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A study of the Brachythecium asperrimum-frigidum species complex Hoisington, Barbara Ludlow 1979

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A STUDY OF THE BRACHYTHECIUM ASPERRIMUM-FRIGIDUM SPECIES COMPLEX by BARBARA LUDLOW HOISINGTON B.A., University of Vermont, 1976 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE i n THE FACULTY OF GRADUATE STUDIES (Department of Botany) We accept t h i s thesis as conforming to- the required standard THE UNIVERSITY OF BRITISH COLUMBIA September, 1979 © , Barbara Ludlow Hoisington, 1979 I n p r e s e n t i n g t h i s t h e s i s i n p a r t i a l f u l f i l m e n t o f t h e r e q u i r e m e n t s f o r a n a d v a n c e d d e g r e 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 , I a g r e e t h a t t h e L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e a n d s t u d y . I f u r t h e r a g r e e t h a t p e r m i s s i o n f o r e x t e n s i v e c o p y i n g o f t h i s t h e s i s f o r s c h o l a r l y p u r p o s e s may be g r a n t e d by t h e Head o f my D e p a r t m e n t o r by h i s r e p r e s e n t a t i v e s . I t i s u n d e r s t o o d t h a t c o p y i n g o r p u b l i c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l n o t be a l l o w e d w i t h o u t my w r i t t e n p e r m i s s i o n . D e p a r t m e n t o f . Botany  The U n i v e r s i t y o f B r i t i s h C o l u m b i a 2075 W e s b r o o k P l a c e V a n c o u v e r , C a n a d a V6T 1W5 D a t e k September 1Q79 Abstract • • , T h i s t h e s i s describes a study of s i x North American, d i o i c o u s Brachytheeium species possessing a rough s e t a . The phenetic v a r i a b i l i t y of the s i x species was analysed i n order to c l a r i f y the r e l a t i o n s h i p s , between species and to provide .simpler, more r e l i a b l e means of i d e n t i -f y i n g specimens. As a consequence of t h i s i n v e s t i g a t i o n , the taxonom.ic i d e n t i t i e s ' o f B. .asperrimum (Mitt'.' ex. C. Muell.) S u l l . and B. f r i g i d u m (C. Muell.) Besch. have been e s t a b l i s h e d through numerical analyses of the v a r i a t i o n i n the gametophytic fe a t u r e s of the two species.. Two types of analyses were used: a p r i n c i p a l components a n a l y s i s and a c l u s t e r a n a l y s i s . The phenetic v a r i a t i o n and h a b i t a t s p e c i f i c i t y of each species-, were analysed anr< documented and synonymy anc? ranges presented. A. t e s t of the.features considered d i a g n o s t i c f o r the two.species has been performed and'the r e s u l t s of t h i s t e s t used to devise a r e l i a b l e key. The r e l a t i o n s h i p s of the two species to the other North American d i o i c o u s species possessing rough setae were determined using a p r i n c i p a l components a n a l y s i s and. a c l u s t e r a n a l y s i s . B. asperrimum ( M i t t . e^ x. C. M u e l l . ) . S u l l . i s shown to be c l o s e l y r e l a t e d to B. b o l a n d e r i (Lesq.) Jaeg. and Sauerb.. B. f r i g i d u m (C. Muell.) Besch. i s shown to be c l o s e l y r e l a t e d to B. r i v u l a r e B.S.G. and B. n e l s o n i i Grout. A dendrogram and d i s -cussion of the h y p o t h e t i c a l , c l a d i s t i c r e l a t i o n s h i p s of the s i x species are provided. 111. Table of Contents T i t l e Page i Abstract i i Table of Contents i i i L i s t of Tables v L i s t of Figur e s and I l l u s t r a t i o n s v i i i Acknowledgement x i v I n t r o d u c t i o n 1 Part One: An A n a l y s i s of the Phenetic V a r i a b i l i t y of B. f r i g i d u m (C. Muell . ) Besch sensu Robinson . 10 I . Data c o l l e c t i o n 10 ( 1 . ) 1. The s e l e c t i o n of specimens 10 ( 1 . ) 2 . The choice of c h a r a c t e r s to be measured 11 ( 1 . ) 3 . The measurement and s c o r i n g of chara c t e r s 12 I I . A n a l y s i s of Data 20 ( 2 . ) 1. General d i s c u s s i o n of methods 20 . ( 2 . ) 2 . R e s u l t s of. the Data A n a l y s i s 23 P a r t Two: Brachythecium asperrimum ( M i t t . ex_ C. Muell.) S u l l . and B. f r i g i d u m (C. Muell.) Besch. r e d e f i n e d 39 Key to the stem leaves of B. asperrimum ( M i t t . _ex C. Muell.) S u l l . ' and B. fr i g i d u m (C. Muelj./) Besch - kO B. f r i g i d u m (C. Muell.) Besch kl B. . asperrimum ( M i t t , ex C. Muell.) S u l l • •]. i v . Table of Contents, continued Part Three: The R e l a t i o n s h i p s of the S i x North American Dioicous Br achy thee ium species w i t h Rough Setae 62 P r o v i s i o n a l Key to the d i o i c o u s Brachythecium species of North America with Rough Setae 7k Summary 82 L i t e r a t u r e C i t e d 83 Appendix A 8k Appendix B 91 Appendix C Appendix D. ... 95 Appendix E 97 Appendix F • 98 V. L i s t of Tables Table l a . The concepts of the s i x North American d i o i c o u s Brachythecium species w i t h rough setae according to Grout (1928-19^0, Moss F l o r a of North America, Volume 3 ) 1 S. Flowers (1973? Mosses: Utah and the VJest) and E. Lawton (1971* Moss F l o r a  of the P a c i f i c Northwest) h Table l b . The concepts of the s i x North American d i o i c o u s Brachythecium species w i t h rough setae, continued 5 Table l c . The concepts of the s i x North American d i o i c o u s Brachythecium species w i t h rough setae, continued 6 Table l d . The concepts of the s i x North American d i o i c o u s Brachythecium species w i t h rough setae, continued 7 Table l e . The concepts of the s i x North American d i o i c o u s Brachythecium species w i t h rough setae, continued 8 Table l f . The concepts of the s i x North American d i o i c o u s Brachythecium species with.rough setae, continued 9 Table 2. The h a b i t a t s and geographic areas, represented i n the sample of s i x t y - e i g h t O.T.U.'s chosen f o r data a n a l y s i s . (See Appendix A f o r a complete l i s t of specimens) 11 Table 3 a . Measurement of Characters: c r i t e r i a used, type and s c o r i n g v i . L i s t of Tables, continued Table 3b. Measurement of Characters: c r i t e r i a used, type and s c o r i n g of d a t a , c o n t i n u e d 1+ Table 3 c . Measurement of Characters: c r i t e r i a used, type and s c o r i n g of data, continued 15 Table k. The c o n t r i b u t i o n of the v a r i a b l e s to the v a r i a t i o n shown along the f i r s t three major components (eigenvectors) i n the o r d i n a t i o n of B. asperrimum ( M i t t , ex C. M u e l l . ) S u l l 30 Table 5« The c o n t r i b u t i o n of the v a r i a b l e s to the v a r i a t i o n shown along the f i r s t three major components (eigenvectors) i n the o r d i n a t i o n of B. f r i g i d u m (C. Muell.) Besch 33 Table 6 . The c o n t r i b u t i o n of the v a r i a b l e s to the v a r i a t i o n shown along the f i r s t three major components (eigenvectors) i n the o r d i n a t i o n of the 68 O.T.U.'s . (combined specimens of B. asperrimum ( M i t t . _ex C. Muell.) S u l l . and B. f r i g i d u m (C. Muell.) Besch 3k Table 7- The r e s u l t of a t e s t of d i a g n o s t i c f e a t u r e s of Brachythecium asperrimum ( M i t t . ex_ C. Muell.) S u l l . and B. f r i g i d u m (C. M u e l l . Besch 37 V l l . L i s t of Tables, continued Table 8 a . A d d i t i o n a l c h a r a c t e r s used i n the exemplary o r d i n a t i o n o f 6 s p e c i e s . Measurement of Characters: c r i t e r i a used, type and s c o r i n g of data 63 Table 8 b . A d d i t i o n a l c h a r a c t e r s used i n the exemplary o r d i n a t i o n of 6 s p e c i e s . Measurement of Characters: c r i t e r i a used, type and s c o r i n g of data, continued Gh Table 9. The c o n t r i b u t i o n s to the v a r i a t i o n of the J>2 v a r i a b l e s measured f o r the comparative o r d i n a t i o n of s i x species....'71 V l l l . L i s t of Figur e s and I l l u s t r a t i o n s F i g . 1. Character s t a t e s of l e a f shape used i n the s c o r i n g of ranked data. (a) broadly d e l t o i d ; (b) d e l t o i d ; ( c , d) broadly ovate; (e, f ) ovate, continued on the next page (Figure 2) 16 F i g . 2 . Character s t a t e s of l e a f shape used i n the s c o r i n g of ranked data, continued. (g, h, i , j ) narrowly ovate; (k, 1, m) l a n c e o l a t e 17 F i g . 3« Character s t a t e s of the l e a f p l i c a t i o n used i n the s c o r i n g of ranked data. (a, b, c) plane; (d,e) s l i g h t l y w r i n k l e d ; ( f ) s t r o n g l y w r i n k l e d ; (g, h, i ) b i p l i c a t e ; ( j , k) s t r o n g l y p l i c a t e 18 F i g . k. Character s t a t e s of the l e a f apex shape used i n the s c o r i n g of ranked data, (a) long and slender; (b,c) long-acuminate; (d, e) short-acuminate; ( f ) acute 19 F i g . 5> Features of the l e a f apex used i n the s c o r i n g of d i c h o t -omous data, ( a , b) f a l c a t e , (c) t w i s t e d . 19 F i g . 6 . The f i r s t and second axes of the f i r s t o r d i n a t i o n of 68 O.T.U.'s showing the h a b i t a t of each O.T.U.. 0= B. asper-r.imum ( M i t t , ex C. Muell.) S u l l . ; © = B. f r i g i d u m (C. Muell.) I X . L i s t of Figures and I l l u s t r a t i o n s , continued F i g . 7« The r e s u l t s of a c l u s t e r a n a l y s i s of 68 O.T.U.'s of B. frig i d u m (C. Muell.) Besch. Distance along the v e r t i c a l s c a l e r e f e r s to taxonomic d i s t a n c e . Groups of two or more O.T.U.'s along the base of the dendrogram i n d i c a t e specimens considered to be i d e n t i c a l by the c l u s t e r a n a l y s i s . 25 F i g . 8. The f i r s t and second axes of the o r d i n a t i o n of 68 O.T.U.'s showing the h a b i t a t of each O.T.U.. o = B. asperrimum ( M i t t . ex. C. Muell.) S u l l . ; o = B. fri g i d u m (C. Muell.) Besch. 28 F i g . 9 » The f i r s t and second axes of the o r d i n a t i o n of B. asperrimum ( M i t t . ex. C. Muell.) S u l l . showing the h a b i t a t s of each O.T.U. 29 F i g . 10. The f i r s t and second axes of the B. fri g i d u m (C. Muell.) Besch. o r d i n a t i o n , showing the h a b i t a t s of the specimens. 32 F i g . 11. The range of Brachythecium frigidum.' (C. Muell.) Besch. ..... kj> F i g . 12. Brachythecium f r i g i d u m (C. Muell.) Besch.: V a r i a t i o n i n shoot s i z e and branching p a t t e r n . (Continued on page h*?, F i g . 13)- Two wet h a b i t a t forms showing h o r i z o n t a l primary stem and e r e c t , v a r i o u s l y branched secondary stems 44 F i g . 13. Brachythecium frig i d u m (C. Muell.) Besch.: V a r i a t i o n i n shoot s i z e and branching p a t t e r n . (Continued on page 46, X. L i s t of Fig u r e s and I l l u s t r a t i o n s , continued F i g . 13 . (continued) F i g . 14). (a)Secondary stem of a l a r g e wet form, (b) e n t i r e shoot of d r i e r , smaller (mat) form V? F i g . l * t . Brachythecium f r i g i d u m (C. Muell.) Besch.: V a r i a t i o n i n shoot s i z e and branching p a t t e r n . (a) l a r g e , s p a r s e l y branched secondary stem of a wet ( t u f t ) form; (b) small aquatic form showing a t y p i c a l l y small s i z e and sparse f o l i a t i o n kS F i g . 15. B. f r i g i d u m (C. Muell.) Besch.: v a r i a t i o n i n the s i z e , shape and p l i c a t i o n of branch leaves 4 7 F i g . 16. B. f r i g i d u m (C. Muell.) Besch.: v a r i a t i o n i n the s i z e , shape and p l i c a t i o n of stem leaves k8 F i g . 17. B. f r i g i d u m (C. Muell.) Besch. (a) median l e a f c e l l s ; (b) c e l l s of the l e a f apex; ( c , d) two stem leaves showing the appearance of the i n f l a t e d a l a r c e l l s under low power m a g n i f i c a t i o n (kX o b j e c t i v e , 10X o c u l a r ) ^9 F i g . 18. B. f r i g i d u m (C. Muell.) Besch.: v a r i a t i o n i n the develop-ment of the a l a r c e l l s , (a) t y p i c a l stem l e a f a r e o l a t i o n w i t h numerous round to oblong c e l l s , (b) branch l e a f areo-l a t i o n showing the fewer number of i n f l a t e d c e l l s . . . 5 0 F i g . 19. The range of B. asperrimum ( M i t t . ex. C. Muell.) S u l l 5^ I j i g . 2 0 . Brachythecium asperrimum ( M i t t . ex. C. -Muell.) S u l l . : j V a r i a t i o n i n shoot s i z e and_ branching p a t t e r n . (Continued j on page 56 , F i g . 2 1 ) . Male p l a n t of a s t o l o n i f e r o u s I ' 1 • • • x i . L i s t of Figures and I l l u s t r a t i o n s , continued F i g . 20. (continued) ( e p i p h y t i c ) form 55 F i g . 21. Brachythecium asperrimum ( M i t t . _ex. C. Muell.) S u l l . : v a r i a t i o n i n shoot s i z e and branching p a t t e r n . (Continued on page 57» F i g . 2 2 ) . (a) female p l a n t of a s t o l o n i f e r o u s ( e p i p h y t i c ) form; (b) smaller p l a n t from r o t t i n g wood 56 F i g . 2 2 . Brachythecium asperrimum ( M i t t . _ex. C. Muell.) S u l l . : v a r i a t i o n i n shoot s i z e and branching p a t t e r n , continued 57 F i g . .23. B. asperrimum ( M i t t . ex. C. Muell.) S u l l . : v a r i a t i o n i n the s i z e and shape of branch leaves 5 3 F i g . 2k. B. asperrimum ( M i t t . ex. C. Muell.) S u l l . : v a r i a t i o n i n the s i z e and shape of stem leaves 59 F i g . 2 5 . B. asperrimum ( M i t t . ex. C. Muell.) S u l l . (a) median l e a f c e l l s ; (b) c e l l s of the l e a f apex; ( c , e) a l a r c e l l s of two leaves showing the t y p i c a l marginal f i l e of quadrate or r e c t a n g u l a r c e l l s ; (d) sketch of a stem l e a f showing the appearance of the a l a r r e g i o n under low power m a g n i f i c a t i o n (kX o b j e c t i v e , 10X o c u l a r . ) 60 Fig. 26 . Character s t a t e s of the siz-e and p o s i t i o n of a l a r c e l l s : used.in the s c o r i n g of ranked data: (a, b, c) not forming decurrent a u r i c l e s ; (d, e) forming small a u r i c l e s ; ( f ) forming l a r g e a u r i c l e s which do not extend to the c o s t a ; (g) forming l a r g e a u r i c l e s which extend to the costa 65 x i i . L i s t of Figures and I l l u s t r a t i o n s , continued F i g . 2?.- Character s t a t e s of the concavity of leaves used i n the s c o r i n g of ranked data: (a) not or s l i g h t l y concave; (b) s t r o n g l y concave but l a c k i n g the a p i c a l spoon-shaped depression; ( c , d) s t r o n g l y concave w i t h an a p i c a l spoon-shaped depression 6 6 F i g . 2 8 . Character s t a t e s of the l e a f apex shape used i n the s c o r i n g of ranked data: (a) a b r u p t l y a p i c u l a t e ; (b) not a p i c u l a t e (the other four character s t a t e s are on page 1 8 , Figure k.) 6 7 F i g . 2 9 . Character s t a t e s of the l e a f apex used i n the s c o r i n g of dichotmous data: (a) ab r u p t l y narrowed; (b, c) g r a d u a l l y narrowed .67 F i g . 3 0 . The f i r s t and second axes of the comparative o r d i n a t i o n of s i x species 6 9 F i g . 3 1 •• The f i r s t and t h i r d axes of the comparative o r d i n a t i o n of s i x species 7 0 F i g . . R e s u l t s of the c l u s t e r a n a l y s i s comparing s i x Brachythecium species 72 F i g . 3 3 - Brachythecium b o l a n d e r i (Lesq.) Jaeg. & Sauerb. (a., b) stem le a v e s ; (c,d) branch le a v e s ; (e) a p i c a l c e l l s ; ( f ) a l a r c e l l s of stem l e a f 76 x i i i . L i s t of Figur e s and I l l u s t r a t i o n s , continued F i g . J>k. Brachythecium n e l s o n i i Grout (a) a p i c a l c e l l s ; (b) stem l e a f ; (c) a l a r c e l l s ; (d) branch l e a f 77 F i g . 35 • Brachythecium hylotapetum B. Hig. & N. Hig. (a, b) branch l e a v e s ; (c) stem l e a f ; (d) a p i c a l c e l l s 7.8 F i g . 3 6 . Brachythecium hylotapetum B. Hig. & N. H i g . A l a r c e l l s of stem l e a f 79 F i g . 37* Brachythecium r i v u l a r e B.S.G. (a, b) branch leaves; (c) stem l e a f ; (d) a p i c a l c e l l s 80 F i g . 3 8 . Brachythecium r i v u l a r e B.S.G.. A l a r c e l l s of a stem l e a f ' 8 l x i v . Acknowledgment I g r a t e f u l l y acknowledge the a s s i s t a n c e of Dr. Gary B r a d f i e l d and Mr. Norm Kenkel f o r t h e i r help w i t h the s t a t i s t i c a l analyses. I would l i k e to thank the c u r a t o r s o f the f o l l o w i n g h e r b a r i a f o r the loan of m a t e r i a l s : the U n i v e r s i t y of Michigan herbarium (MICH), the U n i v e r s i t y o f Tennessee herbarium (-TENN), the herbarium o f the New York B o t a n i c a l Garden (NY), the U n i v e r s i t y of Colorado herbarium (COLO), the N a t i o n a l Herbarium o f the Smithsonian I n s t i t u t i o n (US), and the herbarium of the U n i v e r s i t y of B r i t i s h Columbia (UBC). I would a l s o l i k e to thank Dr. Fred Hermann, who k i n d l y l e n t me s p e c i -mens from h i s personal herbarium. I g r a t e f u l l y acknowledge the a s s i s t a n c e o f Mrs. L e s l i e Bohm, fo r her drawings of the growth forms of Brachythecium asperrimum ( M i t t . ex. C. Muell.) S u l l . and B. f r i g i d u m (C. Muell.) Besch.. I am g r e a t l y i n debt to Dr. W. B. S c h o f i e l d , f o r h i s encouragement, p r o f e s s i o n a l advice and f r i e n d s h i p . I owe more than can be expressed to my mother, Mrs. C e c i l B. Ho i s i n g t o n , f o r her encouragement, care and c r i t i c i s m s of the manuscript, and e s p e c i a l l y f o r t y p i n g the manuscript. INTRODUCTION The primary question addressed by t h i s t h e s i s i s whether or not Brachythecium asperrimum (M i t t . , ex. C. Muell.) S u l l . and B. f r i g i d u m (C. Muell.)' Besch. represent v a r i a t i o n s of the same sp e c i e s . I was f i r s t introduced to the problem by Dr. W. B. S c h o f i e l d of the Univer-s i t y of B r i t i s h Columbia, who c a l l e d my a t t e n t i o n to c e r t a i n d i s c r e p -ancies i n the l i t e r a t u r e . In 1963 Dr. H. Robinson s t a t e d that B.- asperrimum ( M i t t . ex. C. Muell.) S u l l . was merely one of many forms of B. f r i g i d u m (C. Muell.) Besch. (The B r y o l o g i s t 66:136-139) . In 1971 i n her Moss F l o r a of the P a c i f i c Northwest, Dr. E. Lawton r e t a i n e d the two taxa as d i s t i n c t s p e c i e s , separating them on the b a s i s of h a b i t a t and growth form. A survey of the l i t e r a t u r e revealed f u r t h e r a m b i g u i t i e s and d i s c r e p a n c i e s i n the concept of the two taxa according to A. J . Grout (1928-19^0, Moss F l o r a of North America, Volume 3 ) , S. Flowers (1973, Mosses: Utah and the West), and E. Lawton (1971? Moss F l o r a of the P a c i f i c  Northwest.) Furthermore, these d i s c r e p a n c i e s were al s o found i n the de-s c r i p t i o n s of the other North American d i o i c o u s species of Brachythecium with rough setae, namely: B. b o l a n d e r i (Lesq.) Jaeg. & Sauerb., B. h y l o - tapetum B. Hig. & N. Hig., B. n e l s o n i i Grout;, and B. r i v u l a r e B.S.G. The three authors' concepts of the s i x species are presented i n Tables l a . -I f . , (pages k - 9 ) . These s i x species a r e t h e s u b j e c t o f t h i s i n v e s t i -g a t i o n . The vague or c o n t r a d i c t o r y concepts of Brachythecium species may be a t t r i b u t a b l e to species v a r i a b i l i t y , to the authors' taxonomic miscon-c e p t i o n s , or to a l a c k of p r e c i s e d e f i n i t i o n s of the terms employed. Whatever the cause, the r e s u l t i s that there i s today no c l e a r concept 2. or do'cumentation of the extent of the v a r i a b i l i t y of Brachythecium s p e c i e s , nor any workable scheme f o r the i d e n t i f i c a t i o n of specimens. The purposes of t h i s i n v e s t i g a t i o n are: 1. ) to assess the phenetic v a r i a t i o n of Brachythecium f r i g i d u m (C. Muell.) Besch. sensu Robinson; 2. ) to analyse t h i s v a r i a t i o n and determine i f B. asperrimum ( M i t t . ex. C. Muell.) S u l l . and B. f r i g i d u m (C. Muell.) Besch. e x i s t as p h e n e t i c a l l y separable e n t i t i e s , and, i f so: 3 . ) to r e d e f i n e each s p e c i e s , analyse and document i t s phenetic v a r i a b i l i t y , h a b i t a t s p e c i f i c i t y and range, k.) to a r r i v e at a r e l i a b l e means of i d e n t i f y i n g specimens of the two tax a and 5 . ) to assess the r e l a t i o n s h i p of the two taxa to the other North American d i o i c o u s Brachythecium species w i t h rough setae. In.order to achieve these o b j e c t i v e s , I choose to employ the methods of numerical taxonomy. Among the p r i n c i p l e s of numerical taxonomy are: (1.) that a n a t u r a l c l a s s i f i c a t i o n i s based on a c l e a r understanding and documentation of the v a r i a b i l i t y of a l a r g e number of characters and (2.) that no character i s given p r i o r i t y over another i n the-' \ establishment of c l a s s i f i c a t i o n s . . 3 . Former treatments of Brachythecium have assigned p r i o r i t y t o , or have omitted, c e r t a i n c h a r a c t e r s . T h i s has r e s u l t e d i n taxonomic schemes based on an incomplete or weighted assesment of the taxa's phenetic v a r i a b i l i t y . For example, B. rutabulum (Hedw.) B.S.G. and B. r i v u l a r e B.S.G. have been t r e a t e d as c l o s e l y r e l a t e d species because both species have concave and decurrent stem leaves. How-ever, the former i s autoicous w i t h a s e t a that v a r i e s i n roughness and the l a t t e r i s d i o i c o u s w i t h a s e t a rough throughout. M u l t i v a r i a t e analyses r e q u i r e d e f i n i t i o n s of the char a c t e r s measured, provide documentation of the v a r i a b i l i t y of these c h a r a c t e r s , and provide at l e a s t a p a r t i a l l y o b j e c t i v e means of d i s p l a y i n g the t o t a l v a r i a b i l i t y i n a simpler form. An understanding of the taxa's v a r i a b i l i t y w i l l r e v e a l f e a t u r e s which are more constant f o r each, taxon. These f e a t u r e s may then be used i n the c o n s t r u c t i o n of keys to p h e n e t i c a l l y separate the taxa . The a p p l i c a t i o n of the p r i n c i p l e s and methods of numerical taxonomy consequently r e s u l t s i n a more r e l i a b l e means of i d e n t i f y i n g specimens and a c l e a r e r concept of the phenetic v a r i a t i o n of species and r e l a t i o n s h i p s between taxa . Table l a . The concepts of the s i x North American d i o i c o u s Brachythecium species w i t h rough setae, according to A.J. Grout (1928-19^+0, Moss F l o r a  of North America, Volume 3)? S. Flowers (1973? Mosses: Utah and the West) and E. Lawton ( 1971 i Moss F l o r a of the P a c i f i c Northwest.) CHARACTERS SPECIES GROUT FLOWERS LAWTON LEAF MARGIN SERRULATION <| ASPERRIMUM ; BOLANDERI FRIGIDUM HTLOTAPETUM NELSONII RIVULARE Branches, strongly ser-rate above Stem, less strongly • serrate, often nearly entire Stem leaves, margLns entire or finely serrate Branch leaves, nore sharply serrate Stem leaves, serrate at apex Branch leaves, usually strongly serrate Branch leaves, ser-rate a l l around Stem leaves, less distinctly serrate, often nearly entire Stem leaves, serrulate a l l around B ran ch le ave s , mo re strongly serrate Branch leaves denticu-late a l l around Stem leaves, serrate in upper H or well towards the base Stem, margins serrate to serrulate at the apex, serate to entire below Branch, serrate at apex often serrulate to below the middle Stem, entire Branch, serrate Stem leaves slightly serrulate at extreme apex Branch leaves often serrate above Stem leaves entire to distantly serrulate above Branch leaves, apex more strongly serrate denticulate to entire below Stem leaves, denticu-late Branch leaves, dentate above with small pointed teeth Stem leaves serrate with small distant teeth, denticulate toward the base, sometimes entire throughout Branch leaves, more strongly serrate Stem, serrulate MEDIAN LEAF CELLS ^ " ASPERRIMUM BOLANDERI FRIGIDUM HTLOTAPETUM NELSONII RIVULARE Linear, reaching 90 long, 10-15:1 Linear, 6-8 wide 45-125 X 6-9 Rhomboidal-fusiform 36 X 6 30-45 X 5-7 Linear to oblong 7-10:1 60-125 X 7-9 75-140 X 9-11 Much as in B. rivulare 60-90 X 6-10 rarely 100 10-15:1 75-125 X 8-11 Table l b . The concepts of the s i x North American d i o i c o u s Brachythecium species w i t h rough setae, continued. CHARACTERS SPECIES AUTHORS C R O W L A W T O H f ASPERRIMUM Ovate-lanceolate. Soulier, narrower Smaller, ovate lanceolate BOLAKDERI Ovate-lanceolate Smaller BRANCH LEAF SHAPES < FRIGIDUM Ovate tr> ovate-lanceolate Smaller, more narrowly ovate-lanceolate to elongata triangular Smaller HTLOTAPETUM Smaller NELSONII Similar to stem leaves but smaller And usually propor-tionately narrower Narrower to oaaller ^ elongate- triangular to ovate-lanceolate, the apex more slender Smaller RIVULARE Ovate to ovate-lanceolate Smaller, narrower, ovate-lfinceolate to ovate oblong Breadly ovate-lanceolate ASPERRIMUM Sotaawhat concave Stem, concave below Branch, less concave Coucsve BOLANDER1 Not concave • CONCAVITY OF LEAVLS •< FRIGIDUM Concave and excavate below HTLOTAPETUM Concave NELSONII Slightly concave with margins turned Inward toward the apex Concave at the base RIVULARE Strongly concave Strongly concave, excavate at the base Deeply concave r ASPERRIMUM Strongly plicate Plane to atrongly plicate Deeply plicate BOLANDERI Not plicate STEM LKAP PLICATION - FRICLDUM Vary atrongly plicate Slightly to atrongly Pllcatt Deeply plicate HTLOTAPETUM Rot, or only weakly plicate NELSONII Not plicate Not plicate RIVULARE More or l*t9fl p l i c a t e Sllghtlv or riot at a l l plicate Scarcely plicate Table l c . The concepts of the s i x North American d i o i c o u s Brachythecium species w i t h rough setae, continued. | CHARACTERS SPECIES AUTHORS GROUT FLOWERS LAWTON LEAF APEX SHAPE "S ASPERRIMUM BOLANDERI FRICIDUM KTLOTAPETUJi KELSONII RIVULARE Abruptly to gradually lc •nd narrowly acuminate n«]Gradually and slenderly acuminate, no tret lines abruptly long and slenderly cuspidate Acuminate Branch leaves gradually acuminate Lower, stem leaves l i t t l e different Acuminate Stem, mostly short acuminate, occasionally acute Acuminate Often abruptly aplculate Slenderly long-acuminate Usually rather shortly and abruptly acuminate, but sometimes long and •lender Slenderly long acuminata Acute to short acuminate Acute to shortly acuminate Stem, abruptly acuta or shortly acuminate Branch, acumen longer TWISTED OR FALCATE APEX < ASPERRIMUM BOLAKDERl FRICIDUM BTLOTAPETU! NELSONII RIVULARE Somewhat falcate •ccund Tips often twisted, straight or sometimes falcate ( ' \ ' Acuminate with ap«x twisted Tlpa often twisted, straight, or occasionally falcate-secund Twisted Tips often twisted / BRANCH LEAP PLICATION < 1 i ( I ASPERRIMUM BOLANDERI FRICItllM H Y L O T A f ETU MSLSONI1 K I V U L A R E Plicate Plane or slig h t l y pllcatt Plicate or plane Not plicit« Strongly plicate 1*88 strongly pllcstc than stem leaven Often so«*ewUaV plfcate Ofttiti lightly pl (rare 7-T a b l e I d . The c o n c e p t s o f . t h e s i x N o r t h A m e r i c a n d i o i c o u s B r a c h y t h e c i u m s p e c i e s w i t h r o u g h s e t a e , c o n t i n u e d . CHARACTERS SPECIES ' AUTH ORS CROUT FLOWERS LAWTON STEM LEAF SIZE <; ASPERRIMUM BOLANDERI S FRICIDUM HTLOTAPETUM NELSONII RIVULARE 1.5-2 mm. X 0 . 6-1 mn. , 1.8-3 X 0 . 6 - l . S mn. Upper reaching 1.2 mm. 0 . 6-1.2 X 0.4-0.6 mm.I 2-2.5 X 0.8-1.2 mm. 2-2.5 (3) mn. X 0.8-1.3 (1.6) ran. 2-3 X 0.8-1.5 ma. sometimes to 3.5 mm. 2.3-3 . 6 X 1.2-2ma. l,5-2ma. X 0.6 era. 1.5-2 ma. X 0.9-1.2 ma. 1.6-2.4 X 0.8-1.3 mm. Reaching 2 X 1.4 tm. 1.5-2.5 ma. X 0.8-1.3 ma. 1.6-2.8 X 1-1.5 mm. BRANCH LEAF SI ASPERRIMUM BOLANDERI 7.ES FRICIDUM KYLOTAPETU> NELSONII ^ RIVULARE 1 . 6 X 0.45-0.6 inn. Smaller and narrower 1-2 X 0.4-0.8 mm. 0 . 6-0.8 X 0.25-0.3 mm. Smaller than etem lvo 1.2 X 0 . 6 mm. Smaller and more narrowly ovate, lanceolate to elongate triangular Smaller then stem lvs Smaller then stem lvs 2-2.5 X 1-1.5 aa. Smaller and usually proportionately narrower Narrower and smaller Smaller Reaching 1.5 X 0.6mm. Smaller and narrower 1.2-1 . 6 X 0.5-1 mm. STEM LEAF SKAT < ASPERRIMUM BOLANDERI E FRICIDUM HYLOTAPETUI NELSONII ^ RIVULARE Varying from broadly ovate to elongated triangular-ovate Broadly ovate to ovate, lanceolate or elongate, triangular Broadly ovate, d e l t o i d ovate or ovate lanceolate Lover etem leaves • n a i l e r , upper reaching 1.2 mm. otherwise l i t t l e d i f f e r e n t than branch leaves Ovate-lanceolate Triangular-ovate Broad ovate to ovate, lanceolate Broadly ovate to deltoid-ovate Broadly-ovate lanceolate Triangular-ovate T y p i c a l l y d e l t o i d , varying to hroadly ovate-lanceolate Triangular-ovate Broadly ovate Ovate to triangular Deltoid-ovate 8 . Table l e . The concepts of the, s i x North American d i o i c o u s Brachythecium species w i t h rough setae, continued. j CHARACTERS SPECIES AUTHORS /«" CROUT FLOWERS LAWTON < SIZE, AREA, ALAR CE DECVRRENCT " ' '"' ASPERRIKL1 BOLANDERI rRICIDUM AS •i L i t t l e d i f f e r e n t i a t e d Few to numerous A row of short i l n f l a t e d c e l l s extending across the base attaching the leaf to stem Decurrent Few, a few shorter c e l l s running up the margin Not decurrent Not d i f f e r e n t i a t e d Ranging from a email group of 4-5 c e l l s to aa m«ny an 10 c e l l o across the base and along the margin, sometimes extending Into the decurrent wings Branches, alar c e l l s not particularly-d i f f e r e n t i a t e d Sometimes t Inflated Decurrent HYLOTAPETUM Dif f e r e n t i a t e d Somewhat decurrent NELSONII RIVULARE Large area Numeroua, forming auricles and usually extending S way or more to the costa Forming d e f i n i t e a u r i c l e s Decurrent Large area Stem, large area, decurrent Branches, fewer, - decurrent, often strongly to Decurrent a u r i c l e s < 1X.NCTH, STRJ3CTH COSTA ASPERRI Costate to beyond the middle Costa alender, S - 3/4 the length of the blade Strong to beyond middle of leaf BOLANDERI E « « n d l o g the middle Extending beyond the middle FRICIDU HTLOTAP NELSON! 1 Stout, extending 2/3 - 3/4 length of leaf Coata extending 3/4 - 4/5 the length of the blade, often Into the baBe of th acumen Strong, extending beyond the middle :TUM Usually to above the middle, sometimes shorter snd forked Stout at the bane rapidly narrowing 1 In the lower portion Rather wide at the base extending S- 3/4 the length o f the blade Extending above the middle 1 Coeta often forking RIVULARE S - 3/4 Che leaf length, occasionally forked Ending above the middle of the leaf Table l f . The concepts of. the s i x North American d i o i c o u s Brachythecium species w i t h rough setae, continued. CHARACTERS SPEClf.S AUTHORS CROUT FLOWERS , LAWTON HABITATS < "OlSPERRIKUM BOLAHDERI On moist rock* s o i l •nd decaying wood On decayed logs, bases of trees, and on damp s o i l or rocks, less commonly ln vet places Vat s o i l s , rotten logs tree trunks, pendent from branches-lowlands to 2,000 m. On shaded ground On s o i l FRICIDUM rTfLOTAPETUM NELSONII RIVULARE \ — -On decaying wood In moist places Crowing on wet or damp s o i l , humus or rotten wood on brook banks, around springs and In seepage areas, sometines submerged On s o i l or sand in very vet places, often In or near streams*-to about 3300 m. On s o i l , humus, rotten wood, and l i t t e r ln the f o r e s t , or in open places to about 2000 m. Crowing on humus On vet, damp, or rather dry s o l i , hunua and rotten logs, usually In shady places at high elevations In vet places in the mountains, 2,000 D. to 3,000 m. On stones ln beds of brooks and ln other places that are always nuilst and occasionally sub-merged but not always under water. Submerged in springs and slow flowing brooks, or growing on wet s o l i , rocks, humus or rotten logs, on brook bands, ln vet meadows and seepage areas from volleys to higher mountains Oi\ wet ground or atone* in or near strcama, lowlands to about 2,400 n. SIZE, (INFLATION) ALAR CELLS < ZJ 1 ASPERRIMUM BOLAHDERI FRICIDUM HYLOTAPET KELSONII RIVULARE i L i t t l e differentiated Stem leaves, not greatly enlarged or oblong and considerably en-larged , sonetimes Inflated Branch leaves, scarcely d i f f e r -entiated Sometimes ± inflated " ' — - Quadrate Not differentiated Gradually or suddenly enlarged, often Inflated Hyaline, quadrate to rectangular Short and broad, hyaline, sometiocs * Inflated Enlarged and Inflated Suddenly enlarged and usually Inflated Extreme alar c e l l s abruptly enlarged Abruptly enlarged and usually Inflated Plainly Inflated 10 • PART ONE: AN ANALYSIS OF THE PHENETIC VARIABILITY OF B^ FRIGIDUM (C. MUELL.) BESCH. SENSU ROBINSON. I . Data C o l l e c t i o n Data were c o l l e c t e d i n three steps, each of which i s d i s -cussed i n d e t a i l i n the f o l l o w i n g s e c t i o n s : (1.) the s e l e c t i o n of specimens, (2.) the choice of characters to be measured and, (3-) the measurement and s c o r i n g of c h a r a c t e r s . (1.) 1. The S e l e c t i o n of Specimens The sample chosen f o r a numerical taxonomic study should i d e a l l y i n c l u d e specimens e x h i b i t i n g the range of the species' morphological v a r i a b i l i t y . In order to achieve such a sample, specimens were chosen from a wide range of h a b i t a t s and locations... The r e s u l t a n t -sample i s shown i n Table 2 (page l l ) . When s e v e r a l c o l l e c t i o n s from a s p e c i f i c h a b i t a t and l o c a t i o n were a v a i l a b l e , recent c o l l e c t i o n s were p r e f e r r e d to older c o l l e c t i o n s as a source of i n t a c t m a t e r i a l s u i t a b l e f o r d i s s e c t i o n . Specimens c o l l e c t e d by s e v e r a l c o l l e c t o r s were.preferred over s e v e r a l c o l l e c t i o n s by the same c o l l e c t o r since the taxonomic conceptions or b i a s e s of a c o l l e c t o r w i l l i n f l u e n c e h i s perception of taxa i n the f i e l d . See Appendix A f o r a complete l i s t of the s i x t y - e i g h t specimens chosen. 11. Table 2 . The h a b i t a t s and geographic areas represented i n the sample of s i x t y - e i g h t OTU's chosen f o r data a n a l y s i s . (See Appendix A f o r a complete l i s t of specimens.) [ Mexico • Wyoming Montana Alberta Nevada Utah Idaho .Alaska California British Columbia Washington Oregon v ery wet areas wet rocks wet s o i l wet wood wood near water r o t t i n g wood epiphyte d r i e r rocks d r i e r s o i l maritime rocks w 1 'M -\ i w m is 1 'H ESS i (1 . ) 2 . The Choice of Characters to Be Measured Sneath and Sok a l , (1973, page 9 0 ) , recommend the use i n numer-i c a l taxonomy of "kinds of characters from a l l p a r t s of the body and from a l l stages of the l i f e c y c l e . Second, use a l l c h a r a c t e r s v a r y i n g w i t h i n the group s t u d i e d , not merely conventional d i a g n o s t i c c h a r a c t e r s . " In mosses, t h i s would e n t a i l the use of chara c t e r s p e r t a i n i n g to the spores, protonema, gametophore and sporophyte.' However, they a l s o s t r e s s the use of characters which can be unambiguously a p p l i e d i n the i d e n t i f i c a t i o n of specimens. I would the r e f o r e h e s i t a t e to i n c l u d e i n f o r m a t i o n on spores or protonema, and would p r e f e r the use of game-to p h y t i c f e a t u r e s over sporophytic f e a t u r e s , p a r t i c u l a r l y i n analyses of d i o i c o u s s p e c i e s , which f r e q u e n t l y l a c k sporophytes. Among the 12. a v a i l a b l e gametophytic f e a t u r e s , c e r t a i n f e a t u r e s are known to vary w i t h s u b s t r a t e c o n d i t i o n s and/or are d i f f i c u l t to q u a n t i f y , e.g. r h i z o i d placement, growth form (branching p a t t e r n and l e n g t h of branche and l e n g t h of i n t e r n o d e s (the number of le a v e s per centimeter of stem ) Such f e a t u r e s were not chosen. E s s e n t i a l l y nonvariable c h a r a c t e r s , e.g. l e a f c o n c a v i t y , margin s e r r u l a t i o n and the degree of l e a f t a p e r i n g , were a l s o e l i m i n a t e d . The remaining c h a r a c t e r s are a l l measurements of microscopic f e a t u r e s of the l e a v e s . These are l i s t e d and d e f i n e d i n the f o l l o w i n g s e c t i o n . (1.) 3- The Measurement and Scoring of Characters The o p e r a t i o n a l taxonomic u n i t s , (OTU'S), i n t h i s i n v e s t i g a -t i o n were i n d i v i d u a l shoots s e l e c t e d from each specimen. In order to more completely assess the v a r i a t i o n of each OTU, data were taken from both stem and branch l e a v e s . Although the d i s t i n c t i o n between branch and stem l e a f f e a t u r e s i s commonly made i n Brachythecium taxonomy, the terms "branch" and "stem" are r e l a t i v e to each other s i n c e branches integrade developmentally and m o r p h o l o g i c a l l y w i t h stems. For the purposes of t h i s i n v e s t i g a t i o n , a stem i s any s u b d i v i s i o n of the shoot a x i s which bears branches,.while a branch i s defined as any unbranched s u b d i v i s i o n . The e n t i r e v a r i a t i o n , from younger (branch), leaves to o l d e r (stem), l e a v e s was considered to be d i a g n o s t i c , r a t h e r than the f e a t u r e s of stem and branch leaves considered i n d i v i d u a l l y . Shoots were immersed i n hot water and the branches removed from the stem. Leaves were s t r i p p e d from the stem and from s e v e r a l s i z e s of branches and mounted while wet i n Hoyer's medium. Characters 13. of f i v e stem and f i v e branch leaves were measured. The same set of characters was measured f o r both stem and branch leaves. Three types of data were taken: continuous ( q u a n t i t a t i v e ) data, ranked data, and dichotomous data.(the presence or absence of a t r a i t ) . Tables 3 a . -3 c- (pages 13 -15-) present a complete l i s t of the cha r a c t e r s , the method of measuring continuous data and the c r i t e r i a used i n de c i d i n g between the character s t a t e s of the ranked and dichotomous data. Table 3a. Measurement of Characters: c r i t e r i a used, type and scoring-of data. Type of data and sc o r i n g method Character Method of measuring or c r i t e r i a used . ^ u^ iritJ.±.a_Li_ve (continuous) data: l e a f l e n g t h leaves were measured from the attachment expressed i n micrometers of the co s t a to the l e a f apex. l e a f width measured at the widest part of the l e a f , u s u a l l y at or j u s t above the l e a f base len g t h of cos t a measured from point of attachment of the costa to the t i p of the costa,- expressed as a r a t i o of c o s t a l e n g t h : l e a f l e n g t h median l e a f c e l l l e n g t h measured along the longest a x i s of the c e l l 1 ' median l e a f c e l l width measured at the widest part of the c e l l Ik.. Table 3 b. Measurement of c h a r a c t e r s : c r i t e r i a used, type and s c o r i n g of d a t a , continued' Type of data and s c o r i n g method Character Method of measuring or c r i t e r i a u^ed Ranked data: i n d i v i d u a l leaves were scored as having one of s e v e r a l character s t a t e s . l e a f shape leaves were scored as one of s i x types of l e a f shape: 1. broadly d e l t o i d ( f i g . l a ) 2. d e l t o i d ( f i g . l b ) 3. broadly ovate ( f i g s , l c , Id) k. ovate ( f i g s , l e , l f . ) 5. narrowly ovate ( f i g s . 2g, 2h, 2i, 2j) 6. l a n c e o l a t e ( f i g s . 2 k, 21, 2m) l e a f p l i c a t i o n leaves were scored as having one of f i v e types of p l i c a t i o n : 1. plane ( f i g s . 3a, 3b, 3c) 2. s l i g h t l y w r i n k l e d ( f i g s . 3d, 3e) 3- s t r o n g l y w r i n k l e d ( f i g . 3f) k. b i p l i c a t e - w i t h two p l i c a t i o n s (one each side of the lamina ( f i g s . 3h, 31) sometimes w i t h only one ( f i g . 3g) 5. s t r o n g l y p l i c a t e : w i t h two or more p l i c a t i o n s on each side of the c o s t a ( f i g s 3j, 3k) shape of l e a f apex l e a f apices were scored as one of four shapes: 1. long and slender ( f i g . • ka) j i 2. long-acuminate ( f i g s . • 4b, kc) j 15. Table 3 c. Measurement of characters: c r i t e r i a used, type and s c o r i n g of data , continued. Type of data and s c o r i n g method Character Method of measuring or c r i t e r i a used Ranked data: (continued) shape of l e a f apex (continued) 3. short acuminate ( f i g s . kd, ke) k. acute ( f i g . kf) Dichotomous characters (equivalent to presence-absence data) apex . f a l c a t i o n apex w i t h a d i s t i n c t curve, or f a l c a t e ( f i g . 5a, 5b) scored as 1.; apex s t r a i g h t scored as 0. apex t w i s t i n g apex t w i s t e d ( f i g . 5c) scored as 1, apex s t r a i g h t scored as 0. margin r e f l e x e d leaves w i t h two marginal . p l i c a t i o n s ( r e g a r d l e s s of other p l i c a t i o n s ) ( f i g . l f ) scored as 1. leaves l a c k i n g r e f l e x e d margins scored as 0. -decurrency of leaves leaves w i t h a prominent., decurrent area of en-l a r g e d a l a r c e l l s scored as 1. leaves l a c k i n g decurrencies scored as 0. •16. F i g . 1. Character s t a t e s of l e a f shape used i n the s c o r i n g of ranked data, (a) broadly d e l t o i d ; (b) d e l t o i d ; ( c , d) broadly ovate; (e, f ) ovate, continued on the next page ( f i g u r e 2). F i g . 2. Character s t a t e s of l e a f shape used i n the s c o r i n g of ranked data, continued. (g, h, i , . j ) narrowly ovate; (k, 1, m) l a n c e o l a t e . F i g . 3. Character s t a t e s of the l e a f p l i c a t i o n used i n the s c o r i n g of ranked data. (a, b, c) plane; (d, e) s l i g h t l y w r i n k l e d ; ( f ) s t r o n g l y w r i n k l e d ; (g, h, i ) b i p l i c a t e ; ( j , k) s t r o n g l y p l i c a t e . 19. ( f ) acute F i g . 5. Features of the l e a f apex used i n s c o r i n g dichotomous data, (a, b) f a l c a t e ; (c) t w i s t e d . 20. I I . A n a l y s i s of Data (2.) .1. General d i s c u s s i o n of methods P r i o r to the a c t u a l a n a l y s i s of data, the f i v e values measured f o r each O.T.U. f o r each of the stem and branch l e a f t r a i t s were averaged. The r e s u l t i n g data matrix c o n s i s t e d of 2k v a r i a b l e s (12 stem l e a f t r a i t s and 12 branch l e a f t r a i t s ) measured f o r 68 O.T.U.'s. The data were analysed by two methods: p r i n c i p a l components a n a l y s i s and c l u s t e r a n a l y s i s . Both methods s i m p l i f y the es t i m a t i o n of phenetic s i m i l a r i t i e s among O.T.U.'s according to t h e i r d i s t r i b u t i o n i n phenetic space. P r i n c i p a l components a n a l y s i s was used to study continuous v a r i a t i o n among the O.T.U.'s; c l u s t e r a n a l y s i s was used to study discontinuous v a r i a t i o n . O r d i n a t i o n may be c a r r i e d out on e i t h e r raw or standardized data, depending on the nature of the o r i g i n a l ..variables.. The raw data were standardized as part of the o r d i n a t i o n program to produce a mat r i x of c o r r e l a t i o n c o e f f i c i e n t s (see Sneath and Sokal 1973, page 2k5), or the data were standardized according to the method of Gower and then used to produce a covariance matrix. In simplest terms, the-second method takes each value i n . t h e columns of the.data m a t r i x , (columns correspond to v a r i a b l e s ) , s u b t r a c t s the sma l l e s t value i n the column, then d i v i d e s by the column range. The r e s u l t s of the o r d i n a t i o n s obtained using both types of s t a n d a r d i z a t i o n s were- e s s e n t i a l l y s i m i l a r . . In the d i s c u s s i o n of the r e s u l t s which f o l l o w s ( S e c t i o n (2) 2.), the type of matrix ( c o r r e l a t i o n or covariance) 2 1 . used f o r each o r d i n a t i o n w i l l be s p e c i f i e d . For a complete d i s c u s s i o n of p r i n c i p a l components a n a l y s i s , the reader i s r e f e r r e d to Sneath and Sokal (1973, pages 2^5 - 2^7). P r i n c i p a l components a n a l y s i s i s a commonly employed type of o r d i n a t i o n . An o r d i n a t i o n i s the arrangement of u n i t s ( i n t h i s case O.T.U.'s) i n an A-space of n dimensions (or axes), where n = the number of v a r i a b l e s . Each O.T.U. w i l l have a unique combination of values f o r each v a r i a b l e ( t r a i t ) , and i t s p o s i t i o n i n the A-space w i l l depend upon that unique combination. O.T.U.'s which are more s i m i l a r w i l l . b e c l o s e r together i n the A-space. Arranged i n t h i s manner, the O.T.U.'s may be thought of as forming a f o o t b a l l - s h a p e d cloud of p o i n t s , each po i n t r e p r e s e n t i n g a-separate O.T.U. The f i r s t o r d i n a t i o n a x i s , c a l c u l a t e d by p r i n c i p a l components a n a l y s i s , i s the l i n e through the longest a x i s of the cloud of p o i n t s . T h i s l i n e - i s the f i r s t eigenvector which i s a measure - of the great e s t v a r i a b i l i t y among the O.T. U.'s The second eigenvector i s perpendicular (orthogonal) to the f i r s t eigenvector, and i t represents the second l a r g e s t amount of v a r i a t i o n among the O.T.U.'s.. By p l o t t i n g the p o s i t i o n s of O.T.U.'s along the f i r s t three o r d i n a t i o n axes, taken i n p a i r s , c onsiderable i n s i g h t i n t o the main taxonomic r e l a t i o n s h i p s can be gained. In c l u s t e r a n a l y s i s , the set of p o i n t s r e p r e s e n t i n g the O.T.U.'s i s p a r t i t i o n e d i n t o groups of p o i n t s based upon the.closeness (phenetic s i m i l a r i t y ) of the p o i n t s i n phenetic space. I n i t i a l l y , the computer 2 2 . creates c l u s t e r s of p a i r s of p o i n t s . The mathematical centers of these two-point c l u s t e r s are then c a l c u l a t e d , and these centers become new p o i n t s which are then p a i r e d to other p o i n t s or centers of p o i n t c l u s t e r s . This process i s continued u n t i l a l l the p o i n t s i n the phenetic space are inc l u d e d i n one l a r g e c l u s t e r . The e a r l i e s t c l u s t e r s of p o i n t s are taxonomically- the most s i g n i f i c a n t since they represent those p o i n t s which are c l o s e s t i n phenetic space. Later c l u s t e r s are s u c c e s s i v e l y l e s s s i g n i f i c a n t as the distance between c l u s t e r s , and the number of p o i n t s per c l u s t e r , i n c r e a s e s . Various c l u s t e r i n g methods d i f f e r i n the method of computing the di s t a n c e s between p o i n t s or c l u s t e r s , and i n the c l u s t e r i n g a l g o r i t h m (the r u l e s by which c l u s t e r s are formed). The c l u s t e r a n a l y s i s program used i n t h i s i n v e s t i g a t i o n employs a Eu c l i d e a n distance measure and Ward's Method (or Minvar) c l u s t e r -i n g a l g o r i t h m . A complete d i s c u s s i o n of t h i s program i s contained i n the U n i v e r s i t y of B r i t i s h Columbia's Computing Center Manual "UBC C GROUP: H i e r a r c h i c a l Grouping A n a l y s i s w i t h Optional C o n t i g u i t y C o n s t r a i n t " ( O s t e r l i n , P a t t e r s o n and Whitaker, November, 1971> revised:' March 1978) which may be obtained from the U.B.C. Computing Center. For a Complete d i s c u s s i o n of the d i f f e r e n t methods of c l u s t e r a n a l y s i s , the reader i s r e f e r r e d to Sneath and Sokal (1973, pages 201 - 2^5). 23. ( 2 . ) 2 . R e s u l t s of the Data A n a l y s i s The f i r s t o r d i n a t i o n was performed on a covariance matrix of the f u l l set of the 68 O.T.U.'s (see Appendix B. f o r the covariance matrix and the mean, variance and standard d e v i a t i o n of the 2k v a r i a b l e s ) . F i g u r e 6 (page 2k) shows the s c a t t e r p l o t of the f i r s t and second axes ( e i g e n v e c t o r s ) . The f i r s t a x i s accounts f o r 37.69% of the t o t a l v a r i a t i o n of the O.T.U.'s. The second a x i s accounts f o r 8.79%- The numbers on the s c a t t e r p l o t correspond to the num-bers assigned to the specimens l i s t e d i n Appendix A. The s c a t t e r p l o t i n Figure 6 i n d i c a t e s a separation of O.T.U.'s i n t o two l o o s e l y defined groups. In order to v e r i f y and define t h i s s e paration f u r -t h e r , a c l u s t e r a n a l y s i s of.the same data was executed. The r e s u l t s of t h i s c l u s t e r a n a l y s i s , are shown i n the phenogram on page 25 ( F i g . 7 ) . The phenogram shows a p a r t i t i o n i n g of the 68 O.T.U.'s i n t o two groups which correspond e s s e n t i a l l y to the separation shown i n F i g . 6 . The combined evidence of the o r d i n a t i o n and the phenogram i n d i -cate that a p o s s i b l e taxonomic se p a r a t i o n could be recognized be-tween O.T.U.'s (32, k?>, k8, 5 0 , 3k, 55 , 56, 57, 59 , 61, 6^, 6 5 , 66,. 67, 68) and the remaining O.T.U.'s. The 68 specimens were then reexamined and compared to the type c o l l e c t i o n s of the two spec i e s . The former group of 15 specimens were considered to r e -present Brachythecium asperrimum ( M i t t . ex. C. Muell.) S u l l . and the l a t t e r group of remaining specimens to represent B. f r i g i d u m (C. Muell.) Besch.. 2h. 38 • 14 e •40 0 5 9 • 5 e 3 3 061 • 58 - « c 9 •17 • 53 •12 • 3 7 •19 , *0 7oe-ie •15 #44 • 39 0 6 3 •25 22 P«28 •41 .10 « M 9 e 4 6 r w | axisl -1 °64 J, h •23 0 5 4 . 6 065 0 5 5 0 5 6 ©66 • 36 « o 5 7 «42 n O50 043 •1 0 4 7 » 2 9 • f o o 6 0 - ^ w .16 »1 •3 • 51 •35 «4 • 52 •2 F i g . 6 . The f i r s t and second axes of the f i r s t o r d i n a t i o n of 68 O.T.U.'s o = B. asperrimum ( M i t t . ex. C. Muell.) S u l l . ; o = B. frigidum-(C. Muell.) Besch. 25---450 --100 --50 + 25 00 LU u 4-20 5 I— 00 4-15 5 u i u i ( J u i Oi -N Ul Ol cn ro 8! Iii O 01 03 .vl LO ro ro Ui ro U> Ul U / & Ul Ui O i ro O ro U l ro "ro s» CD • - u l W 4 i -r» ro " O U . ^ - CD JO u> U l XJ JO o> o 01 ' ro 0J O i .U 1 .ro k ,U> UJ xo O T U ' S F i g . 7. The r e s u l t s of a c l u s t e r a n a l y s i s of 68 O.T.U.'s of B. fr i g i d u m (C. Muell.) Besch. Distance along the v e r t i c a l s c a l e r e f e r s to taxonomic d i s t a n c e . Groups of two or more O.T.U.'s along the base of the dendrogram i n d i c a t e specimens considered to be i d e n t i c a l by the c l u s t e r a n a l y s i s . 26. Brachythecium asperrimum ( M i t t , ex. C. Muell.) S u l l . and Brachythecium  f r i g i d u m (C. Muell.) Besch. were formerly separated on the b a s i s o f hab-i t a t and growth form (Lawton, 1971)- Growth form o f t e n v a r i e s according to h a b i t a t (or s u b s t r a t e ) . I f the two taxa were each r e s t r i c t e d to s p e c i f i c h a b i t a t s , and showed an i n t e r g r a d a t i o n of gametophytic fea t u r e s from one h a b i t a t type to another, they could then be con-s i d e r e d to represent h a b i t a t forms ( or ecomorphs) of the same t a x o n As Figure 8 (page 28) shows, phenetic v a r i a t i o n of the two taxa r e v e a l s no i n t e r g r a d a t i o n o f f o r m s f r o m one h a b i t a t t y p e to another. Each species i s represented from a range of h a b i t a t s , and O.T.U.'s of the two species from the same h a b i t a t are p h e n e t i c a l l y d i s t i n c t . Therefore i t appears that the two taxa c a n n o t c o n s i s t -e n t l y or r e l i a b l y be separated on the b a s i s of h a b i t a t and growth f o r m a l o n e . I n order to devise a new means of separating the two ta x a , the phenetic v a r i a t i o n of each taxon must be analysed. A key to the two taxa then may be based on more r e l i a b l e d i a g n o s t i c f e a t u r e s . In order to f u r t h e r define and c l a r i f y the phenetic v a r i a t i o n and h a b i t a t s p e c i f i c i t y of each sp e c i e s , the two groups of O.T.U.'s repr e s e n t i n g the two taxa were o r d i n a t e d s e p a r a t e l y . Both o r d i -n a t i o n s were based on a c o r r e l a t i o n m atrix. (See Appendix C, Table 1) f o r the c o r r e l a t i o n matrix of the 15 O.T.U.'s of Brachythecium asperrimum ( M i t t . ex. C. Muell.) S u l l . The f i r s t a x i s accounts f o r 24.26% of the v a r i a t i o n . The second a x i s accounts f o r 19-88% of the v a r i a t i o n . The h a b i t a t of each O.T.U. i s i n d i c a t e d . V a r i a t i o n along the second a x i s , and to some extent along the f i r s t a x i s , has c o n t r i b u t e d to a somewhat d i s t i n c t s e paration of the p o i n t s i n t o two groups. R o t t i n g wood and epiphyte specimens are l o c a t e d p r i m a r i l y i n the upper two quadrants, while s o i l . a n d rock specimens are l o c a t e d i n the lower two quadrants. Table k (page 30) shows the c o n t r i b u t i o n of the v a r i a b l e s to the v a r i a t i o n along the f i r s t three component axes ( e i g e n v e c t o r s ) . The f i v e major c o n t r i b u t i o n s to the v a r i a -t i o n along the second a x i s are the branch and stem l e a f median c e l l l e n g t h and width and the branch l e a f a l a r c e l l s . The •dimensions of median l e a f c e l l s are extremely, v a r i a b l e , even w i t h i n a s i n g l e . l e a f , i n a l l Brachythecium species. Ex-2 8 . • O R • V W • W S o R W o W S «RV» o W S x OJ l 1 ^ W N W . : w o o d n e a r w a t e r W W : w e t w o o d R W r r o t t i n g w o o d E : . epiprtyte W R : w i t r o c k s # W R W S : w e t s o i l V W : v e r y w e t a r e a s , W f t M R : m a r i t i m e r o c k s > w j » D R : d r i e r r o c k s O S : d r i e r s o i l s •WR •VW • Rfcy •W5 A . v w *«vvR • W W W W vw».y** , "WWW ••vw , • » nvR « v ^ w >^/6 a y f e 1 | ORVV O R W o£ »WR «AS ° E °WR o D S •vw o E «WNW °OR • 1 •WR ' «VW • W R * W R • •VW »vw •vw 1 ^ • V W • W R • W R * « * W R --1 .WS Fig.. 8 . The f i r s t and second axes of the o r d i n a t i o n of 68 O.T.U.'s showing the h a b i t a t s of each O.T.U.. o = B. asperrimum ( M i t t . ex. C.Muell.) S u l l . ; • = B. fr i g i d u m (C. Muell.) Besch. 29. •RW W N W . : wood near wat*r W W : wet wood RW:rottr>g wood E : «pcr<yta W R : wet rocks W5:wet soil V W : very v.et areas MR:maritime rocks D R : drier rocks O S : drier soi I s • O S • WR «VYS F i g . 9. The f i r s t and second axes of the o r d i n a t i o n of B. asperrimum ( M i t t . ex. C. Muell.) S u l l . showing the h a b i t a t s of each O.T.U. 3 0 . Table 4. The c o n t r i b u t i o n of the v a r i a b l e s to the v a r i a t i o n shown along the f i r s t three major components (eigenvectors) i n the o r d i n -a t i o n of B. asperrimum ( M i t t . ex. C. Muell.) S u l l . COMPONENTS 1 2 ' 3 LEAF LENGTH 0.2140 -0.1047 - 0 . 2 3 6 8 LEAF WIDTH 0.3480 -O.O541 . -0.1757 COSTA RATIO 0.2530 -0.1969 -0.3224 MEDIAN CELL LENGTH -0.1515 0.3585 -0.2136 STEM MEDIAN CELL WIDTH . 0.1517 0.3020 0.1129 LEAF LEAF SHAPE O .366I - 0 . 0 9 5 0 O .0060 TRAITS ACUMINATION 0.2878 - 0 . 0 0 0 8 0.0764 MARGIN REFLEXED -0 .0346 -0 .0555 0.0655 PLICATION 0.1454 ' -O.O329 0.0679 APEX TWIST 0.2361 0.1358 - 0 . 0 3 5 8 APEX FALCATE 0.0130 0.1394 0.3178 ALAR CELLS 0.0267 0.0151 - 0 . 4 3 8 4 LEAF LENGTH 0.2165 0.1943 -0.1751 LEAF WIDTH 0.2878 0.2474 0.1084 COSTA RATIO 0.2948 -0.1108 0.0733 MEDIAN CELL LENGTH - 0 . 0 6 2 2 0.3556 -0.1818 BRANCH MEDIAN CELL WIDTH 0.0739 0.3060 0.1070 LEAF LEAF SHAPE 0.1775 0.2051 0.3265 TRAITS ACUMINATION 0.2726 - 0 . 0 9 3 3 ' - 0 . 0 6 9 4 MARGIN REFLEXED 0.0377 0.1478 - 0 . 2 6 5 9 PLICATION 0.2034 0.2864 0.1776 APEX TWIST 0.1849 -0 .1343 0.2088 APEX FALCATE 0.0492 •' 0.2614 - 0 . 3 0 0 6 ALAR CELLS -0.1268 0.3224 - 0 . 0 3 9 5 31. cept i n cases where a d i s t i n c t d i f f e r e n c e e x i s t s , the median l e a f c e l l dimensions are probably more a source of confusion than of taxonomically d i a g n o s t i c i n f o r m a t i o n . A d d i t i o n a l c o n t r i b u t i o n s to the v a r i a t i o n along the second a x i s , and to the v a r i a t i o n along the f i r s t a x i s , are made by the le n g t h , width, shape and acumination of the stem and branch leaves. This o r d i n a t i o n i s based on f a r too narrow a sample to be c o n c l u s i v e , yet i t may be a p r e l i m i n a r y i n d i c a t i o n of the smaller and. f i n e r nature of e p i p h y t i c and r o t t i n g wood specimens when compared to specimens found on s o i l and r o c k s . Figure 10 (page 32) shows the f i r s t and second axes of the o r d i n a t i o n of B. fr i g i d u m (C. Muell.) Besch. The f i r s t a x i s accounts f o r 19.^6% of the v a r i a t i o n . The second a x i s accounts f o r 10.62% of the v a r i a t i o n . There i s no apparent phenetic separa-t i o n between the O.T.U.'s found on va r i o u s types of h a b i t a t s . The primary purpose of the separate o r d i n a t i o n s of B. asperrimum ( M i t t . ex. C. Muell.) S u l l . and B. fr i g i d u m (C. Muell.) Besch. was to assess the v a r i a t i o n i n the t r a i t s of the i n d i v i d u a l s p e c i e s . The v a r i a b l e s which show the highest c o n t r i b u t i o n to the v a r i a t i o n along the f i r s t a x i s of the o r d i n a t i o n of 68 O.T.U.'s ( r e f e r to Table k, page 23) are those t r a i t s which account f o r the phenetic s e p a r a t i o n between the two sp e c i e s . I f these same t r a i t s show a reduced c o n t r i -b u t i o n to the v a r i a t i o n i n the o r d i n a t i o n of the i n d i v i d u a l s p e c i e s , the t r a i t s can. then be considered more s t a b l e w i t h i n each s p e c i e s . Characters which vary between species and are l e s s v a r i a b l e among i n d i v i d u a l species are p o t e n t i a l l y u s e f u l d i a g n o s t i c f e a t u r e s . 32. •WR -0.5 • W S • R W • W R • W S vw« •www •RW • V W •ws • W M W • W W • W S • V W • W R w o " V W • W R •wS •OR •MR • W W • V W •WW •RW -0.5+ •vw • V W • W R v f * * ! ^ »VW •vw WNW.: wood near wattr WW: wet wood RW: rotting wood E:.epcryte WR: wet rocks WS:wet soil VW^very wet areas MR: maritime rocks DR: drier rocks DS: drier soi I s • W R • W R ..Mil. • V W • V W W S f + Q S • W S • W S *WR • W R • V W •vw • W R • W R • W R »WR •vw F i g . 1 0 . The f i r s t and second axes of the B. fri g i d u m (C. Muell.) Besch: o r d i n a t i o n , showing the h a b i t a t s of the specimens. A comparison of the c o n t r i b u t i o n s of the v a r i a t i o n along the f i r s t a x i s of the three o r d i n a t i o n s r e v e a l s c e r t a i n f e a t u r e s which show a r e -duced c o n t r i b u t i o n to the v a r i a t i o n of one or both of the i n d i v i d u a l species o r d i n a t i o n s (Tables k and 5 , pages JO and 3 3 ) when compared to the o r d i n a t i o n of the two species together (Table 6, page 3 ^ ) . Of par-t i c u l a r i n t e r e s t are the stem and branch l e a f p l i c a t i o n s and a l a r c e l l s s ince the l i t e r a t u r e r e p o r t s of these t r a i t s i n the two species are par-t i c u l a r l y c o n t r a d i c t o r y ( r e f e r to Tables l b , l c , l e , l f , pages 5 t 6 , 8 , 9)-3 3 " . Table 5. The c o n t r i b u t i o n of the v a r i a b l e s to the v a r i a t i o n shown along the f i r s t three major components (eigenvectors) i n the o r d i n -a t i o n of B. fr i g i d u m (C. Muell.) Besch. COMPONENTS 1 2 3 LEAF LENGTH 0.3589 -0.1655 -0.1552 LEAF WIDTH O.3687 0.1546 -0 .1380 COSTA RATIO 'O.I568 0.2425 -0.1771 MEDIAN CELL LENGTH 0.2906 -0.1013 . -0 .1485 STEM MEDIAN CELL WIDTH 0.0588 O.3667 -O.0521 LEAF LEAF SHAPE 0.3590 - 0 . 0 8 7 3 0.0201 TRAITS ACUMINATION 0.0774 0 . 3 6 4 9 - 0 . 2 0 4 4 MARGIN REFLEXED - 0 . 0 6 4 0 - 0 . 0 4 4 3 0.1132 PLICATION 0.2416 - 0 . 0 4 6 5 0.2252 APEX TWIST 0.0325 -0 .0359 - 0 . 0 4 6 1 APEX FALCATE -0.0132 - o . 2 4 o 6 -0.1688 ALAR CELLS 0.2468 0.0797 - 0 . 0 6 2 2 LEAF LENGTH 0.3394 -0.1557 0.0204 LEAF WIDTH O.2851 0.0640 0.0445 COSTA RATIO 0.0747 0.0598 - 0 . 2 2 9 3 MEDIAN CELL LENGTH 0.2516 -0.2871 -0 .2097 BRANCH MEDIAN CELL WIDTH O.0399 0.4019 0.0352 LEAF LEAF SHAPE 0.1818 - 0 . 2 0 9 8 0.3221 TRAITS ACUMINATION 0.1068 0.3644 - 0 . 0 5 8 5 MARGIN REFLEXED O.0820 0.1216 0.4003 PLICATION 0.1617 0.0227 0.4870 APEX TWIST O.0763 0.0869 0.1186 APEX FALCATE -0 .1009 - 0 . 2 3 3 3 -0.1335 ALAR CELLS 0.0543 0.0698 0.3519 3 4 . Table 6 . The c o n t r i b u t i o n of the v a r i a b l e s to the v a r i a t i o n shown along the f i r s t three major components (eigenvectors) i n the o r d i n -a t i o n of the 68 O.T.U.'s (combined specimens of B. asperrimum ( M i t t . ex. C. Muell.) S u l l . and B. frigid u m (C. Muell.) Besch.-COMPONENTS 1 2 3 LEAF LENGTH 0.0961 - 0 . 3 9 6 8 -0.1055 LEAF WIDTH 0.1639 -0.2213 0.0072 COSTA RATIO 0.1894 -0.0479 -0.0397 MEDIAN CELL LENGTH 0.1593 -0.2614 -O.0323 STEM MEDIAN CELL WIDTH 0.1423 0.0513 O.0625 LEAF LEAF SHAPE 0.1583 -0.1661 - 0 . 0 6 0 1 TRAITS ACUMINATION 0.2185 0.0807 - 0 . 0 0 3 0 MARGIN REFLEXED O.I36O 0.2722 0.1912 PLICATION 0.3171 -0 .0236 0.1770 APEX TWIST 0.1801 O.1167 -0 .5407 APEX FALCATE - 0 . 0 0 0 7 -0.1250 0.0414 ALAR CELLS 0.3633 -0.1852 O.2038 LEAF LENGTH O .I787 -O.3817 - 0 . 0 7 3 3 LEAF WIDTH 0.1652 -0.1473 -O.0615 COSTA RATIO 0.1487 -O.0326 - 0 . 0 3 5 2 MEDIAN CELL LENGTH O.1560 -0 .3664 0.0211 BRANCH MEDIAN CELL WIDTH O.1678 0.1016 - 0 . 0 0 8 4 LEAF LEAF SHAPE 0.1044 - 0 . 0 3 3 3 -0.0157 TRAITS ACUMINATION 0.2505 0.1371 - 0 . 0 0 8 5 MARGIN REFLEXED 0.2744 O.2511 0.3751 PLICATION O.3108 0.1314' 0.0555 APEX TWIST 0.2458 0-2593 -0.6167 APEX FALCATE -©.•0601 •• -0 .0472 0.1574 ALAR CELLS 0.2876 0.2554- : O.1232 3 5 . Once the p r e l i m i n a r y evidence of phenetic s e p a r a t i o n of taxa i s obtained, the c o n s t r u c t i o n of a taxonomic scheme to i d e n t i f y specimens and r e l i a b l y separate the taxa remains. Before t h i s can be'accomplished, c e r t a i n questions need to be answered: (1.) What de f i n e s a species?. 5 (2.) At what l e v e l of confidence does the taxonomist want to make statements about the taxa? Since taxonomy i s the foundation of many other s c i e n t i f i c endeavors, i n c l u d i n g ecology, f l o r i s t i c s and phytogeography, the taxonomic scheme should be based on a working species d e f i n i t i o n . , which i s of p r a c t i c a l a p p l i c a t i o n i n many areas of res e a r c h . P a r t i c u l a r l y i n botany, the concept of a b i o l o g i c a l species i s of l i t t l e or no p r a c t i c a l value. Species are d i s t i n g u i s h e d by e x i s t i n g p h y s i c a l f e a t u r e s , not by breeding c a p a c i t i e s or presumed (or demon-st r a t e d ) l i n e s of o r i g i n and descent. In other words, we r e l y on phenetic, r a t h e r than genetic or phylogenetic, evidence to define and recognize s p e c i e s . The most u s e f u l d e f i n i t i o n w i l l i n c l u d e microscopic and macroscopic f e a t u r e s to provide f o r the r e c o g n i t i o n of a species both i n the l a b o r a t o r y and i n the f i e l d . For a l l s p e c i e s , e s p e c i a l l y v a r i a b l e s p e c i e s , a nucleus of s e v e r a l char-a c t e r s i s p r e f e r r e d to a s i n g l e c h a r a c t e r . A working species concept would then be: a b i o l o g i c a l e n t i t y w i t h a unique combin-a t i o n of s e v e r a l f e a t u r e s which i s r e c o g n i z a b l e nine times out of ten i n the f i e l d . A 90% confidence l e v e l (or nine times out of / ten) i s an a r b i t r a r y g u i d e l i n e . I d e a l l y , a 100% confidence l e v e l i s o p t i m a l , but r a r e l y a c h ievable. However, i f i n d i v i d u a l I I ' ' . • • ' . j I 3 6 . d i a g n o s t i c f e a t u r e s can be demonstrated.at a 90% confidence l e v e l or h igher, then the combination of three or more of these f e a t u r e s w i l l approach a 100% confidence l e v e l . I n d i v i d u a l specimens may l a c k one or more f e a t u r e s , but w i l l s t i l l be r e c o g n i z a b l e by the remaining f e a t u r e s . In order to t e s t the r e l i a b i l i t y of t r a i t s considered to be d i a g n o s t i c of B. asperrimum ( M i t t . ex. C. Muell.) S u l l . and B. f r i g i d u m (C. Muell.) Besch., a sample of f i f t y specimens which were not used i n the o r d i n a t i o n s was s e l e c t e d from the U n i v e r s i t y of B r i t i s h Columbia herbarium (U.B.C ). I n d i v i d u a l s were recorded as ( + ) or •(-) according to the presence or absence of four f e a t u r e s . The s e l e c t i o n of these f e a t u r e s was based on the author's concepts of the two s p e c i e s , derived from the type specimens, the informa-t i o n provided by the numerical analyses and from the examination of hundreds of specimens. Since stem leaves are developmentally mature, they are more l i k e l y to stow the'.full'"complement of p l i c a t i o n and a r e o l a t i o n than are branch l e a v e s . For t h i s reason, stem leaves are p r e f e r r e d as a b a s i s f o r t e s t i n g the r e l i a b i l i t y of f e a t u r e s and as a b a s i s f o r keys l e a d i n g to the i d e n t i f i c a t i o n of specimens. The r e s u l t s of the t e s t (Table 7, page 37 ) shows that the two species may be d i s t i n g u i s h e d -microscopically by a combination of the degree of p l i c a t i o n and the development of a l a r c e l l s and m a c r o s c o p i c a l l y by the p o s i t i o n i n g of the leaves on the stem. The presence of a t w i s t e d apex i s a good secondary charac-t e r i s t i c of B. f r i g i d u m (C. Muell.) Besch. Although not always present i n B. f r i g i d u m (C. Muell.) Besch., i t i s a l t o g e t h e r Table 7. The r e s u l t s of a t e s t of d i a g n o s t i c f e a t u r e s of Brachythecium asperrimum ( M i t t . ex. C. Muell. ) S u l l . and B. frigi d u m (C. Muell.) Besch. STEM LEAF TRAITS leaves s t r o n g l y p l i c a t e , w i t h at l e a s t two l a m i n a l p l i c a t i o n s tone e i t h e r side of the c o s t a (scored as +) apex of at l e a s t some leaves w i t h a d i s t i n c t t w i s t (scored as +) leaves w i t h ( > 10), i n -f l a t e d (round) a l a r c e l l s • • (scored as +) leaves e r e c t -spreading to im b r i c a t e (+) leaves wide-spreading to open or l o o s e l y complanate (-) SPECIES B. FRIGIDUM 96% (+) k°/o ( - ) 36% (+) 64% ( - ) 100% (+) 96% (+) Wo ( - ) B. ASPERRIMUM 100% (-) 100% (-) 92% (-) 8% ( + ) 96% (+) 4% ( - ) 38. l a c k i n g i n B. asperrimum ( M i t t . ex. C. Muell.) S u l l . In Part Two, the taxonomic i d e n t i t i e s of the two species a r e discussed i n d e t a i l , i n c l u d i n g : (1.) a key to the species (2.) a r e d e s c r i p t i o n of each species (3) an i n v e s t i g a t i o n of t h e i r synonyms and (k.) a d i s c u s s i o n of the cause of the former confusion between the two s p e c i e s . 39-PART TWO: BRACHYTHECIUM ASPERRIMUM (MITT. EX. C.MUELL.) SULL. AND B. FRIGIDUM (C. MUELL.) BESCH. REDEFINED I I KEY TO THE STEM LEAVES OF. BRACHYTHECIUM ASPERRIMUM (MITT SULL. AND B. FRIGIDUM (C. MUELL.) BESCH. 1. Stem leaves deeply p l i c a t e , most wi t h at l e a s t two l a m i n a l (not mar-g i n a l ) p l i c a t i o n s . o n e i t h e r side of the c o s t a '. .-.B.. fr i g i d u m 1. Stem leaves.plane, w r i n k l e d or w i t h only one l a m i n a l p l i c a t i o n on e i t h e r side of the costa....-.............2 2. I n f l a t e d a l a r c e l l s i n a w e l l de-f i n e d , o f t e n decurrent, group of ten or more o v a l or r o u n d . c e l l s ; i n f l a t e d c e l l s p l a i n l y , v i s i b l e under low power m a g n i f i c a t i o n (4X o b j e c t i v e , 10X o c u l a r ) , apex sometimes t w i s t e d , . l e a v e s erect-spreading to. i m b r i c a t e , wet or dry...............'. B. frigidum 2. Enl a r g e d : a l a r c e l l s l e s s numerous, ' ( u s u a l l y 6-9), these being more oblong or r e c t a n g u l a r i n o u t l i n e r a t h e r than round, not p l a i n l y v i s i b l e under low power m a g n i f i c a t i o n , apex not t w i s t e d ; leaves wide-spreading, or. open- and somewhat complanate. B.; asperrimum 41. Brachythecium frig i d u m (C. Muell.) Besch., Mem. Soc. S c i . Nat. Cherbourg 16:248 1872 NEOTYPE: C. A. Purpus 3722 January 1909, I x t a c c i h u a t l , ' M e x i c o , near the g l a c i e r s . (US!, NY!, FH! ) Designated by H. R. Robinson, 1963 B r y o l o g i s t 66: I 3 6 . Basionym: Hypnum fr i g i d u m C. M u e l l . , Bot. Z e i t 14:4-56 1856. Brachythecium gemmascens C. M u e l l . & Kindb. ex Macoun, Cat. Canad. P l . 6:198 1892. B. columbico-rutabulum Kindb. ex Macoun, Cat. Canad. P l . 6:198 I 8 9 2 . B. lamprochryseum C. M u e l l . & Kindb. ex Macoun, Cat. Canad. P l . 6:199 ~~ 1892. B. lamprochryseum var. giganteum Grout, Mem. Torrey Bot. C l u b . 6 : l 8 l 1897. , B. washingtonianum Eaton, ex Grout B r y o l o g i s t 2:12 1899. B. r i v u l a r e var. laxum Grout, B r y o l o g i s t 4 :48 1901. P l a n t s yellow-green, green or Olive-green, o f t e n g l o s s y . V a r i a b l e i n s i z e and growth form. Most t y p i c a l l y forming deep mats i n wet areas composed of erect secondary stems a r i s i n g from a t h i c k , r h i z o i d l e s s primary stem. Secondary stems e x h i b i t i n g subpinnate to subdendroid branching, w i t h the t e r t i a r y d i v i s i o n s o f t e n crowded near the t i p s of the secondary stems. In d r i e r areas (and at higher a l t i t u d e s ) o f t e n occuring as somewhat loose to dense mats, the primary stems h o r i z o n t a l and subpinnately to i r r e g u l a r l y branched. Secondary stems i n wet area forms may exceed 1 dm. i n h e i g h t . In smaller mat forms the primary stems may only reach 1-2 cm. i n l e n g t h . Leaves commonly erect-spreading to i m b r i c a t e and crowded, r a r e l y wide-spreading, the t i p s o f t e n somewhat f a l c a t e . Stem leaves (and l a r g e r branch leaves) broadly d e l t o i d , broadly ovate or ovate, reaching 3-0 mm. long, commonly 2 .5 - 1 .8 ( 1 .6) mm. long X ( .7) 1-0 -1.4 mm. wide, u s u a l l y deeply p l i c a t e ( w i t h four or more l a m i n a l p l i -c a t i o n s , not i n c l u d i n g the r e f l e x e d margins), l e s s commonly with only two l a m i n a l p l i c a t i o n s or w i t h many shor t e r p l e a t s ( s t r o n g l y w r i n k l e d ) . Smaller (stem and branch) leaves broadly ovate, ovate or d e l t o i d , commonly w i t h two l a m i n a l p l i c a t i o n s , or s t r o n g l y w r i n k l e d , r a r e l y plane. Leaf apices s h o r t - t o long-acuminate, r a r e l y acute, w i t h the t i p s o f t e n t w i s t e d . Margins commonly s e r r u l a t e to d e n t i c -u l a t e , more sharply so i n branch leaves. Median l e a f c e l l s commonly ten times as long as wide, 70 - 100 (132.5) microns long X ( 6 . 5 ) 7 .5 - 8 .5 (10) microns wide. A p i c a l c e l l s o f t e n s h o r t - to l o n g -rhombic (as i n B. r i v u l a r e B.S.G.). Ba s a l c e l l s g r a d u a l l y enlarged, those of the extreme base commonly forming a conspicuous row of i n f l a t e d , h y a l i n e c e l l s . A l a r c e l l s somewhat v a r i a b l e ; i n l a r g e r leaves forming a w e l l - d e f i n e d and often d e c u r r e n t a r e a o f 1 0 - 2 0 round or oblong c e l l s bordered above by quadrate to r e c t a n g u l a r c e l l s and at the margin by long-rectangular c e l l s ; a l a r c e l l s of smaller leaves l e s s w e l l developed, o f t e n forming only one or two 42. rows of i n f l a t e d c e l l s near the base of the l e a f , the leaves then s l i g h t l y or not decurrent. D i o i c o u s , male p l a n t s not d i f f e r i n g g r e a t l y i n s i z e or growth, form from female p l a n t s . P e r i c h a e t i a l b r a c t s sheathing at the base, ecostate or w i t h a f a i n t nerve, t i p s squarrose and very long acumin-ate. Sporophyte dark brown to' r e d d i s h brown. Seta rough throughout w i t h h i g h , sharp, p a p i l l a e , (1.2) 2.0 - 2-5 (3-0) c m - long. Sporangium h o r i z o n t a l , the urn curved and zygomorphic. Annulus c o n s i s t i n g of . two rows of cells.-. C i l i a long and nodose. Operculum conic.. Range: common along the P a c i f i c coast of North America from the .Aleutian I s l a n d s to.Southern C a l i f o r n i a , o ccuring eastward i n the mountains' of western Nevada, Montana, Wyoming, Utah and probably Colorado. Apparently d i s j u n c t as an a l p i n e species i n Mexico. Not yet reported from Ar i z o n a or New Mexico, although i t should be looked, f o r i n the hi g h mountains of those s t a t e s . (See map, f i g u r e 11, , page 41 ). H a b i t a t s : Most f r e q u e n t l y on mud, wet r o c k s and wood i n s e e p -age a r e a s , a r o u n d s p r i n g s , e t c ; o c c a s i o n a l l y submerged o r ernergent .Less common on somewhat d r i e r s o i l s , r o t t i n g wood and tree bases i n swampy woods. Probably r e s t r i c t e d to areas which are wet during most or a l l of the year. Occuring from sea. l e v e l to 6,000' i n c o a s t a l r e gions and from 4,500' to 12,600' i n l a n d (from 8,700'.to 13,700' i n Mexico). I l l u s t r a t i o n s : . Figures 12 - 18, (pages 44-50) • Notes on synonymy: The f o l l o w i n g specimens have been examined and determined as B. f r i g i d u m (C. Muell.) Besch: Holzinger & Blake I898., B. r i v u l a r e var.. laxum, Musci Americae S e p t e n t r i o n a l i s E x s i c c a t i ' (published by Renauld and Cardot), N.W. Montana, Lake McDonald, Flathead County ..(FH); Macoun 13 May I89O, B. columbico-rutabulum, Canadian Cryptogams; # 108, wet l o g s , Revelstoke, B.C. (FH); J . Macoun 291,1 June 1908, B. gemmascens, d'et. Kindberg, . Sea's Farm, V i c t o r i a , Vancouver I s l a n d , on stones i n small brook (MICH). Note: Dr. E. Lawton (1971) erroneously c i t e d the three species • j u s t mentioned, as w e l l as B. washingtonianum. Eaton ex Grout as synonyms of B. asperrimum. •. A3-F i g . 11.. The range of Brachythecium frig i d u m (C. Muell.) Besch. kk. 3.0 m m . F i g . 12. Brachy.thecium frig i d u m (C. Muell.) Besch.: .Variation i n shoot s i z e and branching p a t t e r n . (Continued: on page k5 F i g • 13) > Two wet h a b i t a t forms showing h o r i z o n t a l primary stem and e r e c t , . v a r i o u s l y branched secondary stems.-45-F i g . 13. Brachythecium f r i g i d u m (C. Muell.) Besch.: V a r i a t i o n i n shoot s i z e and branching pattern.- (Continued on page 46., Fig.- 14) (a) Secondary stem of a l a r g e wet form,, (b) e n t i r e shoot of d r i e r , smaller (mat) form 4 6 . F i g . 14. Brachythecium. frig i d u m (C. Muell.) Besch.: V a r i a t i o n i n shoot s i z e and branching p a t t e r n . • ( a) Large, s p a r s e l y branched-secondary stem of a wet ( t u f t ) form; (b) small aquatic form showing a t y p i c a l l y small s i z e and sparse f o l i a t i o n . 48. F i g . 16. B. f r i g i d u m (C. Muell.) Besch.: v a r i a t i o n i n the s i z e , shape and p l i c a t i o n of stem le a v e s . 49, d . F i g . 17- B. f r i g i d u m (C. Muell.) Besch. (a) median l e a f c e l l s ; (b) c e l l s of the l e a f apex; (c, d) two stem leaves showing the appearance of the i n f l a t e d a l a r c e l l s under low power m a g n i f i c a t i o n (4X o b j e c t i v e , 10X o c u l a r ) . . • 5 0 . F i g . 18. B. f r i g i d u m (C. Muell.) Besch.: v a r i a t i o n i n the develop-ment of the a l a r c e l l s . (a) t y p i c a l stem l e a f a r e o l a t i o n w i t h numerous round to oblong c e l l s , (b) branch l e a f a r e o l a t i o n showing the fewer number of i n f l a t e d c e l l s . 51. Brachythecium asperrimum ( M i t t , ex C. Muell.) S u l l . Icones Muscorum Supplement. 100. 1894 TYPE: L y a l l 1 8 5 8 - 9i Hypnum asperrimum, Columbia, N.W. America, Douglas. (NY) Basionym: Hypnum asperrimum M i t t ex C. M u e l l . , Bot. Z e i t . 16:170 1858. Hypnum v a l l i u m S u l l . & Lesq., Musci Bor.-Amer. ed. 2:84 I 8 6 5 . Brachythecium v i l l a r d i Ren. & Card, e_x R o e l l . , Bot. C e n t r a l be. 44:142 1890. B. spurio-rutabulutn C.Muell. & Kindb. e_x Macoun, Cat. Canad. P l . 6:197 1892. B. s u b i n t r i c a t u m Kindb., Rev. B r y o l . 22:86 l895« B. subasperrimum Card. & Ther., Bot. Gazette. 37=377 1904. B. p a c i f i c u m Jenn., B r y o l o g i s t 16:95 1913-Shiny, pale green or yellow-green , small to moderately s i z e d p l a n t s , forming loose mats (or s t o l o n i f e r o u s i n e p i p h y t i c forms); branching commonly subpinnate, l e s s o f t e n i r r e g u l a r , the branches i n s t o l o n i f e r o u s forms o f t e n exceeding 5 cm. i n l e n g t h . Both stem and branch leav e s somewhat d i s t a n t and l o o s e l y overlapping, wide-spreading to open, o f t e n somewhat co m p l a n a t e - f o l i a t e , r a r e l y f a l -cate at the t i p s . Stem leaves (and l a r g e r branch leaves) broadly ovate to ovate, ( 2 . 5 ) 2 . 2 - 1 . 5 mm. long X ( 1 .2) 1.0 - .6 ( .5) n™. wide, commonly plane except f o r the r e f l e x e d margins or wi t h a few l a m i n a l w r i n k l e s , r a r e l y w i t h 2 l a m i n a l p l i c a t i o n s . Branch leaves ovate, narrowly ovate or l a n c e o l a t e , ( 1 . 8 ) 1.5 - «9 mm. long X ( . 2 ) .5 - • 7 mm. wide, commonly plane or s l i g h t l y w r i n k l e d . Leaf apices long and acuminate to g r a d u a l l y l o n g and slender i n smaller l e a v e s , the apices not t w i s t e d . Median l e a f c e l l s reaching 128 microns l o n g , commonly 50 - 80 microns long X ( 5 . 0 ) 6 . 0 - 7-5 ( 8 . 5 ) microns wide. A l a r c e l l s somewhat v a r i a b l e i n s i z e and number; l a r g e r stem leaves w i t h a l o o s e l y defined group of 6 - 9 o v a l or r e c t a n g u l a r , enlarged but not d i s t i n c t l y i n f l a t e d , c e l l s , bordered at the margin by a f i l e of r e c t a n g u l a r or quadrate c e l l s which i s o f t e n long-decurrent; smaller (stem and branch) leaves w i t h fewer enlarged c e l l s or w i t h only the marginal f i l e of quadrate to re c t a n g u l a r c e l l s . D i o i c o u s , the male p l a n t s f i n e r and more d i s t a n t l y f o l i a t e than the female p l a n t s , e s p e c i a l l y i n s t o l o n i f e r o u s forms. P e r i -c h a e t i a l b r a c t s sheathing at the base, ecostate or with a very f a i n t nerve, the apex very long acuminate and squarrose. Sporo-phyte dark brown or r e d d i s h brown, seta rough throughout w i t h h i g h , sharp p a p i l l a e , ( 1 . 0 ) 2 . 0 - 2.6 (3«0) mm. long. Sporangium h o r i z o n t a l , or i n c l i n e d , the urn curved and zygomorphic. Annulus c o n s i s t i n g of two rows of c e l l s . C i l i a l o n g , nodose to subappen-d i c u l a t e , operculum conic to s u b - r o s t r a t e . 52 Range: Endemic t o t h e P a c i f i c c o a s t o f N o r t h A m e r i c a , ex-t e n d i n g from t h e A l e u t i a n I s l a n d s t o t h e m i d d l e o f C a l i f o r n i a ( P l a c e r v i l l e , E l d o r a d o County and Noyo, Mendocino C o u n t y ) . See map, page 54. H a b i t a t s : most common on t r e e and shrub bases, r o t t i n g twigs and l o g s or humus i n moist woods and swampy areas, l e s s common as a true epiphyte, f a i r l y common on moist s o i l , r a r e r on r o c k s . Ab-sent from both very dry or very web areas. Probably occur from near sea l e v e l to 2 , 0 0 0 ' . I l u s t r a t i o n s : See F i g u r e s 20 - 25 , pages 55 to 60 . Notes on Synonymy: The f o l l o w i n g specimens have been examined and determined as B. asperrimum ( M i t t , ex C. Muell.) S u l l . : J . Macoun I 896 B. s p u r i o -rutabulum, v i c i n i t y of Ottawa, Ontario ( s i c ) (NY); J . Macoun Cana-dian Musci # 65I, B. spurio-rutabulum, Burrard I n l e t and Vancouver I s l a n d , B.C. (FH); Bolander # 506 Hypnum v a l l i u m n. sp., "Hab. ad  rupes s c i s t o s a s i n p r o f u n d i s c o n v a l l i b u s C a l i f o r n i a e " (NY); Jaeger Herbarium America B o r e a l i s America Septentr. C a l i f o r n i a Herbariade Thumen, Hypnum v a l l i u m (NY). (There are s e v e r a l specimens of t h i s c o l l e c t i o n at NY, some are B. asperrimum, others are B. f r i g i d u m ("on r o c k " ) , p l u s one c o l l e c t i o n of Scleropodium.); J.W. B a i l e y 3.0 January 1905, North American Musci P l e u r o c a r p i # 221, B. sub- asperrimum, Washington, near Lake Washington, U n i v e r s i t y S t a t i o n , S e a t t l e , on l o g s i n woods (US); J . Macoun # 65O, 2k September I 8 9 3 , B. s u b i n t r i c a t u m , Meeche's Lake, Quebec ( s i c ) , on rocks along a brook (MICH, US, NY). No specimens of B. v i l l a r d i Ren. & Card, or B. pacificum have I been seen. Jennings' d e s c r i p t i o n of B. p a c i f i c u m s t a t e s that the leaves are "not markedly plicate" ( 1 9 1 3 B r y o l o g i s t 1 6 : 9 6 ) , so i t i s l i k e l y that Grout c o r r e c t l y placed i t i n synonymy w i t h B. asperrimum 53-( M i t t , ex C. Muell.) S u l l . (1928 Moss F l o r a of North America, Volume 3, p. 40). (1897 Mem. Torrey Bot. Club 6: I78) B. v i l l a r d i Ren. & Card, i s i n c l u d e d as a synonym of B. asperrimum ( M i t t , ex C M u e l l . ) S u l l . on a p r o v i s i o n a l b a s i s , according to Grout (1897, Mem. Torrey Bot. Club 6 : 1 7 8 ) . Note: the.type d e s c r i p t i o n of B. asperrimum ( M i t t , ex C. Muell.) S u l l . s t a t e s the type l o c a l i t y as "America b o r e a l i s - o c c i d e n t a l i s , ad Columbia flumen: Douglas". For t h i s reason, I have c i t e d the holotype (page 49) as L y a l l ' s c o l l e c t i o n "Columbia. N.W. America. Douglas". L y a l l ' s other c o l l e c t i o n s ("near the 49th p a r a l l e l of La t . , N.W. America. Douglas", "N.W. America. Douglas" and " 49th p a r a l l e l " (NY), although marked "TYPE" are b e l i e v e d to be i s o t y p e s . 55-j F i g . 20. Brachythecium asperrimum ( M i t t , ex C. Muell.) S u l l . : V a r i a t i o n ! i n shoot s i z e and branching p a t t e r n . (Continued on page 56 , F i g . 21) Male pl a n t of a s t o l o n i f erous ( e p i p h y t i c ) form. I i ' ' • 1 56. b. F i g . 21. Brachythecium asperrimum ( M i t t , ex C. Muell) S u l l . : V a r i a t i o n i n shoot s i z e and branching p a t t e r n . (Continued on page 5 .7 , F i g . 22) (a) Female pla n t of a s t o l o n i f e r o u s . ( e p i p h y t i c ) form; (b) smaller p l a n t from r o t t i n g wood. 57. 3.0 m m. /' F i g - 22. Brachythecium asperrimum ( M i t t . _ex C. Muell.) S u l l . : V a r i a t i o n ,' i n shoot s i z e and branching p a t t e r n , continued, (a) l a r g e r p l a n t from r o t t i n g wood, (b) l a r g e r mat form from wet s o i l . i I 5 8 . F i g . 23. B. asperrimum ( M i t t , ex C. Muell.) S u l l . V a r i a t i o n i n the s i z e and shape of branch le a v e s . 59-1 F i g . 24. B. asperrimum ( M i t t , ex C. Muell.) S u l l . V a r i a t i o n i n the s i z e and shape of stem le a v e s . 60. F i g . 25. B. asperrimum ( M i t t , ex C. Muell.) S u l l . (a) median l e a f c e l l s ; (b) c e l l s of the l e a f apex; (c,e) a l a r c e l l s of two leaves showing the t y p i c a l marginal f i l e of quadrate or r e c t a n g u l a r c e l l s ; (d) sketch of a stem leaf-showing the appearance of the a l a r r e g i o n under low power m a g n i f i c a t i o n (kX o b j e c t i v e , 10X o c u l a r . ) 61. Having described B. asperrimum ( M i t t . ex. C. Muell.) S u l l . and B. f r i g i d u m (C. Muell.) Besch., i t would be h e l p f u l to d i s c u s s the probable causes of confusion between the two species. Most of the confusion has r e s u l t e d from specimens i d e n t i f i e d on the b a s i s of growth form and h a b i t a t . Using these c r i t e r i a , mat forms of B. f r i g i d u m (C. Muell.) Besch. have been r e f e r r e d to B. asperrimum ( M i t t . ex. C. Muell.) S u l l . Consequently, d e s c r i p t i o n s of B. asperrimum ( M i t t . ex. C. Muell.) S u l l . probably have been based on m a t e r i a l of both species. M i t t e n ' s type d e s c r i p t i o n of B. asperrimum ( M i t t . ex. C. Muell.) S u l l . as " d i s t i n c t e p l u r i e s s u l c a t o - p l i c a t a " ( i n C. M u e l l e r , Bot.. Z e i t . 16:170, I 8 5 8 ) misrepresents that species as d i s t i n c t l y p l i c a t e , which may have f u r t h e r obscured the d i s t i n c t i o n between i t and B. f r i g i d u m (C. Muell.) Besch.. 6 2 . PART THREE: THE RELATIONSHIPS OF THE SIX NORTH AMERICAN DIOICOUS  BRACHYTHECIUM SPECIES WITH ROUGH SETAE. The o b j e c t i v e s of t h i s part of the i n v e s t i g a t i o n are: (1) to explore the phenetic r e l a t i o n s h i p s of Brachythecium  asperrimum ( M i t t . ex. C. Muell.) S u l l . , B. b o l a n d e r i (Lesq.) Jaeg. & Sauerb., B. hylotapetum B. Hig. & N. Hig., B. n e l s o n i i Grout, B. frigi d u m (C. Muell.) Besh., and B. r i v u l a r e B.S.G., (2) to introduce a p r o v i s i o n a l key to the s i x species and (3) to di s c u s s the h y p o t h e t i c a l c l a d i s t i c r e l a t i o n s h i p s of the s i x species. In order to demonstrate the phenetic r e l a t i o n s h i p s of the s i x species, two types of analyses were performed: a p r i n c i p a l components a n a l y s i s and a c l u s t e r a n a l y s i s . The b a s i c procedure followed was e s s e n t i a l l y the same as the procedure o u t l i n e d i n Part One, Sections (1 . ) 2 . , ( 1 . ) 3« and ( 2 . ) 1 . , d i f f e r i n g i n the f o l l o w i n g e s s e n t i a l s . The comparison of the s i x species was made on an exemplary b a s i s . I t does not s a t i s f y assumptions which would be 'required f o r s t a t i s t i c a l t e s t i n g , nor does i t r e f l e c t the v a r i a t i o n of the s i x species. A sample of nine specimens was s e l e c t e d from m a t e r i a l a v a i l a b l e i n the U n i v e r s i t y of B r i t i s h Columbia herbarium. (UBC). The specimens chosen were co n s i d -ered t y p i c a l o f each s p e c i e s . See Appendix E f o r a l i s t of the specimens. In a d d i t i o n to the t r a i t s measured f o r the a n a l y s i s of B. fr i g i d u m (C. Muell.) Besch. sensu Robinson (see Tables 3a - 3 c, pages 13 - 15, f i v e supplemental t r a i t s , p l u s one a d d i t i o n a l character s t a t e of a s i x t h t r a i t , were measured 63-Table 8 a. A d d i t i o n a l c h a r a c t e r s used i n the exemplary o r d i n a t i o n of 6 species; Measurement of c h a r a c t e r s : c r i t e r i a used, type and s c o r i n g of data. Type of Data and Scoring Method Character C r i t e r i a Used Ranked data: i n d i - s i z e and p o s i t i o n leaves were recorded as v i d u a l leaves were of a l a r c e l l s having one of four char-scored as having ac t e r s t a t e s : one of s e v e r a l 1) a l a r c e l l s not i n -character s t a t e s . f l a t e d , a l a r c e l l s not forming decur-ren t a u r i c l e s . F i g . 26a, 26b, 26c. 2) a l a r c e l l s i n f l a t e d and forming small a u r i c l e s . F i g . 26d, 26e 3) a l a r c e l l s i n f l a t e d and forming l a r g e , long-decurrent a u r i c l e s which ex-tend to the c o s t a . F i g . 26 g. l e a f concavity leaves recorded as having one of three character s t a t e s : 1) not or s l i g h t l y concave. F i g . 27a 2) s t r o n g l y concave, but l a c k i n g a spoon-shaped depression at the l e a f apex. F i g s . 27c,-27d. margin s e r r a t i o n . leaves were recorded as having one of four char- . ac t e r s t a t e s : 1) not s e r r a t e (plane) 2') s e r r a t e i n the upper 1A of the l e a f 3) s e r r a t e i n the upper i 1/2 of the l e a f 4) s e r r a t e throughout 64. Table 8 b. A d d i t i o n a l characters used i n the exemplary o r d i n a t i o n of' 6 species. Measurement of characters: c r i t e r i a used, type and sc o r i n g of data, continued. Type of Data and Scoring Method Character C r i t e r i a Used Ranked data: (continued)' i n d i v i d u a l leaves were scored as having one of se v e r a l charac-t e r s t a t e s . Dichotomous data: (equivalent to presence-absence data) Shape of l e a f apex apex g r a d u a l l y or abrup t l y narrowed b a s a l c e l l s i z e one a d d i t i o n a l character s t a t e was inc l u d e d (the other four are l i s t e d i n Tables 3b - 3c, pp. 13-14) 5) apex a b r u p t l y a p i c u l a t e . F i g . 28a. l e a f apices were recorded as g r a d u a l l y narrowed (0) F i g . 29b, 29c; a b r u p t l y narrowed ( l ) F i g . 29a. b a s a l c e l l s (the row of c e l l s nearest the point of attachment of the l e a f ) were recorded as enlarged, but not d i s -t i n c t l y i n f l a t e d and h y a l i n e (0) or i n f l a t e d and h y a l i n e ( l ) . f o r use i n the comparative analyses. These t r a i t s are l i s t e d and de-f i n e d i n Tables 8a - 8b, pages 63- 64. The comparative o r d i n a t i o n was based on a c o r r e l a t i o n matrix (see Appendix F, Tables l a and l b f o r the c o r r e l a t i o n matrix and Table 2 f o r the mean, variance and standard d e v i a t i o n of the v a r i a b l e s ) . Figure 30 (page 69) shows the v a r i a t i o n along the f i r s t and second component axes. The f i r s t a x i s accounts f o r 32.69% of the v a r i a t i o n , the second a x i s f o r 23.46%. Figure 31 (page 70) shows the v a r i a t i o n along the f i r s t and t h i r d component axes. The t h i r d , component a x i s accounts f o r 20.17% of the v a r i a t i o n . The c o n t r i b u t i o n s of the v a r i a b l e s ( t r a i t s ) to the v a r i a t i o n along the three component 6 5 -F i g . 2 6 . Character s t a t e s of the s i z e and p o s i t i o n of a l a r c e l l s used i n the s c o r i n g of ranked data: (a,b,c,) not forming decurrent a u r i c l e s ; (d,e) forming small a u r i c l e s ; ( f ) forming l a r g e a u r i c l e s which do not extend to the cost a ; (g) forming l a r g e a u r i c l e s which extend to the c o s t a . 66. C. F i g . 27. Character s t a t e s of the concavity of leaves used i n the s c o r i n g of ranked data: (a) not or s l i g h t l y concave; (b) s t r o n g l y concave but l a c k i n g the a p i c a l spoon-shaped depression; (c,d) st r o n g -l y concave w i t h an a p i c a l spoon-shaped depression. 6 7 . F i g . 28. Character s t a t e s of the l e a f apex shape used i n the s c o r i n g of ranked data: (a) a b r u p t l y a p i c u l a t e ; • (b) not a p i c u l a t e (the other four character s t a t e s are on page 18, Figure 4.), F i g . 29. Character s t a t e s of the l e a f apex used i n the s c o r i n g of dichotomous data: (a) a b r u p t l y narrowed; (b,c) g r a d u a l l y narrowed. 6 8 . axes are l i s t e d i n Table 9 (page 7 6 ) . A comparison of the con-t r i b u t i o n of the t r a i t s to the v a r i a t i o n along the separate axes r e v e a l s the f o l l o w i n g trends i n the phenetic v a r i a t i o n of the s i x species: 1) the species are arranged along the f i r s t a x i s according to the s i z e s , median l e a f c e l l dimensions and concavity of both the branch and stem l e a v e s . This trend i s shown below: BOLANDERI—ASPERRIMUM—» NELSONII > RIVULARE—•> HYLOTAPETUM Leaf s i z e s : smaller \ l a r g e r Median l e a f c e l l dimensions: shorter and narrower ) longer and wider Leaf concavity: not concave ) s t r o n g l y concave • 2) s i x species are d i s t r i b u t e d along the second a x i s l a r g e l y as a r e s u l t of the v a r i a t i o n i n t h e i r a l a r c e l l s . /BOLANDERI RIVULARE > NELSONII > FRIGIDUM ) € ASPERRIMUM VHYLOTAPETUM Strongly a u r i c u l a t e •) l e s s a u r i c u l a t e > not a u r i c u l a t e 3) the v a r i a t i o n along the t h i r d a x i s demonstrates the unique-ness of B. fr i g i d u m (C. Muell.) Besch.: I t i s the only species w i t h s t r o n g l y p l i c a t e l e a v e s . A phenogram of these species (Figure 32, page j^) summarizes the three trends. The f i r s t two-point c l u s t e r s are of the two c o l l e c t i o n s of B. hylotapetum B. Hig. & N. Hig. , B. r i v u l a r e B.S.G., and B. n e l s o n i i Grout. The' l a s t two-point c l u s t e r i s of-69, F i g . 30. The f i r s t and second axes of the comparative o r d i n a t i o n of s p e c i e s . s i x 70. . . . . • FRIGlttUW +1-1 1 i axtsl u .pntANTteai 1 - i I F i g . 31.' The f i r s t and t h i r d axes of the comparative o r d i n a t i o n of s i x s p e c i e s . 71. Table 9- T n e c o n t r i b u t i o n s .'.to. the v a r i a t i o n "of the 32 v a r i a b l e s measured f o r the comparative o r d i n a t i o n of' s i x species. • COMPONENTS 1 2 3 LEAF LENGTH 0.1902 0.1734 0.0841 LEAF WIDTH 0.2814 O.O835 0.0145 COSTA RATIO - 0 . 0 2 3 8 0.2201 0.1935 MEDIAN CELL LENGTH 0.2955 0.0535 .0 .0721 MEDIAN CELL WIDTH 0.2885 - 0 . 0 9 9 3 O.OI65 APEX TAPERED 0.2244 0.1899 -0.1468 STEM LEAF SHAPE , 0 . 0 9 0 6 -0.1533 0.3032 LEAF ACUMINATION -O.IO69 -0 .3222 0.0727 TRAITS MARGIN REFLEXED - 0 . 0 6 9 2 O.O698 0.3151 PLICATION 0.1052 0.0904 0.3523 APEX TWIST 0.0887 0.2748 O.0518 APEX FALCATE O.OO63 - 0 . 0 8 5 8 -0.1072 ALAR CELLS 0.0978 - 0 . 3 3 9 2 -O.OO65 MARGIN SERRATED -O.I766 0.2122 0-1377 BASAL CELTS 0.1946 -0.1376 0.1784 CONCAVITY 0.2500 0.0442 -0.1001 LEAF LENGTH 0.2217 0.0879 0.0163 LEAF WIDTH 0.2624 -0.1513 - 0 . 0 9 4 5 COSTA RATIO - 0 . 0 6 4 5 - 0 . 0 5 0 3 0.1123 MEDIAN CELL LENGTH 0.2845 0.0133 -0.0618 MEDIAN CELL WIDTH 0.2113 -0.2317 0.0777 APEX TAPERED 0.1722 0.1000 -O.1983 BRANCH LEAF SHAPE 0.2153 -0.1109 O.2115 LEAF ACUMINATION -0 .1083 - 0 . 2 8 0 6 0.1572 TRAITS MARGIN REFLEXED 0.0124 - 0 . 0 0 0 1 0.3814 PLICATION OO.0365 -0.0192 0.3823 APEX TWIST O.1716 0.2367 0.1751 APEX FALCATE -0.1359 O . I I 8 5 -0 .0377 ALAR CELLS O.0725 -0 .2947 -0.1027 MARGIN SERRATED -0.1657 0.1059 0.2217 BASAL CELLS -O.0302 -0.3239 O.0919 CONCAVITY 0.2452 0.0002 -0.0162 72. i 1 -< 81-14 68- 53 h-56-17 27-80 (-26-84 1—11. 97 10-27 h-3-24 00 LU CJ z < r — 00 Q > TJ I c g ro O > z o m 33 O T U ' S F i g . 32. E e s u l t s of the c l u s t e r a n a l y s i s comparing s i x Brachythecium s p e c i e s . B. asperrimum ( M i t t , ex C. Muell.) S u l l . and B. b o l a n d e r i (Lesq.) Jaeg. & Sauerb. Subsequent c l u s t e r s , i n order of phenetic c l o s e -ness, are between B. r i v u l a r e B.S.G. and B. n e l s o n i i , Grout, followed by the a d d i t i o n of B. fr i g i d u m (C. Muell.) Besch. and f i n a l l y by the a d d i t i o n of B. hylotapetum B. H i g . and N. Hig. I t i s t h e r e f o r e apparent that B. asperrimum ( M i t t . ex. C. Muell.) S u l l . i s more c l o s e l y r e l a t e d - t o B. b o l a n d e r i (Lesq.) Jaeg. & Sauerb., whereas B. fr i g i d u m (C. M u e l l . ) Besch. i s more c l o s e l y r e l a t e d to B. r i v u l a r e B.S.G. and B . . n e l s o n i i Grout. B. b o l a n d e r i (Lesq.) Jaeg. and Sauerb. and B. hylotapetum B. Hig. and N. Hig. represent the two extremes of the group. Reviewing the phenetic r e l a t i o n s h i p s of the s i x s p e c i e s , i t i s i n t e r e s t i n g to consider t h e i r h y p o t h e t i c a l c l a d i s t i c r e l a t i o n -7 3 . s h i p s . B. r i v u l a r e B.S.G. may be considered as the h y p o t h e t i c a l parent species of the group. I t i s the most widespread s p e c i e s , and probably the most v a r i a b l e . The p o s s i b l e d e r i v a t i o n s of the other f i v e species from B. r i v u l a r e B.S.G. are shown i n the dendrogram below. B O L A N D E R I / A S P E R R I M U M V^-HY LOTA P E T U M • F R I G I D U M I N J | ^ N E L S O N I ! Rl VU L ARE Although each of the s i x species has a unique combination of ch a r a c t e r s , the group i s u n i f i e d by s e x u a l i t y , s e t a c o n d i t i o n and trends i n the v a r i a t i o n of gametophytic f e a t u r e s . The v a r i a b i l i t y of the s i x species and t h e i r phenetic closeness to B. r i v u l a r e B.S.G. appear to be c o r r e l a t e d to t h e i r d i s t r i b u t i o n . Both B. n e l s o n i i Grout and B. r i v u l a r e B.S.G. are widespread i n the northern hemi-sphere, whereas the other four species a r e : r e s t r i c t e d to western North America. Of these four s p e c i e s , B. -frigidum (C. Muell.) Besch. i s the most widespread and v a r i a b l e , f o l lowed by B. asper- rimum ( M i t t . _ex C. Muell.) S u l l . . B. b o l a n d e r i (Lesq.) Jaeg. 8c Sauerb. (reported' only from C a l i f o r n i a and Oregon) and B. h y l o - tapetum B. Hig. & N. Hig. (reported from B r i t i s h Columbia, A l b e r t a , Washington, Oregon,'Idaho and western Nontana) are the most r e s t r i c t e d i n range, and probably the l e a s t v a r i a b l e species i n the group. 7k. The extent of the v a r i a t i o n of the group as a whole i s not yet thoroughly understood or documented, however, a p r o v i s i o n a l key to the s i x species i s introduced i n order to f a c i l i t a t e the i d e n t i f i c a t i o n of specimens. PROVISIONAL KEY TO THE NORTH AMERICAN DIOICOUS BRACHYTHECIUM SPECIES WITH ROUGH SETAE 1. Stem leaves deeply p l i c a t e (with two or more l a m i n a l p l i c a t i o n on e i t h e r s i d e of the c o s t a , not i n c l u d i n g r e f l e x e d margins) B. f r i g i d u m (C. Muell.) Besch. ( F i g ' s 12 - 18) 1. Stem leaves plane, w r i n k l e d or w i t h not more than one l a m i n a l p l i c a t i o n on e i t h e r s i d e of the c o s t a 2 . 2 . Stem leaves w i t h fewer than 10 i n f l a t e d a l a r c e l l s , stem leaves not a u r i c u l a t e or. prominently decurrent 3« 2 . Stem leaves w i t h more than 10 i n f l a t e d (round or oblong) a l a r c e l l s , the stem leaves a u r i c u l a t e and decurrent 5« 3- Stem leaves g e n e r a l l y exceeding 2-5 nim. lo n g , deeply concave, a b r u p t l y a p i c u l a t e , w i t h the apex o f t e n t w i s t e d B. hylotapetum B. Hig. &'N. H i g . ( F i g ' s . 35 - 36) i 3- Stem leaves s m a l l e r , not concave, g r a d u a l l y acuminate. . . . k. 75-4. Largest stem leaves 1.0 - 1.5 mm. long X .4 - .5 mm. wide, median l e a f c e l l s (35) 40 - 60 micrometers long X 5 - 6 (7) micrometers wide B. b o l a n d e r i (Lesq.) Jaeg. & Sauerb. ( F i g . 33) 4 . Largest stem leaves 2 . 0 mm. long or longer, .6 - 1.0 mm. wide, median l e a f c e l l s 50 - 80 ( - 100 ) micrometers long X 6 . 0 - 7 .5 ( 8 . 5 ) micrometers wide B. asperrimum ( M i t t . ex. C. Muell.) S u l l . ( F i g . ' s 20 - 25) i 5 . ' Stem leaves not concave 6 5- Stem leaves moderately to strongly.concave 7 6 . Stem leaves g r a d u a l l y accuminate, w i t h small a u r i c l e s composed of up to 20 i n f l a t e d c e l l s , leaves short decurrent . . B. frigi d u m (C. Muell.) Besch. ( F i g . ' s 12 - 18) 6 . Stem leaves a b r u p t l y long-acuminate, w i t h prominent a u r i c l e s which commonly extend to the co s t a , leaves long-decurrent . B. n e l s o n i i Grout ( F i g . 34) 7- Largest stem leaves exceeding 2 .5 mm. long, deeply concave w i t h a spoon-shaped depression below at the l e a f apex, the apex a b r u p t l y a p i c u l a t e and o f t e n t w i s t e d , a u r i c l e s small and not prominently de-current B. hylotapetum B. Hig. & N. Hig. ( F i g . ' s 35-36) 7. Largest stem leaves g e n e r a l l y not exceeding 2 . 0 mm. lo n g , s t r o n g l y concave but without a spoon-shaped depression below the apex, a u r i c l e s prominent (not u s u a l l y extending to the costa and long -decurrent . . . . . . . . . . . . . . B. r i v u l a r e B.S.G. ( F i g . ' s 37-38) 76. Q. f; F i g . 33- Brachythecium b o l a n d e r i (Lesq.) Jaeg. & Sauerb. (a, b) stem leaves; (c,d) branch leaves; (e) a p i c a l c e l l s ; ( f ) a l a r c e l l s of stem l e a f . 77-F i g . J>k. Brachythecium n e l s o n i i Grout (a) a p i c a l c e l l s ; (b) stem l e a f ; (c) a l a r c e l l s ; (d) branch l e a f . .78-F i g . 35- Brachythecium hylotapetum B. Hig. & N. Hig. (a,b) branch leaves; (c) stem l e a f ; a p i c a l c e l l s . 79-1 F i g . 36. Brachythecium hylotapetum B. Hig. 8c N. Hig. A l a r c e l l s of stem l e a f . ,8o. F i g ' 37. Brachythecium r i v u l a r e B.S.G. (a, b) branch leaves; (c) stem l e a f ; (d) a p i c a l c e l l s . 8 1 . F i g . 38. Brachythecium r i v u l a r e B.S.G. A l a r c e l l s of a stem' l e a f . 82. Summary This t h e s i s describes a.study of s i x North American, d i o i c o u s Brachythecium species possessing a rough s e t a . The phenetic v a r i a b i l i t y of the s i x species.was analysed i n order to c l a r i f y the r e l a t i o n s h i p s between species and to provide s i m p l e r , more r e l i a b l e means of i d e n t i -f y i n g specimens. As a consequence of t h i s i n v e s t i g a t i o n , the taxonomic i d e n t i t i e s of B. asperrimum ( M i t t . ex. C. Muell.) S u l l . and B. f r i g i d u m (C. Muell.) Besch. have been e s t a b l i s h e d through numerical analyses of the v a r i a t i o n i n the gametophytic fe a t u r e s of the two s p e c i e s . Two types of analyses were used: a p r i n c i p a l components a n a l y s i s and a c l u s t e r a n a l y s i s . The phenetic v a r i a t i o n ' and h a b i t a t s p e c i f i c i t y of each species were a n a l y s e d and d o c u m e n t e d and synonymy and r a n g e s presented. A t e s t of the f e a t u r e s considered to be d i a g n o s t i c f o r the two species W a s performed and the r e s u l t s of t h i s t e s t used to devise a r e l i a b l e key. The r e l a t i o n s h i p s of the two species to the other North American d i o i c o u s species possessing rough setae were determined us i n g a p r i n c i p a l components a n a l y s i s and a c l u s t e r a n a l y s i s . B. asperrimum ( M i t t . ex. C. Muell.) S u l l . i s shown to be c l o s e l y r e l a t e d to B. b o l a n d e r i (Lesq.) Jaeg. and Sauerb.. B. f r i g i d u m (C. Muell.) Besch. i s shown to be c l o s e l y r e l a t e d to B. r i v u l a r e B.S.G. and B. n e l s o n i i Grout. A dendrogram and d i s -c ussion of the h y p o t h e t i c a l c l a d i s t i c r e l a t i o n s h i p s of the s i x species are provided. 8 3 . L i t e r a t u r e C i t e d Flowers, S. 1973- Mosses:. Utah and the West. ( E d i t o r ) A. Holmgren. Brigham U n i v e r s i t y P r e s s . Provo, Utah. 567 pages. Grout, A.J. 1897. A r e v i s i o n of the North American Isotheciaceae. Memoirs of the Torrey B o t a n i c a l Club 6:131-210. Grout, A.J. 1928-1940. Moss F l o r a of North America North of Mexico. P u b l i s h e d by the author, Newfane, Vermont, Volume 3« Jennings, O.E. 1913- Brachythecium p a c i f i c u m , new s p e c i e s . The_.Bryologist 16:95-96. Lawton, E. 1971- Moss F l o r a of the P a c i f i c ! Northwest, H a t t o r i B o t a n i c a l Laboratory, Nichinan, Japan, 362 pages. O s t e r l i n , D., J.M. P a t t e r s o n and R.A. Whitaker 1978. UBC C-Group Manual, published by the U n i v e r s i t y of B r i t i s h Columbia Computing Center, 2075 Wesbrook M a l l , Vancouver, B r i t i s h Columbia, Canada, V6T 1W5. Mimeographed, 21 pages. Robinson, H. 1963. The nomenclature and d i s t r i b u t i o n of three species of Brachythecium. The B r y o l o g i s t 66:136-139. Sneath, P.H.. and R.R. S o k a l , 1973- Numerical Taxonomy; The P r i n c i p l e s and P r a c t i c e of Numerical C l a s s i f i c a t i o n , W. H. Freeman and Company, San F r a n c i s c o , 5^7 pages. Appendix A. A l i s t of the s i x t y - e i g h t specimens used i n the a n a l y s i s of the v a r i a t i o n of Brachythecium f r i g i d u m (C. Muell.) Besch.. sensu Robinson. h i ? r Vvap ii i m co l lec tor and co l l ec t ion 0 date of col lect ion county/state, province or country location of col lect ion habitat of co l lect ion data it annotation or ver i f icat ion date of work 1-2-11 ttf Vai IUI 11 M ICH T E N N T E N N A . J . Sharpe »152 A . J . Sharpe 04272 23 July 1944 21 October 1945 DF, Mexico Pue t Mexico Above Contreras Above Huejotzlngo 8700', cold spring 13,700' near Los Arevas on Ixtachuatl FH1CME1 FTENME3 FTENME5 det. A. J . Sharpe as B. frigidum det. A. J . Sharpe as B. frigidum 15 Jan. 18 Feb. 18 Feb. 1979 1979 1979 3 - U S E . K . Ba l l s I A110 4 January 1938 DF, Mexico OJos de Agua, Nevada de Toluca 12,00', boggy slopes^elow and by spring FNALME1 °e't . E .K. Balls as B. frigidum 25 Feb. 1979 A- U S E . K . Ba l l s 13 A p r i l 1938 Popo, Mexico State Paraje Provincial growing in shade of deep Darrancas on moist rocks FNALME2 det. E . K . Balls as B. frigidum 25 Feb. 1979 5- W T U F. J . Hermann # 25518 28 August 1973 Fremont C o . , Wyoming Burrough's Creek Road a l t . 8400', Absaroka Range 14 ml, N. on wet boulder in stream on spruce-f ir slope FWASWY1 det. F.J.Hermann as B. frigidum 10 Jan. 1979 6- W T U , U S F . J . Hermann t 25578 29 August 1973 Teton Co. , Wyoming Along Hidden Fal l s t r a i l , NW of Jenny Lake a l t . 7000', Teton Range, par t ia l ly submerged in shallow stream FWASWV2 FNALWY2 det. F . J . Hermann as B. frigidum 10 Jan.1979 25 Feb. 1979 7- B I H B . L . Holslngton * 2702678 2 June 1978 Swift Creek, Wyoming Bridger National Forest Campground on rock in stream FBLHWY1 det. B . L . Holslng-ton as B. frigldui 8- W T U T . C . Frye 20 August 1925 Mineral Co . , Montana At. Regis, on mud at margin of pool AVASM02 det. T . C . Fyre as B. asperrimum 18 Dec. 1978 9- W T U T . A . Bunaer 19 July 1905 Montana Swan River Valley, east of Flathead and West of the in moist places, on logs in mountain streams WWASMOl det. T.A.Bunser afl 3. vashingtonianun 8 Jan. 1979 Continental Divide 10- C O L O S.Flowora *10. 159 24 June 1967 Lake Co. , Montana Flathead Lake, Bear Dance Recreation area, 3200', on wet rocks in booklet (sic) by roadside LC0LM01 uet. S. Flowers as B. lamprochryseura 9 Nov. 1978 foundation 11. U B C W.B. Schofleld 1 12003 25 July 1960 Lincoln Co . , Montana Leigh Lake T r a i l , Cabinet Mtns. wet stones in streamlet FUBCM05 det. BL Hoislngtor as B. frigidum 22 Jan. 1979 12- U B C W.B. Schofleld 1 115A1 18 June 1960 Flathead C o . , Montana Bear Dance, Flathead Lake jcommon, forming mats on weepy spring slope LUBIJI01 ri«t. WB Schofleld B. lamprochryseum 18 Feb. 19B t BL Holslngton 18 March 19?1 13- F J H . U S F . J . Hermann 017340 6 June 1962 Plumas Co . , Callfornls Feather River Canyon :alt. ca. 2500", 1 ml. SW 5lorric-, wvt face of bluf b e s i d e Small waterfall AFJ!ICA1 ANALCA3 4at. F. J . Herman ft. anperriaum 23 Jan. 22 Feb. 1979 1979 Appendix A. p. 2, A l i s t of the s i x t y - e i g h t specimens used i n the a n a l y s i s of the v a r i a t i o n of Brachythecium f r i g i d u m (C. Muell.) Besch. sensu Robinson. # herbarium collector and collection # date of collection county/state, province or country location of collection habitat of collection data 8 annotation or verification date of work 14-15-16-17. 1 8 -19-2 0 -21-2 2 2 3 -W T U , U S W T U , U S . M I C H W T U , U S N Y WTU.US.NY F . J . Hermann II 18128 F . J . Hermann * 20559 F . J . Hermann t 20561 P. Train t 3231 E. Lawton t 2696 W T U.US.NY E- L"«<»> C O L O . ' 2 9 8 6 T E N N C O L O , U S I s - Flowers * 396 C O L O C O L O , N Y jS. Flowers iiC 1145 JS. Flowers C O L O , N Y P. "owe™ 0 2240 21 August 1962 [29 August 1966 29 August 1966 8 July 1939 18 June 1955 118 July 1955 Glacier Co., Montana Alberta » September 1925 jsalt Lake Co., Utah lAlberta brmsby Co., Nevada Elko Co., Nevada Nye Co., Nevada 21 July 1928 2 July 1957 2 July 1939 below Logan Pass, Going-to-the-Sun Highway, Glacier National Park Jaterton Lakes National Park, pn north shore of Cameron Lake Waterton Lakes National Park, bn north shore of Cameron Lake creek entering Marlette from Mt. rim along road, Virginia ity Hater Supply fenced area, Jiierra Nevada Range jlumboldt National Forest, Ruby Mountains, Angel Lake alt. ca. 6000', submerged [on rocks in Siyeh Creek -lalt. 5450', overflow of |slugglsh streamlet in wil-low bog, submerged in poo'. WWASM01 IANALM03 IFWASAL3 JFMICAL1 1FNALAL1 lalt. 5450', overflow of tFWAS AL4 sluggish streamlet in wil-tFNALAL2 [low bog, emergent in pool [ Cache Co., Utah Salt Lake County. Utah Summit Co., Utah attached to granite boulders on moist crack Elevation 8000', on 6 o i l pjiyabe National Forest, Toquima] Range, east of Mt. Jefferson, Pine Creek, near campground North Fork, City Creek Canyon, Mlllvll le, Smith's Arboretu Wasatch Mtns; Big Cottonwood Canyon above Maxfleld Mine, Upper Provo River, Stewart's Ranch along the creek on soil LNYGNE1 FVASNE1 P1ALNE1 LNYGNE3 FVASNE4 FNYGNE1 FC0LNE1 LC0LNE4 t-TENTJEl ALNE3 NYCNE2 on dead log over stream AC0LUT1 6000' ANALUT1 4600' submerged on. clean gravel in ditch ca. 7,000', submerged wit tips emergent in slow, cold brooklet 6,000', wet rocks det. as B. det. det. as B. F.J.Hermann [18 Dec. asperrimum -22 Feb. • i i F.J . Hemann l^O Jan. frigidum -12 Jan. ! 6 Mar. I • F.J . Hermann1'10 Jan. 1979 frlRidun | 6 Mar. 1979 1978 1979 1979 1979 1979 [det. P.Train as | l2 Mar. 1979 B. lamprochryseum1 ! BI Hoisington J20 Mar. 1979 jdet. E. Lawton as ;10 Jan. 1979 |B. frlRldm >18 Feb. 1979 I B.L. Hoisington J12 Mar. 1979 3 Jun. 1979 det. las B. frigidum pet. E. Lawton as j 9 Nov. 1978 p. lamprochryseum 21 Jan. 1979 8 Feb. 1979 2 March 1979 net. S. Flowers as fe. asperrimum COLUT22 (iLCOLUT6 LNYGUT3 IC0LUTI3 LNVCUT9 .0 Jan. 1979 .0 Mar. 1979 9 Nov. 1978 9 Nov. 1978 b Feb. 1979 B.L.Hoisington Si Mar. 1979 let. S. Flowers as I. lamprochryseum let. S. Flowers as >. lamprochryseum det. S. Flowers B. latnproclirvscum 9 Nov. 1978 9 Nov. 1978 12 Mar. 1979 9 Nov; 1978 12 Mar. 1979 OO Appendix A. p. 3 , A l i s t of the s i x t y - e i g h t specimens used i n the a n a l y s i s of the v a r i a t i o n of Brachythecium frigidum (C. Muell.) Besch. sensu Robinson. h c r b a r i u m collector and col I act Ion Jt date of collecCion eounty/fffc^te, province or country location of collection Vakftat ef collection d a t a * annoratftin or vcr! flcatlDn aarc of-vorlc-24- C O L O , NY C. Hermann 20 Jv»e 195? /uaU Co., Vtah Deep CrealC rfaunraliw, Tlwmas Creak, on quartzite rock Along drainage way, 10,000' LC0LUT5 ACOLUT4 d«f. G. Heroiann S. InmprocKryjcum 9 Nov. 197B def. ateo *S-B. Bnpfrrimtia det. Q. Hermann B. laCT!prochry9eua 9 Nov. 1978 12 Mar. 1979 2 5 - C O L O , N Y J.L.Drake * 6699 18 November 1967 Tooele Co., Utah Stansbury Mtns; South Willow Creek, near Middle Forest Camp attached to slender twlge dangling in brooklet Lcourr i FNYCL'T 11 det. J . L . Drake' B. lamprr.chryseua 9 Nov. 1978 5,800' FC0LUT11 det. J .L . Drake as B. frigidum 10 Mar. 197* • 10 Nov. 1979 2e W T U C.H. Jones 1 866 6 May 1928 Latah Co., Idaho Cedar Mountain Dn decaying wood LWASID3 det. C.N. Jones at B. lanprochryseum 10 Jan. 197$ 27. W T U A.M. Steele # M-125 20 June 1973 Custer Co, Idaho Bear Creek Road, TISN, R17E, S34 elevation 8000', Aspect 80 Slope 1Z, clinging to rocks in stream FWASID4 det. A.M. Steele B. frigidum 10 Jan. 1979 2&-W T U T.C. Frye 29 June 1934 Valley Co., Idaho about 30 miles east of Cascade along road to Stanley, near the source of Deadwood River in vet log along River (si c) LWASID2 det. T.C. Frye aa B. lamprochryseum 10 Jan. 1979 2 9 - W T U , NY E.F. Layser t 1509 August 1970 Boundary Co., Idaho South Fork of Salmo River, Sec. 18, T65N, R5W, 5000' seepage areas FWASID6 FNYGID1 det. E.F.. Layser aa B. frigidum 1 B.L.Hoisington 10 Jan. 1979 10 Mar. 197P 23 Mar. 197P 30- W T U T.C. Frye 4 July 1934 Fremont Co., Idaho Trude, along Snake River Dn wet soil close to water FWASID5 det. T.C. Frye as B. frigidum 10 Jan. 1979 31- U B C I.A. Worley * 9738 23 July 1968 • Alaska East shore of Kruzof Island, North of outlet of Fred's Creek forested lava c l i f f s immediate upon sound, overhanging wet c l i f f rocks AUBCAK6 det. I.A. Worley as B. asperrimum 10 Feb. 1979 32- W T U , N Y tf.J. Eyerdam il 543 22 June 1939 Alaska. Port'n T f t*trong, Baranof Island, on bark of decaying spruce logs, alt. 10 -m-. AWASAK1 ANYGAK 1 Jet. W.J.Eyerdam as B. asperrimum 1 BL Hoisington IS Dec. 1978 6 Mar. 1979 8 Mar. 197? 3 3 - U B C I.A. Worley t 12922 13 July 1969 Alaska Upper'end of Lituya Bay, Glacier Bay National Monument Just above Fucus line on exposed rocks LUBCAK2 det. I.A. Worley B. lamprochrvseum 18 Feb. 197? Appendix A. p. 4, A l i s t of the s i x t y - e i g h t specimens used i n the a n a l y s i s of the v a r i a t i o n o Brachythecium f r i g i d u m (C. Muell.) Besch. sensu Robinson. f 34-35-36-37-3S-39-40-41-42-h e r b a r i u m M I C H M I C H , NY, T E N N W T U C O L O U B C U B C C O L O , N Y col lector and collection U date of* collecrfoQ H.T. Shacklette 21 November 1965 U.J, Eyerdam I 889 D. Toren 9 475 S. Mulalk I 8089 L. Holmberg t 960 M.A. Hove if 122 S. Flowers « 5049 M I C H , U B C , : D - H . N o r r i s T E N N 1 , 8 8 6 7 U B C F.M. Boas t 423 30 May 1945 June 1972 6 June 1946 county/state, province or country Lake Co., California Tulare Co., Calif. 9 September 1968 Siskiyou Co., Calif. 25 July 1894 29 August 1957 18 May 1968 9 May 1962 Siskiyou Co., Calif. Tuolomne Co., Calif. Del Norte Co., Calif. British Columbia location of cotlec+Ton Near Cyril Cove, Ber1*3 3<a Coast, AmchUICa Island, j Aleutian chain ! Part Vita, Raspberry Island, Kodlak group near Hell's Peak Sequoia National Park ridges south of Cydone Gap, Preston Peak Quad. Sec 8, TI7N R8W, h&fcltat-of £olll«cr(o*i . [attacked1 to breccia in botfon of rapidly flowing tundra stream nn damp ground under alders ' Slsson Sierra Nevada Mtns; Middle Tuolomne River, near Cliff House, along Knopki Creek, near Smith River, ca. 18 miles east of Gasquet on Highway 199 Weston Lake, Salt6pring Island data -ff rVMICAK5 I . M I C A K 8 LT£NAK3 1 L N Y G A K 8 1700', on sedge tussock iniAWASCA6 land near running water rotten logs ln open to closed forests !of noble f i r , western white pine, brewer spruce and Alaskan yellow cedar, |on moist diffusely l i t rock, elev. ca. 5600' on log ln stream WC0LCA2 UUBCCA6 U.UBCCA2 Forkon damp soli near brook inFC0LCA6 [woods, 4600' |on moist diffusely l i t shale roots in creek F N Y G C A 4 AMICCA1 ATENCA4 AUBCCA4 LUBCBC79 annotation or verification der. HT ShackleUi 15 Jan. 199 as B. onpcfrlaum det. W.J. Eyerdam B. lamprochryseum ann. H. Koblnbon 6 July 1959 det. D. Toren as B. asperrimum I EL Hoisington det. S. Mulalk as B. asperrimum I B.L. Hoisington dare of-wsrk 12 Jan. 1979 21 Jan. 1979 27 May 19 79 18 Dec. 197a 27 Mar. 1979 9 Nov. 1978 28 Mar. 19S det. L. Holmberg ;10 Feb. 1979 as B. asperrimum I B.L. Hoisington)18 Mar. 1979 det. M.A. Howe as B. lamprochryseum ! B.L. Hoisington det. S.Flowers as 3. frlRldum ! B.L.Hoisington det. D.H.Norris as B. asperrimum ! B.L.Hoisington det. F.M. Boas as |B. lamprochryseum ! B.L.Hoisington 18 Feb. 1979 28 Mar. 19B 9 Nov. 1978 10 Mar. 1979 29 Mar. 1979 12 Jan. 1979 18 Jan. 1979 10 Feb. 1979 31 Mar. 1979 14 Feb. 1979 4 April 19B Appendix A. p. 5, A l i s t of the s i x t y - e i g h t specimens used i n the a n a l y s i s , of the v a r i a t i o n of Brachythecium fri g i d u m (C. Muell.) Besch. sensu Robinson. h e r b a r i urr collector and ] collection t date of collection county/state, province or country location of collection habitat of collection data t annotation or verification date of work 43- U B C W.B.Scbofield 1 16183 15 Oct. 1961 Vancouver Island, British Columbia Elk F a l l s , near Campbell Siver over shaded boulder AUBCBC36 det. WBSchofield as B. asperrimum ! BLHoisington 22Jan 1979 6 April 1979 44- F J H . U B C F.J. Hermann # 19081 24 July 1963 Yakima Co., Washingtoi s along Raven's Roost Road,22 ml | NE Chinook Pass on spruce-pine slope, a l t . ca. 4000', mucky bank of streamlet FFJHWA2 FUBCWA1 det. F.J.Hermann as B. fricidum ! BLHoisington 23 Jan 1979 22 Jan. 1979 10 April 19B 4 S B L H B.L. Hoisington 23 May 1978 Washington | ONP, Sol Due Campground on rotting wood in stream on seepy slope FBLHWA3 det. .B.L.Hoising-ton as B. fri g i d u 10 April 193 4 6 - U S F.J.Hermann # 18879 16 July 1969 Onion Co., Oregon 1 opposite Catherine Creek, ea. 10 miles SE of Union on steep roadside embank-ment forming dense beds or boggy edge of rivulet FNAL0R3 det. F.J. Hermann as B. frigidum ! B.L.Hoisington 6 Mar. 1979 12 April 19B 47- U B C U.B. Schofield f 21947 17 August 1963 Union Co., Oregon Humbug Mtn State Park wet c l i f f i n streamlet LUBC0R1 det. WBSchofield B. lamprochrvseua 18 Feb. 1979 - ! B.L.Hoisington 14 April 193 4 8 - B L H B.L. Hoisington t 14260578 26 May 1978 Oregon Susan Creek, campground on s o i l on rock i n stone wall ABLH0R2 det.BLHoisington as B. asperrimum 14 April 19 B 4 9 u e c W.B Schofield and J.G. Codfre M734 16 February 1978 J British Columbia Lynn Creek, North Vancouver seepage slope FUBCBC15 det WBSchofield as B. fripidum ! B.L.Hoisington 22 Jan. 1979 6 April 1979 5 0 -51-52-U B C M I C H . T E N T E N N W.B.Schofield and F.M. Boas # 19033 Rzedovski t 19457 1 19471a. • 21 July 1962 28 March 1965 British Colucbls Mexico: Canada de Conteras Taylor River, ca. 17 ml. If. of Port Albeml, Vancouver Island alrededores del 4* Dinamo, DF, on alder ladera, andesltlca con vegetacion de bosque de Abies roligiosa 3100 =. AUBCBC92 i FMICME2 FTENME1 FTENME2 et. WBSchofield as B. asperrimum ! B.L.Hoisington det. Rzedowskl as B. frigidum 30 Jan. 1979 6 April 1979 15 Jan. 1979 18 Feb. 1979 18 Feb. 1979 53- U B C W.B. Schofield 1 12068 26 July 1960 Sanders Co., Montana Snake Creek, Bull River, Cabinet Mountains on boulders ln stream FUBCM04 LUBCM02 det. BLHoisington as B. fricidum det. B. lamprochrvseua 22 Jan. 1979 26 Jan. 1979 18 Feb. 1979 5 4 - U B C I.A.Worley t 7391 27 May 1968 Alaska Trout Creek Timber Sales Area | about 4 miles SW of Mt. Francis Kutiiusko Island, 56 K , 133 45'y 1 >ase of Salmon-berry shrul in stream, opening In forest AUBCAK3 1 det. I.A. Worley as B.asperrimum 10 Feb. 1979 CA CO Appendix A. p. 6, A l i s t of the s i x t y - e i g h t specimens used i n the a n a l y s i s of the v a r i a t i o n of Brachythecium frig i d u m (C. Muell.) Besch. sensu Robinson. # h e r b a r i u m co 11 «cror and cot lecr-fon^ date of collection cownty/atate, province or country location of collect-Ion habitat of Collection data # annotation or verl f 1 cat ion date, of-•ttrk: 5 5 - U B C W.U. Schoficld i !).W. Jaraicuon II 40436 1 November 1961 Brltliili Columbia about IS mlli;a W. of Popkum, Franer Hlver Valley on boulders In alder thicket AUBCBC97 det. WBSchofield as B, aMpt-rrlcj.n 1 B.L.Holsington 30 3 Jan. 19 7) Apr. 1979 5 6 - C O L O S. Flowers It 8025 3 August 1950 British Columbia Stillwater, 15 miles cast of Powell River ca. 250', on rotten logs in shady woods AC0LBC1 det. S. Flowera as B. aiperrittum 1 B.L.Hciislngton 9 U lov. 1978 Apr. 1979 5 7 - U S . N Y L.C.Smith * 15 21 January 1936 Alaska Sitka end vicinity up Indian River Trail growing on Small alder or salmonberry bush ANALAK1 ANYCAK4 det. LC. Smith as B. asperrlrcura ! B.L.Holsington 25 6 27 Feb. 19B Mar. 19 7) Mar. 19B 58- C O L O , U B C , T E N N , US, NY. W.B. Schofleld 29 December 1960 Santa Cruz Co., Calif. canyon of Boulder Creek at Boulder Creek abundant over lower shelves and on wet canyon walls ACOLCAl ATENCA5 AUBCCAIO AL'BCCAll ANALCA9 A.VYGCA16 ANYGCA17 9:iov. 1978 18 Jan. 19B 10 Feb. 19» 10 Feb. 197) 22 Feb. 19B 10 Mar. 19B 10 Mar. 19B 59 - U B C W.B. Schofleld II 28463 27 February 1966 British Columbia Sumas Mountain, near Kllgard on earth of slope AUBCBC23 det. WBSchofield as B. asperrimum ! B.L.Holsington 22 7 Jan. 19 B Apr. 19S 6 0 - U B C W.B. Schofieldk F.M. Boas * 19093 28 July 1962 Clallam Co., Washlngto i Hurricane Ridge, Olympic National Park damp earth near stream LUBCWA3 det. WBSchofield B. lamprochryseum 18 Feb 1979 1 B.L.Holsington 9 Mar 1979 61- U B C W.B. Schofleld?, B.L. Bohm II 59626 . 9 April 1976 British Columbia Kanaka Creek, east of Haney on leanning (sic) trunk of Acer clrcinatum AUBCBC89 det. WBSchofield as B. asperrimum ! B.L.Holsington 30 6 Jan 1979 Apr 1979 6 2 - U B C W.B. Schofleld II 57753 28 May 1975 British Columbia Upper Deena River, Moresby I . , Queen Charlotte Islands seepy muck flat on forest slope AUBCBC99 det. WBSchofield as B.' asperrimum 1 B.L.Holsington 30 6 Jan 1979 Apr 1979 6 3 - M I C H , U B C R.R. Ireland & A. J . Sharp II 9216 17 October 1964 King Co., Washington 4 miles east of Black Diamond, Green River Gorge, on boulders beside water FMICWA1 FUBCWA2 det. R.R.Ireland as B. frigidum 1 B.L.Holsington 12 Jan 1979 22 Jan 1979 1 J Apr. 1979 . 64 - B L H B.L. Holsington 1240578 24 May 1978 Clallam Co..Washington Olympic National Park, Graves Campground on root arm near river FBLHWA2 let. B.L.Holsing-ton as B. frigidum 10 Apr. 1979 i Appendix A. p.7, A l i s t of the s i x t y - e i g h t specimens used i n the a n a l y s i s of the v a r i a t i o n of Brachythecium f r i g i d u m (C- Muell.) Besch. sensu Robinson. h e r b a r i u m collector and collection 0 date of collection i county/state, province or country location of collection habitat of collection data S annotation or verification date of work 65-6 6 -6 7 68-M I C H U B C U B C B L H W.J. Eyerdam t 3022 W.B. Schofleld J . & G. Godfrey t 67994 W.B,Schofleld & J . and G. Godfre t 67078 B.L. Holsington t 260578 30 March 1948 22 March 1978 23 March 1978 y 26 May 1978 Snohomish Co., Wash. Clatsop Co, Washington Multnoman Co., Oregon Oregon Mt. Pilchuck, Cold Basin Saddle Mtn. State Park. Latourell Falls, Columbia Rivei Gorge 5 miles from dsporc on Rte. 38 on alder trees sloees (?) and peak, earth by tra i l , (drier, pers. comm. WBSchofield) log on slope side of road on tree AMICWA1 AUBC0R3 < AUBC0R2 ABLH0R1 det. W.J. Eyerdam as B. asperrimum ! B.L. Holsington et. WBSchofield a: B. asperrimum ! B.L.Hoisington-det. WBSchofield as B. asperrimum ! B.L.Holsington det. B.LUoisingtoi aB B. asperrimum' 12 Jan. 19S 10 Apr 1979 8 Feb. 19 » 12 Apr 1979 8 Feb. 1979 14 Apr 1979 14 Apr 19B rH • C •H o Tj CD U CD O cq 4-> IQ • U rH •H rH CD CU .d • +-> o <M O s X X ! •H •H u br -t-> •H a H £ CU e o • H 3 O •H 0) X ! 3 4-> > >: o Xi o O a cu u Xi m E H O rH CO d) • r H !=> •9 ' ' cu fH C E H • O O W cq co - H vo ^0 X . o •H <t-t K x i o CD E) W Pn o a ft-H (1) < -P W STEM LEAF TRAITS ERANCH LEAP TRAITS LEAF LENGTH 0 . 0 4 3 8 LEAF WIDTH 0 . 0 2 7 5 0 . 0 3 6 9 COSTA RATIO 0 . 0 1 3 4 0 . 0 2 6 0 O .O56O MEDIAN CELL LENGTH 0 . 0 2 9 0 0 . 0 2 0 9 0 . 0 1 4 5 0 . 0 5 3 2 MEDIAN CELL WIDTH 0 . 0 0 0 3 0 . 0 1 8 0 0 . 0 1 5 2 0 . 0 1 4 0 LEAF SHAPE 0 . 0 2 2 2 0 . 0 2 7 0 0 . 0 2 4 4 0 . 0 1 7 9 ACUMINATION 0 . 0 0 4 5 0 . 0 2 2 5 0 . 0 3 3 8 O.OI65 MARGIN REFLEXED -C.OO67 O.OO58 0 . 0 0 5 3 0 . 0 1 1 0 PLICATION 0 . 0 2 1 7 . 0 . 0 3 4 1 0 . 0 3 6 3 0 . 0 2 8 5 APEX TWIST 0 . 0 1 5 1 0 . 0 1 1 9 0 . 0 2 0 6 0 . Q 1 9 2 APEX FALCATE - 0 . 0 0 1 0 -O.OOO6 - 0 . 0 0 4 6 -O .OO3I ALAR C E L L S 0 . 0 2 9 5 0 . 0 4 4 3 0 . 0 4 2 7 O . O 3 8 6 LEAF LENGTH 0 . 0 2 8 9 0 . 0 2 3 0 0 . 0 1 4 9 0 . 0 2 9 1 LEAF WIDTH 0 . 0 1 3 0 0 . 0 2 1 8 0 . 0 1 6 6 0 . 0 1 4 8 COSTA RATIO 0 . 0 0 8 2 0.0144 0 . 0 2 5 2 0 . 0 0 7 9 MEDIAN CELL LENGTH 0 . 0 2 2 1 0 . 0 1 3 7 0 . 0 1 5 4 0 . 0 3 4 5 MEDIAN CELL WIDTH 0 . 0 0 2 2 0 . 0 1 8 3 0 . 0 2 0 9 0 . 0 1 3 0 LEAF SHAPE 0 . 0 0 6 6 0 . 0 0 7 4 0 . 0 0 7 3 0 . 0 0 6 8 ACUMINATION 0 . 0 0 6 2 0 . 0 2 4 6 0 . 0 3 7 1 0 . 0 1 7 4 MARGIN REFLEXED 0 . 0 0 2 2 0 . 0 1 8 0 0 . 0 2 5 5 0 . 0 2 0 6 PLICATION O .OO91 0 . 0 2 3 9 0 . 0 2 9 1 0 . 0 2 4 3 APEX TV; 1ST 0 . 0 0 7 3 0.0146 0 . 0 2 4 4 0 . 0 1 8 5 APEX FALCATE - 0 . 0 0 6 2 - 0 . 0 0 7 4 -O.OO88 - 0 . 0 0 3 9 AIJAIR CZ.1I1.JS - 0 . 0 0 0 9 0 . 0 1 5 2 0.0242 0 . 0 2 0 5 0 . 1 3 9 0 0 . 0 3 2 0 0 . 0 2 9 4 0 . 0 2 8 5 0.0420 O . O 3 1 9 . 0 1 3 4 . 0 5 4 7 .0493 • 0 5 3 6 . 0 3 6 5 - 0 . 0 0 4 8 0 . 0 7 0 9 0. 0. 0. 0. 0. • C - 5 9 3 • 0 3 7 9 . 0 1 9 3 , C A 2 5 , 0 1 0 8 0 1 6 7 0 1 6 7 C2<57 0 3 3 8 0 . 0 2 3 1 •O.OO83 0 . 0 1 8 2 0 . 0 4 3 6 0 . 0 1 9 0 0 . 0 2 4 2 C . 0 2 2 3 0 . 0 2 1 6 0 . 0 1 9 7 0 . 0 2 3 9 0 . 0 3 0 0 0 . 0 2 5 1 - 0 . 0 0 2 9 0 . 0 2 1 0 0 . 0 3 6 9 0 . 0 1 5 4 0 . 0 1 5 3 . 0 0 8 5 • 0 3 1 3 . 0 2 1 1 . 0 2 8 3 . 0 1 8 7 - 0 . 0 0 3 1 0 . 0 1 5 9 0. 0. 0. 0. 0. 0 . 0 5 8 8 0 . 0 1 0 7 0 . 0 1 1 5 0 . 0 1 0 8 0.0144 0.0226 0.0143 -O.OO3I 0.0246 O.G405 0 . 0 0 7 1 0 . 0 2 8 0 r> .—"^ f V.U£GG 0 . 0 3 3 0 0 . 0 2 7 3 - 0 . 0 1 3 9 0 . 0 3 3 1 0 . 0 3 4 5 0 . 0 1 0 0 0 .0333 O.C246 0.0201 0 .0516 0 .0133 0 .0093 0 .0274 0 .0843 0.0244 0 .0335 0.0408 0 .0284 c.0096 0.0156. 0 . 0 2 6 8 0 .0136 0 .CO25 O.OC35 - 0 . 0 0 2 9 0 .0016 0 .0308 0 .0360 0 .0452 0 .0263 0.0098 0 .0209 c . 0 1 7 2 - 0 . 0 0 2 1 0 .0189 0 .0179 0.0204 0 .0078 0 .0139 0 .0129 0 .0279 ' 0 . 0 0 7 5 0 .0038 0 .0177 O.OI59- 0 .0091 0 .0260 0 .0119 0 .0256 0 .0131 0 .0043 0 .0174 0.CO84 0 .0102 0 .0238 0 .0219 0 . 0 4 6 0 0.0270 0 .0233 0 .0132 . 0 . 0 3 1 8 0 .0348 0 .0308 0 .0289 0.0420 0.0271 0 .0161 0 .0234 0 . 0 2 8 6 0 .0158 - 0 . 0 0 6 7 - 0 . 0 0 7 7 - 0 . 0 0 9 7 0 . 0 0 0 0 0 .0228 0 .0157 O.C275 0 .0156 0 . 0 3 2 9 0 . 0 0 9 4 0 . 0 1 5 5 0 . 0 2 8 1 0 . 0 1 6 0 - 0 . 0 0 2 7 0 . 0 1 6 5 0 . 0 6 2 8 0 . 0 4 2 0 0.0439 0 . 0 4 0 9 -O.OO9I 0 . 0 3 6 9 0.11O6 0.0599 0.0260--0.0075 0.0596 0 . 0 9 9 0 0 .0274 0 . 1 0 1 5 0 .0023 - 0 . 0 0 . 5 0.0616 0 .0680 0 .0258 - 0 . 0 0 0 1 0 .0277 0.0143 0 . 0 0 c 5 0 .0253 0 .0164 0.0016 0 .0275 0 .0189 O.CC-o 0.0241 0.0072 0.0112 0.0268 0 .0163 -o.ocoo 0 .0280 0 .0106 0.0-023 0 . 0491 0.0237 O . C C i i 0 . 0 5 7 2 0 .0103 - 0 . C 0 C 3 0 .0675 0 .0329 ' - 0 . 0 0 0 3 0 . 0 2 9 4 0 .0522 - 0 . 0 0 1 S - 0 . 0 0 7 4 0 .0003 0 .0226 0 . 0 4 8 5 0 . 0 2 6 9 - 0 . 0 0 2 3 0 . 0 8 6 8 0 . 0 4 2 6 0 . 1 2 5 1 - O . O I 7 6 - 0 . 0 1 9 7 0 . 0 6 1 2 0 . 0 4 8 4 0 . 0 7 7 7 - 0 . 0 0 0 8 0.111!! 92. Appendix B. Table 2. The mean, variance and standard d e v i a t i o n of the 2k v a r i a b l e s ( t r a i t s ) used to estimate the phenetic v a r i a -b i l i t y of 68 O.T.U.'s of Brachythecium f r i g i d u m (C. Muell.) Besch. sensu Robinson. Note: these values correspond to standardized v a r i a b l e s . STANDARD MEAN VARIANCE DEVIATION LEAF LENGTH 0.390 O.V58IE-OI* 0.2093E+00 LEAF WIDTH 0.^50 O.3686E-OI 0.1920E+00 COSTA RATIO 0.616 O.56OIE-OI 0.2367E+00 STEM MEDIAN CELL LENGTH O.kkk 0.532^-01 0.2307E+00 LEAF MEDIAN CELL WIDTH 0.k91 0.3^5E-01 O.1856E+OO TRAITS LEAF SHAPE 0.511 0.338ifE-01 0.1840E+00 ACUMINATION O . 5 6 2 0 . 5 1 5 6 E - 0 1 0.2271E+00 MARGIN REFLEXED O . 5 6 8 0.8V31E-01 0.290^+00 PLICATION OA85 0.9900E-01 0.31^+00 APEX TWIST 0.32k 0.1015E+00 0.3186E+00 APEX FALCATE O.096 0.6162E-01 0.2J+82E+00 ALAR CELLS O.509 0.1390E+00 0.3729E+00 • LEAF LENGTH 0.520 0.6930E-01 0.2632E+00 . LEAF WIDTH 0.512 o.^359E-ol 0.2088E+00 COSTA RATIO 0.593 0.3685E-01 0.1920E+00 BRANCH MEDIAN CELL LENGTH 0.^93 0.5882E-01 0.2^252+00 LEAF MEDIAN CELL WIDTH o.k9k 0.i+053E-01 0.2013E+00 TRAITS LEAF SHAPE 0A50 0.3286E-01 0.1813E+00 ACUMINATION 0.659 0.6276E-01 0.2505E+00 MARGIN REFLEXED 0.532 0.1106E+00 0.3325E+00 PLICATION 0.611 0.8678E-01 O.29A6E+OO APEX TWIST 0.3^6 0.1251E+00 0.3536E+00 APEX FALCATE 0.152 0.7773E-01 0.2788E+00 ALAR CELLS 0.7V7 0.1118E+00 0.33^+00 *Note: E denotes the exponential power to which each value i s r a i s e d , e.g. "E + 0 5" i s equivalent to " X 10-7". 93-Appendix C. Table 15 O.T.U.'s of B. 1. The c o r r e l a t i o n matrix of t h e . o r d i n a t i o n of asperrimum ( M i t t . e_x C. Muell.) S u l l . O O O O O O O O O O O O H " W P H ' O f W H I U W Q H O Q ^ H C ^ M H ^ W O V t l O y C O O v O v o C J i V J - O O ^ r u O o v O O - v ) 0 O O O O O O O O O O O W - P - r v o i - - - r o P r o c r - Q r o f o c n v n O C o v n O o .p - v n O O -0 O V J O v n c o 0 - - • - - 0 0 O O O O O O O O O p l - ' V n v n P V J v O t - j r o P - v n i - , Q 4 - v n v o c o r o o i o r o v n M O 1 1 1 1 1 O O O O O O O O O l - ' Jp- c r C S V n V I V I C M U 1 W Q - r r o o c r v V r t O - ^ - i ^ o o ^ J Q f u - p - v O I - 1 4r v n 0 0 0 0 0 O O O O O O O O I - ' V n op- -p-vi M - r l y w o H H H i r Q v o o ^ - v J V - J r o ^ j v i Q - u v n v o v o c n - s j o v o o O O O O O O O t - 1 w ro o v i o o w 0 4- JP- c o g O O J 0 r u v n o - F - ^ i - o -p - v o r u 0 O O O O O O M f H V J V I H O O r o 1— V O V-' a - , V J O 0~> O - J V n M C P . Q - v j : 4~ O 0> V J o 0 o 0 0 0 M r o M o V J M 0 r o v n 6 - c - - J - Q O 00 Q c n ^1 Q C o H' O v o v O O O O O O H 1 H M H P Q o r o 4- - o Q 1-- OVJ - v ] o r o 4- i - " - p - O r o - r r V n V J 1-9 o H t-1 tn H G tS (5 8 u a c i M M Hj fe fe > " " ro ci f l 0 Pl o O B B W M H •*! fe fe s> W M M n n E a K M M i-3 i-3 O t< r1 O X G G ,~ M G O O !t> 0 2: . . . . . W M H 1-] O ••I o M r-" r H i-H H H - J C - f O 3: Pj Cl X 1-3 O O O O O O O O O O O O O O O O O O O O O O O l - 1 V J l H M O H r o M O O ^ H M l 6 H l o v i o O V I N I V O ro op- H* V J - O v O 4 r v o p V n - O V n O - i O V n O v o u i O i v o v n r o t - ' ^ o o I-J w f o f o C B C o H o v i v j r o v i r j ^ V I O ^ V J V J C o v D H \ J 1 C O H V J V n r o V J v n c n - v l p V J 01 v n O O O O O O O O O O O O O O O O O O O O O O W r o H V J W H r u c o r u r o s i Q -p-vi o o v - j v n v o 4-- r o C O C A O r o - v i c i 4r- 00 c r v v n -N3 V J i — O H C O M V D H H H V l - ^ - O O M H V I H O V l H I - ' O V l V l f -v i rv v i f vi v i O C ^ H D o i - " r u - p - O v V J V I V n - P - O CM.H v n V J r o - P - O O - P - V n 00 v n o w n v n v i O I O O O O O O O O O O O O O O O O O O O O O M V I M H I J Q V l l - ' W W V l O H O I M V I ^ O P - O O M J M J 1 H - - J H r o - P - O V J as o v o - o 0 V J r o v o p - ^ D O H V 1 V l \ O V I u M ^ O ) V n r o r o Q O V J - o r o V J i O c c v i y o ^ x ) O M - v J v O V i r o 4 - r o C o o o V n - 0 ' r o - x j -0 00 4- v n - v ] c o 1 O O O O O O O O O O O O O O O O O O O O O . . . . . . . , f M P O M O M ^ 4 - M r o o * 5 P ~ c r \ i-> - o 00 w 00 4~ v o 00 r o v n o w n P A O O O ^ J ^ O V I W V I V ! O l V I V I M H - O W O V J P V I g V O r o V n V I O V n v n Q O o V J O V J J> - v l J-- I v Q p - p p v o r o - 0 - 0 r o o O O O O O O O O O O O O O O O O O O O H' r o r o o -0 v i r o v i v n M 0 v n v i M v u v i i H o r o > j o o o c n v a v j r o v o v j v o v o r o v j v i O M V r o ON w r o g v i N i ^ > ] H p r o r o v i o r o o > 0S0 O«0 C O C O P - C O V J V J O o V I O O O O O O O O O O O O O O O O O O t J V J o p-ro o O M J J H V I V I P H V n O - O V J h - ' v j j C o l - ' O v O v O O o - O ^3 r o - P - C O CX> CT* O O v O C7\ o o r o 4 - v i v i c o c r \ p c r * v i v i o H o v i op-1—' P ( - O V n J p V l v O ( V I * X ) H P V I TO! r o 0 P - P M O O i O O O O O O O O O O O O O O O O O ^ - 1 M O v n O W Q U l P f l ) O C O Q M O 4- 4> V J O O J 5 - v n H ' 00 o 0 0 0 0 1-' O H H P f u O H O l P P p r u c B H N i a i P u u j C I M V I VIJ V n i - J O CTxvo O C u V l C o O C A V J V i ^ o M u v i N i o o i P v i r u I I I I I I I I I O O O O O O O O O O O O O O O l J >-• l - J M O V J I V VtJ • r - r o 4:- o c r u o v i p - r o r o H % i V J P- v n i- 1 v n o > O v n r o P ^ J 00 V l 0 0 V I I ' V J H N ) V I 4 - H V l O l O O O O O O O O O O O O o o V J r o 0 - r VH M M H - r o o V i y- 1 v i v O V n 1-- r o \ - J e n c r . ^ l 00 r*J 4- V J M V J c o o i 4- 1—1 v n VO 4^0vo P O U J O i - \ ] ^ - v l \ 0 O O O O O O O O O O O O O O I - J O K O Q ( W H H O ( V O V J v O 4 - ( v o r o v n as < O O O t - i o 4- r o P O U I P O K OMOI o u i H H 00 v n v o 4- 4~ c r * v o 4- Q ' V i r \ j - o v o VKJ'O v n r . i r o V n Q U I H 1 00 c o M o v v n V J v o O O O O O O O O O O O O r o 0 i ~ * v n K-1 i - j V J v i H r o o r o H v i D Oi Cv p - 4- VO O Vn CUV.-I C l r v i - * i i r v v .-j p - - v l O v o o w . o v n Vjl V J x l l-- 1 V I P-VJ OV Cn •• ' ON J« 9h. Appendix C. Table 2 . The mean, variance and standard d e v i a t i o n of the 28 v a r i a b l e s measured i n . t h e o r d i n a t i o n of 15 O.T.U.'s of B. asperrimum ( M i t t , ex C. Muell.) S u l l . STANDARD MEAN VARIANCE . DEVIATION LEAF LENGTH. 15^1.000 0 . i t 8 l 8 E + 0 5 * 0 .2195E+03 LEAF WIDTH 0 .1882E+05 0 .1372E+03 - COSTA RATIO 0 . 5 0 8 0 . 5 7 5 3 E - 0 2 0 . 7585E-01 STEM MEDIAN CELL LENGTH 6 5 . 9 2 9 0 .7788E+02 0 .8825E+01 LEAF MEDIAN CELL WIDTH 6 . ^ 9 3 0 .V330E+00 O.658IE+OO TRAITS LEAF SHAPE Z.klh 0 .3783E+00 0 .6150E+00 ACUMINATION I . 8 7 1 0 .8990E-01 0 .2998E+00 MARGIN REFLEXED 0 . 3 2 9 0 .7143E-01. 0 .2673E+00 PLICATION 0 . 1 5 7 0 .3187E-01 O.I785E+OO APEX TWIST O .071 0 . 1 6 0 ^ - 0 1 0 .1267E+00 APEX FALCATE 0 . 0 2 9 0 . 5 2 7 5 E - 0 2 0 .7263E-01 . ALAR CELLS 0 . 0 5 7 0 .2725E-01 0 .1651E+00 LEAF LENGTH 1 7 7 5 . 7 1 ^ 0 .1157E+06 0.3401E+03 LEAF WIDTH 7 9 3 . 9 2 8 0 .4059E+05 0 .2015E+03 COSTA RATIO . 0 . ^ 9 9 0 . 5 2 3 1 E - 0 2 0 .7233E-01 BRANCH MEDIAN CELL LENGTH 7 2 . 1 1 4 0 .8076E+02 O.8987E+OI LEAF MEDIAN CELL WIDTH 6.^00 O.^IE+OO 0 .63^9E+00 TRAITS LEAF SHAPE 3 ^ 5 7 0 .3350E+00 0 .5788E+00 ACUMINATION 1 . 6 0 0 0 .8616E-01 0 .2935E+00 MARGIN REFLEXED O .186 O^^OE-Ol 0 .2538E+00 PLICATION 0 . 6 5 7 0 .4026E+00 "0.63it-5E+00 APEX TWIST 0 . 0 2 9 0 . 5 2 7 5 E - 0 2 0 . 7263E-01 APEX FALCATE 0.1^3 0 .3956E-01 0 .1989E+00 ALAR CELLS 0 . 3 5 7 0 .8110E-01 0 .2848E+00 *Note: E denotes the exponential power to which each value i s r a i s e d , e.g. "E+05" i s equivalent to "X l o 5 " . 95-Appendix D. Table 1. The c o r r e l a t i o n matrix of the o r d i n a t i o n of •:k3 O.T.U.'s of B. fri g i d u m (C. Muell.) Besch. o o o o o o o o o o o o o r o Q rv> rv> V H M O O M \-> OOVN - p - OA T J H 0 > H OS - t r v n O O O O O O O O O O O H 1 S H j i - » v > i r o o v > i Q c r » r \ > a A Q OO -tr r o h - 1 OA w OA v n v o V n Q s i \ j i c o H OA - o o v o O \ P O Vj1 V H C3A OOVJ1 V M AJ1 - p " V n OA O O O O O O O O O O O O I - ' O O H M H H f - U W U O V j l O I A I O C O O i - W A j o o H H S 1 H N V ^ 4 - - \ D Q W O v O H C o f O * " V M O O O r l r W H o tn W 1 0 W >-3 1-3 ' ^5 S P > I M i j j 1-3 > £ H o W . . . . 1-3 5 fe > • § " £ p g « g H I tg o • TJ n > I t-1 1^ S K £i b . _ _ > H I I-I fe > 1-3 z a co a a H > f 5 t o O t o H £ • O O > 2 W H ' O W P ^ EE H3 W 1-3 O T l i-3 .. M > C I—1 O J > H I 1-3 M > O H t l H o W t'1 5 S H O EG Hi ss i-9 O K Hi so O O O O O O O O O ) - ' r o o o i — 1 H * V n VM . _ . _ M ^ O O H O O A O -P- \£> r u VM o n Q r u o V P A O *A3 v nI I I I I o o o o o o o o o o o o O O O O O O O O O O O H ' W O - P " M r u V n O O O O Q O . _ . V n COVM - o f H c o - o OA o o o H ' v > J r o v n o a A - p - v o O A O r v > r u v H I j l U I W O O O O H A l M ) H U I ffi O ^ I V M - O _» H o o v n r u 1 J H J 1 A ] V O H ^ O O H O O O O M ) O O ' O O O O O 1—' O O O O H ' I A J H ' I - ' I VM v n r u c o M o r u v o i H \ J 1 H c r \ - o r o f \ ) I o o - o o A o o r u - p - r u i o O i o o o o o o o o o o O I O O O O O O O H 1 o r u o r u v M o - p - V j i O A - p - J r C A l - ' . C - H ' O A r-- OS\D C O O A C O - < ] O -A3 V M V M V O - - 3 H V n V n - f - O A O H ' 0 0 V M O i 8 V M ' v n o o - P - o r u O A V M - p - V M r o r o r u i - ' v o o o o A v n v n r u H J v n i - ' v n o o a A H J v > i V M ! - J v O H U 1 - p " O V M r o H " -C" H 6 o o o o o M V M M O H ' l - i Q • C T i H ?-»0 - O H O - p - TA v n V h i v n M o o o o o o o o o o O O O O O O O O O M Q H ' O Q Q r u O M O H ' H ' O Q O A V M O A O A O l - ( 7 i A l ^ V I ^ ON r u M H * r u v n VM r o OA v n H » o OA Os - C - V n - P " S\ as J" c o - s 3 A O O o P P O H V l U M O H i 4^ O r u r u J - M O O N ] w r u J - j - a w n o o H r o N H - O W I o m S c o v n O O O O O O H O O H " 4?" H 1 O Q H W O Ol C - OO Q - o v o v n O - P - VrJ Q V M 4 ^ - P " OS V M P " O o o v r i OA VM o c o < O O A - V J v n VfJ i 4 j l W H 4 ^ - P - I o o O O H -i i O O f o V n v n r u O o A i y i O H O CA H v n r o O O O O I-" O O H -W H O O O O O O O O O O O O O O O O O I O O O H O _ _ _ 4J1 H - 0 V H V M . - . _ . C O O A v O - p - M V n O A O O I A j h - ' ^ O OA o o v s o OA o v n o o v n os rv a^ O O O O O O V n o r - ^ - P -- ' - ' 1 - P - - C - O A M r P - O o V d W H H H . O - P - O O c o o r o - o o - p - H - ' c o o O * v n O H W H H o - c - r o - o ^ o o v n O A O o O I I I I I o o o o o o o Q H J O O O H ' H J - P - l - O r o O J - 4 r r o \ o v n i - ' c o - < i ^ j H ' O A v n v o - p - O A V u - p - o c o r o v j - p - r o - p -- - o r o r o r o 4 ^ H A O - p - v n v i A O ^ o O O O O O O O O O O O O W o H * O O r o Q c ON r\) v n r o r o - p - J r - ( W O H C i l W H H ( - p - v n v n V J v n \ o V>J < O O O O O O M O H P M O Q - f O W O N W H H - O U 3 ^ V O 4 ^ P - V j J - O N i C D 4 ^ 0 A O A V D - O O A v O P J O A H ' V n ^ J O O V n O A O - O V U V n O A O 4 r - J T 4 ^ O O O O O O O O O O O O i j l O O U l O H ( V n OA CA - o o H< ( o o r o v n O C D O M r o o r o ^ - \ O \ J J ( r m j rt 35-vn CA - - . l is-} m o n H J c o r o 4 > v - J J^- OA r o r o v n r o \ j i v i v o C B r o o c u y i A ] o o \ j j p - N j I I I I 4 I I I O O O O O O O O O O O O O O O O O H H 1 h J CA O 1— Q r \ i ~ - j i - I-- . - o OA - v i (..-j v n i - * O co o o v n v O \ o o O O O O H ' C J O 1 -' O I 1 1 J C } l J O H ' • P - o J - o ^ } < i o r u . r o ^ o r o r u V O C T A O A - P - O O ^ O H C U V T M O 4r-r o - v j o o o o r o o A j ^ r o . V H H V O H o o o o o V - J 1—' V M V Q O A " N ] I—' I—1 O r o o v n c o o o V J o o o b o o o o <b o o o o O O H P M H P H O O H M 3\V1 O O V I H O Q U o w r o o w O H v n v o c o O H C M N i o y i i - O A V U H 1 - O - P - - P - 0 A O C O C O A ) o <S a i -M O O O P - H ru o OA p - v n O O M j I A l O O O O O O O O O O O O O O O H O O V j I O M H O O H O O O O O O M A l P U 1 O C O H ' J M j l H - P - V i J O - P - - - 0 r u O A 3 C J | l - 1 V n * . n H C A o o O O O H ' O A O A \ 0 - p - t j r 0 4 ~ N o 4 - A j O O O A O O O O O O O O O O O O O O h-' -P" O H ' H * O O l J M O Q H * r o f~ P - V l l M V-J i - J V M C o V n O AO r - H - » x - ( 1 A i.A) 4 - - o r - v n r o v n • - * SD 0\ l \ l C o O n V n t " V.-J v o - N ! V M O AM Q r u o O o 96. Appendix D. Table 2. The mean,-variance and standard d e v i a t i o n of the 28 v a r i a b l e s measured i n the o r d i n a t i o n of 43 O.T.U.'s of B. fr i g i d u m ( C . Muell.) Besch. STANDARD MEAN VARI A N C E D E V I A T I O N L E A F LENGTH 1642.593 0.9071E+05* 0.3012E+03 • L E A F WIDTH 710. W t . 0.2023E+05 0.1422E+03 COSTA R A T I O 0.617 0.3257E-02 0.5707E-01 MEDIAN C E L L LENGTH 79-943 0.1901E+03 0.1379E+02 STEM MEDIAN C E L L WIDTH . 7.652 0.4179E+00 0.6464E+00. L E A P L E A F SHAPE 3.200 0.3321E+00 0.5762E+00 T R A I T S ACUMINATION 2.833 0.9923E-01 0.3150E+00 MARGIN R E F L E X E D O.63O 0.7005E-01 0.2647E+00 P L I C A T I O N 2.159 0.773^+00 0.8794E+00 A P E X TWIST O.389 0.1033E+00 ' 0.3214E+00 A P E X F A L C A T E O.OH-1 0.1114E-01 0.1055E.00 ALAR C E L L S 0.626 0.1012E+00 0.3181E.OO L E A F LENGTH 2021 . 3 3 3 011025E+06 0.3201E+03 L E A F WIDTH 1001.296 0.3023E+05 0.1739E+03 COSTA R A T I O 0.631 0.3937E-02 0.6275E-01 MEDIAN C E L L LENGTH 87.150 0.1821E+03 0.1349E+02 BRANCH MEDIAN C E L L WIDTH 7.880 0.4343E+00 0.6590E+00 L E A F L E A F SHAPE A . 030 0.4489E+00 0.6700E+00 T R A I T S ACUMINATION 2.504 0.1249E+00 0.3534E+00 MARGIN R E F L E X E D 0.622 0.8403E-01 0.2899E+00 P L I C A T I O N 2.907 0.5943E+00 0.7709E+00 A P E X TWIST 0.341 0.7944E-01 0.2819E+00 A P E X F A L C A T E O.O78 0.2478E-01 0.1574E+00 ALAR C E L L S 0 . 8 4 8 0.7085E-01 0.2662E+00 *Note: E denotes the exponential power to which each value i s r a i s e d , e.g. "E+05" i s equivalent to "X 10 5". o-Appendix E. The sample of 9 specimens used i n the exemplary o r d i n a t i o n of the s i x North American rough setaed diocous Brachythecium species. A l l specimens are l o c a t e d at the U n i v e r s i t y o f B r i t i s h Columbia h e r b a r i u m T U B C ) . # collector and collection $ date of collection county/state , province or country location of collection habitat of collection data S annotation or verification date of work 1- r.A. MacFaddea t 8014 2 Hay 1929 San Diego County, California, U.S.A. Moreno Dam Base of oak tree Det. A.J. Grout B. bolanderi ] B.L. Holsington 30 May 1979 -2- W.B. Schofleld f 56826 7 August 1974 British Columbia, Canada slopes of Mt. Matier, trail foi l . 3 lakes, e. of Pemberton boulder on subalpine slope among alders Det. B.L. Hoislng ton, 23 May 1979 B. nelsonli 30 May 1979 3-W.B. Schofleld » 56732 22 July 1974 British Columbia, Canada two waterfall area from Kathlyn Glacier on Hudson Bay Mts, Smithers among shrubs in spray of waterfall Det. 0. Ng ! B.L. Holsington April 1978 B. nelsonli 30 May 1979 4-W.B. Schofleld b.W. Jamleson |r 58892 20 September 1975 British Columbia, Canada Just W. of Balfour, W. arm of Kootenay Lake Cm forest floor humus 1 B. L. Hoisingtoi 27 May 1979 B. hylotapetum 30 May . 1979 5-W.B. Schofleld jf 56259 17 July 1974 British Columbia, Canada Bear Lake Campground, ca. 45 mi n. of Prince George log In forest Det. B.L.Holsing-ton, 20 May 1979 B. hylotapetum 30 May 1979 6- W.B. Schofleld t 24817a 25 July 1964 British Columbia, Canada Takakia Lake, about 10 miles S. of Moresby Logging Camp, Moresby Island, Queen Charlotte Islands seepy area of hemlock thicket Det. B.L.Holsing-ton, 21 May 1979 B. rivulare 30 May 1979 7- Y-B. Schofleld #28065 24 August 1965 British Columbia, Canada trail from Garibaldi Station to Garibaldi Lake, wet boulders in creek near Rubble Creek B. asperrimum Ann. 15 May 1979 B.L. Holsington B. rivulare 30 May 1979 8- W.B. Schofleld B.C. Godfrey i 67634 16 February 1978 British Columbia Canada Lynn Creek, North Vancouver seepage slope 1 B.L. Holsington 22 January 1979 B. frigidum 30 May 1979 9. W.B. Schofleld fc.M. Boas V 19033 21 July 1962 British Columbia Canada Taylor River, ca. 17 mi, W. of Port Albemi, Vancouver Island on alder I B.L. Holsington 30 January 1979 30 May 1979 Appendix F. Table l a . P a r t one of the c o r r e l a t i o n m a t r i x of the exemplary o r d i n a t i o n of s i x species of Brachythecium (continued i n Table l b ) . LEAF LENGTH LEAF WIDTH CCSTA RATIO MEDIAN CELL LENGTH MEDIAN CELL WIDTH APEX TAPERED STEM' LEAF SKAFS LEAF ACUHir.'ATION "TRAITS MARGIN REFLEXED PLICATION APEX TWIST APEX FALCATE ALAR CELLS MARGIN SERRATED BASAL CELLS CONCAVITY LEAF LENGTH LEAF WIDTH CCSTA RATIO MEDIAN CELL LENGTH MEDIAN CELL WIDTH APEX TAPERED BRANCH LEAF SHAPE LEAF ACUMINATION TRAITS MARGIN REFLEXED PLICATION APEX TWIST APEX FALCATE ALAR CELLS MARGIN SERRATED BASAL CELLS CONCAVITY 1.0000 0.7919 0.5609 O.6445. 0.3271 0.5382 -0.0537 -O.5945 O.JS53 0.5550 0.2754 - C . O O 9 5 -O.I896 -0.2243 0.2207 0.3247 0.9447 • 0.3372 -O.O607 0.5973 0.0519 O.3923 O.4084 -0.5346 0.2703 0.3360 0.6226 O.2478 -0.2035 -0.3073 -0.3323 0.3812 -0 1.0000 0.0505 0.8823 0.7395 0.7448 0.1354 O.493O 0 . 0 4 2 4 0.4245 0.3623 1974 0.1178 0.3901 0.5794 0.6902 0.8072 0.7339 -0.2802 0.8112 0.4895 O.5665 O.6063 -0.4712 0.0837 0.1671 0.6169 -0.1974 0.0210 -0.4812 -0.2430 0.6921 1.0000 0.1055 1.0000 -0.3112 0.8783 1.0000 0.0417 0.6871' C.5365 1.0000 0.0290 0.3907 0.4788 -0.2304 -0.4211 -0.4053 -0.0783 -0.8012 0.7354 -0.0461 -0.3275 -0.4470 0.5773 0.5C28 0.2678 0.0622 0.2396 0.4107 0.1763 C.6505 0.1516 -0.1148 C.0088 O.O606 -O.6OO6 0.1361 0.5233 -0.2425 0.3621 -0.3919 . -C.6078' -0.1543 -0.0248 O.6898 ' 0.7677 0.0313 -O.3OO8 0.8024 • 0.8039 0.6933 0.4022 0.6775 0.4650 0.5519 -0.4499 0.7028 •  0.8678 ' 0.4631 -0.0340 -O.238O -0.0992 -0.3621 O.OO69 0.8903 ; 0.8255- 0.6953 -0.5234 0.5990 0.8702 • 0.1027 -0.0257 0.4167 0.3783 O.8562 0.0699 O.685I • 0.7279 0.2178 -0.4012 -0.3749 -0.0599 -O.8343 0.5533 0.2003 0.0487 -O.3031 0.5189 0.2785 0.1241 -0.3137 0.4892 0.7112 0.4061 0.6174 0.4169 -0.3785 -0.6282 -0.3032 -0.4561 0.C136 O.3686 -0.0875 0.2401 -0.3737 -0.4436 -0.3789 -0.2610 -0.2026 0.0944 -0.5882 -O.3277 0.7633 0.8216- 0.5370 1.0000 0.4018 0.3465 0.6748 0.0096 -0.1250 0.4285 0.0000 0.6971 0.03C2 -0.0673 0.1951 0.0607 0.1274 0.6540 -0.2843 0.7903 0.4929 0.7749 ' 0.7761 O.2898 -0.5750 0.2164 0.2397 0.5214 0.1571 1.0000 0.0557 1.0000 -0.1833 . 0.6972 1.0000 -0.7426 • -0.1130 0.3976 1.0000 0.0135 -0.1843 -0.2485 -0.3352 0.7166 -O.2528 -0.1168 -0.6157 -0.2573 O.3282 0.2702 0.5483 0.2193 0.1142 0.4594 -0.0371 -0.3935 -0.4797 0.0104 0.4095 -0.4557 0.2067 0.3789 0.06E9 0.0673 -0.4114 -0.0241 -0.1478 0.1337 0.1617 0.143C -C.0490 -0.3922 -0.2044 0.1563 O.1280 0.3574 -0.2170 0.2398 -0.1343 -0.5897 -O.506O -0.1163 0.3475 0.0848 0.2118 0.6881 . O.C6C2 0.8817 0.1877 0.0256 -0.5552 0.1348 0.6109 0.9057 0.0953 0.2053 0.8142 0.9091 0.0429 -0.6738 0.2194 0.7433 0.8515 -0.1079 0.4792 -0.1763 -0.3352 0.5252 -0.3589 -0.2992 -0.6348 0.0127 0.3535 0.3817 O.4386 0.8601 0.1252 -0.0039 -0.7550 -0.2678 -0.3370 0.1832 0.3558 ON Appendix F. Table l b . P a r t two of the c o r r e l a t i o n m a t r i x of the exemplary o r d i n a t i o n of s i x species of Brachythecium. . 1.0000 0.2143 _ 0 . 3 9 7 5 _ 0 . 0 ? ' 0 . 2 2 0 8 0 .1889 0.0*432 0 .1496 0 .1501 - O.OS25 0 .3922 O.0898 - 0 . 1 2 8 7 - 0 . 1 2 5 0 - 0 . 2 2 3 9 --0.2741 - 0 . 1 2 5 0 0 .6086 - 0 . 3 3 0 0 0 .3032 - 0 . 2 5 0 0 1 .0000 - 0 . 7 3 4 4 ' 0 . 4 7 9 5 0 .0996 0 .0598 0 .6819 0 .0853 0 .2131 0 .7998 - 0 . 0 2 8 0 0 .5131 0 .6036 0 . 0 0 0 0 0 .0853 - 0 . 4 3 6 4 -0.1*286 O.8501 -O.4503 0 .8142 0.2449 1.0000 - 0 . 4 9 2 6 1 .0000 -0 .4360 0 .5400 1 .0000 -0 .5143 0 .3102 0 .3924 1 .0000 - 0 . 8 3 0 5 1 0 .5387 0 .6391 0 .5536 1.0000 0.0278 0 .2052 -O.1388 - 0 . 1 1 0 0 - 0 . 2 6 0 3 1.0000 -0.6529 0 .5769 0.7941 0.7321 ••• 0 .7083- -O.3069 1.O000 - O . 6 O O 3 0 . 7 6 7 3 ' 0 .5297 0 .2011 0 .7961- O.0839 0 .5213 1.0000 - 0 . 2 4 9 5 - 0 . 2 0 2 5 0 .3509 0 .4912 0.4471 - 0 . 4 2 1 6 0 .5307 0 .0436 1 .0000 - 0 . 3 2 3 6 0 .6492 0 .2337 0.4645 0.5721 -O.063I 0.5145 0 . 7 4 3 9 " 0.2466 -0 .0947 0 .3147 -C.3405 -0.4893 - 0 . 0 6 1 9 0 .5053 - 0 . 4 5 7 2 0.4120 - 0 . 7 5 1 7 0 .3180 0 .3873 - 0 . 3 2 5 3 0 .1207 -J .2099 0.1731 - 0 . 0 9 7 4 0 .1759 - 0 . 3 4 3 2 0.2004 0.4S81 -O.2289 0 .2055 -0.0348 0 .1487 -O.0308 0 .2593 -0.4000 0.2980 0 .3337 0 .4659 0.4440 O.0696 - 0 . 0 3 7 4 0 .4O35 0 .0922 0 . 2 8 4 2 0 . 0 7 9 5 - 0 . 3 8 7 3 - 0 . 3 2 5 3 0 .1694 - 0 . 3 8 2 7 0 .0070 - 0 . 2 0 0 5 - 0 . 6 3 1 7 - 0 . 3 4 3 2 -0 .7073 0 .1979 -O.0691 0 .0873 0 .5647 - 0 . 0 9 3 9 " 0 .2324 0 .5281 O . 3 O O 6 0 .8395 - 0 . 1 3 5 6 - 0 . 3 9 1 2 - 0 . 5 6 2 7 -O.7629 '• 0 .4837 -0.7016l' - 0 . 3 1 1 9 - 0 . 5 1 7 7 -0 .3985 0.2505 - 0 . 4 2 8 3 -0.1494 0 .2435 - 0 . 0 0 1 2 -0 .1489 0 .4561 - 0 . 2 3 7 8 -O.4543 0.6455 - . 9 1 2 9 0.4265 O.6S37 • 0.2008 0 . 6 8 0 2 » 0.6686' 0 .1961 1.0000 0 .1143 0 .6286 -0.6434 ' 0 .4291 -0 .5837 0.4210 -0 .1839 0 .4356 0 . 3 5 9 2 1. 0. 0. - 0 , - 0 . 0, 0. 0. -0. 0000 2980 3356 4922 1287 2967 3072 6865 0832 1.0000 0 .9702 O.43C8 -C I250 - 0 . 1 3 5 3 0 .4580 0 .3032 - 0 . 1 4 2 9 1.0000 0 .4326 -0.0746 -0.1391 • 0 .3739 O .3258 - 0 . 0 3 5 5 1 .0000 -0.2741 - 0 . 5 2 9 7 0 . 2 7 3 2 - 0 . 5 6 0 4 0 .4756 1 .0000 - 0 . 4 0 5 8 - 0 . 0 3 7 6 - 0 . 2 4 2 5 - 0 . 3 5 7 1 1.0000 - 0 . 5 9 7 5 0 .7873--0 .0451 1.0000 - 0 . 2 0 9 1 - 0 . 2 3 2 8 1 .0000 - 0 . 3 1 1 8 1 .0000 100. Appendix.F. Table 2. Mean, variance and standard d e v i a t i o n of the v a r i a b l e s measured i n the exemplary o r d i n a t i o n of s i x species of Brachythee ium. STANDARD MEAN VARIANCE DEVIATION LEAF LENGTH 1575-555 0.1725E+06* 0.^15^+03 LEAF WIDTH 702.778 0.2781E+05 0.1668E+03 COSTA RATIO 0.523 0.2Mf8E-02 O.^VTE-Ol STEM MEDIAN CELL LENGTH 89.833 0.6550E+03 0.2559E+02 LEAF MEDIAN CELL WIDTH 7.578 O.8869E+OO 0.9418E+00 TRAITS APEX TAPERED ' 0-333 0.1700E+00 0.irl23E+00 LEAF SHAPE 3.111 0.1111E+00 0-333^E+00 ACUMINATION 2.356 0.1528E+01 0.1236E+01 MARGIN REFLEXED 0.111 0.51HE-01 0.2261E+00 PLICATION O.689 0.1081E+01 O.IO^OE+Ol APEX TWIST. 0.311 0.1211E+00 0.3^802+00 APEX FALCATE 0.022 O.Mt¥fE-02 O.6667E-OI ALAR CELLS 0.800 O.^OOE+OO 0.7000E+00 MARGIN SERRATED 2.200 O.89OOE+OO 0.9^3^+00 BASAL CELLS 0.600 0.1500E+00 0.3873E+00 CONCAVITY 0.622 0 . 5 1^+00 0.7172E+00 LEAF LENGTH I926.III 0.1482E+06 0.38^+03 - LEAF WIDTH IO65.OOO 0.9227E+05 0.3038E+03 COSTA RATIO 0.523 0.ir322E-02 0.657^E-01 BRANCH MEDIAN CELL LENGTH 86.722 0.3306E+03 O.I818E+02 LEAF MEDIAN" CELL WIDTH 7.356 O.l/^SE+Ol 0.1161E+01 TRAITS APEX TAPERED 0.^67 0.2600E+00 0.5099E+00 LEAF SHAPE 3.867 0.3100E+00 0.5568E+00 ACUMINATION 1.822 0.151^+01 0.1231E+01 MARGIN REFLEXED 0.111 O.llllE+OO 0.3333E+00 PLICATION 0.600 0.1010E+01 0.1005E+01 APEX TWIST 0.311 0.1811E+00 0.if256E+00 APEX FALCATE o.o¥f 0.1778E-01 0.1333E+00 ALAR CELLS 1.200 0.1230E+01 0.1109E+01 MARGIN SERRATED 1.^ 89 0.1531E+01 0.1237E+01 BASAL CELLS O.667 0.1700E+00 0.ifl23E+00 CONCAVITY O.667 0.4900E+00 0.7000E+00 *Note: E denotes the exponential power to,which each value i s r a i s e d , e.g.."E+06M i s equivalent to "X 10 6". 

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