i s t a k e n f o r P. c u s p i d a t u m a r e P o t t i a b r y o i d e s , w h i c h has e x e r t e d , e l l i p s o i d c a p s u l e s , smoother upper l e a f c e l l s and i s r a r e , and P t e r y q o n e u r u m k o z l o v i i , w h i c h has l a m e l l a t e l e a v e s , l a c k s a g o l d e n hue ( i t i s more r e d d i s h ) and i s r e s t r i c t e d t o t h e b o r d e r s o f a l k a l i n e d e p r e s s i o n s . O c c a s i o n a l l y , some s t e r i l e s p e c i m e n s of P. c u s p i d a t u m have b u l g i n g c e l l s o v e r t h e upper c o s t a . T h i s c a n r e s u l t i n some c o n f u s i o n w i t h t h e genus C r o s s i d i u m . The c h a r a c t e r s n o t e d i n t h e key s h o u l d r e s o l v e most p r o b l e m s . C o n t r a r y t o some r e p o r t s i n t h e l i t e r a t u r e ( e . g . , Tan 1980), P. c u s p i d a t u m i s r a r e l y , i f e v e r , an e p h e m e r a l o r a n n u a l s p e c i e s . T h r e e p o p u l a t i o n s , two n e a r Vaseaux L a k e i n t h e s o u t h Okanagan ( S i t e s 73 and 74) and one n e a r Cache C r e e k ( S i t e 3 4 ) , were o b s e r v e d o v e r a p e r i o d o f t h r e e y e a r s . D u r i n g t h i s p e r i o d , new p l a n t s grew e a c h w i n t e r and s p r i n g f r o m buds t h a t a p p e a r a l o n g t h e stem below t h e main c l u s t e r of l e a v e s . O n l y t h e o l d e r l e a v e s and t h e s p o r o p h y t e s d i e d down e a c h y e a r , n o t t h e whole p l a n t . The d e s c r i p t i o n by D u r i n g (1979) o f Phascum a s a p a u c i p e r e n n i a l may be t h e b e s t t e r m t o use i n t h i s c a s e . The o n l y t r u e e p h e m e r a l s i n t h e s t e p p e a r e t h e two s p e c i e s o f R i c c i a and, p o s s i b l y , F u n a r i a m u h l e n b e r g i i and F. h y q r o m e t r i c a . 81 PHASCUM VLASSOVII L a z a r . L y d i a e a v l a s s o v i i L a z a r . P l a n t s y e l l o w - ( g o l d e n - ) g r e e n , s c a t t e r e d a s i n d i v i d u a l s on s o i l ( o f t e n c l a y - r i c h ) ; l e a v e s t w i s t e d when d r y , s l i g h t l y s p r e a d i n g and somewhat r e c u r v e d when wet, o v a t e t o o v a t e -l a n c e o l a t e , t a p e r i n g t o a s t o u t , o r a n g e awn, u s u a l l y somewhat c o n c a v e and p r o w - s h a p e d above; m a r g i n s s t r o n g l y r e c u r v e d a t m i d l e a f , r e c u r v e d t o p l a n e above m i d l e a f , m a r g i n a l c e l l s o c c a s i o n a l l y more e l o n g a t e t h a n c e l l s n e a r e r c o s t a , u s u a l l y smooth, r a r e l y w e a k l y p a p i l l o s e , n e v e r c o l u m n a r - p a p i l l o s e l i k e a d j a c e n t l a m i n a l c e l l s ; c o s t a s t o u t , o f t e n w i d e r n e a r apex t h a n below, e x c u r r e n t a s l o n g , smooth, o r a n g e awn; u p p e r l a m i n a l c e l l s 10-15 m i c r o m e t e r s w i d e , q u a d r a t e t o i r r e g u l a r l y i s o d i a m e t r i c , o r n a m e n t e d w i t h t a l l , h o l l o w , a p i c a l l y p a p i l l o s e p r o j e c t i o n s ( F i g . 16), e s p e c i a l l y t o w a r d s t h e c o s t a , c e l l s g r a d u a l l y l a r g e r t o w a r d s t h e m i d d l e o f t h e l e a f ; b a s a l c e l l s s h o r t t o l o n g r e c t a n g u l a r , t h i n - w a l l e d and smooth. S p o r o p h y t e s common, immersed i n l e a v e s , o n l y immature s p o r o p h y t e s c o l l e c t e d , immature s p o r e s 18-25 m i c r o m e t e r s w i d e , w e a k l y p a p i l l o s e ; c a p s u l e s a p i c u l a t e ; c a l y p t r a t i n y , c u c u l l a t e . R e p r e s e n t a t i v e s p e c i m e n s : B89041, B89042 I l l u s t r a t i o n s : F i g . 16, S a v i c z - L j u b i t z k a j a and S m i r n o v a 1970 D i s t r i b u t i o n : A: r a r e , 2 s i t e s : s i t e 52, s o u t h s i d e of t h e Thompson R i v e r a t t h e D a l l e s ; s i t e 72, e a s t s i d e o f P e n t i c t o n ( F i g . 74) B: s o u t h e r n B r i t i s h C o l u m b i a C: w e s t e r n N o r t h A m e r i c a / w e s t e r n E u r a s i a ( U k r a i n e , 82 A r m e n i a , C. A s i a ) ; new t o N o r t h A m e r i c a Remarks : M i c r o s c o p i c a l l y , t h i s e l e g a n t l y o r n a m e n t e d moss c a n n o t be m i s t a k e n f o r any o t h e r t a x o n . P. v l a s s o v i i d i f f e r s f r o m P. c u s p i d a t u m not o n l y i n l e a f c e l l o r n a m e n t a t i o n , but a l s o i n i t s g e n e r a l l y s m a l l e r s t a t u r e ( i n c l u d i n g s m a l l e r l e a f c e l l s , s h o r t e r l e a f l e n g t h and s m a l l e r s p o r e s ) . F u r t h e r m o r e , P. v l a s s o v i i has s t i f f e r , more e r e c t l e a v e s and s t o u t e r , more g o l d e n c o s t a e t h a n P. c u s p i d a t u m . POTTIA F u r n r . P o t t i a i s c h a r a c t e r i z e d by s m a l l , u s u a l l y i n c o n s p i c u o u s g a m e t o p h y t e s t h a t b e a r s h o r t s p o r o p h y t e s w i t h p e r i s t o m e s a b s e n t o r , i f p r e s e n t , composed o f 16 r e d u c e d , i m p e r f e c t l y d i v i d e d o r i r r e g u l a r l y b i f i d t e e t h ( S m i t h 1978a). P o t t i a i s c l o s e l y r e l a t e d t o Desmatodon and Phascum. F o r f u r t h e r d i s c u s s i o n o f t h e s e r e l a t i o n s h i p s , see t h e e a r l i e r s e c t i o n on Desmatodon and t h e r e m a r k s i n f o l l o w i n g c h a p t e r . F o u r s p e c i e s o f P o t t i a have been f o u n d i n t h e s t u d y r e g i o n . Of t h e s e , o n l y P. h e i m i i was c o l l e c t e d a t more t h a n one l o c a l e . T h i s r a r e n e s s p o s s i b l y r e s u l t s f r o m b e i n g o v e r l o o k e d , r e f l e c t i n g t h e i r m i n u t e s i z e and, i n some c a s e s , t h e i r m a c r o s c o p i c r e s e m b l a n c e t o t h e u b i q u i t o u s moss, P t e r y g o n e u r u m ovaturn. Key t o t h e S p e c i e s o f POTTIA 1 p l a n t s a l o n g b o r d e r s o f s e a s o n a l l y wet h a b i t a t s ; l e a f m a r g i n s u s u a l l y t o o t h e d above by p r o j e c t i n g ends of e l o n g a t e d b o r d e r c e l l s ; o p e r c u l u m r e m a i n i n g a t t a c h e d t o c a p s u l e by c o l u m e l l a ( s y s t y l i u s ) 2 83 p l a n t s o f d r y h a b i t a t s ; l e a v e s u s u a l l y n o t t o o t h e d (sometimes t o o t h e d i n P. w i l s o n i i ) ; o p e r c u l u m c o m p l e t e l y d e h i s c e n t .... .3 l e a v e s l o n g awned; l e a f b o r d e r s i n d i s t i n c t ; u p p e r l e a f c e l l s smooth t o w e a k l y p a p i l l o s e P. n e v a d e n s i s l e a v e s n o t awned; l e a v e s b o r d e r e d by a band o f smooth, e l o n g a t e y e l l o w i s h c e l l s ; u p p e r l e a f c e l l s c o n s p i c u o u s l y p a p i l l o s e P. h e i m i i l e a v e s l a n c e o l a t e ; upper c e l l s smooth t o w e a k l y p a p i l l o s e ; c a p s u l e s s h o r t e x s e r t e d ; c a p s u l e s e l l i p s o i d and c l e i s t o c a r p o u s P. b r y o i d e s l e a v e s o v a t e - l a n c e o l a t e , u p p e r l e a f c e l l s d i s t i n c t l y p a p i l l o s e ; c a p s u l e s w e l l e x s e r t e d above l e a v e s , s h o r t -c y l i n d r i c a l and s t e g o c a r p o u s P. c f . w i l s o n i i POTTIA BRYOIDES ( D i c k s . ) M i t t . P l a n t s g r e e n t o p a l e brown, f o r m i n g s m a l l open c o l o n i e s on s o i l ; l e a v e s b r o a d l y l a n c e o l a t e , t a p e r e d above, p r o w - s h a p e d and c o n c a v e , l o o s e l y t w i s t e d when d r y ; m a r g i n s w e a k l y r e c u r v e d a l m o s t t o apex; c o s t a e x c u r r e n t a s smooth awn, d a r k r e d on b a c k ; u p p e r l e a f c e l l s e l o n g a t e - r e c t a n g u l a r t o a l m o s t i s o d i a m e t r i c ( h e x a g o n a l ) , most c e l l s 20-30 m i c r o m e t e r s i n l o n g e s t d i m e n s i o n , u s u a l l y w e a k l y p a p i l l o s e ; b a s a l c e l l s u n e v e n l y r e c t a n g u l a r , smooth, l a r g e r t h a n upper c e l l s . S p o r o p h y t e s common; c a p s u l e s s h o r t e x s e r t e d , e l l i p s o i d and c l e i s t o c a r p o u s , a l t h o u g h w i t h a t h i n band o f d i f f e r e n t i a t e d c e l l s n e a r t h e t o p o f t h e c a p s u l e . R e p r e s e n t a t i v e s p e c i m e n : B89043 I l l u s t r a t i o n s : S a v i c z - L j u b i t z k a j a and S m i r n o v a 1 970, S m i t h 1978a, Wareham 1939 84 D i s t r i b u t i o n : A: r a r e , S i t e 101 o n l y , 8 km s o u t h of Cawston, S i m i l k a m e e n V a l l e y ( F i g . 78) B: w e s t e r n N o r t h A m e r i c a : B r i t i s h C o l u m b i a , (?) S a s k a t c h e w a n , A r i z o n a , C a l i f o r n i a ; new t o B r i t i s h C o l u m b i a and p r o b a b l y new t o Canada C: w e s t e r n N o r t h A m e r i c a / w e s t e r n E u r a s i a Remarks : S m i t h (1978a) n o t e d t h a t P o t t i a i s c l o s e l y r e l a t e d t o Phascum t h r o u g h P o t t i a b r y o i d e s ( a n d P. r e c t a , a E u r o p e a n t a x o n ) b e c a u s e o f t h e more o r l e s s c l e i s t o c a r p o u s and emergent c a p s u l e s , and c o n s i d e r e d r e c o g n i t i o n of two g e n e r a i n s t e a d o f one a s somewhat a r b i t r a r y . As n o t e d i n t h e p r e s e n t s t u d y , t h e r e s e m b l a n c e o f t h e g a m e t o p h y t e s o f P. b r y o i d e s t o t h o s e o f Phascum c u s p i d a t u m i s c e r t a i n l y s t r i k i n g , and i f s p o r o p h y t e s were l a c k i n g , i t wou l d be d i f f i c u l t t o d i s t i n g u i s h them. Phascum d i f f e r s , u s u a l l y , i n b e i n g more p a p i l l o s e , s m a l l e r ( b o t h i n l e a f and c e l l s i z e ) , l a c k i n g a r e d c o s t a a n d h a v i n g a l o n g e r awn. However, a l l o f t h e s e c h a r a c t e r s o v e r l a p somewhat between t h e two t a x a . POTTIA HEIMII (Hedw.) F u r n r . e x. Hampe Desmatodon h e i m i i (Hedw.) M i t t . P l a n t s g r e e n t o y e l l o w - g r e e n ( s ometimes r e d d i s h t i n g e d ) , o c c a s i o n a l l y e n c r u s t e d w i t h w h i t e s a l t s , a s i s o l a t e d i n d i v i d u a l s o r , more commonly, i n open c o l o n i e s a l o n g m a r g i n s o f a l k a l i n e d e p r e s s i o n s o r s e e p a g e a r e a s . S p o r o p h y t e s common; c a p s u l e s 85 s y s t y l i u s . R e p r e s e n t a t i v e s p e c i m e n s : B89044, B89045 I l l u s t r a t i o n s : Crum and A n d e r s o n 1981, F l o w e r s 1973, Lawton 1971, I r e l a n d 1982, Wareham 1939 D i s t r i b u t i o n : A: o c c a s i o n a l , more common i n t h e s o u t h Okanagan V a l l e y ( F i g . 79) B: m a i n l y t h e n o r t h e r n p o r t i o n o f t h e c o n t i n e n t : s c a t t e r e d i n t h e A r c t i c , s o u t h t o C a l i f o r n i a i n t h e west, and t o s o u t h e r n Quebec i n t h e e a s t C: c i r c u m p o l a r ; b i p o l a r (New Z e a l a n d , S o u t h A m e r i c a , A n t a r c t i c a ) Remarks : P o p u l a t i o n s o f P o t t i a h e i m i i a r e v e r y d i s t i n c t i v e , p a r t i c u l a r l y b e c a u s e of t h e i r h a b i t a t and t h e abundance of t h e c h a r a c t e r i s t i c a l l y s y s t y l i u s s p o r o p h y t e s . M i c r o s c o p i c a l l y , t h e t o o t h e d upper l e a f b o r d e r composed o f smooth and e l o n g a t e y e l l o w i s h c e l l s c h a r a c t e r i z e s o n l y P. h e i m i i . The l e n g t h o f t h e s e t a e , p a r t i c u l a r l y between p o p u l a t i o n s o f P. h e i m i i , i s h i g h l y v a r i a b l e . R a r e l y , p o p u l a t i o n s a r e f o u n d w i t h o u t s y t y l i o u s c a p s u l e s . The t a x o n o m i c p o s i t i o n o f P. h e i m i i i s c o n t r o v e r s i a l . I t i s o f t e n c o n s i d e r e d a Desmatodon, but a number o f c h a r a c t e r s , i n c l u d i n g l a r g e r c e l l s i z e , l a c k o f a p e r i s t o m e and l a r g e r s p o r e s i z e , f o r example, a r e more c h a r a c t e r i s t i c o f t h e genus P o t t i a . F o r f u r t h e r d i s c u s s i o n see Desmatodon and t h e f o l l o w i n g c h a p t e r . A number o f f e a t u r e s , i n c l u d i n g t h e s y s t y l i u s c a p s u l e s , t h e 86 l a c k of a p e r i s t o m e , t h e wea k l y m o d i f i e d upper b o r d e r o f t h e l e a f and t h e h a b i t a t , c l o s e l y a l l y P. n e v a d e n s i s t o P. h e i m i i , f u r t h e r s u p p o r t i n g t h e p l a c e m e n t o f P. h e i m i i i n P o t t i a and not Desmatodon. POTTIA NEVADENSIS C a r d , e t T h e r . P l a n t s u s u a l l y p a l e g o l d e n - g r e e n , i n open t o compact c o l o n i e s a l o n g m a r g i n s of a l k a l i n e d e p r e s s i o n s ; l e a v e s e r e c t t o s l i g h t l y t w i s t e d . S p o r o p h y t e s common; c a p s u l e s s y s t y l i o u s . R e p r e s e n t a t i v e s p e c i m e n : B89046 I l l u s t r a t i o n s : F l o w e r s 1973, Lawton 1971, Wareham 1939 D i s t r i b u t i o n : A: r a r e , s i t e 81 o n l y , W h i t e L a k e a r e a ( F i g . 80) B: s c a t t e r e d i n w e s t e r n N o r t h A m e r i c a : s o u t h e r n B r i t i s h C o l u m b i a , A l b e r t a ( ? ) , S a s k a t c h e w a n ( ? ) , W a s h i n g t o n , Oregon, Nevada, U t a h , A r i z o n a C: w e s t e r n N o r t h A m e r i c a n endemic Remarks : P o t t i a n e v a d e n s i s was o v e r l o o k e d d u r i n g t h e c o u r s e o f t h e p r e s e n t s u r v e y b u t was d i s c o v e r e d l a t e r i n t h e UBC h e r b a r i u m (W.B. S c h o f i e l d , 33382; May 7, 1967). I t s s o l e l o c a t i o n , t h e W h i t e L a k e s i t e , i s r a t h e r i n t e r e s t i n g c o n s i d e r i n g t h a t two o t h e r t a x a , P t e r y q o n e u r u m l a m e l l a t u m and E n t o s t h o d o n " r u b i g i n o s u s , h a v e , w i t h i n t h e s t u d y a r e a , t h e i r o n l y l o c a t i o n s t h e r e a s w e l l . P. l a m e l l a t u m was a l s o f o u n d o r i g i n a l l y d u r i n g a h e r b a r i u m s e a r c h b u t was l a t e r f o u n d i n t h e f i e l d , whereas P. n e v a d e n s i s was n o t r e l o c a t e d i n t h e f i e l d . I t was p r o b a b l y 87 o v e r l o o k e d b e c a u s e o f i t s r e l a t i v e l y s m a l l s i z e c ombined w i t h i t s h a b i t o f g r o w i n g among a number of o t h e r mosses, i n c l u d i n g F u n a r i a h y q r o m e t r i c a , a d i m i n u t i v e f o r m o f T o r t u l a r u r a l i s , Bryum c a e s p i t i t i u m and P o t t i a h e i m i i . S i n c e b o t h P. h e i m i i and P. n e v a d e n s i s have s y s t y l i u s c a p s u l e s , t h e r e i s f u r t h e r c o n f u s i o n s i n c e t h e awns of P. n e v a d e n s i s , d i s t i n c t i v e m i c r o s c o p i c a l l y , a r e not r e a d i l y a p p a r e n t i n t h e f i e l d . P. n e v a d e n s i s i s s m a l l e r and l e s s c r i s p e d o r c o n t o r t e d t h a n P. h e i m i i . POTTIA c f . WILSONII (Hook.) B.S.G. P l a n t s s m a l l and i n c o n s p i c u o u s , u s u a l l y y e l l o w - g r e e n , g r o w i n g on c l a y - r i c h s o i l ; l e a v e s 1.0 - 1.5mm l o n g , b r o a d o v a t e -l a n c e o l a t e t o somewhat o b l o n g - l i g u l a t e , u s u a l l y t a p e r e d a b o v e , b r o a d l y c o n c a v e , i n f o l d e d and somewhat t w i s t e d when d r y , s p r e a d i n g when wet; m a r g i n s r e c u r v e d t o a l m o s t r e v o l u t e f r o m n e a r t h e l e a f b a s e t o n e a r t h e apex, c o s t a s t r o n g , e x c u r r e n t a s a l o n g , h y a l i n e , smooth awn, one h a l f t o t h e f u l l l e n g t h o f t h e l e a f ; m e d i a l l a m i n a l c e l l s ( 10)14-16 m i c r o m e t e r s w i d e , i r r e g u l a r l y q u a d r a t e o r r e c t a n g u l a r , p l u r i - p a p i l l o s e t o t h e m a r g i n s , u s u a l l y g r a d i n g i n t o e l o n g a t e , smooth and h y a l i n e c e l l s w h i c h sometimes f o r m t e e t h a l o n g t h e u p p e r l e a f b o r d e r s by p r o j e c t i n g e n d s o f c e l l s ; b a s a l c e l l s smooth, h y a l i n e , l o n g r e c t a n g u l a r a n d t h i n - w a l l e d , sometimes c e l l w a l l s o r a n g e ; p r o b a b l y a u t o i c o u s . S p o r o p h y t e s o c c a s i o n a l ; c a p s u l e s s h o r t -c y l i n d r i c a l ; p e r i s t o m e a b s e n t ; c a l y p t r a r o s t r a t e ; s p o r e s o v a t e -s p h e r i c a l , f i n e l y p a p i l l o s e , 20-25 m i c r o m e t e r s w i d e . 88 R e p r e s e n t a t i v e s p e c i m e n : B89047 I l l u s t r a t i o n s : S m i t h 1978a D i s t r i b u t i o n : A: r a r e , s i t e 70 o n l y , west o f P e n t i c t o n ( F i g . 81) B: s o u t h e r n B r i t i s h C o l u m b i a C: w e s t e r n N o r t h A m e r i c a / w e s t e r n E u r a s i a ; new t o N o r t h A m e r i c a Remarks : B o t h A . J . E . S m i t h and R. Zander i d e n t i f i e d o u r m a t e r i a l as t h e E u r o p e a n P. w i l s o n i i . However, c l o s e e x a m i n a t i o n o f E u r o p e a n m a t e r i a l r e v e a l s t h a t P. w i l s o n i i d i f f e r s i n a number of r e s p e c t s from t h e B r i t i s h C o l u m b i a m a t e r i a l . Most n o t i c e a b l y , l e a v e s o f t h e E u r o p e a n s p e c i m e n s a r e l a r g e r ( u s u a l l y g r e a t e r t h a n 1.5mm) and have s h o r t , brown awns. In c o n t r a s t , our m a t e r i a l has s m a l l e r l e a v e s and v e r y l o n g , h y a l i n e awns. More m a t e r i a l , e s p e c i a l l y i f i n a more m a t u r e s t a t e , w o u l d h e l p i n c o n f i r m i n g i t s s t a t u s . P o t t i a c f . w i l s o n i i was f o u n d i n a p o p u l a t i o n o f P t e r y g o n e u r u m ovaturn, f r o m w h i c h i t d i f f e r s i n l a c k i n g l e a f l a m e l l a e and h a v i n g p a p i l l o s e l a m i n a l c e l l s . PSEUDOCROSSIDIUM W i l l i a m s A number of c h a r a c t e r s , most n o t a b l y t h e s t r o n g l y r e v o l u t e l e a f m a r g i n s , an o r n a m e n t e d u p p e r c o s t a ( s ometimes d i v i d e d i n t o f i l a m e n t s , a l t h o u g h n o t i n our s p e c i e s ) , and h i g h l y d i f f e r e n t i a t e d p e r i c h a e t i a l l e a v e s c l e a r l y s e p a r a t e P s e u d o c r o s s i d i u m f r om b o t h B a r b u l a and T o r t u l a ( W i l l i a m s 1915; 89 Zander 1979a, 1981a), g e n e r a i n w h i c h members of P s e u d o c r o s s i d i u m a r e o f t e n p l a c e d . PSEUDOCROSSIDIUM REVOLUTUM ( B r i d . i n S c h r a d . ) Zander B a r b u l a r e v o l u t a B r i d . i n S c h r a d . B. p l a t y n e u r a C. M u e l l . e t K i n d b . i n Macoun Desmatodon e l l e s m e r e n s i s B r a s s a r d P l a n t s brown t o y e l l o w - g r e e n , u s u a l l y f o u n d a s i n d i v i d u a l s o r s m a l l c o l o n i e s , r a r e l y as e x t e n s i v e t u r f s ( i n s h a d e d c o n d i t i o n s ) , u s u a l l y on s o i l i n s t e p p e ; l e a v e s i m b r i c a t e - t w i s t e d when d r y , s p r e a d i n g when wet ( p l a n t s a p p e a r i n g s t e l l a t e from a b o v e ) , o v a t e - l a n c e o l a t e ( l a n c e o l a t e ) t o o v a t e - t r i a n g u l a r , d e e p l y c h a n n e l l e d above ( F i g . 1 7 ) ; m a r g i n s s t r o n g l y r e v o l u t e t o n e a r b a s e ( F i g . 17); c o s t a s t o u t , u s u a l l y e x c u r r e n t a s m u c r o n a t e or c u s p i d a t e t i p ; upper l a m i n a l c e l l s 10-12 m i c r o m e t e r s , q u a d r a t e t o r o u n d e d ( i r r e g u l a r ) t o t r a n s v e r s e l y - r e c t a n g u l a r , d e n s e l y p a p i l l o s e , o f t e n o b s c u r i n g t h e a r e o l a t i o n ; b a s a l c e l l s smooth, s h o r t - r e c t a n g u l a r t o r e c t a n g u l a r , upper b a s a l c e l l s w i t h c o r n e r t h i c k e n i n g s . S p o r o p h y t e s n o t c o l l e c t e d ( u n r e p o r t e d i n N o r t h A m e r i c a ) . R e p r e s e n t a t i v e s p e c i m e n s : B89048, B89049, B89050 I l l u s t r a t i o n s : F i g . 17, Lawton 1971, Tan e t a l . 1981 D i s t r i b u t i o n : A: w i d e s p r e a d and o f t e n uncommon l o c a l l y ( F i g . 82) B: m a i n l y s o u t h w e s t N o r t h A m e r i c a C: c i r c u m t e m p e r a t e 90 Remarks : M a c r o s c o p i c a l l y , P s e u d o c r o s s i d i u m r e v o l u t u m c a n be m i s t a k e n f o r C e r a t o d o n p u r p u r e u s , Desmatodon c o n v o l u t u s and, i n some c a s e s , Didymodon v i n e a l i s v a r . b r a c h y p h y l l u s . None o f t h e s e t a x a , however, have t h e s t r o n g l y r e v o l u t e l e a f m a r g i n s and t h e abun d a n t l a m i n a l p a p i l l a e t h a t c h a r a c t e r i z e P. r e v o l u t u m . O t h e r d i s t i n c t i v e c h a r a c t e r i s t i c s of P. r e v o l u t u m i n c l u d e t h e s t r o n g l y k e e l e d upper h a l f o f t h e l e a f a s w e l l a s a w e l l d e v e l o p e d a b a x i a l p o r t i o n of t h e c o s t a ( i n c r o s s - s e c t i o n ) w h i c h i s u s u a l l y w i d e r t h a n t h e upper s e c t i o n . The v a r i e t a l name, v a r . o b t u s u l u m ( L i n d b . ) T a n, Zander e t T a y l o r , s u g g e s t e d f o r t h e r e g i o n a l m a t e r i a l (Tan e t a l . 1981), i s n o t u s e d h e r e . The c h a r a c t e r s w h i c h t h e a u t h o r s employed t o d i s t i n g u i s h t h e v a r i e t y a r e t o o v a r i a b l e i n l o c a l m a t e r i a l t o be o f u s e . PTERYGONEURUM J u r . P t e r y g o n e u r u m i s c h a r a c t e r i z e d by c o n c a v e , l o n g awned l e a v e s , e a c h b e a r i n g two t o s i x l a m e l l a e on t h e upper p a r t o f t h e a d a x i a l c o s t a l s u r f a c e . The l o n g awns a r e r e a d i l y v i s i b l e i n t h e f i e l d a nd o f t e n g i v e t h e t y p i c a l l y compact c o l o n i e s a f u z z y a p p e a r a n c e . F u r t h e r m o r e , u n l i k e most o t h e r d r y l a n d t a x a , s p o r o p h y t e s a r e common, a i d i n g i n r a p i d and c o n s i s t e n t i d e n t i f i c a t i o n . F o u r s p e c i e s o f P t e r y g o n e u r u m a r e r e c o g n i z e d h e r e . P t e r y g o n e u r u m ovaturn and P. s u b s e s s i l e a r e common, whereas P. k o z l o v i i i s l e s s common, a l t h o u g h l o c a l l y v e r y a b u n d a n t , and r e s t r i c t e d t o t h e m a r g i n s of a l k a l i n e d e p r e s s i o n s . P t e r y g o n e u r u m 91 l a m e l l a t u m i s r a r e , f o u n d o n l y a t one s i t e , a l s o an a r e a of a l k a l i n e c o n d i t i o n s . P t e r y g o n e u r u m k o z l o v i i i s r e p o r t e d h e r e as new t o N o r t h A m e r i c a and P. l a m e l l a t u m as new t o B r i t i s h C o l u m b i a . P t e r y g o n e u r u m i s one o f t h e few b r y o p h y t e g e n e r a w i t h w h i c h d e t a i l e d e c o l o g i c a l - d i s t r i b u t i o n a l r e s e a r c h has been a t t e m p t e d . P o s p i s i l (1975) s t u d i e d t h e d i s t r i b u t i o n i n C z e c h o s l o v a k i a o f two s p e c i e s , P. ovatum and P. s u b s e s s i l e , and r e l a t e d t h e i r g e o g r a p h i c a l p a t t e r n s t o c l i m a t i c d a t a . He f o u n d P. s u b s e s s i l e t o be r e s t r i c t e d t o t h e warm and m o d e r a t e l y warm r e g i o n s of t h e c o u n t r y , whereas P. ovatum, a l t h o u g h i t a t t a i n s maximum f r e q u e n c y i n t h e s e r e g i o n s , shows a w i d e r r a n g e , f o u n d i n c o o l e r and somewhat w e t t e r a r e a s a s w e l l . In a d d i t i o n , P. ovatum, even i n t h e warmest r e g i o n s , i s more a b u n d a n t t h a n P. s u b s e s s i l e . The p r e s e n t s t u d y s u p p o r t s P o s p i s i l ' s o b s e r v a t i o n s . P t e r y g o n e u r u m  ovatum i s much more common t h a n P. s u b s e s s i l e t h r o u g h o u t t h e d r y s t e p p e , and a l s o grows i n open a r e a s a t h i g h e r e l e v a t i o n s where P. s u b s e s s i l e has n o t y e t been c o l l e c t e d . P o s p i s i l a l s o n o t e d t h a t P. s u b s e s s i l e e x h i b i t s a n a r r o w e r r a n g e o f pH t o l e r a n c e t h a n P. ovatum . A l t h o u g h i n t h e p r e s e n t s t u d y t h i s has n o t been e x a m i n e d , P. s u b s e s s i l e t e n d s t o f o r m l a r g e p o p u l a t i o n s o n l y i n a r e a s o f h i g h pH, s u c h a s f r o s t b o i l s o r l a c u s t r i n e b a n k s , whereas P. ovatum c a n form e x t e n s i v e p o p u l a t i o n s on a l m o s t any m i n e r a l s o i l s u b s t r a t u m i n t h e s t e p p e . Key t o t h e S p e c i e s o f PTERYGONEURUM 1 c a p s u l e s c l e a r l y e x s e r t e d w e l l above t h e l e a v e s , s t e g o c a r p o u s ; c a l y p t r a c u c u l l a t e and c o v e r i n g much o f t h e 92 c a p s u l e 2 1 1 c a p s u l e s immersed i n l e a v e s o r p a r t l y e x s e r t e d , s t e g o c a r p o u s o r c l e i s t o c a r p o u s , c a l y p t r a m i t r a t e and c o v e r i n g much o f t h e c a p s u l e o r t i n y and c o v e r i n g o n l y t h e t o p o f t h e c a p s u l e 3 2 l e a v e s smooth on t h e back; c a p s u l e s o v o i d t o e l l i p t i c , s h o r t e x s e r t e d ; p e r i s t o m e a b s e n t ; c e l l s of o p e r c u l u m i n v e r t i c a l rows; v e r y common P. ovaturn 2 1 l e a v e s o f t e n w e a k l y p a p i l l o s e on back; c a p s u l e s o b l o n g -c y l i n d r i c t o c y l i n d r i c , l o n g e x s e r t e d ; p e r i s t o m e sometimes p r e s e n t , t h i n and f r a g i l e ( o f t e n r e m a i n i n g i n t h e l i d ) ; c e l l s o f o p e r c u l u m i n o b l i q u e rows; r a r e P. l a m e l l a t u m 3 c a p s u l e s s t e g o c a r p o u s , p a r t i a l l y e x s e r t e d t o immersed; o p e r c u l u m r o s t r a t e ; l e a f awns u s u a l l y s e r r u l a t e ; l a m e l l a e u s u a l l y 4, >8 c e l l s h i g h P. s u b s e s s i l e 3 1 c a p s u l e s c l e i s t o c a r p o u s , immersed i n l e a v e s ; apex o f c a p s u l e w i t h o n l y a s h o r t a p i c u l u s ; l e a f awns smooth; l a m e l l a e u s u a l l y 2, u s u a l l y <8 c e l l s h i g h P. k o z l o v i i PTERYGONEURUM KOZLOVII L a z a r . P. smardaeanum Vanek S m a l l (2-3mm t a l l ) , more o r l e s s b u l b i f o r m p l a n t s , g r e e n t o y e l l o w - g r e e n when young, p a l e r and b r o w n i s h when m a t u r e , g r o w i n g i n t u f t s o r r a t h e r compact t u r f s among g r a s s e s a l o n g t h e m a r g i n s o f a l k a l i n e d e p r e s s i o n s ; b a s a l l e a v e s 0.75 t o 1.0mm l o n g , o v a t e -l a n c e o l a t e t o l a n c e o l a t e , c o n c a v e , u s u a l l y n a r r o w l y t a p e r i n g t o t h e apex; m a r g i n s e n t i r e o r s l i g h t l y t o o t h e d a t t h e apex, w e a k l y r e c u r v e d t o p l a n e ; c o s t a p a l e brown, i n c r o s s - s e c t i o n w i t h two g u i d e c e l l s a n d a b a x i a l l a y e r composed o f wide-lumened s t e r e i d s s u r r o u n d i n g a s m a l l e r a r e a o f n a r r o w l y lumened s t e r e i d s , B e g l e i t e r - l i k e c e l l s a r e p r e s e n t between t h e s t e r e i d s and t h e 93 g u i d e s ; a d a x i a l s u r f a c e composed o f 2-4 c e l l s f o r m i n g base of t h e l a m e l l a e ; l a m e l l a e u s u a l l y 2, r a r e l y 3 i n number, ( 2 ) 4 - 6 ( 8 ) c e l l s h i g h , i r r e g u l a r and u s u a l l y s t r o n g l y p a p i l l o s e ; c o s t a e x c u r r e n t as a h y a l i n e , smooth t o s l i g h t l y t o o t h e d awn; upper l a m i n a l c e l l s 10-15 x 15-25 m i c r o m e t e r s t o 15-20 x 35-45 m i c r o m e t e r s , s m ooth.to s l i g h t l y p a p i l l o s e , r a r e l y s t r o n g l y p a p i l l o s e , rhombic t o o b l o n g t o i r r e g u l a r l y q u a d r a t e o r r e c t a n g u l a r , u s u a l l y l o n g e r a nd n a r r o w e r a t t h e apex o f t h e l e a f , w eakly c o l l e n c h y m a t o u s ; b a s a l c e l l s l a r g e r , h y a l i n e , t h i n -w a l l e d and s h o r t - t o l o n g - r e c t a n g u l a r , a p p e a r i n g w e a k l y i n f l a t e d i n some l e a v e s ; upper and p e r i c h a e t i a l l e a v e s u s u a l l y t h r e e o r f o u r , s i m i l a r t o b a s a l l e a v e s , e x c e p t l o n g e r ( a b o u t 1.5mm l o n g ) , b ecoming p a l e r and d y i n g w i t h t h e m a t u r a t i o n o f t h e s p o r o p h y t e ; b u l b i l s p r e s e n t on t h e r h i z o i d s ; a u t o i c o u s . S p o r o p h y t e s common; c a p s u l e s l a r g e , 0.8mm t o 1.0mm a t m a t u r i t y , w i t h t i n y a p i c u l u s a t t h e apex; c a l y p t r a c u c u l l a t e , 1/3 t o 1/2 t h e l e n g t h o f t h e c a p s u l e ; most e x o t h e c i a l c e l l s r e c t a n g u l a r , w i t h a d i f f e r e n t i a t e d band o f o f s m a l l e r , q u a d r a t e c e l l s u s u a l l y n e a r t h e apex of t h e c a p s u l e ; s e t a s h o r t , <0.2mm; s p o r e s l a r g e , 30-45 m i c r o m e t e r s , w e a k l y o v a t e , o c c a s i o n a l l y s p h e r i c a l , r o u g h l y p a p i l l o s e . R e p r e s e n t a t i v e s p e c i m e n s : B89051, B89052, B89053 I l l u s t r a t i o n s : Vanek 1952 D i s t r i b u t i o n : A: s c a t t e r e d , b u t u s u a l l y l o c a l l y a b u n d a n t , m a i n l y i n t h e s o u t h Okanagan V a l l e y ( F i g . 84) B: s o u t h e r n B r i t i s h C o l u m b i a 94 C: w e s t e r n N o r t h A m e r i c a / w e s t e r n E u r a s i a ( s o u t h e r n C z e c h o s l o v a k i a , U k r a i n e ) ; new t o N o r t h A m e r i c a Remarks : In E u r o p e , l i t t l e i s known o f t h i s moss. I t a p p e a r s t o be r a t h e r w i d e s p r e a d t h e r e , a l t h o u g h i t i s u s u a l l y c o n s i d e r e d t o be a h y b r i d o f Phascum .cuspidatum and P t e r y q o n e u r u m ovatum ( B o r o s 1953, W i j k e t a l . 1959-1969), o r of P. c u s p i d a t u m and P t e r y g o n e u r u m s u b s e s s i l e ( C o r l e y e t a l . 1981). T h e r e i s no e x p e r i m e n t a l e v i d e n c e a v a i l a b l e t o s u p p o r t e i t h e r c l a i m . PTERYGONEURUM LAMELLATUM ( L i n d b . ) J u r . P. a r c t i c u m S t e e r e P l a n t s g r e e n t o p a l e g r e e n , s c a t t e r e d on c l a y - r i c h s o i l , u s u a l l y among g r a s s e s a t edge of s a l i n e d e p r e s s i o n s . S p o r o p h y t e s common. R e p r e s e n t a t i v e s p e c i m e n s : B89054, B85666, B85667 I l l u s t r a t i o n s : Wareham 1939, F l o w e r s 1973 D i s t r i b u t i o n : A: r a r e , S i t e 81 o n l y , W h i t e L a k e a r e a ( F i g . 85) B: a c r o s s a r c t i c ; s c a t t e r e d s o u t h w a r d : s o u t h e r n B r i t i s h C o l u m b i a , U t a h , A r i z o n a C: w e s t e r n N o r t h A m e r i c a ~ w e s t e r n E u r a s i a Remarks : P t e r y q o n e u r u m l a m e l l a t u m w o u l d p r o b a b l y have been o v e r l o o k e d i f i t h a d not been f o r a c o l l e c t i o n made i n 1962 by W.B. S c h o f i e l d i n t h e W h i t e L a k e a r e a ( l a b e l l e d a s P. o v a t u m ) . Upon f i n d i n g t h i s e a r l y c o l l e c t i o n , s p e c i a l a t t e n t i o n was g i v e n 95 t o t h e W h i t e L a k e a r e a ( S i t e 81) i n o r d e r t o f i n d more m a t e r i a l . Two s m a l l p o p u l a t i o n s were f o u n d , b o t h on c l a y r i c h f l a t s n e a r d e p r e s s i o n s . A l t h o u g h many d e p r e s s i o n a l f l a t s s i m i l a r t o t h o s e a t W h i t e Lake were c a r e f u l l y c h e c k e d l a t e r , P. l a m e l l a t u m has n o t been f o u n d e l s e w h e r e . Two o t h e r r a r e mosses, E n t o s t h o d o n  r u b i g i n o s u s , and P o t t i a n e v a d e n s i s , were f o u n d a l s o o n l y i n t h e W h i t e Lake a r e a . PTERYGONEURUM OVATUM (Hedw.) D i x . Uk P o t t i a c a v i f o l i u m E h r h . ex F u e r n r . S m a l l g r e y - g r e e n t o g r e e n p l a n t s , o f t e n a p p e a r i n g r e d d i s h -brown when d r y , b e c a u s e of c o l o u r a t i o n of d o r s a l c o s t a l s u r f a c e , s c a t t e r e d o r a s s m a l l c o l o n i e s ( s o m e t i m e s as e x t e n s i v e t u r f s i n a r e a s of i n f r e q u e n t d i s t u r b a n c e ) on s o i l , u s u a l l y i n sunny s i t e s . S p o r o p h y t e s v e r y common; c a p s u l e s red-brown, o f t e n w r i n k l e d ; p e r i s t o m e a b s e n t . R e p r e s e n t a t i v e s p e c i m e n s : B89055, B89056, B89057 I l l u s t r a t i o n s : F r o n t i s p i e c e , F i g . 13, Lawton 1971, F l o w e r s 1973, Crum and A n d e r s o n 1981 D i s t r i b u t i o n : A: common t h r o u g h o u t ( F i g . 86) B: s o u t h e r n B r i t i s h C o l u m b i a t o t h e p r a i r i e s , s o u t h t o M e x i c o ; r a r e i n t h e e a s t ; d i s j u n c t i n t h e A r c t i c C: c i r c u m t e m p e r a t e ; b i p o l a r ( A u s t r a l i a , New Z e a l a n d ) Remarks : F e r t i l e p o p u l a t i o n s o f P t e r y g o n e u r u m ovatum, one o f t h e most common b r y o p h y t e s i n t h e s t u d y r e g i o n , a r e e a s y t o 96 r e c o g n i z e , m a i n l y b e c a u s e of t h e abundance o f t h e d i s t i n c t r e d -brown c a p s u l e s . S t e r i l e c o l l e c t i o n s , a l t h o u g h uncommon, a r e not d e t e r m i n e d r e a d i l y s i n c e t h e y may be c o n f u s e d w i t h P. s u b s e s s i l e w i t h w h i c h P. ovatum i s o f t e n f o u n d . A l t h o u g h P. ovatum t e n d s t o have a smoother awn, i t c a n be s h a r p l y s e r r a t e , e s p e c i a l l y a t t h e u p p e r end, t h u s making i d e n t i f i c a t i o n o f s t e r i l e c o l l e c t i o n s u n c e r t a i n . P. ovatum i s n o t p a p i l l o s e , e x c e p t on t h e l a m e l l a e , b u t s t i l l c a n be c o n f u s e d w i t h P. l a m e l l a t u m w h i c h i s u s u a l l y w e a k l y p a p i l l o s e . E x a m i n a t i o n of t h e s o i l under t u r f s of P. ovatum shows t h a t r h i z o i d s a r e abu n d a n t and d e e p l y embedded i n t h e s o i l and a c t as b i n d i n g a g e n t s , t h u s r e d u c i n g a c c e l e r a t e d s o i l e r o s i o n . T h i s has been n o t e d a l s o by F l o w e r s ( 1 9 7 3 ) . PTERYGONEURUM SUBSESSILE ( B r i d . ) J u r . S i m i l a r t o P. ovatum, a l t h o u g h i t t e n d s n o t t o form l a r g e p a t c h e s e x c e p t on l a c u s t r i n e b a n k s . S p o r o p h y t e s a r e common, but a r e l e s s a p p a r e n t t h a n i n P. ovatum, n o t o n l y b e c a u s e t h e y a r e immersed b ut a l s o b e c a u s e t h e y t e n d t o become c o n t o r t e d a f t e r m a t u r i t y and a r e h i d d e n among t h e l e a v e s . R e p r e s e n t a t i v e s p e c i m e n s : B89058, B89059 I l l u s t r a t i o n s : F l o w e r s 1973, Lawton 1971, Crum and A n d e r s o n 1981 D i s t r i b u t i o n : A: w i d e s p r e a d , b u t u s u a l l y i n c o n s p i c u o u s ( F i g . 87) B: m a i n l y w e s t e r n N o r t h A m e r i c a : s o u t h e r n B r i t i s h C o l u m b i a and A l b e r t a s o u t h t o M e x i c o 97 C: c i r c u m t e m p e r a t e Remarks : P t e r y g o n e u r u m s u b s e s s i l e , a l t h o u g h u s u a l l y v e r y d i s t i n c t i v e , c a n be e a s i l y o v e r l o o k e d , b o t h b e c a u s e of i t s s m a l l s i z e and b e c a u s e i t i s u s u a l l y h i d d e n among o t h e r mosses, i n p a r t i c u l a r P. ovatum. On f l a t s a d j a c e n t t o a l k a l i n e d e p r e s s i o n s , a form of P. s u b s e s s i l e o c c a s i o n a l l y o c c u r s w h i c h d i f f e r s f r o m t y p i c a l m a t e r i a l i n h a v i n g a t i n y , c u c u l l a t e c a l y p t r a w h i c h c o v e r s o n l y t h e t o p p a r t o f t h e o p e r c u l u m , more o r l e s s s moother l e a f awn's and more d e e p l y immersed c a p s u l e s . A l s o , some c o l l e c t i o n s have smoother and s m a l l e r s p o r e s . A l l o f t h e s e m o r p h o l o g i c a l c h a r a c t e r s , a s w e l l a s t h e d i s t i n c t h a b i t a t ( t y p i c a l P. s u b s e s s i l e d o e s n o t grow i n a l k a l i n e c o n d i t i o n s ) , i n d i c a t e t h a t t h e u n u s u a l f o r m w a r r a n t s a t a x o n o m i c r a n k , p o s s i b l y a s a v a r i e t y . STEGONIA V e n t . S t e g o n i a i s c l o s e l y r e l a t e d t o P o t t i a , and i s o f t e n i n c l u d e d i n t h a t genus, p a r t i c u l a r l y by N o r t h A m e r i c a n a u t h o r s ( f o r example, F l o w e r s 1973 and Lawton 1971). I t i s s e g r e g a t e d h e r e f o l l o w i n g t h e r e a s o n i n g o f V e n t u r i ( 1 8 8 3 ) . STEGONIA L A T I F O L I A (Schwaegr.) V e n t . v a r . P I LIFERA ( B r i d . ) B r o t h . P o t t i a l a t i f o l i a v a r . p i l i f e r a ( B r i d . ) C . M u e l l . S m a l l g r e e n p l a n t s f o r m i n g open c o l o n i e s , m i x e d w i t h o t h e r 98 mosses o v e r s o i l . S p o r o p h y t e s o c c a s i o n a l , and immature. R e p r e s e n t a t i v e s p e c i m e n : B89060 I l l u s t r a t i o n s : F l o w e r s 1 973, Lawton 1971, Wareham 1939 D i s t r i b u t i o n : A: r a r e , S i t e 6 o n l y , n e a r B i g Bar n o r t h o f L i l l o o e t ( F i g . 92) B: m a i n l y n o r t h - w e s t e r n N o r t h A m e r i c a : t h e a r c t i c r e g i o n s s o u t h i n t o U t a h ; G r e e n l a n d C: more o r l e s s c i r c u m a r c t i c - a l p i n e Remarks : When s t e r i l e , t h e b u d - l i k e p l a n t s o f S t e g o n i a l a t i f o l i a v a r . p i l i f e r a ( t h e y r e s e m b l e s m a l l c a b b a g e s ) a r e s i m i l a r t o p l a n t s of T o r t u l a b r e v i p e s . However, when d r y , S t e g o n i a i s n o t t w i s t e d and i s g r e e n , u n l i k e t h e t w i s t e d and o f t e n r e d d i s h - b r o w n p l a n t s of t h e T o r t u l a . M i c r o s c o p i c a l l y , t h e upper l e a f c e l l s o f T. b r e v i p e s a r e s t r o n g l y p a p i l l o s e , i n c o n t r a s t t o t h e smooth c e l l s o f S t e g o n i a . The l o c a l c o l l e c t i o n has l e a v e s w i t h c o m p l e t e l y r e v o l u t e m a r g i n s , w h i c h have n e v e r been r e p o r t e d f o r e i t h e r v a r i e t y o f S t e g o n i a l a t i f o l i a . P o s s i b l y , i t s h o u l d be g i v e n v a r i e t a l s t a t u s , c o n s i d e r i n g t h a t t h e o n l y c o n s i s t e n t d i f f e r e n c e between t h e o t h e r two v a r i e t i e s i s t h e p r e s e n c e or a b s e n c e o f an awn. TORTULA Hedw. F o u r s p e c i e s o f T o r t u l a a r e r e c o g n i z e d i n t h e s t u d y r e g i o n : T. b r e v i p e s , T. c a n i n e r v i s (= T. b i s t r a t o s a F l o w e r s ) , T. i n t e r m e d i a and T. r u r a l i s . T h e s e t a x a a r e c h a r a c t e r i z e d by 99 p l u r i p a p i l l o s e , awned l e a v e s ( t h e awns a r e sometimes h i g h l y r e d u c e d i n T. i n t e r m e d i a and T. c a n i n e r v i s ) and t h e l o n g -c y l i n d r i c a l c a p s u l e s w i t h t a l l , w e l l d e v e l o p e d p e r i s t o m e s . Some w o r k e r s d i v i d e T o r t u l a i n t o two g e n e r a , T o r t u l a and S y n t r i c h i a , t h e l a t t e r genus d i f f e r i n g by h a v i n g a l o n g e r b a s a l p e r i s t o m e t u b e and an o v e r a l l more r o b u s t g r o w t h h a b i t (Agnew and V o n d r e c e k 1975, Chen 1941, Monkemeyer 1927). As n o t e d i n t h e f o l l o w i n g c h a p t e r , t h e r e i s m e r i t t o t h i s d i v i s i o n . However, b e c a u s e of t h e r e g i o n a l f a m i l i a r i t y w i t h t h e name T o r t u l a , as w e l l as t h e n e e d f o r more t a x o n o m i c r e s e a r c h , o n l y t h e s i n g l e g e n e r i c name i s u s e d h e r e . R e c e n t r e v i s i o n s of T o r t u l a i n c l u d e K ramer's s t u d y of T o r t u l a s e c t . R u r a l e s of t h e e a s t e r n H o l a r c t i c (Kramer 1980) and t h e s t u d i e s o f M i s c h l e r ( p e r s . comm.), w h i c h i n c l u d e a s u r v e y o f t h e M e x i c a n s p e c i e s o f T o r t u l a a s w e l l as a r e v i s i o n of T o r t u l a s e c t . R u r a l e s i n N o r t h A m e r i c a . Key t o t h e S p e c i e s of TORTULA 1 s m a l l p l a n t s , u s u a l l y w e l l u nder 0.5 cm. t a l l , g r o w i n g a s s o l i t a r y i n d i v i d u a l s o r s m a l l c o l o n i e s on c l a y - r i c h s o i l ; l e a f awn smooth; l e a f m a r g i n s r e v o l u t e ; s p o r o p h y t e s common T. b r e v i p e s 1 1 p l a n t s v a r i a b l e i n s i z e , b u t o f t e n r o b u s t , f o r m i n g mats o v e r s o i l o r o u t c r o p s , o f t e n i n l i t t e r ; l e a f awn t o o t h e d ; l e a f m a r g i n s r e c u r v e d t o p l a n e ; s p o r o p h y t e s r a r e .2 2 l e a v e s b i s t r a t o s e t o t r i s t r a t o s e , a t l e a s t i n p a r t , r a r e l y u n i s t r a t o s e and i f so t h e n l e a v e s o v a t e , w i t h b r o a d l y c u c u l l a t e a p i c e s T. c a n i n e r v i s 2 1 l e a v e s u n i s t r a t o s e ( r a r e l y b i s t r a t o s e p a t c h e s i n T. i n t e r m e d i a ) ; l e a v e s u s u a l l y s p a t h u l a t e , w i t h o u t c u c u l l a t e a p i c e s 3 100 3 l e a v e s s t r o n g l y t w i s t e d when d r y , s q u a r r o s e when wet, not c o n s p i c u o u s l y c o n s t r i c t e d i n t h e m i d d l e ; l e a f awn w e l l d e v e l o p e d on most l e a v e s ; u p p e r l e a f c e l l s u s u a l l y >12 m i c r o m e t e r s ; common i n most h a b i t a t s , o f t e n a s r o b u s t mats o r as s c a t t e r e d , s m a l l i n d i v i d u a l s T. r u r a l i s 3 1 l e a v e s l o o s e l y t w i s t e d when d r y , e r e c t t o w e a k l y r e c u r v e d when wet, o f t e n c o n s t r i c t e d a t m i d - l e a f ; awn s h o r t o r a b s e n t on many l e a v e s ; upper l e a f c e l l s m o s t l y <12 m i c r o m e t e r s ; common o n l y on o u t c r o p s , u s u a l l y a s s m a l l (<1cm t a l l ) , compact mats r e s e m b l i n g Grimmia s p p., o c c a s i o n a l l y a s r o b u s t p a t c h e s on s o i l a t h i g h e r e l e v a t i o n s T. i n t e r m e d i a TORTULA BREVIPES ( L e s q . ) B r o t h . P l a n t s , red-brown t o d a r k g r e e n and c r i s p e d when d r y , u s u a l l y b r i g h t g r e e n and w i t h s p r e a d i n g l e a v e s when wet; s c a t t e r e d on c l a y - r i c h s o i l a s i s o l a t e d i n d i v i d u a l s o r as s m a l l t u r f s , most common on l a c u s t r i n e t e r r a c e s , uncommon i n s t e p p e . S p o r o p h y t e s common, w i t h l o n g , d a r k r e d - b r o w n , somewhat c u r v e d c a p s u l e s ; p e r i s t o m e w e l l d e v e l o p e d , w i t h t a l l b a s a l t u b e . R e p r e s e n t a t i v e s p e c i m e n s : B89061, B89062 I l l u s t r a t i o n s : F l o w e r s 1973, Lawton 1971, S t e e r e 1 939b D i s t r i b u t i o n : A: t h r o u g h o u t , b u t r a r e l y a b u n d a n t l o c a l l y ( F i g . 93) B: w e s t e r n N o r t h A m e r i c a : s o u t h e r n B r i t i s h C o l u m b i a t o M e x i c o ; new t o Canada and B r i t i s h C o l u m b i a C: w e s t e r n N o r t h A m e r i c a n endemic Remarks : S t e r i l e p l a n t s o f T o r t u l a b r e v i p e s a r e i n c o n s p i c u o u s i n d r y c o n d i t i o n , n o t o n l y b e c a u s e o f t h e i r s m a l l s i z e and e a r t h y c o l o u r a t i o n , b u t a l s o b e c a u s e t h e y a r e o f t e n p a r t i a l l y b u r i e d by 10.1 f i n e s o i l o r h i d d e n among l e a v e s of n e i g h b o u r i n g mosses. F o r t u n a t e l y , s p o r o p h y t e s a r e common and h e l p i n l o c a t i n g p o p u l a t i o n s . A l s o , when wet, t h e s p r e a d i n g , b r i g h t g r e e n l e a v e s a r e p r o n o u n c e d . T o r t u l a b r e v i p e s c a n be c o n f u s e d w i t h o n l y a few o t h e r t a x a i n t h e s t u d y r e g i o n , i n p a r t i c u l a r S t e g o n i a l a t i f o l i a v a r . p i l i f e r a , Desmatodon spp. o r C r o s s i d i u m s p p . w i t h p o o r l y d e v e l o p e d c o s t a l f i l a m e n t s . T. b r e v i p e s has h y a l i n e awns and r e v o l u t e l e a f m a r g i n s wheras s p e c i e s o f C r o s s i d i u m u s u a l l y have c o l o u r e d awns and r e c u r v e d m a r g i n s . F u r t h e r m o r e , c a r e f u l o b s e r v a t i o n o f t h e a d a x i a l c o s t a l anatomy r e v e a l s t h a t T. b r e v i p e s has a more o r l e s s e v e n l y f u s e d r i d g e o f c e l l s , w h i c h C r o s s i d i u m n e v e r h a s . D i f f e r e n c e s between T. b r e v i p e s , t h e Desmatodon s p p . and S t e g o n i a , have been d i s c u s s e d e a r l i e r . T. b r e v i p e s i s m o r p h o l o g i c a l l y s i m i l a r t o T. m u r a l i s , a t a x o n common on r o c k s u r f a c e s ( p a r t i c u l a r l y w a l l s ) i n c o a s t a l a r e a s o f B r i t i s h C o l u m b i a . B. M i s h l e r ( p e r s . comm.) s u g g e s t e d t h a t t h e two t a x a a r e c o n s p e c i f i c . TORTULA CANINERVIS ( M i t t . ) B r o t h . T. b i s t r a t o s a F l o w e r s T. d e s e r t o r u m B r o t h . P l a n t s b l a c k t o d a r k g r e e n , u s u a l l y a s s m a l l , open and p a r t i a l l y b u r i e d p o p u l a t i o n s , o c c a s i o n a l l y a s compact t u r f s on s o i l i n e x p o s e d s t e p p e and l a c u s t r i n e t e r r a c e s . S p o r o p h y t e s r a r e , c o l l e c t e d o n l y o n c e . R e p r e s e n t a t i v e s p e c i m e n s : B89063, B89064 102 I l l u s t r a t i o n s : F l o w e r s 1951, 1973; Kramer 1980 D i s t r i b u t i o n : A: f a i r l y w i d e s p r e a d , o c c a s i o n a l l y l o c a l l y a b u n d a n t ( F i g . 94) B: w e s t e r n N o r t h A m e r i c a : s o u t h e r n B r i t i s h C o l u m b i a t o M e x i c o ; d i s j u n c t p o p u l a t i o n s i n t h e a r c t i c ; new t o B r i t i s h C o l u m b i a C: c i r c u m t e m p e r a t e Remarks : A l o n g w i t h t h e key c h a r a c t e r s , T o r t u l a c a n i n e r v i s i s d i s t i n g u i s h e d f r o m T. r u r a l i s , w i t h w h i c h i t i s e a s i l y c o n f u s e d , by t h e l e s s t w i s t e d , s u b - e r e c t (when d r y ) , b l u n t and c u c u l l a t e l e a v e s and i t s r e d u c e d s i z e ( o n l y r a r e l y does T. c a n i n e r v i s grow t a l l e r t h a n 1.0 cm.). O c c a s i o n a l l y , some l e a v e s o f T. c a n i n e r v i s a r e u n i s t r a t o s e . E x a m i n a t i o n of o t h e r l e a v e s on t h e same p l a n t u s u a l l y r e v e a l s t h e b i s t r a t o s e c h a r a c t e r , a l t h o u g h some c o l l e c t i o n s , c o m p l e t e l y u n i s t r a t o s e and i n t e r m e d i a t e i n m o r p h o l o g y , have been l e f t unnamed. Kramer (1980) d e s c r i b e d T. c a n i n e r v i s s s p . s p u r i a (Amann) W. Kramer as u n i s t r a t o s e but n o t e d t h a t a number of c h a r a c t e r s i n c l u d i n g t h e o v a t e l e a v e s and t h e d u m b - b e l l s h a p e d l a m i n a l p a p i l l a e d i s t i n g u i s h i t from s i m i l a r t a x a . To d a t e , I have been u n a b l e t o f o l l o w Kramer's k e y s i n d e t e r m i n i n g t h e unnamed c o l l e c t i o n s . TORTULA INTERMEDIA ( B r i d . ) De N o t . T. r u r a l i s v a r . c r i n i t a De N o t . P l a n t s o f t e n f o r m i n g d a r k g r e e n mats, o f t e n a s s c a t t e r e d 103 i n d i v i d u a l s g r o w i n g i n o t h e r moss mats, on r o c k , o r , r a r e l y on s o i l ; i n s h a d e d s i t u a t i o n s o r h i g h e l e v a t i o n s t e p p e . S p o r o p h y t e s n o t s e e n . R e p r e s e n t a t i v e s p e c i m e n s : B89065, B89066 I l l u s t r a t i o n s : F l o w e r s 1973, Kramer 1980, S t e e r e 1939b D i s t r i b u t i o n : A: f a i r l y w i d e s p r e a d , but uncommon l o c a l l y ( F i g . 95) B: s c a t t e r e d a c r o s s t h e s o u t h e r n h a l f of t h e c o n t i n e n t C: c i r c u m t e m p e r a t e Remarks : A l o n g w i t h t h e key c h a r a c t e r s , a number o f a d d i t i o n a l f e a t u r e s h e l p s e p a r a t e T o r t u l a i n t e r m e d i a from T. r u r a l i s . T y p i c a l l y , T. i n t e r m e d i a d i f f e r s i n h a v i n g t h e c e l l s i n t h e upper h a l f o f t h e l e a f f o r m i n g more o r l e s s e v e n , d i a g o n a l rows r a d i a t i n g f r o m t h e c o s t a , more p l a n e u p p e r l e a f m a r g i n s and a more e m a r g i n a t e l e a f a pex. F u r t h e r m o r e , T. i n t e r m e d i a i s o f t e n more compact and much s m a l l e r t h a n T. r u r a l i s . A few c o l l e c t i o n s i n c l u d e b o t h t a x a g r o w i n g t o g e t h e r . The r e s e m b l a n c e o f T. i n t e r m e d i a t o T. r u r a l i s has l e d t o i t s f r e q u e n t p l a c e m e n t i n synonomy w i t h T. r u r a l i s (Lawton 1971; Crum and A n d e r s o n 1981; B. M i s h l e r , p e r s . comm.). B e c a u s e o f t h e d i f f e r e n c e s n o t e d above, however, I am c o n v i n c e d t h a t i t i s d i s t i n c t enough t o be r e t a i n e d as a s e p a r a t e s p e c i e s and t h u s s u p p o r t t h e d e c i s i o n s o f S t e e r e (1939b, 1940), F l o w e r s ( 1 9 7 3 ) , S m i t h (1978a) and Kramer ( 1 9 8 0 ) . 104 TORTULA RURALIS (Hedw.) G a e r t n . T. p a p i l l o s i s s i m a (Copp.) B r o t h . T. r u r a l i f o r m i s ( B e s c h . ) I n g h . P l a n t s t y p i c a l l y d a r k g r e e n t o r e d d i s h - b r o w n o r b l a c k when d r y , l i g h t e r g r e e n when wet; v e r y common and v a r i a b l e , e i t h e r i n s m a l l t u f t s o r a s s c a t t e r e d i n d i v i d u a l s ( l e s s t h a n 0.5 cm t a l l ) , o r , i n more p r o t e c t e d s i t e s , as l a r g e , l o o s e mats; o v e r s o i l o r r o c k , o f t e n i n l i t t e r . S p o r o p h y t e s o c c a s i o n a l on l a r g e r p l a n t s . R e p r e s e n t a t i v e s p e c i m e n s : B89067, B89068, B89069 I l l u s t r a t i o n s : Crum and A n d e r s o n 1981, F l o w e r s 1973, Lawton 1 971 D i s t r i b u t i o n : A: w i d e s p r e a d and e x t r e m e l y common ( F i g . 96) B: a c r o s s t h e c o n t i n e n t , a l t h o u g h more common i n t h e west C: c i r c u m p o l a r ; b i p o l a r ( S o u t h A f r i c a , S o u t h A m e r i c a : P e r u and A r g e n t i n a ) Remarks : U n l i k e T o r t u l a i n t e r m e d i a , c o n s i s t e n t c h a r a c t e r s c o u l d n o t be f o u n d t o j u s t i f y r e c o g n i t i o n o f T. p a p i l l o s i s s i m a ( F l o w e r s 1953) or T. r u r a l i f o r m i s , even t h o u g h some c o l l e c t i o n s a p p r o a c h e d t h e d e s c r i p t i o n s o f t h e s e t a x a . T o r t u l a r u r a l i s i s e x t r e m e l y common. I t t e n d s t o be most abundant and r o b u s t i n s h a d e d , u n d i s t u r b e d s i t e s s u c h a s under A r t e m e s i a s h r u b s i n t h e s t e p p e o r o v e r o u t c r o p f a c e s i n more f o r e s t e d a r e a s . T o r t u l a r u r a l i s can be c o n f u s e d w i t h T. c a n i n e r v i s , T. i n t e r m e d i a and, a t l e a s t i n t h e f i e l d , P o l y t r i c h u m p i l i f e r u m 105 ( s e e t h e d i s c u s s i o n s a c c o m p a n y i n g t h e s e t a x a ) . TRICHOSTOMOPSIS C a r d . Zander (1978b, 1981b) i n c l u d e d T r i c h o s t o m o p s i s w i t h i n t h e c l o s e l y r e l a t e d genus Didymodon. However, t h e h i g h l y d i f f e r e n t i a t e d , h y a l i n e and t h i n - w a l l e d b a s a l c e l l s , t h e s t r o n g l y c r i s p e d l e a v e s and t h e u n u s u a l b l a c k i s h c o l o u r a t i o n ( Z a n d e r 1981b) a r e a number of d i s t i n c t i v e c h a r a c t e r s t h a t s u p p o r t t h e r e t e n t i o n o f T r i c h o s t o m o p s i s as a genus ( R o b i n s o n 1970). TRICHOSTOMOPSIS AUSTRALASIAE (Hook, e t G r e v . ) H. R o b i n s . T. f a y a e G r o u t T. umbrosa ( C . M u l l . ) H. R o b i n s . Didymodon a u s t r a l a s i a e (Hook, e t G r e v . ) Zander P l a n t s b l a c k t o d a r k g r e e n o r , o c c a s i o n a l l y , brown, r a r e l y l i g h t g r e e n , u s u a l l y s c a t t e r e d i n s m a l l , open c o l o n i e s on s o i l , most common i n s h a d e d m i c r o h a b i t a t s i n t h e s t e p p e ; l e a v e s s t r o n g l y c r i s p e d t o s u b - e r e c t when d r y , p l a n t s o f t e n r e s e m b l i n g s m a l l f i s t s on t h e s o i l . S p o r o p h y t e s n o t c o l l e c t e d . R e p r e s e n t a t i v e s p e c i m e n s : B89070, B89071 I l l u s t r a t i o n s : Lawton 1971, G r o u t 1928-1941, Zan d e r 1981b D i s t r i b u t i o n : A: w i d e s p r e a d , but i n c o n s p i c u o u s l o c a l l y ( F i g . 97) B: w e s t e r n N o r t h A m e r i c a : s o u t h e r n B r i t i s h C o l u m b i a t o M e x i c o ; new t o Canada, B r i t i s h C o l u m b i a and W a s h i n g t o n S t a t e 106 C: w e s t e r n N o r t h A m e r i c a / w e s t e r n E u r a s i a ; b i p o l a r ( A u s t r a l i a , New Z e a l a n d , S o u t h A f r i c a , S o u t h A m e r i c a ) Remarks : A l t h o u g h u s u a l l y d i s t i n c t i v e , s m a l l , l e s s c r i s p e d and l i g h t e r g r e e n p l a n t s o f T. a u s t r a l a s i a e can be c o n f u s e d m a c r o s c o p i c a l l y w i t h Didymodon t a x a . The t r i a n g u l a r t o l a n c e o l a t e l e a f shape, t h e d i s t i n c t i v e c o s t a l r i d g e i n c r o s s -s e c t i o n , t h e l a r g e r a v e r a g e c e l l s i z e ( u s u a l l y >12 v s . <10 m i c r o m e t e r s ) and t h e t h i n - w a l l e d b a s a l c e l l s , a r e f e a t u r e s w h i c h w i l l s e p a r a t e T r i c h o s t o m o p s i s f r o m Didymodon. WEISSIA Hedw. WEISSIA BRACHYCARPA C M . W. h e d w i g i i Crum W. m i c r o s t o m a (Hedw.) C M . P l a n t s g r e e n , a n d s t r o n g l y c r i s p e d , a p p e a r i n g f i s t - l i k e , i n s m a l l open c o l o n i e s on s o i l , u s u a l l y i n shaded r e c e s s e s n e a r b o u l d e r s , common on b o u l d e r s l o p e s i n P i n u s p o n d e r o s a - s t e p p e t r a n s i t i o n a l a r e a s , r a r e i n s t e p p e . S p o r o p h y t e s o c c a s i o n a l ; c a p s u l e s o v a t e , s h o r t e x s e r t e d . R e p r e s e n t a t i v e s p e c i m e n s : B89072, B89073 I l l u s t r a t i o n s : Crum and A n d e r s o n 1981 D i s t r i b u t i o n : A: o c c a s i o n a l , p r i m a r i l y a l o n g t h e p e r i p h e r y o f a r e a ( F i g . 98) B: s c a t t e r e d : o c c a s i o n a l i n e a s t , New B r u n s w i c k s o u t h t o N. C a r o l i n a , west t o I n d i a n a ; i n t h e west: T e x a s and s o u t h e r n 107 B r i t i s h C o l u m b i a ; new t o B r i t i s h C o l u m b i a C: p r o b a b l y c i r c u m t e m p e r a t e Remarks : W e i s s i a b r a c h y c a r p a d i f f e r s f r o m o t h e r N o r t h A m e r i c a n s p e c i e s of W e i s s i a i n h a v i n g s h o r t - e x s e r t e d c a p s u l e s w h i c h a r e c o n s t r i c t e d a t t h e mouth and w i t h t h e mouth c o v e r e d by an e v a n e s c e n t membrane. The o n l y o t h e r l o c a l t a x o n w h i c h forms f i s t - l i k e s t r u c t u r e s when d r y i s T r i c h o s t o m o p s i s a u s t r a l a s i a e , b ut i t i s u s u a l l y b l a c k i s h i n c o l o u r and u s u a l l y f o u n d o n l y i n t h e s t e p p e . M i c r o s c o p i c a l l y , W. b r a c h y c a r p a d i f f e r s i n b e i n g s t r o n g l y p a p i l l o s e and h a v i n g i n c u r v e d l e a v e s . GRIMMIACEAE The f a m i l y G r i m m i a c e a e i s r e p r e s e n t e d i n t h e s t u d y a r e a by t h r e e g e n e r a : C o s e i n o d o n , Grimmia and S c h i s t i d i u m . R e p r e s e n t a t i v e s of two o t h e r g e n e r a have been f o u n d i n a r e a s a d j a c e n t t o t h e s t u d y a r e a b u t a r e n o t i n c l u d e d : R a c o m i t r i u m  c a n e s c e n s (Hedw.) B r i d . , f o u n d i n a p i n e s t a n d n o r t h of V e r n o n and a l o n g V a s e a u x C r e e k i n t h e s o u t h Okanagan and S c o u l e r i a  a q u a t i c a Hook., w h i c h i s common on wet, u s u a l l y s h a d e d r o c k a l o n g t h e F r a s e r and Thompson R i v e r s s o u t h o f A s h c r o f t and L i l l o o e t . O c c a s i o n a l l y , C o s c i n o d o n and S c h i s t i d i u m a r e i n c l u d e d i n Grimmia (Crum and A n d e r s o n 1981, F l o w e r s 1973, Lawton 1971). They a r e k e p t s e p a r a t e h e r e f o l l o w i n g t h e t r e a t m e n t s o f D e g u c h i (1978) and C h u r c h i l l ( 1 9 8 1 ) . 108 COSCINODON S p r e n g . C o s e i n o d o n i s c h a r a c t e r i z e d by c o m p a r a t i v e l y r o b u s t p l a n t s , w i t h l a r g e - c a m p a n u l a t e and p l i c a t e c a l y p t r a e . Key t o t h e S p e c i e s o f COSCINODON I l e a v e s p l i c a t e a bove; c a p s u l e s s h o r t - o v o i d , immersed o r emmergent; p e r i s t o m e o r n a t e - c r i b r o s e ; r a r e C. c r i b r o s u s I I l e a v e s n o t p l i c a t e . a b o v e ; c a p s u l e s c y l i n d r i c a l , e x s e r t e d ; p e r i s t o m e n o t o r n a t e - c r i b r o s e ; v e r y common C. c a l y p t r a t u s COSCINODON CALYPTRATUS (Hook.) C. J e n s . Grimmia c a l y p t r a t a Hook. P l a n t s d a r k g r e e n t o g r e e n , u s u a l l y h o a r y w i t h l o n g l e a f awns, g r o w i n g a s l o o s e c u s h i o n s on b o u l d e r s and, l e s s commonly, o u t c r o p s . S p o r o p h y t e s common, d a r k ( r e d d i s h - ) brown when young, p a l e when o l d o r d r y . R e p r e s e n t a t i v e s p e c i m e n s : B89074, B89075 I l l u s t r a t i o n s : F l o w e r s 1973, Lawton 1971 D i s t r i b u t i o n : A: w i d e s p r e a d , o f t e n l o c a l l y a b u n d a n t ( F i g . 35) B: w e s t e r n N o r t h A m e r i c a : A l a s k a s o u t h t o C a l i f o r n i a and i n l a n d t o A l b e r t a and C o l o r a d o C: w e s t e r n N o r t h A m e r i c a n endemic Remarks : Cose i n o d o n c a l y p t r a t u s i s a v e r y common moss, a l t h o u g h i t i s f a r more a b u n d a n t on b o u l d e r s t h a n on more e x t e n s i v e r o c k o u t c r o p f a c e s , h a b i t a t s o f t e n d o m i n a t e d by s p e c i e s o f Grimmia, 109 i n p a r t i c u l a r G. a l p e s t r i s , G. l a e v i g a t a and G. o v a l i s . In t h e f i e l d , i t s h o a r y a p p e a r a n c e , a b u n d a n t , s t r a w c o l o u r e d s p o r o p h y t e s c a p p e d by l a r g e , p l i c a t e c a l y p t r a e and t h e c u s h i o n -f o r m i n g g r o w t h h a b i t a r e u s u a l l y d i s t i n c t i v e . Some s t e r i l e c o l l e c t i o n s c a n be c o n f u s e d w i t h G. p u l v i n a t a w h i c h has a s i m i l a r g r o w t h h a b i t and c o l o u r and i s e q u a l l y h o a r y . F u r t h e r m o r e , t h e l e a v e s of b o t h t a x a a r e u s u a l l y o v a t e -l a n c e o l a t e and have r e c u r v e d m a r g i n s . C o s e i n o d o n c a l y p t r a t u s , however, has a l e a f awn w h i c h t a p e r s f r o m a b r o a d base t o t h e apex, whereas G. p u l v i n a t a has a l e a f awn w h i c h t a p e r s o n l y s l i g h t l y from b a s e t o apex, and C. c a l y p t r a t u s has t h i n - w a l l e d b a s a l l e a f c e l l s i n c o n t r a s t t o t h e t h i c k - w a l l e d c e l l s o f G. p u l v i n a t a . The l a r g e and d i s t i n c t i v e s p o r o p h y t e s a s w e l l a s t h e l o n g awned, m o s t l y u n i s t r a t o s e l e a v e s s h o u l d r e a d i l y s e p a r a t e C. c a l y p t r a t u s from Grimmia t a x a . COSCINODON CRIBROSUS (Hedw.) S p r u c e Grimmia c r i b r o s a Hedw. P l a n t s d a r k g r e e n t o g r e e n , u s u a l l y h o a r y , g r o w i n g as l o o s e c u s h i o n s on b o u l d e r s . S p o r o p h y t e s common; c a p s u l e s immersed t o e m e r g e n t . R e p r e s e n t a t i v e s p e c i m e n s : B89076, B89077 I l l u s t r a t i o n s : Crum and A n d e r s o n 1981, D e g u c h i 1978, S m i t h 1978a, J o n e s 1933 D i s t r i b u t i o n : A: r a r e , 2 s i t e s : s i t e 45, s o u t h s i d e o f Kamloops; s i t e 106, e a s t o f Keremeos, s o u t h s i d e o f S i m i l k a m e e n R i v e r 110 ( F i g . 36) B: w i d e s p r e a d b u t e x t r e m e l y s c a t t e r e d : A l a s k a , E l l e s m e r e I s l a n d , B r i t i s h C o l u m b i a , A r i z o n a i n t h e west, i n t h e e a s t O n t a r i o ( T h u n d e r Bay D i s t r i c t ) , Quebec (Gaspe) s o u t h t o N. C a r o l i n a and T e n n e s s e e ; G r e e n l a n d ( s e e I r e l a n d and Crum 1976) C: c i r c u m p o l a r Remarks •: C o s c i n o d o n c r i b r o s u s i s a d i s t i n c t i v e moss. The p l i c a t e u p p e r l e a f l a m i n a e a r e u n l i k e t h o s e o f any o t h e r b r y o p h y t e i n t h e s t u d y a r e a . The immersed c a p s u l e s g i v e t h e p l a n t s t h e a p p e a r a n c e of G. anodon, b u t t h e l e a f c h a r a c t e r s , a l o n g w i t h t h e n o n - v e n t r i c o s e c a p s u l e s and s t r a i g h t s e t a e , r e a d i l y s e p a r a t e i t from t h e l a t t e r t a x o n . GRIMMIA Hedw. A l t h o u g h o n l y Grimmia i n t h e b r o a d s e n s e i s u s e d h e r e , t h e r e i s m e r i t i n t h e r e c o g n i t i o n of t h r e e g e n e r a (G r i m m i a, G u e m b e l i a and Rhabdogrimmia) i n s t e a d o f one. However, f o r t h e r e a s o n s n o t e d i n t h e f o l l o w i n g c h a p t e r , t h e s i n g l e genus i s u s e d h e r e . T r a d i t i o n a l l y , G. montana and G. t e n e r r i m a a r e r e c o g n i z e d a s s p e c i e s , b u t h e r e a r e i n c l u d e d i n G. a l p e s t r i s . Key t o t h e S p e c i e s of GRIMMIA I uppe r l e a f l a m i n a b i - t o t r i - s t r a t o s e , r a r e l y one c e l l t h i c k i n s p o t s ; l e a f m a r g i n i n r o l l e d t o p l a n e , u s u a l l y t h e same t h i c k n e s s a s t h e l a m i n a ; c a p s u l e s uncommon, smooth t o i r r e g u l a r l y w r i n k l e d , e x s e r t e d on s t r a i g h t s e t a e 2 I I u p p e r l a m i n a u n i s t r a t o s e t o , r a r e l y , b i s t r a t o s e i n s p o t s ; 111 l e a f m a r g i n p l a n e t o w e a k l y r e c u r v e d , o f t e n b i - s t r a t o s e ; c a p s u l e s common ( e x c e p t i n G. t r i c h o p h y l l a ) , immersed and smooth t o w r i n k l e d or e x s e r t e d on a r c u a t e s e t a e and o f t e n l o n g i t u d i n a l l y r i b b e d 4 l e a v e s b r o a d l y o v a t e - l a n c e o l a t e , u s u a l l y e v e n l y t a p e r e d f r o m b a s e t o apex ( i e . l o n g - t r i a n g u l a r ) ; b a c k s o f l e a v e s r a r e l y w i t h d i s t i n c t c o s t a l r i d g e , u s u a l l y a l m o s t f l a t ; most l e a v e s l y i n g f l a t on a s l i d e ; b a s a l c e l l s t o w a r d s t h e m a r g i n u s u a l l y w i d e r t h a n l o n g and t h i c k e n e d a t t h e c o r n e r s G. l a e v i g a t a ( s e e a l s o S c h i s t i d i u m h e t e r o p h y l l u m T l e a v e s o v a t e - l a n c e o l a t e t o l a n c e o l a t e , t a p e r e d f r o m an o v a t e b a s e ; c o s t a f o r m i n g a d i s t i n c t r i d g e on t h e back o f t h e l e a f ( e x c e p t i n G. o v a l i s w h i c h i s sometimes a l m o s t f l a t ) ; many l e a v e s l y i n g on t h e i r s i d e s on a s l i d e ; b a s a l c e l l s t o w a r d m a r g i n s u s u a l l y l o n g e r t h a n wide and w i t h c r o s s w a l l s t h i c k e r t h a n l o n g i t u d i n a l w a l l s , r a r e l y w i t h t h i c k e n e d c o r n e r s 3 p l a n t s i n l o o s e t u f t s or mats; many l e a v e s e r e c t - p a t e n t when wet and o v e r 2mm l o n g ; b a s a l c e l l s n e a r t h e c o s t a e l o n g a t e ( u s u a l l y > 4:1) and u n e v e n l y t h i c k - w a l l e d G. o v a l i s p l a n t s u s u a l l y i n t i g h t , compact c u s h i o n s ; most l e a v e s e r e c t t o w e a k l y s p r e a d i n g when wet and l e s s t h a n 1.5mm l o n g ; b a s a l c e l l s n e a r c o s t a u s u a l l y <4:1 and t h i n - t o t h i c k - w a l l e d , r a r e l y u n e v e n l y t h i c k e n e d G. a l p e s t r i s l e a v e s o v a t e - l a n c e o l a t e t o o b l o n g ; l e a f m a r g i n s u s u a l l y p l a n e o r w e a k l y r e c u r v e d ; m i d - l e a f c e l l s u s u a l l y e v e n l y t h i c k e n e d ; c a p s u l e s v e n t r i c o s e , g l o b o s e and smooth (sometimes w r i n k l e d when d r y ) , immersed 5 l e a v e s l a n c e o l a t e t o o v a t e - l a n c e o l a t e ; m a r g i n s u s u a l l y c o n s p i c u o u s l y r e c u r v e d ; m i d - l e a f c e l l s o f t e n u n e v e n l y t h i c k e n e d ; c a p s u l e s n ot v e n t r i c o s e , c y l i n d r i c a l and u s u a l l y l o n g i t u d i n a l l y r i b b e d , e x s e r t e d 6 l e a v e s o v a t e - l a n c e o l a t e , u s u a l l y p a r t i a l l y b i s t r a t o s e and u s u a l l y awned; upper c e l l s m o s t l y i s o d i a m e t r i c t o w e a k l y l o n g i t u d i n a l l y r e c t a n g u l a r , 8-10(12) m i c r o m e t e r s i n d i a m e t e r ; c a p s u l e s common, p e r i s t o m e a b s e n t ; g r o w i n g on r o c k , f a i r l y common G. anodon l e a v e s b r o a d l y o v a t e - l a n c e o l a t e t o o b l o n g , u n i s t r a t o s e o r , r a r e l y , b i s t r a t o s e i n s p o t s a l o n g t h e l e a f m a r g i n s , many l e a v e s a w n l e s s e s p e c i a l l y i n s m a l l e r p l a n t s ; u p p e r c e l l s ( o v a t e - ) r e c t a n g u l a r and >15 m i c r o m e t e r s l o n g ; c a p s u l e s o c c a s i o n a l , r a r e l y m a t u r e ; p e r i s t o m e p r e s e n t ; g r o w i n g on 112 l a c u s t r i n e s i l t b a n ks; r a r e G. p l a g i o p o d i a 6 l e a v e s somewhat c o n t o r t e d when d r y , l a n c e o l a t e ; awns s h o r t t o n e a r l y a b s e n t ; gemmae sometimes p r e s e n t on l e a v e s G. t r i c h o p h y l l a 6 1 l e a v e s e r e c t , n o t c o n t o r t e d when d r y , u s u a l l y o v a t e -l a n c e o l a t e ; awns l o n g ; gemmae a b s e n t G. p u l v i n a t a GRIMMIA ALPESTRIS (Web. e t Mohr) S c h l e i c h . ex. Nees. G. t e n e r r i m a Ren. e t C a r d . G. montana Hampe P l a n t s d a r k g r e e n (sometimes g r e e n ) t o a l m o s t b l a c k , sometimes g l a u c o u s - g r e y , o f t e n h o a r y from t h e awns, commonly a s compact, f i r m c u s h i o n s , o c c a s i o n a l l y a s s h o r t t u r f s , on r o c k s u r f a c e s , r a r e l y on h a r d - p a c k e d s o i l , i n sunny l o c a t i o n s . S p o r o p h y t e s o c c a s i o n a l ; c a p s u l e s e x s e r t e d and s h o r t - c y l i n d r i c a l . R e p r e s e n t a t i v e s p e c i m e n s : B89078, B89079 I l l u s t r a t i o n s : Crum and A n d e r s o n 1981, F l o w e r s 1 973, J o n e s 1933, Lawton 1971, S a v i c z - L j u b i t z k a j a and S m i r n o v a 1970, S m i t h 1 978a D i s t r i b u t i o n : A: w i d e s p r e a d and u s u a l l y common ( F i g . 56) B: m a i n l y w e s t e r n : s o u t h e r n B r i t i s h C o l u m b i a t o Wyoming, s o u t h t o C a l i f o r n i a ; s c a t t e r e d i n t h e e a s t : M i c h i g a n , New H a m p s h i r e , N e w f o u n d l a n d , G r e e n l a n d C: c i r c u m t e m p e r a t e Remarks : F o r r e a s o n s n o t e d i n t h e n e x t c h a p t e r , G. montana and G. 113 t e n e r r i m a a r e i n c l u d e d i n G. a l p e s t r i s . T h i s i s c o n t r a r y t o t h e o p i n i o n o f most o t h e r r e s e a r c h e r s . A l l c h a r a c t e r s u s e d by o t h e r w o r k e r s ( u p p e r l e a f c e l l s i z e , l e a f shape and l e n g t h , c o l o u r , c a p s u l e s i z e and shape and t h e s i z e o f t h e beak on t h e c a l y p t r a ) v a r i e d between and, o f t e n , w i t h i n most p o p u l a t i o n s i n t h e s t u d y a r e a . Grimmia a l p e s t r i s i s e a s i l y c o n f u s e d w i t h s m a l l p l a n t s o f G. o v a l i s , a l t h o u g h G. a 1 p e s t r i s n e v e r has t h e l o n g , t h i c k e n e d p a r a c o s t a l c e l l s i n d i c a t i v e o f t h e l a t t e r s p e c i e s . Grimmia  a l p e s t r i s n e v e r grows as l a r g e as G. o v a l i s and r a r e l y forms l o o s e t u r f s , w h i c h a r e t y p i c a l f o r G. o v a l i s . I n a d d i t i o n , G. a l p e s t r i s has v a r i a b l e but g e n e r a l l y l a r g e r u p p e r l e a f c e l l s t h a n G. o v a l i s (8-11 m i c r o m e t e r s compared t o a r o u n d 8 m i c r o m e t e r s f o r G. o v a l i s ) . GRIMMIA ANODON B.S.G. P l a n t s d a r k g r e e n , o f t e n w i t h a b r o w n i s h t i n g e , u s u a l l y a s s m a l l , f l a t c u s h i o n s o r t u r f s on r o c k . S p o r o p h y t e s common; c a p s u l e s g l o b o s e and immersed; s e t a e c u r v e d . R e p r e s e n t a t i v e s p e c i m e n s : B89080, B89081 I l l u s t r a t i o n s : Crum and A n d e r s o n 1981, F l o w e r s 1973, I r e l a n d 1982, Lawton 1971 D i s t r i b u t i o n : A: s c a t t e r e d , b u t f a i r l y common i n t r a n s i t i o n a l s i t e s ( F i g . 57) B: p r i m a r i l y i n w e s t e r n N o r t h A m e r i c a : A l a s k a t o t h e Yukon, s o u t h t o A r i z o n a and C a l i f o r n i a ; r a r e i n t h e e a s t : 1 1 4 I l l o n o i s , , O n t a r i o and G r e e n l a n d ( s e e I r e l a n d and M i l l e r 1982) C: c i r c u m p o l a r ; b i p o l a r : S o u t h A m e r i c a Remarks : Grimmia anodon c a n be c o n f u s e d w i t h S c h i s t i d i u m apocarpum, w i t h w h i c h i t o c c a s i o n a l l y grows. The l a t t e r s p e c i e s h a s , however, s m a l l e r , more c y l i n d r i c a l and p e r i s t o m a t e c a p s u l e s t h a t a r e n o t v e n t r i c o s e . F u r t h e r m o r e , t h e l e a v e s of G. anodon a r e o v a t e - l a n c e o l a t e , o f t e n l o n g awned, somewhat c o n c a v e and b o r d e r e d by p l a n e t o , r a r e l y , i n d i s t i n c t l y r e c u r v e d m a r g i n s . In c o n t r a s t , S. apocarpum has l a n c e o l a t e , s h o r t - a w n e d and u s u a l l y s t r o n g l y k e e l e d l e a v e s w i t h d i s t i n c t l y r e c u r v e d m a r g i n s . G. p l a g i o p o d i a a l s o c l o s e l y r e s e m b l e s G. anodon b ut s h o u l d be r e a d i l y s e p a r a t e d by t h e key c h a r a c t e r s . In t h e s t u d y a r e a , G. p l a g i o p o d i a was c o l l e c t e d o n l y on h a r d - p a c k e d l a c u s t r i n e s i l t s ( a l t h o u g h s a x i c o l o u s e l s e w h e r e ) and G. anodon was n o t . GRIMMIA LAEVIGATA ( B r i d . ) B r i d . G. l e u c o p h a e a G r e v . P l a n t s d a r k g r e e n t o b l a c k i s h , u s u a l l y f o r m i n g r e a d i l y f r i a b l e t u r f s o r , r a r e l y , l o w - g r o w i n g c u s h i o n s on r o c k . S p o r o p h y t e s r a r e ; c a p s u l e s e x s e r t e d . R e p r e s e n t a t i v e s p e c i m e n s : B89082, B89083 I l l u s t r a t i o n s : F l o w e r s 1973, Lawton 1971, S a v i c z - L j u b i t z k a j a and S m i r n o v a 1970, S m i t h 1978a D i s t r i b u t i o n : A: s c a t t e r e d , more common i n Okanagan V a l l e y , o f t e n l o c a l l y 115 ( F i g . 58) B: a c r o s s s o u t h e r n p a r t o f t h e c o n t i n e n t : B r i t i s h C o l u m b i a t o Quebec s o u t h t o New M e x i c o and F l o r i d a ; p o s s i b l y new t o B r i t i s h C o l u m b i a C: c i r c u m t e m p e r a t e ; b i p o l a r ( S o u t h A m e r i c a , S o u t h A f r i c a , New Z e a l a n d , A u s t r a l i a ) Remarks : Grimmia l a e v i g a t a i s d i s t i n c t i v e and s h o u l d not be c o n f u s e d w i t h any o t h e r moss e x c e p t f o r S c h i s t i d i u m h e t e r o p h y l l u m . I t i s o f t e n abundant on sunny o u t c r o p f a c e s and i s g e n e r a l l y s t e r i l e . F r u i t i n g m a t e r i a l w h i c h a p p e a r s t o be t h a t o f G. l a e v i g a t a i s u s u a l l y t h a t of G. o v a l i s , w i t h w h i c h i t f r e q u e n t l y grows. G. o v a l i s d i f f e r s f r o m G. l a e v i g a t a i n h a v i n g n a r r o w e r , e r e c t -p a t e n t l e a v e s , a n a r r o w l y c h a n n e l l e d u p p e r l e a f apex and l o n g m a r g i n a l b a s a l c e l l s . Once c o l l e c t e d , t h e t u r f s o f G. l a e v i g a t a r e a d i l y f r a g m e n t i n t o s m a l l clumps o r i n d i v i d u a l s t r a n d s . O f t e n t h e b a s e s o f t h e b r a n c h e s a r e a l m o s t b a r e , w i t h t h e l e a v e s m a i n l y clumped and b u d - l i k e on t h e u p p e r p a r t of t h e stem. I t i s s e p a r a t e d from S. h e t e r o p h y l l u m o n l y w i t h g r e a t d i f f i c u l t y s i n c e b o t h t a x a a r e u s u a l l y s t e r i l e ( i f f o u n d f r u i t i n g t h e n S. h e t e r o p h y l l u m has immersed c a p s u l e s ) . The l e a f a r e o l a t i o n i n b o t h s p e c i e s i s r e m a r k a b l y s i m i l a r , e s p e c i a l l y a l o n g t h e m a r g i n s o f t h e l e a f b ase where many c e l l s a r e t h i c k e n e d i n t h e c o r n e r s and a r e w i d e r t h a n l o n g , a d i s t i n c t c h a r a c t e r i s t i c o f o n l y t h e s e two grimmia'ceous t a x a i n t h e s t u d y . A l s o , b o t h s p e c i e s have t r i a n g u l a r , e v e n l y t a p e r e d l e a v e s , 1 1 6 u s u a l l y w i t h d e c u r r e n t awns. However, v e g e t a t i v e p l a n t s c a n be s e p a r a t e d by a few ambiguous c h a r a c t e r s : t h e a b a x i a l c o s t a l s u r f a c e of G. l a e v i g a t a i s u s u a l l y f l a t and r a r e l y r a i s e d above t h e l a m i n a l s u r f a c e , whereas S. h e t e r o p h y l l u m has a more t e r e t e and p r o m i n e n t l y r i d g e d c o s t a ; t h e l e a v e s of G. l a e v i g a t a a r e b r o a d e r and u s u a l l y d a r k e r g r e e n ( b l a c k i s h ) and a r e n o t as i n f o l d e d when d r y ; G. l a e v i g a t a t e n d s t o have a l a r g e r c e l l s i z e (7-9 m i c r o m e t e r s ) t h a n S. h e t e r o p h y l l u m (5-6 m i c r o m e t e r s ) ; and t h e l e a v e s a l o n g t h e stem of S. h e t e r o p h y l l u m t e n d t o be l a r g e r t h a n t h o s e o f G. l a e v i g a t u m , w h i c h u s u a l l y n a r r o w c o n s i d e r a b l y f r o m t h e apex o f t h e stem t o t h e b a s e . A l t h o u g h r e p o r t e d f r o m B r i t i s h C o l u m b i a ( I r e l a n d e t a l . 1980, J o n e s 1933) no e a r l i e r c o l l e c t i o n s c o u l d be l o c a t e d . I t i s p o s s i b l e t h a t t h e c o l l e c t i o n s f r o m t h e p r e s e n t s t u d y a r e t h e o n l y ones f o r t h e p r o v i n c e . GRIMMIA OVALIS (Hedw.) L i n d b . G. commutata Heub. P l a n t s d a r k g r e e n t o b l a c k i s h , u s u a l l y as l o o s e t o , l e s s commonly, compact t u r f s on r o c k i n sunny l o c a t i o n s . S p o r o p h y t e s r a r e ; c a p s u l e s e x s e r t e d , u s u a l l y o v a t e and r e l a t i v e l y l a r g e . R e p r e s e n t a t i v e s p e c i m e n s : B89084, B89085 I l l u s t r a t i o n s : F l o w e r s 1973, Lawton 1971, S a v i c z - L j u b i t z k a j a and S m i r n o v a 1970, S m i t h 1978a D i s t r i b u t i o n : A: f a i r l y w i d e s p r e a d , common i n s o u t h e r n p o r t i o n ( F i g . 59) B: w e s t e r n N o r t h A m e r i c a : B r i t i s h C o l u m b i a t o Montana s o u t h 117 t o A r i z o n a ; p r o b a b l y n o t i n t h e e a s t ( s e e t h e r e m a r k s ) C: c i r c u m t e m p e r a t e Remarks : S m a l l e r f o r m s o f Grimmia o v a l i s a r e e a s i l y c o n f u s e d w i t h G. a l p e s t r i s ( s e e t h e d i s c u s s i o n t h e r e ) . I t a l s o c a n be m i s t a k e n f o r G. l a e v i g a t a , w i t h w h i c h i t o f t e n grows. T h e r e i s some u n c e r t a i n t y c o n c e r n i n g t h e p r e s e n c e o f G. o v a l i s i n e a s t e r n N o r t h A m e r i c a . I r e l a n d (1982) and Crum and A n d e r s o n (1981) do not i n c l u d e i t , y e t i t i s r e p o r t e d i n I r e l a n d e t a l . (1980) and by J o n e s (1933) f r o m Quebec and by B r a s s a r d (1983) f r o m N e w f o u n d l a n d . I t a p p e a r s t h a t t h e e a s t e r n c o l l e c t i o n s r e f e r r e d t o by t h e l a t t e r r e s e a r c h e r s a r e m i s i d e n t i f i c a t i o n s o f G. a f f i n i s , a moss which c l o s e l y r e s e m b l e s G. o v a l i s and t h a t i s f r e q u e n t l y c o n f u s e d w i t h i t ( D e g u c h i 1978, S m i t h 1978a, Tan 1980). Grimmia a f f i n i s d i f f e r s i n h a v i n g r e c u r v e d l e a f m a r g i n s , e s p e c i a l l y i n t h e lower h a l f o f t h e l e a f , somewhat i r r e g u l a r l y t h i c k e n e d m i d - l e a f c e l l s , u s u a l l y l o n g e r and t h i n n e r - w a l l e d b a s a l m a r g i n a l c e l l s and i n b e i n g a u t o i c o u s ( c o l l e c t i o n s o f G. a f f i n i s u s u a l l y have abundant s p o r o p h y t e s , whereas G. o v a l i s p r o d u c e s them o n l y o c c a s i o n a l l y ) . A l l c o l l e c t i o n s o f G. o v a l i s t h a t I have examined from e a s t e r n Canada, i n c l u d i n g B r a s s a r d ' s c o l l e c t i o n s , have p r o v e d t o be G. a f f i n i s . Grimmia a f f i n i s i s r a r e l y c o l l e c t e d i n B r i t i s h C o l u m b i a (Tan 1980) and has not y e t been f o u n d n e a r t h e s t u d y a r e a . 118 GRIMMIA PLAGIOPODIA Hedw. P l a n t s b l a c k i s h - g r e e n , u s u a l l y a s low c u s h i o n s o r t u r f s on h a r d - p a c k e d l a c u s t r i n e s o i l s . S p o r o p h y t e s uncommon, o f t e n p o o r l y d e v e l o p e d and h i d d e n among the long-awned p e r i c h a e t i a l l e a v e s . R e p r e s e n t a t i v e s p e c i m e n s : B89086, B89087 I l l u s t r a t i o n s : Crum and A n d e r s o n 1981, F l o w e r s 1973, J o n e s 1933, Lawton 1971 D i s t r i b u t i o n : A: r a r e , 5 s i t e s o n l y : s i t e 35 e a s t o f Cache C r e e k ; s i t e s 48, 51 and 52 n e a r Kamloops; s i t e 68 a t Sage Mesa n e a r P e n t i c t o n ( F i g . 60) B: m a i n l y i n w e s t e r n N o r t h A m e r i c a : a r c t i c A l a s k a , B r i t i s h C o l u m b i a s o u t h t o C a l i f o r n i a ; s c a t t e r e d i n t h e mid-west and e a s t : N e b r a s k a , W i s c o n s i n , M i c h i g a n , O n t a r i o , O h i o , M a r y l a n d ; G r e e n l a n d C: c i r c u m t e m p e r a t e ; b i p o l a r ( S o u t h A m e r i c a , A n t a r c t i c a ) Remarks : Grimmia p l a g i o p o d i a i s one o f t h e most d i s t i n c t i v e o f a l l t h e l o c a l d r y l a n d G r i m m i a c e a e . Not o n l y i s i t s h a b i t a t u n u s u a l (on s o i l ) , b u t t h e wide, o f t e n o b l o n g and a w n l e s s v e g e t a t i v e l e a v e s and t h e u p p e r l e a f a r e o l a t i o n ( l a r g e o v a t e - r e c t a n g u l a r c e l l s ) a r e , i n c o m b i n a t i o n , c h a r a c t e r i s t i c o n l y o f G. p l a g i o p o d i a . In a d d i t i o n , G. p l a g i o p o d i a has a u n i q u e p y r a m i d a l g r o w t h h a b i t , c r e a t e d by t h e s t r o n g l y i m b r i c a t e , c o n c a v e l e a v e s , f o r m i n g t h r e e - s i d e d b u d - l i k e p l a n t s , a f e a t u r e e s p e c i a l l y e v i d e n t i n young p l a n t s . The c l o s e l y r e l a t e d G. anodon i s t h e o n l y o t h e r moss w h i c h c a n d e v e l o p t h e s e 119 " p y r a m i d s " b u t i t d o e s so o n l y o c c a s i o n a l l y , and t h e y a r e n e v e r a s w e l l d e v e l o p e d as w i t h G. p l a q i o p o d i a . The p r e d o m i n a t e l y u n i s t r a t o s e l e a v e s , a l o n g w i t h t h e wide b a s e d and somewhat d e c u r r e n t awns, s h o u l d s e p a r a t e i t from s t e r i l e c o l l e c t i o n s of G. anodon o r S c h i s . t i d i u m apocarpum. G. p l a q i o p o d i a was c u l t u r e d s u c c e s s f u l l y on a, r o o f t o p a t U.B.C. d u r i n g t h e y e a r s 1980-1983. Even tho u g h s p o r o p h y t e s ( a l w a y s p o o r l y d e v e l o p e d ) were r a r e l y c o l l e c t e d i n t h e f i e l d , a b u n d a n t , w e l l d e v e l o p e d s p o r o p h y t e s were p r o d u c e d i n t h e c u l t u r e s , p o s s i b l y b e c a u s e of t h e i n c r e a s e d m o i s t u r e a v a i l a b l e . The c u l t u r e d s p o r o p h y t e s s u b s t a n t i a t e d t h e i d e n t i t y o f G. p l a q i o p o d i a . GRIMMIA PULVINATA (Hedw.) Sm. P l a n t s g r e e n t o d a r k g r e e n , f o r m i n g h o a r y c u s h i o n s on r o c k . S p o r o p h y t e s common; c a p s u l e s e x s e r t e d and r i b b e d ; o p e r c u l u m w i t h l o n g r o s t r a t e beak; c a l y p t r a m i t r a t e . R e p r e s e n t a t i v e s p e c i m e n s : B89088, B89089 I l l u s t r a t i o n s : Crum and A n d e r s o n 1981, F l o w e r s 1973, J o n e s 1933, Lawton 1971, S m i t h 1978a D i s t r i b u t i o n : A: o c c a s i o n a l , m a i n l y i n t h e Okanagan V a l l e y ( F i g . 61) B: p r i n c i p a l l y w e s t e r n : B r i t i s h C o l u m b i a s o u t h t o M e x i c o , i n l a n d t o U t a h ; s c a t t e r e d e a s t w a r d : M i s s o u r i , Iowa, Oklahoma, K a n s a s , I n d i a n a , I l l i n o i s , O n t a r i o , M i c h i g a n C: c i r c u m t e m p e r a t e ; b i p o l a r ( S o u t h A m e r i c a , A u s t r a l i a , New Z e a l a n d , S o u t h A f r i c a ) 120 Remarks : Grimmia p u l v i n a t a , a l t h o u g h l o c a l l y a b u n d a n t , has a narrow ra n g e i n t h e s t u d y a r e a , where i t i s f o u n d i n t h e more m e s i c , t r a n s i t i o n a l s i t e s . The r i b b e d c a p s u l e s and c u r v e d s e t a e ( e s p e c i a l l y when wet) r e a d i l y s e p a r a t e i t from o t h e r g r i m m i a c e o u s t a x a e x c e p t G. t r i c h o p h y l l a ( s e e t h e key f o r d i f f e r e n c e s ) . V e g e t a t i v e l y , however, i t i s s i m i l a r i n l e a f shape and g e n e r a l h a b i t t o C o s e i n o d o n c a l y p t r a t u s . F o r t u n a t e l y b o t h s p e c i e s f r u i t f r e e l y (C. c a l y p t r a t u s has a l a r g e , p l i c a t e c a l y p t r a and a smooth c a p s u l e ) . F o r f u r t h e r d i f f e r e n c e s s e e t h e d i s c u s s i o n u nder C. c a l y p t r a t u s . GRIMMIA TRICHOPHYLLA G r e v . P l a n t s y e l l o w - g r e e n t o g r e e n , f o r m i n g l o o s e mats o r t u f t s on r o c k i n s h a d e d s i t e s . S p o r o p h y t e s n o t c o l l e c t e d . R e p r e s e n t a t i v e s p e c i m e n s : B89090, B89091 I l l u s t r a t i o n s : J o n e s 1933, Lawton 1971, S m i t h 1978a D i s t r i b u t i o n : A: r a r e ( F i g . 62) B: m a i n l y w e s t e r n N o r t h A m e r i c a : B r i t i s h C o l u m b i a and A l b e r t a s o u t h t o C a l i f o r n i a ; s t a t u s u n c e r t a i n i n e a s t : p r o b a b l y N e w f o u n d l a n d and p o s s i b l y Quebec C: c i r c u m t e m p e r a t e ; b i p o l a r ( S o u t h A m e r i c a , A u s t r a l i a , New Z e a l a n d ) Remarks : A l t h o u g h Grimmia t r i c h o p h y l l a i s r a r e i n t h e s t u d y a r e a , i t 121 i s common and a b u n d a n t on s h a l l o w s o i l o v e r o u t c r o p s a l o n g t h e C o l u m b i a R i v e r between Oregon and W a s h i n g t o n . I t i s a f a i r l y d i s t i n c t i v e moss and i s u n l i k e l y t o be c o n f u s e d w i t h o t h e r g r i m m i a c e o u s t a x a , m a i n l y b e c a u s e of i t s l i g h t c o l o u r , s l i g h t l y c o n t o r t e d l e a v e s and s i n u o u s l e a f a r e o l a t i o n . However, m a c r o s c o p i c a l l y , a t l e a s t i n x e r i c h a b i t a t s , i t s t r o n g l y r e s e m b l e s C e r a t o d o n p u r p u r e u s , but d i f f e r i n g i n h a v i n g w e a k l y r e c u r v e d l e a f m a r g i n s and h y a l i n e awns. SCHISTIDIUM B r i d . S c h i s t i d i u m i s c h a r a c t e r i z e d by immersed, n o n - v e n t r i c o s e c a p s u l e s w i t h w e l l d e v e l o p e d p e r i s t o m e s and c o l u m e l l a e t h a t r e m a i n a t t a c h e d t o t h e o p e r c u l a when t h e y f a l l . Two s p e c i e s , S. apocarpum and S. h e t e r o p h y l l u m , have been f o u n d i n t h e s t u d y r e g i o n . A r e c e n t s u r v e y by Bremer (1980a, 1980b, 1981) i s i m p o r t a n t i n t h e s t u d y o f S c h i s t i d i u m , a l t h o u g h she i n c l u d e s o n l y one s p e c i e s f r o m t h e s t u d y r e g i o n , S. apocarpum. S c h i s t i d i u m  h e t e r o p h y l l u m was o m i t t e d s i n c e she had n o t s e e n any m a t e r i a l of i t . The c o m b i n a t i o n , S c h i s t i d i u m h e t e r o p h y l l u m , i s new. Key t o t h e S p e c i e s o f SCHISTIDIUM 1 p l a n t s u s u a l l y i n compact t u f t s composed o f s h o r t b r a n c h e s ( a b o u t 1cm l o n g ) ; l e a v e s u n i s t r a t o s e t o i r r e g u l a r l y b i s t r a t o s e a b o v e ; m a r g i n s r e c u r v e d ; m a r g i n a l b a s a l c e l l s u s u a l l y l o n g e r t h a n wide S. apocarpum 122 1 1 p l a n t s u s u a l l y as l o o s e t u r f s composed of l o n g b r a n c h e s ( u s u a l l y much l o n g e r t h a n 1cm); l e a v e s b i s t r a t o s e a bove; m a r g i n s p l a n e , n o t r e c u r v e d ; many m a r g i n a l b a s a l c e l l s q u a d r a t e t o t r a n s v e r s e l y e l o n g a t e S. h e t e r o p h y l l u m SCHISTIDIUM APOCARPUM (Hedw.) B.S.G. S. apocarpum v a r . c o n f e r t u m ( F u n c k ) M o e l l e r Grimmia a p o c a r p a Hedw. G. c o n f e r t a Funck S. a t r o f u s c u m (Schimp.) L i m p r . P l a n t s g r e e n t o b l a c k i s h , o f t e n w i t h b r o w n i s h t i n g e , f o r m i n g l o o s e t o compact t u f t s o r s m a l l mats on s h a d e d o r , o c c a s i o n a l l y , sunny r o c k s u r f a c e s , c o l l e c t e d once on b u r n t wood. S p o r o p h y t e s common; c a p s u l e s immersed and o f t e n h i d d e n . R e p r e s e n t a t i v e s p e c i m e n s : B89092, B89093 I l l u s t r a t i o n s : Bremer 1980b, Crum and A n d e r s o n 1981, F l o w e r s 1973, Lawton 1971 D i s t r i b u t i o n : A: s c a t t e r e d , more common i n p e r i p h e r a l s i t e s ( F i g . 90) B: a c r o s s N o r t h A m e r i c a : A l a s k a t o G r e e n l a n d , s o u t h f r o m M e x i c o t o G e o r g i a C: c o s m o p o l i t a n Remarks : S c h i s t i d i u m apocarpum, a l t h o u g h h i g h l y v a r i a b l e , i s most l i k e l y t o be c o n f u s e d w i t h o n l y G. anodon o r G. p l a g i o p o d i a , two t a x a a l s o h a v i n g immersed c a p s u l e s ( s e e t h e d i s c u s s i o n s w i t h t h e s e t a x a ) . S. apocarpum i s c h a r a c t e r i z e d by a m y r i a d o f m o r p h o l o g i c a l 123 f o r m s , none of w h i c h a r e r e c o g n i z e d h e r e ( f o l l o w i n g Bremer 1980b). SCHISTIDIUM HETEROPHYLLUM ( K i n d b . ) M c i n t o s h comb. nov. i n ed. B a s i onym: Grimmia h e t e r o p h y l l a K i n d b . G. c o s e i n o d o n t o i d e s K i n d b . P l a n t s g r e e n t o d a r k ( b r o w n i s h ) g r e e n , u s u a l l y s h i n y , f o r m i n g l o o s e t u r f s ( r a r e l y a s low, compact t u f t s ) o v e r r o c k o u t c r o p s . S p o r o p h y t e s r a r e , d e e p l y h i d d e n i n l o n g awned p e r i c h a e t i a l l e a v e s . R e p r e s e n t a t i v e s p e c i m e n s : B89094, B89095, B89096 I l l u s t r a t i o n s : J o n e s 1933, Lawton 1971 D i s t r i b u t i o n : A: s c a t t e r e d t h r o u g h o u t ( F i g . 91) B: w e s t e r n N o r t h A m e r i c a : B r i t i s h C o l u m b i a , W a s h i n g t o n , Idaho, C a l i f o r n i a C: w e s t e r n N o r t h A m e r i c a n endemic Remarks : S c h i s t i d i u m h e t e r o p h y l l u m was an i n t e r e s t i n g d i s c o v e r y i n t h e l a t t e r s t a g e s o f t h e f i e l d r e s e a r c h . U n t i l t h a t t i m e , I had c o l l e c t e d i t o n l y i n s t e r i l e c o n d i t i o n , i d e n t i f y i n g i t as G. l a e v i g a t a b e c a u s e o f t h e r e m a r k a b l e v e g e t a t i v e s i m i l a r i t i e s o f t h e two t a x a . However, t h e immersed s p o r o p h y t e s and t h e c h a r a c t e r i s t i c c o l u m e l l a ( i t f a l l s w i t h t h e o p e r c u l u m ) r e a d i l y s e p a r a t e f e r t i l e c o l l e c t i o n s . T h e s e c h a r a c t e r s a l s o n e c e s s i t a t e d t h e change f r o m Grimmia t o S c h i s t i d i u m . 124 FUNARIACEAE W i t h few e x c e p t i o n s , t h e F u n a r i a c e a e i s c h a r a c t e r i z e d by l a r g e , l a x and t h i n - w a l l e d l a m i n a l c e l l s and by an a n n u a l or b i e n n i a l l i f e h i s t o r y . S p o r o p h y t e s a r e common and t h e c a p s u l e s a r e u s u a l l y p y r i f o r m . Two g e n e r a , E n t o s t h o d o n and F u n a r i a , a r e p r e s e n t i n t h e s t u d y a r e a . F u n a r i a i s c h a r a c t e r i z e d by i n c l i n e d a nd a s y m m e t r i c c a p s u l e s , whereas E n t o s t h o d o n has e r e c t and s y m m e t r i c a l c a p s u l e s . ENTOSTHODON Schwaegr. ENTOSTHODON RUBIGINOSUS ( W i l l i a m s ) G r o u t P l a n t s g r e e n t o p a l e g r e e n , t o re d - b r o w n , s c a t t e r e d o v e r s o i l a l o n g t h e m a r g i n o f an a l k a l i n e l a k e . S p o r o p h y t e s common; c a p s u l e s e r e c t - p y r i f o r m , s h r u n k e n below t h e mouth and w r i n k l e d . R e p r e s e n t a t i v e s p e c i m e n : B89097 I l l u s t r a t i o n s : Lawton 1971, W i l l i a m s 1913 D i s t r i b u t i o n : A: r a r e , s i t e 81 o n l y , W h i t e L a k e a r e a ( F i g . 52) B: s o u t h e r n B r i t i s h C o l u m b i a , Montana, A r i z o n a , New M e x i c o C: w e s t e r n N o r t h A m e r i c a n endemic Remarks : E n t o s t h o d o n r u b i g i n o s u s i s d i s t i n c t from most o t h e r s p e c i e s o f E n t o s t h o d o n i n h a v i n g e i t h e r a r u d i m e n t a r y p e r i s t o m e o r l a c k i n g a p e r i s t o m e a l t o g e t h e r . I t i s one o f t h e r a r e s t o f t h e b r y o p h y t e s i n t h e s t u d y . I have made two a d d i t i o n a l c o l l e c t i o n s o f an E n t o s t h o d o n f r o m j u s t o u t s i d e t h e s t u d y a r e a : TM4482 from 125 j u s t s o u t h of P r i n c e t o n and TM7015 from n o r t h of Kamloops. B o t h l a c k p e r i s t o m e s b ut have e x o t h e c i a l c e l l s o f t h e c a p s u l e w h i c h a r e n o t as l o n g (many a r e more o r l e s s i s o d i a m e t r i c ) o r as t h i c k - w a l l e d a s t h e White Lake c o l l e c t i o n s . The i l l u s t r a t i o n s i n Lawton (1971) i n d i c a t e t h a t t h e s e may f a l l w i t h i n t h e v a r i a t i o n o f E . r u b i g i n o s u s . As d i s c u s s e d i n d e t a i l i n t h e n e x t c h a p t e r , E. r u b i g i n o s u s may be c o n s p e c i f i c w i t h o t h e r s p e c i e s o f E n t o s t h o d o n , i n c l u d i n g E. t u c s o n i i ( B a r t . ) G r o u t , f rom s o u t h - w e s t e r n N o r t h A m e r i c a . FUNARIA Hedw. Key t o t h e S p e c i e s of FUNARIA I l e a v e s a c u t e and d e e p l y c o n c a v e ; c o s t a sometimes p e r c u r r e n t ; c a p s u l e s d e e p l y r i b b e d and w r i n k l e d when d r y ; a n n u l u s p r e s e n t ; l e a f y p l a n t s l a r g e ( l e a v e s o f t e n >3mm); f o u n d i n s e a s o n a l l y wet s i t e s F. h y g r o m e t r i c a I I l e a v e s u s u a l l y l o n g - a c u m i n a t e , n o t d e e p l y c o n c a v e ; c o s t a u s u a l l y e n d i n g w e l l b e f o r e t h e apex; c a p s u l e s smooth t o l i g h t l y w r i n k l e d when d r y ( t h e b a s e i s u s u a l l y w r i n k l e d ) ; a n n u l u s a b s e n t ; l e a f y p l a n t s s m a l l ( l e a v e s u s u a l l y <2mm); f o u n d i n s t e p p e F. m u h l e n b e r g i i FUNARIA HYGROMETRICA Hedw. P l a n t s g r e e n t o p a l e ( y e l l o w i s h - ) g r e e n , u s u a l l y a s s m a l l c o l o n i e s on s o i l ( e s p e c i a l l y i n s h a d e d , s e a s o n a l l y wet s i t e s ) . S p o r o p h y t e s common; c a p s u l e s i n c l i n e d , p y r i f o r m and d i s t i n c t l y r i b b e d . R e p r e s e n t a t i v e s p e c i m e n s : B89098, B89099 126 I l l u s t r a t i o n s : Crum and A n d e r s o n 1981, F l o w e r s 1973, I r e l a n d 1982, Lawton 1971 D i s t r i b u t i o n : A: s c a t t e r e d t h r o u g h o u t but uncommon l o c a l l y ( F i g . 54) B: t h r o u g h o u t N o r t h A m e r i c a C: c o s m o p o l i t a n Remarks : A l t h o u g h i t t e n d s t o f r e q u e n t s i t e s o f s e a s o n a l w a t e r a c c u m u l a t i o n , F u n a r i a h y g r o m e t r i c a i s n o t o f t e n f o u n d on t h e m a r g i n s of a l k a l i n e d e p r e s s i o n s , where i t might be e x p e c t e d . FUNARIA MUHLENBERGII T u r n . P l a n t s g r e e n t o p a l e g r e e n , i n d i v i d u a l s s c a t t e r e d i n s m a l l open c o l o n i e s on s o i l i n t h e s t e p p e o r i n shaded, b u t open s i t e s . S p o r o p h y t e s common; c a p s u l e s i n c l i n e d , p y r i f o r m and somewhat smooth e x c e p t a t t h e b a s e . R e p r e s e n t a t i v e s p e c i m e n s : B89100, B89101 I l l u s t r a t i o n s : F l o w e r s 1973, G r o u t 1928-1941, Lawton 1971, S m i t h 1978a D i s t r i b u t i o n : A: s c a t t e r e d , m a i n l y i n n o r t h - e a s t p o r t i o n o f t h e s t u d y a r e a ( F i g . 55) B: s c a t t e r e d a c r o s s t h e s o u t h e r n h a l f o f t h e c o n t i n e n t ; i n Canada p r o b a b l y o n l y i n B r i t i s h C o l u m b i a , p o s s i b l y i n A l b e r t a and t h e Yukon C: w e s t e r n N o r t h A m e r i c a / w e s t e r n E u r a s i a - N o r t h A f r i c a 127 Remarks : F o r a number o f r e a s o n s , i n c l u d i n g i t s t i n y s i z e , i t s h a b i t o f g r o w i n g a s s c a t t e r e d i n d i v i d u a l s and i t s i n c o n s p i c u o u s c o l o u r a t i o n , even when wet, F u n a r i a m u h l e n b e r g i i i s l i k e l y t o be e a s i l y o v e r l o o k e d and i s p o s s i b l y more common t h a n t h e few c o l l e c t i o n s i n d i c a t e . F o r t u n a t e l y , s p o r o p h y t e s a r e common and a r e most h e l p f u l i n l o c a t i n g t h e moss. T h e r e i s some c o n f u s i o n c o n c e r n i n g t h e s t a t u s o f F. m u h l e n b e r g i i . C r u n d w e l l and Nyholm (1974) s e g r e g a t e F. p u l c h e l l a and F. c o n v e x a f r o m t h e F. m u h l e n b e r g i i c o m p l e x . P o s s i b l y t h e N o r t h A m e r i c a n p o p u l a t i o n s need r e v i e w . ORTHOTRICACEAE ORTHOTRICHUM Hedw. An e m p h a s i s on s p o r o p h y t e c h a r a c t e r s t h a t a r e e i t h e r h i g h l y v a r i a b l e , s u c h a s t h e d e g r e e o f r i b b i n g o f t h e c a p s u l e w a l l , o r t h a t a r e d i f f i c u l t t o o b s e r v e , s u c h as t h e t y p e o f s t o m a t e s on t h e c a p s u l e , makes O r t h o t r i c h u m a t r o u b l e s o m e genus t o s t u d y t a x o n o m i c a l l y . However, a number o f r e c e n t s t u d i e s have been o f g r e a t i m p o r t a n c e i n t h e u n d e r s t a n d i n g o f t h e g r o u p (Crum and A n d e r s o n 1981; H i n n e r i 1976; L e w i n s k y 1977a, 1977b; V i t t 1973). T h e s e works a r e o f a b r o a d s c o p e , p l a c i n g a g r e a t e r e m p h a s i s on g a m e t o p h y t i c c h a r a c t e r s t h a n many e a r l i e r works, a l t h o u g h t h e use o f s p o r o p h y t i c c h a r a c t e r s i s s t i l l e s s e n t i a l i n t h e d e t e r m i n a t i o n o f most s p e c i e s . They p r o v i d e e x c e l l e n t and 128 c o m p l e t e i l l u s t r a t i o n s . V i t t (1973) i n c l u d e s d i s t r i b u t i o n maps o f a l l of t h e s p e c i e s d i s c u s s e d h e r e . O r t h o t r ichum i s composed of two major e v o l u t i o n a r y g r o u p s t h a t a r e d e f i n e d , p r i m a r i l y , by t h e m o r p h o l o g y of t h e s t o m a t a on t h e c a p s u l e w a l l s ( L e w i n s k y 1977a, V i t t 1973). One g r o u p has : s u p e r f i c i a l s t o m a t a ( p h a n e r o p o r e s ) and t h e o t h e r has immersed s t o m a t a ( c r y p t o p o r e s ) . O f t e n t h e s e s t o m a t a l t y p e s c o r r e l a t e w i t h e i t h e r t h i c k - w a l l e d and n o d u l o s e b a s a l l e a f c e l l s ( p h a n e r o p o r e s ) o r w i t h t h i n - w a l l e d and smooth b a s a l l e a f c e l l s ( c r y p t o p o r e s ) . Of t h e f i v e s p e c i e s f o u n d i n t h e s t u d y r e g i o n , two a r e o f t h e f o r m e r t y p e and t h r e e a r e o f t h e l a t t e r t y p e ( s e e t h e k e y ) . A l l f i v e s p e c i e s a r e more or l e s s r e s t r i c t e d t o t h e more m e s i c s i t e s o f t h e s t u d y r e g i o n , and a r e g e n e r a l l y s a x i c o l o u s . O n l y O. l a e v i g a t u m i s w i d e s p r e a d and common, w h i l e t h e o t h e r f o u r a r e uncommon o r r a r e . The f o l l o w i n g key i n c l u d e s t h r e e a d d i t i o n a l s p e c i e s : O. p e l l u c i d u m L i n d b . ( = 0. jamesianum S u l l . ) , 0. a f f i n e B r i d . and 0. s p e c i o s u m , t h a t have been f o u n d i n a r e a s n e a r t h e s t u d y r e g i o n . Key t o t h e S p e c i e s o f ORTHOTRICHUM I c a p s u l e w i t h immersed s t o m a t a ; b a s a l l e a f c e l l s u s u a l l y t h i n -w a l l e d ; p l a n t s u s u a l l y i n compact, l o w - g r o w i n g t u f t s , d a r k g r e e n t o b l a c k i s h t h r o u g h o u t 2 I I c a p s u l e w i t h s u p e r f i c i a l s t o m a t a ; b a s a l l e a f c e l l s u s u a l l y t h i c k - w a l l e d and more o r l e s s n o d u l o s e ; p l a n t s o f t e n i n l o o s e , t a l l t u f t s , u s u a l l y g r e e n i n upper p o r t i o n : 5 129 2 l e a v e s u n i f o r m l y b i s t r a t o s e above, w i t h o c c a s i o n a l t r i s t r a t o s e p a t c h e s ; c a p s u l e s 8 - r i b b e d O. h a 1 1 i i 2 1 l e a v e s u n i s t r a t o s e or b i s t r a t o s e i n p a t c h e s ; c a p s u l e s 8 o r 16 r i b b e d 3 3 c a p s u l e e x s e r t e d , c y l i n d r i c a l , o f t e n w i t h 8 d i s t i n c t l o n g r i b s a l t e r n a t i n g w i t h 8 s h o r t e r , l e s s d e f i n e d r i b s ; l e a v e s u n i s t r a t o s e 0. anomalum 3 1 c a p s u l e immersed t o emergent, o v a t e t o o b l o n g , w i t h 8 o r 16 r i b s ; l e a v e s u s u a l l y b i s t r a t o s e i n p a t c h e s 4 4 c a p s u l e o f t e n o v a t e , immersed and w i t h 16 r i b s , a l t h o u g h 8 o f them a r e sometimes i n d i s t i n c t ; l e a v e s a c u t e 0. c u p u l a t u m 4 1 c a p s u l e s u b - c y l i n d r i c a l , o f t e n c o n s t r i c t e d i n t h e m i d d l e , emergent and w i t h 8 r i b s ; l e a v e s b l u n t ( n o t y e t f o u n d i n s t u d y r e g i o n ) ( 0 . p e l l u c i d u m ) 5 c a p s u l e r i b b e d and e x s e r t e d ; p e r i s t o m e t e e t h r e c u r v e d t o r e f l e x e d , endostome o f t e n w e l l d e v e l o p e d ; u s u a l l y on wood (not y e t f o u n d i n s t u d y r e g i o n ) 6 5 1 c a p s u l e e x s e r t e d and smooth t o w e a k l y r i b b e d or immersed and r i b b e d ; p e r i s t o m e t e e t h i n c u r v e d , e r e c t o r s p r e a d i n g ( r a r e l y r e f l e x e d when o l d ) , endostome a b s e n t o r r u d i m e n t a r y ; u s u a l l y on r o c k 7 6 c a p s u l e s t r o n g l y r i b b e d t o b a s e (0. a f f i n e ) 6 1 c a p s u l e w e a k l y g r o o v e d o r smooth on upper h a l f (0. s p e c i o s u m ) 7 c a p s u l e c y l i n d r i c , e x s e r t e d , u s u a l l y smooth; p l a n t s u s u a l l y l i g h t g r e e n ; v e r y common 0. l a e v i g a t u m 7 1 c a p s u l e g l o b o s e t o o b l o n g , immersed t o emergent, o f t e n r i b b e d ; p l a n t s u s u a l l y d a r k g r e e n ; o c c a s i o n a l 0. r u p e s t r e 130 ORTHOTRICHUM ANOMALUM Hedw. P l a n t s d a r k g r e e n t o b l a c k i s h , a s f l a t , l o o s e t u f t s on sha d e d o u t c r o p s , o f t e n i n f i l t r a t e d w i t h f i n e s i l t s . S p o r o p h y t e s common; c a p s u l e s e x s e r t e d , c y l i n d r i c a l and w i t h 8 l o n g and 8 s h o r t e r r i b s ; s e t a u s u a l l y o r a n g e or r e d . R e p r e s e n t a t i v e s p e c i m e n s : B89102, B89103 I l l u s t r a t i o n s : Crum and A n d e r s o n 1981, F l o w e r s 1973, I r e l a n d 1982, Lawton 1971, V i t t 1973 D i s t r i b u t i o n : A: s c a t t e r e d and r a r e , 5 s i t e s : s i t e 21 n e a r S p e n c e s B r i d g e ; s i t e 45 s o u t h s i d e o f Kamloops; s i t e s 76 and 90 i n t h e S o u t h Okanagan; s i t e s 101 and 102 n e a r Keremeos ( F i g . 68) B: m a i n l y a c r o s s c e n t r a l N o r t h A m e r i c a : B r i t i s h C o l u m b i a t o t h e M a r i t i m e p r o v i n c e s , s o u t h t o V i r g i n i a i n t h e e a s t , H a i t i ( s e e V i t t 1973 f o r map) C: c i r c u m p o l a r Remarks : O r t h o t r i c h u m anomalum i s l i k e l y t o be c o n f u s e d w i t h low g r o w i n g , d a r k forms o f 0. l a e v i g a t u m , b u t d i f f e r s i n h a v i n g immersed s t o m a t a , c o n s p i c u o u s l y r i b b e d c a p s u l e s and o r a n g e t o r e d d i s h s e t a e . I t t e n d s t o grow i n more s h a d e d s i t e s t h a n 0. l a e v i g a t u m . The u n i s t r a t o s e l e a v e s and e x s e r t e d , c y l i n d r i c a l c a p s u l e s s e p a r a t e 0. anomalum f r o m o t h e r t a x a w i t h immersed s t o m a t a . 131 ORTHOTRI CHUM CUPULATUM B r i d . P l a n t s d a r k g r e e n t o b l a c k i s h , sometimes w i t h g l a u c o u s hue, f o r m i n g low g r o w i n g , compact t u f t s on f a c e s o f r o c k o u t c r o p s ; S p o r o p h y t e s common; c a p s u l e s immersed t o s l i g h t l y e m ergent, w i t h 16 r i b s . R e p r e s e n t a t i v e s p e c i m e n s : B89104, B89105 I l l u s t r a t i o n s : F l o w e r s 1973, Lawton 1971, V i t t 1973 D i s t r i b u t i o n : A: r a r e , 2 s i t e s : s i t e 13 n o r t h - e a s t of L y t t o n ; s i t e 75 ne a r K a l e d e n i n t h e s o u t h Okanagan V a l l e y ( F i g . 69) B: w e s t e r n N o r t h A m e r i c a : s o u t h e r n B r i t i s h C o l u m b i a s o u t h t o C a l i f o r n i a and e a s t t o S o u t h D a k o t a and T e x a s ( p r i m a r i l y i n t h e Rocky M o u n t a i n s ; see V i t t 1973) C: w e s t e r n N o r t h A m e r i c a / w e s t e r n E u r a s i a - N o r t h A f r i c a Remarks : A l t h o u g h some c a p s u l e s i n c o l l e c t i o n s o f O r t h o t r i c h u m  c u p u l a t u m have o n l y 8 d i s t i n c t r i b s ( c h a r a c t e r i s t i c o f 0. p e l l u c i d u m , w h i c h has n o t y e t been f o u n d i n t h e a r e a , y e t w i t h w h i c h i t i s o c c a s i o n a l l y s y n o n y m i z e d ) , o t h e r s have 16 r i b s , sometimes w i t h t h e a l t e r n a t i n g r i b s l e s s o b v i o u s t h a n t h e o t h e r s . 0. c u p u l a t u m i s a r a r e moss i n s o u t h e r n Canada, h a v i n g been f o u n d o n l y t h r e e t i m e s p r e v i o u s l y , a l l i n B r i t i s h C o l u m b i a ( s e e Tan 1980 and V i t t 1973). 132 ORTHOTRICHUM HALLII S u l l . e t L e s q . P l a n t s d a r k g r e e n , f o r m i n g s m a l l t u f t s on r o c k o u t c r o p s . S p o r o p h y t e s common; c a p s u l e s immersed, 8 r i b b e d . R e p r e s e n t a t i v e s p e c i m e n : B89106 I l l u s t r a t i o n s : F l o w e r s 1973, G r o u t 1928-1941, Lawton 1971, V i t t 1973 D i s t r i b u t i o n : A: r a r e , 2 s i t e s : s i t e 66 n o r t h - w e s t o f Summerland; 10 km n o r t h o f L y t t o n ( f r o m V i t t 1973) F i g . 70 B: w e s t e r n N o r t h A m e r i c a : B r i t i s h C o l u m b i a s o u t h t o A r i z o n a and New M e x i c o , m a i n l y i n d r i e r p a r t s o f t h e Rocky M o u n t a i n s ( s e e V i t t 1973) C: w e s t e r n N o r t h A m e r i c a n endemic Remarks : M a c r o s c o p i c a l l y , O r t h o t r i c h u m h a l l i i most c l o s e l y r e s e m b l e s 0. c u p u l a t u m and 0. p e l l u c i d u m , b u t t h e e v e n l y b i s t r a t o s e and o c c a s i o n a l l y t r i s t r a t o s e l e a v e s r e a d i l y s e p a r a t e 0. h a 1 1 i i f r o m t h e o t h e r two. A l s o , 0. h a l l i i has 8 r i b b e d c a p s u l e s , d i s t i n c t f r o m 0. c u p u l a t u m t h a t u s u a l l y has 16 r i b s . L e w i n s k y ( 1977c) n o t e d t h a t 0. p e l l u c idum i s sometimes c o m p l e t e l y b i s t r a t o s e i n h e r a r e a and, t h u s , may be c o n f u s e d w i t h 0. h a 1 1 i i . She c o n s i d e r e d , f o r one c a s e a t l e a s t , t h a t p a p i l l a h e i g h t w o u l d be a more r e l i a b l e c h a r a c t e r t h a n l e a f t h i c k n e s s ( 0 . p e l l u c i d u m c o n s i s t e n t l y had much h i g h e r p a p i l l a e t h a n 0. h a l l i i ) . B a s e d on t h e few s p e c i m e n s c o l l e c t e d i n t h e p r e s e n t s t u d y , t h e r e a p p e a r s t o be some s u p p o r t f o r t h i s o p i n i o n . 133 ORTHOTRICHUM LAEVIGATUM Z e t t . 0. m a c o u n i i A u s t . P l a n t s ( y e l l o w - ) g r e e n , u s u a l l y d a r k e r g r e e n t o b l a c k i s h below, r a r e l y d a r k g r e e n t h r o u g h o u t , f o r m i n g l o o s e t o somewhat compact t u f t s on open t o p a r t i a l l y s haded o u t c r o p f a c e s , r a r e l y on wood. S p o r o p h y t e s common; c a p s u l e s e x s e r t e d , c y l i n d r i c , u s u a l l y smooth and p a l e b r o w n , o c c a s i o n a l l y w e a k l y r i b b e d . R e p r e s e n t a t i v e s p e c i m e n s : B89107, B89108 I l l u s t r a t i o n s : F l o w e r s 1973, G r o u t 1928-1941, Lawton 1971, V i t t 1973 D i s t r i b u t i o n : A: t h r o u g h o u t , o c c a s i o n a l l y l o c a l l y common ( F i g . 71) B: w e s t e r n N o r t h A m e r i c a : s c a t t e r e d i n w e s t e r n A r c t i c , B r i t i s h C o l u m b i a t o Montana, s o u t h t o C a l i f o r n i a , C o l o r a d o and Nevada C: c i r c u m t e m p e r a t e Remarks : O r t h o t r i c h u m l a e v i g a t u m i s a common moss on open r o c k s u r f a c e s t h a t a r e n o t e x p o s e d t o l o n g p e r i o d s o f s u n l i g h t , h a b i t a t s more c h a r a c t e r i s t i c o f t h e genus G r i m m i a . A l t h o u g h p r i m a r i l y s a x i c o l o u s , t h e s p e c i e s was c o l l e c t e d a l s o on a b r a n c h o f Oregon g r a p e ( B e r b e r i s a q u i f o l i u m P u r s h ) and on d e c a y i n g c o n i f e r wood. 0. l a e v i g a t u m i s h i g h l y v a r i a b l e and c a n be c o n f u s e d w i t h a number o f o t h e r s p e c i e s . D a r k , compact forms r e s e m b l e 0. anomalum (s e e t h e d i s c u s s i o n t h e r e ) , whereas d a r k , more open forms r e s e m b l e 0. r u p e s t r e . However, c l o s e e x a m i n a t i o n s h o u l d 134 r e v e a l t h e more e x s e r t e d , g e n e r a l l y smoother and more c y l i n d r i c a l c a p s u l e s o f 0. l a e v i g a t u m . In a d d i t i o n , 0. r u p e s t r e t y p i c a l l y f o r m s l a r g e r t u f t s t h a n 0. l a e v i g a t u m and i s f o u n d i n more shaded h a b i t a t s . Some p o p u l a t i o n s have w e a k l y g r o o v e d c a p s u l e s and c a n be c o n f u s e d w i t h 0. s p e c i o s u m or 0. a f f i n e ( a l t h o u g h n e i t h e r t a x o n has y e t been f o u n d i n t h e s t u d y a r e a ) . T h e s e t a x a u s u a l l y have s t r o n g l y g r o o v e d c a p s u l e s a s w e l l a s r e f l e x e d exostome t e e t h and w e l l d e v e l o p e d e n d o s t o m e s . The exostome t e e t h o f 0. l a e v i g a t u m a r e u s u a l l y e r e c t t o s p r e a d i n g ( r a r e l y r e f l e x e d ) , and t h e endostome i s a b s e n t or r u d i m e n t a r y . ORTHOTRICHUM RUPESTRE S c h l e i c h . e t Schwaegr. P l a n t s u s u a l l y d a r k g r e e n t o b l a c k i s h , sometimes l i g h t e r g r e e n a b o v e , f o r m i n g l o o s e t u f t s on s h a d e d o u t c r o p s u r f a c e s . S p o r o p h y t e s u s u a l l y common; c a p s u l e s g l o b o s e t o s h o r t c y l i n d r i c a l , r i b b e d t o n e a r l y smooth, em e r g e n t , o f t e n brown. R e p r e s e n t a t i v e s p e c i m e n s : B89109, B89110 I l l u s t r a t i o n s : F l o w e r s 1973, G r o u t 1928-1941, Lawton 1971, L e w i n s k y 1977b, V i t t 1973 D i s t r i b u t i o n : A: s c a t t e r e d , p r i m a r i l y i n t h e s o u t h Okanagan V a l l e y ( F i g . 72) B: m a i n l y w e s t e r n N o r t h A m e r i c a : B r i t i s h C o l u m b i a t o A l b e r t a and S o u t h D a k o t a , s o u t h t o C a l i f o r n i a a n d A r i z o n a ; O n t a r i o C: c i r c u m t e m p e r a t e 135 Remarks : In t h e f i e l d , low g r o w i n g forms o f O r t h o t r i c h u m r u p e s t r e s ometimes r e s e m b l e 0. h a l l i i o r 0. p e l l u c idum. They s h o u l d be s e p a r a t e d r e a d i l y by t h e key c h a r a c t e r s . 0. r u p e s t r e may be c o n f u s e d a l s o w i t h s m a l l , d a r k f o r m s o f 0. l a e v i g a t u m ( s e e t h e d i s c u s s i o n t h e r e ) . BRYACEAE  BRYUM Hedw. Two s p e c i e s o f Bryum have been c o l l e c t e d i n t h e s t u d y a r e a . B. c a e s p i t i c i u m i s f o u n d a t a l m o s t e v e r y c o l l e c t i n g s i t e whereas B. a r g e n t e u m i s somewhat l e s s common, f r e q u e n t i n g more m e s i c s t e p p e s i t u a t i o n s t h a n t h e u b i q u i t o u s B. c a e s p i t i c i u m . Key t o t h e Spec i e s o f BRYUM I p l a n t s u s u a l l y s i l v e r y - w h i t e ( u p p e r p a r t o f l e a f n o t c h l o r o p h y l l o u s ) ; c o s t a e n d i n g below t h e apex t o s h o r t e x c u r r e n t ; m a r g i n s n o t r e c u r v e d ; l e a v e s n o t t w i s t e d ; u p p e r c e l l s u s u a l l y >12 m i c r o m e t e r s wide B. a r g e n t e u m I I p l a n t s g r e e n t o w h i t i s h ( u p per p a r t o f l e a f c h l o r o p h y l l o s e ) ; c o s t a e x c u r r e n t a s l o n g awn; m a r g i n s r e c u r v e d ; l e a v e s t w i s t e d t o e r e c t a b o v e ; upper c e l l s u s u a l l y <12 m i c r o m e t e r s wide B. c a e s p i t i c ium BRYUM ARGENTEUM Hedw. P l a n t s o f t e n f o r m i n g g r o u p s of s h o r t , w h i t i s h (sometimes g r e e n ) c o l u m n s , o c c a s i o n a l l y b u d - l i k e , on s o i l , e s p e c i a l l y n e a r 136 o u t c r o p s . S p o r o p h y t e s n o t s e e n . R e p r e s e n t a t i v e s p e c i m e n s : B89111, B89112 I l l u s t r a t i o n s : Lawton 197-1, F l o w e r s 1 973, Crum and A n d e r s o n 1981 D i s t r i b u t i o n : A: t h r o u g h o u t ( F i g . 30) B: t h r o u g h o u t N o r t h A m e r i c a C: c o s m o p o l i t a n Remarks : Robus t p l a n t s o f Bryum a r g e n t e u m a r e r e c o g n i z e d e a s i l y i n t h e f i e l d , b e c a u s e o f t h e i r c o l u m n - l i k e g r o w th h a b i t and w h i t i s h c o l o u r . S m a l l p l a n t s c an be c o n f u s e d w i t h B. c a e s p i t i c i u m b u t t h e s e s h o u l d be s e p a r a t e d r e a d i l y by c o n s u l t i n g t h e key. B. a r g e n t e u m i s o f t e n f o u n d g r o w i n g w i t h t h e l i v e r w o r t M a n nia f r a g r a n s o v e r t h i n s o i l on s h e l v e s o f r o c k o u t c r o p s . BRYUM CAESPITICIUM Hedw. P l a n t s y e l l o w - g r e e n t o p a l e g r e e n , g r o w i n g a s i n d i v i d u a l s or open c o l o n i e s on s o i l , e s p e c i a l l y i n d i s t u r b e d s i t e s , u s u a l l y m i x e d w i t h o t h e r mosses, i n p a r t i c u l a r C e r a t o d o n p u r p u r e u s . S p o r o p h y t e s common i n sh a d e d s i t e s , r a r e o t h e r w i s e ; c a p s u l e s y e l l o w i s h - b r o w n , i n c l i n e d , w i t h w r i n k l e d neck and u s u a l l y s h r u n k e n below t h e mouth. R e p r e s e n t a t i v e s p e c i m e n s : B89113, B89114 I l l u s t r a t i o n s : Crum and A n d e r s o n 1981, F l o w e r s 1973 D i s t r i b u t i o n : A: t h r o u g h o u t and common ( F i g . 31) 137 B: t h r o u g h o u t N o r t h A m e r i c a , more common i n t h e n o r t h e r n h a l f C: s u b - c o s m o p o l i t a n , r a r e i n t h e t r o p i c s Remarks : Bryum c a e s p i t i c ium and C e r a t o d o n p u r p u r e u s a r e e x t r e m e l y common t h r o u g h o u t t h e s t u d y a r e a and c a n be c o n f u s e d , a t l e a s t m a c r o s c o p i c a l l y . See t h e d i s c u s s i o n under C. p u r p u r e u s f o r comment. LEPTOBRYUM (B.S.G.) W i l s . LEPTOBRYUM PYRIFORME (Hedw.) W i l s . P l a n t s y e l l o w - g r e e n , u s u a l l y as s m a l l d e n s e t o open c o l o n i e s ( r a r e l y as s c a t t e r e d i n d i v i d u a l s ) on s o i l , u s u a l l y a d j a c e n t t o s e a s o n a l l y wet s i t e s . S p o r o p h y t e s o c c a s i o n a l ; c a p s u l e s p e n d e n t , e l o n g a t e - p y r i f o r m , g l o s s y R e p r e s e n t a t i v e s p e c i m e n s : B89117, B89118 I l l u s t r a t i o n s : F l o w e r s 1973, Crum and A n d e r s o n 1981 D i s t r i b u t i o n : A: uncommon, m a i n l y i n t h e s o u t h p a r t o f t h e Okanagan V a l l e y ( F i g . 66) B: t h r o u g h o u t N o r t h A m e r i c a C: c o s m o p o l i t a n Remarks : The e r e c t g r o w t h h a b i t and t h e l o n g , l i n e a r - l a n c e o l a t e l e a v e s a r e t h e most c h a r a c t e r i s t i c f e a t u r e s o f L e p t o b r y u m  p y r i f o r m e . I t i s a l s o a v e r y common moss i n g r e e n h o u s e s . 138 POHLIA Hedw. POHLIA CRUDA (Hedw.) L i n d b . P l a n t s p a l e g r e e n and w i t h m e t a l l i c l u s t r e , r a r e l y d a r k g r e e n , f o u n d as s m a l l , open c o l o n i e s on s o i l , u s u a l l y i n d e e p l y s h a d e d r e c e s s e s s u c h as a d j a c e n t t o b o u l d e r s . S p o r o p h y t e s n o t c o l l e c t e d . R e p r e s e n t a t i v e s p e c i m e n s : B89115, B89116 I l l u s t r a t i o n s : F l o w e r s 1973, Crum and A n d e r s o n 1981 D i s t r i b u t i o n : A: uncommon ( F i g . 75) B: a c r o s s n o r t h e r n h a l f o f c o n t i n e n t , s o u t h t o N. C a r o l i n a i n e a s t and t o M e x i c o i n t h e west, s c a t t e r e d i n c e n t r a l p o r t i o n s C: more o r l e s s c o s m o p o l i t a n Remarks : P o h l i a c r u d a i s an uncommon moss t h a t grows i n t h e most h i d d e n l o c a t i o n s and c a n be r e a d i l y o v e r l o o k e d . The p l a n t s a r e u s u a l l y p a l e and g l o s s y ( m e t a l l i c ) and once o b s e r v e d a r e u s u a l l y e a s y t o r e c o g n i z e . LESKEACEAE  PSEUDOLESKEELLA K i n d b . PSEUDOLESKEELLA TECTORUM ( B r i d . ) B r o t h . L e s k e e l l a t e c t o r u m ( B r i d . ) Hag. P l a n t s d a r k g r e e n t o g r e e n , g r o w i n g a s t i g h t , f l a t mats o v e r b o u l d e r s , u s u a l l y i n s h a d e d s i t u a t i o n s . S p o r o p h y t e s n o t 139 c o l l e c t e d . R e p r e s e n t a t i v e s p e c i m e n s : B89119, B89120 I l l u s t r a t i o n s : Crum and A n d e r s o n 1981, Lawton 1971, Nyholm 1954-1969 D i s t r i b u t i o n : A: o c c a s i o n a l ( F i g . 83 ) B: a c r o s s n o r t h e r n h a l f of t h e c o n t i n e n t , m a i n l y w e s t e r n C: more o r l e s s c i r c u m p o l a r ( s e e L e w i n s k y 1974) Remarks : P s e u d o l e s k e e l l a t e c t o r u m , l i k e Hypnum v a u c h e r i , forms mats on t h e s u r f a c e s o f r o c k s . U n l i k e H. v a u c h e r i , however, P. t e c t o r u m d o e s n o t grow on s o i l i n t h e s t u d y a r e a . In a d d i t i o n , t h e l e a v e s o f P. t e c t o r u m a r e d a r k e r g r e e n , l e s s s h i n y , and m a r k e d l y s m a l l e r t h a n H. v a u c h e r i . AMBLYSTEGIACEAE CONARDIA R o b i n s . CONARDIA COMPACTA (C. M u e l l . ) R o b i n s . A m b l y s t e q i u m compacturn (C. M u e l l . ) A u s t . R h y n c h o s t e g i e l l a compacta (C. M u e l l . ) L o e s k e Hypnum compacturn C. M u e l l . P l a n t s s h i n y y e l l o w - g r e e n , stems and l e a v e s t i n y ( l e a v e s u s u a l l y < 1mm l o n g ) , f o r m i n g compact t o o c c a s i o n a l l y l o o s e mats a l o n g t h e m a r g i n s o f a l k a l i n e d e p r e s s i o n s ; p r o p a g u l a o c c a s i o n a l , a t t a c h e d t o t h e b a c k s o f t h e u p p e r p a r t s o f t h e l e a v e s . 1 40 S p o r o p h y t e s n o t c o l l e c t e d . R e p r e s e n t a t i v e s p e c i m e n s : B89121, B89122 I l l u s t r a t i o n s : Crum and A n d e r s o n 1981, F l o w e r s 1973 (as A m b l y s t e g i u m compacturn ), I r e l a n d 1982 D i s t r i b u t i o n : A: o c c a s i o n a l , m a i n l y i n t h e s o u t h p a r t o f t h e Okanagan V a l l e y ( F i g . 34) B: s c a t t e r e d a c r o s s t h e n o r t h e r n h a l f of t h e c o n t i n e n t C: c i r c u m p o l a r Remarks : R o b i n s o n (1976) c o n s i d e r e d t h e l e a f p r o p a g u l a and t h e u n u s u a l l y t o o t h e d l e a v e s ( t h e t e e t h a r e o f t e n r e c u r v e d o r d o u b l e a t t h e l e a f b a s e ) d i s t i n c t enough t o c r e a t e a new genus b a s e d on t h i s t a x o n . I t s s m a l l s i z e i s r e m i n i s c e n t o f b o t h A m b l y s t e g i u m and R h y n c h o s t e q i e l l a , but t h e a f o r e m e n t i o n e d c h a r a c t e r s , a l o n g w i t h i t s l o n g - a p i c u l a t e o p e r c u l u m ( t o o s h o r t f o r R h y n c h o s t e q i e l l a and t o o l o n g f o r A m b l y s t e g i u m ) , d i s t i n g u i s h i t f r o m t h e s e two g e n e r a . C o n a r d i a has c h a r a c t e r s o f b o t h t h e A m b l y s t e g i a c e a e and t h e B r a c h y t h e c i a e a e (Crum and A n d e r s o n 1981, R o b i n s o n 1976, Tan 1980). BRACHYTHECIACEAE BRACHYTHECIUM B.S.G. Two s p e c i e s of B r a c h y t h e c i u m were c o l l e c t e d i n t h e s t u d y 141 r e g i o n : B. a l b i c a n s and B. c o l l i n u m . B o t h s p e c i e s a r e f o u n d i n sha d e d h a b i t a t s ( s u c h a s n o r t h f a c i n g l a c u s t r i n e b a n k s ) . B. a l b i c a n s i s e x t r e m e l y common, whereas B. c o l l i n u m i s r a r e . Key t o t h e S p e c i e s o f BRACHYTHECIUM I l e a v e s l a r g e ( o f t e n > 2mm), a r r a n g e d l o o s e l y on b r a n c h e s , l a n c e o l a t e w i t h l o n g , a t t e n u a t e a p i c e s ; upper and median c e l l s u s u a l l y > 60 m i c r o m e t e r s l o n g B. a l b i c a n s I I l e a v e s s m a l l ( o f t e n < 1mm), l o o s e l y t o d e n s e l y a r r a n g e d on j u l a c e o u s b r a n c h e s , o v a t e - t r i a n g u l a r , a p i c e s narrow b ut n o t l o n g - a t t e n u a t e ; upper and median c e l l s u s u a l l y <50 m i c r o m e t e r s l o n g B. c o l l i n u m BRACHYTHECIUM ALBICANS (Hedw.) B.S.G. P l a n t s ( p a l e ) y e l l o w - g r e e n , f o r m i n g sometimes e x t e n s i v e mats o v e r s o i l and o c c a s i o n a l l y on o u t c r o p s , o f t e n under l i t t e r , common i n s h a d e d s i t e s . S p o r o p h y t e s common, a t l e a s t i n l a r g e r p o p u l a t i o n s ; c a p s u l e i n c l i n e d t o h o r i z o n t a l ; s e t a smooth. R e p r e s e n t a t i v e s p e c i m e n s : B89123, B89124 I l l u s t r a t i o n s : Crum and A n d e r s o n 1981, F l o w e r s 1973, G r o u t 1928-1941, Lawton 1971 D i s t r i b u t i o n : A: w i d e s p r e a d and o f t e n common l o c a l l y ( F i g . 26) B: w i d e s p r e a d , more common i n t h e west C: c i r c u m p o l a r ; b i p o l a r (New Z e a l a n d ) Remarks : B r a c h y t h e c i u m a l b i c a n s i s common under l i t t e r i n s h a d e d s i t e s , sometimes g r o w i n g w i t h P o l y t r i c h u m j u n i p e r i n u m , C e r a t o d o n 142 p u r p u r e u s , E u r h y n c h i u m p u l c h e l l u m and, on r o c k s u r f a c e s , w i t h H o m a l o t h e c i u m aeneum. I t s p a l e y e l l o w - g r e e n c o l o u r a n d l o n g a t t e n u a t e l e a v e s d i s t i n g u i s h i t f r om most s i m i l a r t a x a , i n p a r t i c u l a r , H. aeneum. A l s o , H. aeneum has much s t r a i g h t e r and more d e e p l y p l i c a t e l e a v e s , and t h e b r a n c h e s a r e u s u a l l y r e g u l a r l y c u r v e d , e s p e c i a l l y n e a r t h e t i p s , a f e a t u r e n o t f o u n d i n B r a c h y t h e c i u m a l b i c a n s . BRACHYTHECIUM COLLINUM (C. M u e l l . ) B.S.G. P l a n t s y e l l o w - g r e e n , a s mats i n shaded t o , o c c a s i o n a l l y , open s i t e s on s o i l , o f t e n n e a r o u t c r o p s or b o u l d e r s . S p o r o p h y t e s o c c a s i o n a l ; c a p s u l e s i n c l i n e d t o h o r i z o n t a l ; s e t a smooth t o w e a k l y r o u g h . R e p r e s e n t a t i v e s p e c i m e n s : B89125, B89126 I l l u s t r a t i o n s : Lawton 1971, F l o w e r s 1973 D i s t r i b u t i o n : A: o c c a s i o n a l , n e v e r l o c a l l y a b undant ( F i g . 27) B: B r i t i s h C o l u m b i a and A l b e r t a s o u t h t o C a l i f o r n i a t o T e x a s , G r e e n l a n d C: more o r l e s s c i r c u m t e m p e r a t e Remarks : B r a c h y t h e c i u m c o l l i n u m c a n be e a s i l y c o n f u s e d w i t h E u r h y n c h i u m p u l c h e l l u m , b e c a u s e o f s i m i l a r l e a f shape and s i z e a s w e l l a s t h e m a r g i n a l s e r r a t i o n , b ut d i f f e r s i n t h e c o n s i s t e n t l y l o n g e r and n a r r o w e r l e a f a p i c e s , y e l l o w - g r e e n c o l o u r and u p p e r l e a f a r e o l a t i o n ( s e e t h e g e n e r i c k e y ) . A l s o , E . p u l c h e l l u m t e n d s t o have l a m i n a l f o l d s on e i t h e r s i d e of t h e 1 43 l e a f b a s e , w h i c h a r e a b s e n t o r p o o r l y d e v e l o p e d i n B. c o l l i n u m . F l o w e r s (1973) l i s t s B. c o l l i n u m as one o f t h e most common d r y l a n d mosses i n U t a h . EURHYNCHIUM B.S.G. EURHYNCHIUM PULCHELLUM (Hedw.) J e n n . E . f a l l a x (Ren. e t C a r d . ) G r o u t P l a n t s g r e e n t o ( d a r k ) y e l l o w - g r e e n , as l o o s e mats on s o i l i n s h a d e d t o open c o n d i t i o n s . S p o r o p h y t e s o c c a s i o n a l , c a p s u l e i n c l i n e d t o h o r i z o n t a l . R e p r e s e n t a t i v e s p e c i m e n s : B89127, B89128 I l l u s t r a t i o n s : Lawton 1971, F l o w e r s 1 973, Crum and A n d e r s o n 1 981 D i s t r i b u t i o n : A: t h r o u g h o u t ( F i g . 53) B: a c r o s s c o n t i n e n t C: c i r c u m p o l a r Remarks : E u r h y n c h i u m p u l c h e l l u m i s most common i n t h e f o r e s t - s t e p p e t r a n s i t i o n a l s i t e s o f t h e s t u d y a r e a . O n l y B r a c h y t h e c ium  c o l l i n u m c a n be c o n f u s e d w i t h i t ( s e e c o m p a r a t i v e n o t e s u n d e r B. c o l l i n u m ) . 144 HOMALOTHECIUM B.S.G. HOMALOTHECIUM AENEUM ( M i t t . ) Lawt. C a m p t o t h e c i u m aeneum ( M i t t . ) J a e g e r e t S a u r b . P l a n t s s h i n y y e l l o w - g r e e n ( u s u a l l y w i t h a m e t a l l i c l u s t r e ) a s l a r g e mats o v e r r o c k s u r f a c e s , o c c a s i o n a l l y on s o i l . S p o r o p h y t e s r a r e ; c a p s u l e s i n c l i n e d . R e p r e s e n t a t i v e s p e c i m e n s : B89129, B89130 I l l u s t r a t i o n s : Lawton 1971 D i s t r i b u t i o n : A: f a i r l y common t h r o u g h o u t ( F i g . 63) B: w e s t e r n N o r t h A m e r i c a C: w e s t e r n N o r t h A m e r i c a n endemic Remarks : H o m a l o t h e c i u m n e v a d e n s e ( L e s q . ) Ren. e t C a r d , i s v e r y s i m i l a r t o H. aeneum, t h e major d i f f e r e n c e b e i n g t h e e r e c t c a p s u l e o f H. nevad e n s e and t h e c u r v e d c a p s u l e o f H. aeneum. V e g e t a t i v e l y , I c o u l d n o t f i n d many h e l p f u l d i f f e r e n c e s (H. aeneum t e n d s t o be l a r g e r ) . A l t h o u g h H. n e v a d e n s e has been r e p o r t e d f r o m t h e s t u d y a r e a (Lawton 1971), i t i s n o t i n c l u d e d h e r e s i n c e e v e r y c o l l e c t i o n I made w i t h s p o r o p h y t e s had c u r v e d c a p s u l e s . 1 45 HYPNACEAE HYPNUM Hedw. Key t o t h e s p e c i e s o f HYPNUM 1 l e a f m a r g i n s p l a n e H. v a u c h e r i 1 1 a t l e a s t one l e a f m a r g i n r e v o l u t e H. r e v o l u t u m HYPNUM REVOLUTUM ( M i t t . ) L i n d b . P l a n t s g r e e n t o b r o w n i s h - g r e e n , f o r m i n g compact mats i n sha d e d a r e a s , u s u a l l y on t a l u