UBC Theses and Dissertations

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UBC Theses and Dissertations

Hyphomycetes decaying the litter of Thuja plicata Donn Dunn, Michael Thomas 1980

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HYPHOMYCETES D E C A Y I N G THE L I T T E R OF T H U J A P L I C A T A DON N H I C H A E L THOMAS DUNN B . S c . ( H o n s . ) , T u l a n e U n i v e r s i t y , 1973 H . S c , T u l a n e U n i v e r s i t y , 1 9 7 5 T H E S I S S U B M I T T E D IN P A R T I A L F U L F I L M E N T THE REQUIREMENTS FOE THE DEGREE OF THE F A C U L T Y OF GRADUATE S T U D I E S ( D e p a r t m e n t o f B o t a n y ) We a c c e p t t h i s t h e s i s a s c o n f o r m i n g t o t h e r e q u i r e d s t a n d a r d THE U N I V E R S I T Y OF B R I T I S H COLOMBIA A u g u s t 1 9 8 0 DOCTOR OF P H I L O S O P H Y x n M i c h a e l T h o m a s Dunn, 1980 In 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 the r e q u i r e m e n t s f o r an advanced degree a t the U n i v e r s i t y o f B r i t i s h Columbia, I agree t h a t the 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 and s t u d y . I f u r t h e r agree t h a t p e r m i s s i o n f o r e x t e n s i v e 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 purposes may be g r a n t e d by the Head o f my Department 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 or 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 not 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 . Department o f , Botany The U n i v e r s i t y o f B r i t i s h Columbia 2075 Wesbrook P l a c e Vancouver, Canada V6T 1W5 J a n 24, 1981 i i A b s t r a c t 0 The p r e s e n t s t u d y was u n d e r t a k e n t o examine t h e l i t t e r d e c a y f u n g i of r e d c e d a r ( T h u j a p i i c a t a D onn). The f u n g i i n v o l v e d i n c o n i f e r l i t t e r d e c a y have not been examined i n t e n s i v e l y and t h e r e has been no s t u d y of r e d c e d a r l i t t e r d e c a y . S i n c e T h u j a i s o f t e n f o u n d i n wet, p o o r l y d r a i n e d a r e a s , i t p r o v i d e d an o p p o r t u n i t y t o o b s e r v e o c c u r r e n c e o f l i t t e r d e c a y f u n g i r e l a t i v e t o s l i g h t d i f f e r e n c e s i n d i s t a n c e from a s t r e a m m a r g i n . T h i s was d e t e r m i n e d by o b s e r v i n g c h a n g e s i n t h e f r e q u e n c y of t h e -fungi a t v a r i o u s d i s t a n c e s f r o m t h e m a r g i n of a s t r e a m and a l s o by o b s e r v i n g s e a s o n a l c h a n g e s i n f u n g a l p o p u l a t i o n s . An o r d i n a t i o n of t h e d a t a was p e r f o r m e d t o d e t e r m i n e i f t h e r e were s p e c i e s a s s o c i a t i o n s w h i c h would c h a r a c t e r i z e t h e r e l a t i v e l y m i n o r h o r i z o n t a l and v e r t i c a l s p a t i a l c h a n g e s i n t h e s i t e s . The s i t e s were d i v i d e d i n t o s u b s i t e s ( h i g h , m i d d l e and low w i t h r e s p e c t t o t h e s t r e a m ) and samples were t a k e n a t e a c h s u b s i t e . Red c e d a r b r a n c h l e t s f r o m t h e L and F l i t t e r l a y e r s were washed and p l a t e d on a s e l e c t i v e medium or p l a c e d i n m o i s t chambers; a l l o b s e r v e d s p e c i e s were i s o l a t e d and i d e n t i f i e d . Most of t h e s e were members of t h e F u n g i I m p e r f e c t i . The f r e q u e n c y of o c c u r r e n c e of t h e more commonly i s o l a t e d s p e c i e s was u t i l i z e d i n a P r i n c i p a l Components A n a l y s i s (PCA) t o d e t e r m i n e a s s o c i a t i o n s of s u b s i t e s , l a y e r s o r s p e c i e s . T h e r e was l i t t l e d i s t i n c t i o n between t h e t h r e e s u b s i t e s , but t h e r e was a g e n e r a l s e p a r a t i o n of t h e h i g h s u b s i t e from t h e low s u b s i t e , t h e m i d d l e s u b s i t e showing a f f i n i t i e s t o b o t h . The L and F l a y e r s r e p r e s e n t e d a d i v i s i o n of t h e l i t t e r l a y e r w h i c h formed more d i s t i n c t c l u s t e r s i n t h e o r d i n a t i o n , e s p e c i a l l y i n t h e S i t e B d a t a . S e a s o n a l d i s t i n c t i o n among t h e samples seemed t o r e f l e c t t h e e x t r e m e s of t h e s e a s o n s . The s p r i n g and summer samples g e n e r a l l y g r o u p e d t o g e t h e r , as d i d t h e f a l l and w i n t e r s a m p l e s . S p e c i e s a s s o c i a t i o n s r e f l e c t e d v a r i o u s c o m b i n a t i o n s of t h e above g r o u p i n g s , d e p e n d i n g on t h e s p e c i e s g r o u p s i n v o l v e d . A g a i n t h e most d i s t i n c t g r o u p s r e p r e s e n t e d e x t r e m e s , e£. , s p e c i e s p r o m i n e n t i n d r y s a m p l e s , e s p e c i a l l y i n t h e h i g h s u b s i t e , or s p e c i e s p r o m i n e n t i n m o i s t c o n d i t i o n s , e s p e c i a l l y i n the low s u b s i t e . No s i n g l e v a r i a b l e p r o v i d e d c l e a r d i s t i n c t i o n among t h e v a r i o u s s u b s i t e s , l a y e r s , s e a s o n s o r s p e c i e s . However, t h e c o m b i n a t i o n o f a l l of t h e s e gave g e n e r a l i n d i c a t i o n s as t o p r o b a b l e f u n g a l a s s o c i a t i o n s . i v T a b l e Of C o n t e n t s A b s t r a c t i i L i s t Of T a b l e s v L i s t Of F i g u r e s v i i Acknowledgements x i I . I n t r o d u c t i o n 1 I I . M a t e r i a l s and Methods 8 1. D e s c r i p t i o n of the Study S i t e 8 2. C l a s s i f i c a t i o n of the L i t t e r 12 3. Treatment of the Samples 14 4. M u l t i v a r i a t e A n a l y s i s of the Data 19 I I I . R e s u l t s And O b s e r v a t i o n s 21 1. Predominant S p e c i e s i n the L L a y e r 37 2. Predominant S p e c i e s i n the F Layer 42 3. Comparison of S p e c i e s Found i n the L and F L a y e r s ... 48 4. S e a s o n a l i t y of Fungi i n the L and F L a y e r s 51 5. Fungal S p e c i e s I s o l a t e d 60 6. M u l t i v a r i a t e A n a l y s i s of the Data 135 IV. D i s c u s s i o n and C o n c l u s i o n s 150 1. D i s c u s s i o n of the S u b s i t e s 150 2. D i s c u s s i o n of the L i t t e r L a y e r s 151 3. D i s c u s s i o n of S e a s o n a l i t y 153 4. S p e c i e s A s s o c i a t i o n s 154 5. Summary 159 V. B i b l i o g r a p h y 162 V L i s t Of T a b l e s T a b l e I . C l i m a t i c Data For Sample Dates At S i t e A 11 T a b l e I I . C l i m a t i c Data For Sample Dates At S i t e B 11 T a b l e I I I . S p e c i e s I s o l a t e d In The High S u b s i t e At S i t e A . 24 T a b l e IV. S p e c i e s I s o l a t e d In The M i d d l e S u b s i t e At S i t e A 26 T a b l e V. S p e c i e s I s o l a t e d In The Low S u b s i t e At S i t e A .... 28 T a b l e V I . S p e c i e s I s o l a t e d From L i v i n g Leaves At Both S i t e s 30 Table V I I . S p e c i e s I s o l a t e d In The High S u b s i t e At S i t e B . 31 T a b l e V I I I . S p e c i e s I s o l a t e d In The M i d d l e S u b s i t e At S i t e B 33 Table IX. S p e c i e s I s o l a t e d In The Low S u b s i t e At S i t e B ... 35 T a b l e X. D i s t r i b u t i o n Of S p e c i e s In The L Layer At S i t e A . 39 T a b l e X I . D i s t r i b u t i o n Of S p e c i e s In The L Layer At S i t e B 39 T a b l e X I I . D i s t r i b u t i o n Of S p e c i e s In The F Layer At S i t e A 43 T a b l e X I I I . D i s t r i b u t i o n Of S p e c i e s In The F L a y e r At S i t e B 44 T a b l e XIV. S p e c i e s R e s t r i c t e d To One Or The Other Main S i t e 47 Table XV. Comparison Of S p e c i e s In.The L And F L a y e r s At S i t e A 49 T a b l e XVI. Comparison Of S p e c i e s In The L And F L a y e r s At S i t e B , 50 T a b l e X V I I . S e a s o n a l D i s t r i b u t i o n Of S p e c i e s In The L Layer At S i t e A 53 v i T a b le X V I I I . S e a s o n a l D i s t r i b u t i o n Of S p e c i e s In The L Layer At S i t e B 54 Tab l e XIX. Season a l D i s t r i b u t i o n Of S p e c i e s In The F Layer At S i t e A 55 Tab l e XX. Se a s o n a l D i s t r i b u t i o n Of S p e c i e s In The F Layer At S i t e B 56 Tab l e XXI. Frequency And V a r i a n c e Of The Most Common S p e c i e s At S i t e A 136 T a b l e X X I I . Frequency And V a r i a n c e Of The Most Common S p e c i e s At S i t e B 137 4 L i s t Of F i g u r e s F i g u r e 1. S p a t i a l Arrangement Of The Samples At S i t e A .... 17 F i g u r e 2. S p a t i a l Arrangement Of The Samples At S i t e B .... 18 F i g u r e 3. Se a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of Acremonium s t r i c t u m 61 F i g u r e 4. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of Acrodontium c r a t e r i f o r m e 63 F i g u r e 5. Se a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of A r t i c u l o s p o r a t e t r a c l a d i a 65 F i g u r e 6. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of C h a l a r a c o n s t r i c t a 67 F i g u r e 7. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of C h a l a r a c y l i n d r o s p e r m a 69 F i g u r e 8. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of C h a l a r a l o n g i p e s • 7 0 F i g u r e 9. Se a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of C h l o r i d i u m v i r e s c e n s v a r . chlamydosporum 72 F i g u r e 10. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of C l a d o s p o r i u m c l a d o s p o r i o i d e s 73 F i g u r e 11. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of C y l i n d r o c a r p o n didymum 7 5 F i g u r e 12. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of C y l i n d r o c a r p o n tenue - - S i t e A 77 F i g u r e 13. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of C y l i n d r o c a r p o n tenue - - S i t e B 78 F i g u r e 14. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of Endophragmia a l t e r n a t a 80 F i g u r e 15. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of F l a g e l l o s p o r a s t r i c t a 82 F i g u r e 16. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of F u s a r i u m l a t e r i t i u m 84 F i g u r e 17. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of G l i o c l a d i u m roseum 85 F i g u r e 18. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of H e l i c o d e n d r o n t r i g l i t z i e n s i s 87 F i g u r e 19. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of H e l i c o o n f u s c o s p o r u m 89 F i g u r e 20. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of H y a l o d e n d r o n l i g n i c o l a 91 F i g u r e 21. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of K r i e g e r i a s e a v e r i 93 F i g u r e 22. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of L i b e r t e l l a sp 94 F i g u r e 23. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of M o r t i e r e l l a r a m a n niana v a r . ramanniana 96 F i g u r e 24. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of Mucor h i e m a l i s f . h i e m a l i s 98 F i g u r e 25. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of Pen i c i 11 ium b r e v i - c o m p a c t u m 100 F i g u r e 26. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of P e n i c i l l i u m c i t r i n u m 101 F i g u r e 27. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of P e n i c i l l i u m f r e q u e n t a n s 103 F i g u r e 28. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of i x Pen i c i 11 i urn n i g r i c a n s 104 F i g u r e 29. Se a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of P e n i c i l l i u m notatum 106 F i g u r e 30. Se a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of P e n i c i11ium r a i s t r i c k i i 107 F i g u r e 31. Se a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of P e n i c i l l i u m verrucosum v a r . c y c l o p i u m 109 F i g u r e 32. Se a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of P e s t a l o t i a m o n o c h a e t i o i d e s I l l F i g u r e 33. Se a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of P o l y s c y t a l u m fecundissimum 113 F i g u r e 34. Seaso n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of R a m i c h l o r i d i u m subulatum 115 F i g u r e 35. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of Selenophoma sp 116 F i g u r e 36. Se a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of Septogloeum sp 118 F i g u r e 37. Se a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of Septonema c h a e t o s p i r a '.120 F i g u r e 38. Se a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of Septonema c h a e t o s p i r a v a r . p i n i 122 F i g u r e 39. Se a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of Spor idesmium flexum 123 F i g u r e 40. Se a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of Tr ichoderma k o n i n q i i 125 F i g u r e 41. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of Tr ichoderma polysporum 126 F i g u r e 42. Seaso n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of X Trichoderma v i r i d e 128 F i g u r e 43. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of Tripospermum m y r t i 130 F i g u r e 44. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of T r i p o s p o r i u m e l e g a n s 131 F i g u r e 45. Se a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of V a r i c o s p o r i u m elodeae 133 F i g u r e 46. Se a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of V e r t i c i l l i u m b u l b i l l o s u m 134 F i g u r e 47. O r d i n a t i o n Of Samples At S i t e A. A x i s I v s . I I ( S c a l e d ) 142 F i g u r e 48. O r d i n a t i o n Of S p e c i e s At S i t e A. A x i s I v s . I I ( S c a l e d ) 143 F i g u r e 49. O r d i n a t i o n Of Samples At S i t e B. A x i s - I v s . I I ( S c a l e d ) 147 F i g u r e 50. O r d i n a t i o n Of S p e c i e s At S i t e B. A x i s I v s . I I ( S c a l e d ) 148 x i A cknowledgements I would l i k e t o thank P r o f e s s o r s G.C. Hughes, J.R. Maze and W.B. S c h o f i e l d f o r s e r v i n g on t h e r e s e a r c h c o m m i t t e e . T h e i r h e l p and s u g g e s t i o n s were g r e a t l y a p p r e c i a t e d . I would a l s o l i k e t o t hank G.E. B r a d f i e l d f o r h i s i n t r o d u c t i o n t o t h e computer and t h e MIDAS program and a l s o f o r s e r v i n g on t h e r e s e a r c h c o m m i t t e e . J . Paden, U n i v e r s i t y of V i c t o r i a , was h e l p f u l i n b e g i n n i n g my i d e n t i f i c a t i o n s of P e n i c i l l i u m i s o l a t e s . My f o n d e s t t h a n k s a r e e x t e n d e d t o my s u p e r v i s o r , R . J . B a n d o n i . H i s s u p p o r t and g u i d a n c e t h r o u g h o u t t h i s s t u d y have been most i m p o r t a n t and a r e g r e a t f u l l y a p p r e c i a t e d . F i n a l l y , I would l i k e t o thank my f a m i l y and f r i e n d s who have h e l p e d , s u p p o r t e d and put up w i t h me t h r o u g h o u t t h e s e y e a r s . 1 I . I n t r o d u c t i o n The p r o d u c t i o n and a c c u m u l a t i o n of l i t t e r i n a f o r e s t i s a p r o c e s s w h i c h i s i m p o r t a n t i n t h e b i o l o g y of t h e e c o s y s t e m . The breakdown of t h i s l i t t e r , and t h e s u b s e q u e n t c y c l i n g of n u t r i e n t s , i s r e s p o n s i b l e f o r much of t h e e n e r g y f l o w i n t o and s u r v i v a l of t h e m a c r o f l o r a p r e s e n t i n t h e f o r e s t . E c o l o g i s t s a r e w e l l aware of t h e s i g n i f i c a n c e of t h e d e c o m p o s i t i o n of p l a n t m a t e r i a l i n i n f l u e n c i n g t h e c h a r a c t e r i s t i c s of f o r e s t e c o s y s t e m s ; s e v e r a l w o r k e r s have r e v i e w e d t h i s s u b j e c t ( B r a y and Gorham, 1964; S i n g h and G u p t a , 1977; W i l l i a m s and G r a y , 1 9 7 4 ) . The o r g a n i s m s i n v o l v e d i n t h e d e c o m p o s i t i o n o f l i t t e r a r e m a i n l y b a c t e r i a , f u n g i and i n v e r t e b r a t e s . The f u n g i , have been r e c o g n i z e d as a major component of t h i s h a b i t a t and t h e i r i m p o r t a n c e i n t h e d e c a y p r o c e s s i s w e l l e s t a b l i s h e d . S o i l f u n g i have been examined w i t h r e s p e c t t o t h e i r e c o l o g y and a number of r e v i e w s have been p u b l i s h e d on s u b s t r a t e r e l a t i o n s h i p s ( G a r r e t t , 1 9 57), s o i l as a s o u r c e of p l a n t p a t h o g e n s ( B u r g e s , 1939; G a r r e t t , 1956, 1 970), and g e n e r a l i n v o l v e m e n t of f u n g i i n t h e d e c a y p r o c e s s ( P a r k i n s o n and Waid, 1960; S z e g i , 1 9 7 2 ) . In s p i t e of t h e i m p o r t a n c e of t h e l i t t e r l a y e r i n the s u p p o r t of t h e a b o v e - g r o u n d v e g e t a t i o n , t h e f u n g i i n v o l v e d i n t h e d e c a y of l i t t e r have o n l y r e c e n t l y come t o be s t u d i e d . The p r o b l e m s i n v o l v e d i n s t u d y i n g t h e d e c o m p o s i t i o n p r o c e s s i n l i t t e r , t h e o r g a n i s m s i n v o l v e d and t h e t y p e s of l i t t e r t h a t have been examined a r e r e v i e w e d e l s e w h e r e ( A n d e r s o n and M a c f a d y e n , 1976; D i c k i n s o n and Pugh, 1 9 7 4 ) . One of t h e b a s i c q u e s t i o n s i n s t u d y i n g t h e l i t t e r l a y e r i s : 2 what f u n g i a r e p r e s e n t and i n v o l v e d i n t h e d e c a y ? A t t e m p t s t o d e t e r m i n e t h i s u s u a l l y i n v o l v e i s o l a t i o n s of t h e f u n g i u s i n g v a r i o u s t e c h n i q u e s . S i n c e s o i l f u n g a l s t u d i e s have been c a r r i e d on l o n g e r t h a n l i t t e r d e c a y s t u d i e s , t h e v a r i o u s s o i l methods a r e numerous and have been th e s u b j e c t of much d i s c u s s i o n (Chou and S t e p h e n , 1968; N i c h o l a s and P a r k i n s o n , 1967; P a r k i n s o n , 1970, 1973; P a r k i n s o n , G r a y and W i l l i a m s , 1971; P h i l l i p s o n , 1971 Warcup, 1 9 6 0 ) . Many of t h e s e t e c h n i q u e s have a l s o been a d o p t e d f o r use i n l i t t e r d e c a y s t u d i e s . The most i m p o r t a n t c o n s i d e r a t i o n i n e i t h e r t y p e of s t u d y i s t h e e x t e n t t o w h i c h t h e t e c h n i q u e s g i v e an a c c u r a t e p i c t u r e of t h e o r g a n i s m s i n v o l v e d i n d e c o m p o s i t i o n . Some of t h e more common t e c h n i q u e s a r e d i s c u s s e d h e r e and t h e i r a p p l i c a b i l i t y e v a l u a t e d . The s o i l d i l u t i o n p l a t e t e c h n i q u e was d e v e l o p e d i n s t u d i e s of m i c r o o r g a n i s m s i n s o i l (Waksman, 1 9 2 7 ) . B e cause of t h e d i l u t i o n p r o c e d u r e , t h i s t e c h n i q u e i n i t i a l l y a p p e a r s t o p r o v i d e a means of q u a n t i f y i n g t h e f u n g i p r e s e n t . U n f o r t u n a t e l y t h i s t e c h n i q u e d o e s not d i s t i n g u i s h between f u n g i t h a t a r e a c t i v e l y g r o w i n g i n t h e s o i l , and s p o r e s or r e s t i n g s t a g e s . Whether g r o w t h has been from a s i n g l e s p o r e , a mass of s p o r e s , o r h y p h a l f r a g m e n t s c a n n o t be d e t e r m i n e d . C o n s e q u e n t l y , c o u n t s of s p e c i e s t h a t p r o d u c e abundant s p o r e s a r e o f t e n e x a g g e r a t e d w i t h r e g a r d t o t h e number of i n d i v i d u a l s i n v o l v e d . F o r a p p l i c a t i o n t o l i t t e r s t u d i e s , t h e d i l u t i o n t e c h n i q u e i s m o d i f i e d s l i g h t l y , w i t h t h e l i t t e r f i r s t b e i n g h o mogenized i n a w a r i n g b l e n d e r , o r o t h e r w i s e c r u s h e d , and t h e n d i l u t e d and p l a t e d o u t . T h i s t e c h n i q u e has been u t i l i z e d i n a number of l i t t e r s t u d i e s , e.g., by B r a n d s b e r g , 1969 (on c o n i f e r s ) , V i s s e r and P a r k i n s o n , 1975 (on 3 aspen) and Verona and R a m b e l l i , 1972 (on Monterey p i n e ) . The t e c h n i q u e has the same drawbacks mentioned e a r l i e r i n t h a t the number of times a s p e c i e s i s i s o l a t e d o n l y r e f l e c t s the number of spores or p i e c e s of hyphae p r e s e n t . T r i b e (1957, 1960, 1966) examined l i t t e r f u n g i t h a t became e s t a b l i s h e d on b u r i e d c e l l u l o s e f i l m u s i n g a t e c h n i q u e t h a t i n v o l v e d p u t t i n g c e l l u l o s e f i l m , u s u a l l y c e l l o p h a n e , on the s o i l or l i t t e r , and l e a v i n g i t f o r a s p e c i f i e d amount of t i m e . O b s e r v a t i o n s were made on the hyphae found growing on the f i l m t o a s s e s s the f u n g a l a c t i v i t y i n the s o i l or l i t t e r . U n f o r t u n a t e l y , i d e n t i f i c a t i o n i s p o s s i b l e o n l y i f the f u n g i s p o r u l a t e on the f i l m . A d d i t i o n a l l y the t e c h n i q u e s e l e c t s f o r c e l l u l o s e decomposers and the s u b s t r a t u m i s a c t u a l l y composed of m o d i f i e d c e l l u l o s e . A more p r o m i s i n g t e c h n i q u e i n v o l v i n g f l u o r e s c e n t a n t i b o d y s t a i n i n g ( P r e e c e , 1971) has been used i n s t u d i e s of the mycobiota of Douglas f i r ( B e r n s t e i n e_t a_l. , 1973). T h i s t e c h n i q u e i n v o l v e s p r e p a r a t i o n of a n t i b o d i e s f o r a s p e c i f i c fungus and attachment of a f l u o r e s c e n t dye t o the a n t i b o d i e s . The m a t e r i a l examined, such as a l e a f , i s " s t a i n e d " w i t h t h i s m i x t u r e and observed u s i n g f l u o r e s c e n c e m i c r o s c o p y . The presence of the s p e c i e s can be d e t e c t e d by the f l u o r e s c e n c e of the a n t i b o d y . Hyphae on the s u r f a c e of the s u b s t r a t u m t h e r e f o r e can be i d e n t i f i e d by t h i s method. S i n c e the p r o d u c t i o n of a n t i b o d i e s i s a complex p r o c e s s and many f u n g i are i n v o l v e d i n decay, r o u t i n e use of t h i s method would not be f e a s i b l e . Furthermore d i f f e r e n t s t r a i n s of the same fungus sometimes do not s t a i n w i t h the a n t i b o d i e s , thus c o n t r i b u t i n g t o f u r t h e r d i f f i c u l t i e s . 4 S e v e r a l i n v e s t i g a t o r s (Edwards and H e a t h , 1963; H e a t h e t a l . , 1964; H e a t h e t a l . , 1966; T u b a k i and Yokoyama, 1971, 1972a, 1972b; Yokoyama and T u b a k i , 1973; Yokoyama e t a l . , 1977) have u s e d the i d e a of b a i t i n g t h e l i t t e r f o r t h e f u n g i p r e s e n t . They p l a c e gas s t e r i l i z e d l e a v e s i n mesh bags and put t h e s e i n t h e l i t t e r l a y e r t o be r e t r i e v e d a t some l a t e r d a t e . The p r o b l e m s h e r e i n c l u d e i n t e r f e r e n c e w i t h i n v e r t e b r a t e movements and s p o r e t r a n s p o r t i f f i n e mesh bags a r e u s e d . I n v e r t e b r a t e s may be i n v o l v e d i n t h e s p r e a d of c e r t a i n f u n g i as w e l l as b e i n g i m p o r t a n t i n t h e p h y s i c a l breakdown and p r o c e s s i n g o f t h e l i t t e r . S t e r i l i z e d l e a v e s , a l t h o u g h e x p e r i m e n t a l l y i d e a l f o r d e t e r m i n i n g th e f u n g i p r e s e n t , a r e n o t n a t u r a l . N o r m a l l y t h e r e a r e a number of f u n g i i n the l e a v e s when t h e y f a l l and t h e s e f u n g i may be v e r y i m p o r t a n t i n p r e p a r i n g t h e l e a v e s f o r s u b s e q u e n t c o l o n i z e r s . T h i s t e c h n i q u e a l s o t e n d s t o f a v o r t h o s e s p e c i e s w h i c h a r e a c t i v e l y s p o r u l a t i n g i n t h e l i t t e r l a y e r when th e l e a v e s a r e p o s i t i o n e d t h e r e . P o s i t i o n i n g d i s t u r b s t h e l i t t e r l a y e r and t h e s t e r i l e l e a v e s p r o b a b l y a r e r a p i d l y c o l o n i z e d by f a s t g r o w i n g f u n g i . A method d e v e l o p e d by H a r l e y and Waid (1955) f o r t h e s t u d y of r h i z o p l a n e f u n g i seems t o o f f e r a p r o m i s i n g a l t e r n a t i v e . They washed p i e c e s of r o o t s i n numerous c h a n g e s of s t e r i l e w a ter t o remove most of t h e e x t e r n a l s p o r e s and hyphae. T h e r e f o r e t h e f u n g i w h i c h were on t h e s u r f a c e o r i n s i d e t h e r o o t , p r e s u m a b l y a c t i v e l y g r o w i n g , grew out and c o u l d be i s o l a t e d . The washed r o o t s , or p i e c e s t h e r e o f , were p l a c e d i n s t e r i l e m o i s t chambers and o b s e r v e d a t l a t e r d a t e s . V a r i a t i o n s of t h i s t e c h n i q u e have been a d o p t e d i n t h e s t u d y of l i t t e r d e c a y o f s c o t s p i n e 5 ( K e n d r i c k and B u r g e s , 1962), b r a c k e n f e r n ( F r a n k l a n d , 1976), s i l v e r f i r ( G o u r b i e r e , 1974a, 1974b, 1975, 1979) and p i n e -hardwood s t a n d s (Watson e t a l . , 1 9 7 4 ) . A m o d i f i c a t i o n of t h i s method i s u s e d i n the p r e s e n t s t u d y . Most l i t t e r d e c a y s t u d i e s have d e a l t w i t h a n g i o s p e r m l i t t e r ( J e n s e n , 1974) w i t h l i t t l e work done i n c o n i f e r f o r e s t s ( M i l l a r , 1 9 7 4 ) . I n c r e a s e d i n t e r e s t has been g e n e r a t e d i n d e c a y of c o n i f e r l i t t e r , p o s s i b l y b e c a u s e of t h e economic i m p o r t a n c e of c o n i f e r f o r e s t s , but a l s o due t o t h e u n i q u e e c o s y s t e m of t h i s l i t t e r . In t h e o r g a n i c h o r i z o n of a c o n i f e r f o r e s t t h e d e c o m p o s i t i o n r a t e i s low, b e c a u s e of t h e p h y s i c a l n a t u r e of t h e l i t t e r and t h e t a n n i n s and o t h e r s u b s t a n c e s p r e s e n t . As a r e s u l t o f t h i s r e l a t i v e l y low r a t e of d e c o m p o s i t i o n , t h e a c c u m u l a t i o n o f l i t t e r i s u s u a l l y great.. Much o f t h e work on t h e m y c o b i o t a of c o n i f e r l i t t e r has i n v o l v e d d e s c r i b i n g t h e f u n g i p r e s e n t and c o m p a r i s o n s of d i f f e r e n t s i t e s or of v a r i o u s s p e c i e s of t r e e s . The i n f o r m a t i o n p r o v i d e d by t h e s e s t u d i e s i n c l u d e s t h e o c c u r r e n c e and f r e q u e n c y of s p e c i e s i s o l a t e d and, o f t e n , s e a s o n a l v a r i a t i o n s . The d i v i s i o n of l i t t e r i n t o v a r i o u s l a y e r s , i n d i c a t i n g s t a g e s i n t h e t i m e s e q u e n c e of d e c a y , i s e s p e c i a l l y s i g n i f i c a n t w i t h r e s p e c t t o c o n i f e r l i t t e r , s i n c e d e c o m p o s i t i o n i s slow and i n c o r p o r a t i o n of l i t t e r i n t o t h e humus may t a k e s e v e r a l y e a r s (Dimock, 1958; K e n d r i c k , 1 9 5 9 ) . S e v e r a l i n v e s t i g a t o r s have f o u n d t h e f u n g i p r e s e n t i n t h e d i f f e r e n t l a y e r s of t h e l i t t e r t o r e p r e s e n t d i s t i n c t e c o l o g i c a l g r o u p s , and t h u s have been a b l e t o i n d i c a t e a s u c c e s s i o n a l p a t t e r n w i t h r e s p e c t t o t i m e (Borowska, 1966; B r a n d s b e r g , 1969; K e n d r i c k , 1963; K e n d r i c k and B u r g e s , 6 1962; Verona and R a m b e l l i , 1972; V i s s e r and P a r k i n s o n , 1975; Watson et a l . , 1974; Wicklow and W i t t i n g h a m , 1974, 1978; Widden and P a r k i n s o n , 1973). In the p r e s e n t s t u d y , the i n t e n t i o n has been t o examine the f u n g i i n v o l v e d i n the decay of red cedar l i t t e r (Thuja p i i c a t a Donn) w i t h emphasis on the major component of the m y c o b i o t a , the hyphomycetes (Fungi I m p e r f e c t i ) . The d i s t r i b u t i o n of s p e c i e s between the l a y e r s of l i t t e r , and s e a s o n a l changes throughout the year a l s o were i n v e s t i g a t e d . A number of r e c e n t workers have examined the breakdown of l i t t e r i n a q u a t i c s i t u a t i o n s (Kaushik and Hynes, 1968, 1971; S e d e l l e t a l . , 1974; Subberkropp et a l . , 1975; Wenner et a l . , 1977). Some i n v e s t i g a t o r s have a l s o s t u d i e d c o n i f e r l i t t e r i n streams and the a c t i v i t i e s - of " a q u a t i c " hyphomycetes i n l i t t e r breakdown and decay ( A r n o l d , 1970; B a r l o c h e r and K e n d r i c k , 1974; B a r l o c h e r and O e r t l i , 1978; B a r l o c h e r e_t a l . , 1978). S i n c e Thuja o f t e n grows a l o n g streams, • i t was i d e a l l y s u i t e d f o r an e x a m i n a t i o n of the p o s s i b l e i n f l u e n c e of d r a i n a g e upon t e r r e s t r i a l l i t t e r decay. The changes i n the mycobiota i n the a r e a a l o n g the stream m a r g i n , and a t s l i g h t l y e l e v a t e d s u b s i t e s were i n v e s t i g a t e d t o observe what e f f e c t t h i s s l i g h t t o p o g r a p h i c a l v a r i a t i o n might have upon the o c c u r r e n c e of decay fung i . P l a n t e c o l o g i s t s have o f t e n used o r d i n a t i o n p r o c e d u r e s t o h e l p them c h a r a c t e r i z e v e g e t a t i o n a l groups and e c o l o g i c a l l y s i g n i f i c a n t a s s o c i a t i o n s (Bray and C u r t i s , 1957; Mueller-Dombois and E l l e n b e r g , 1974; Sneath • and S o k a l , 1973). Such a n a l y s e s n o r m a l l y u t i l i z e the o c c u r r e n c e of s p e c i e s as the b a s i s f o r 7 o r d i n a t i o n . In r e c e n t y e a r s o r d i n a t i o n t e c h n i q u e s a l s o have been u s e d i n s t u d i e s of s o i l f u n g i ( B i s s e t t and P a r k i n s o n , 1979a, 1979b, 1979c; D i c k i n s o n and K e n t , 1972;'Kent,. 1972; M o r r a l l , 1974; Pemadasa and M u e l l e r - D o m b o i s , 1 9 7 9 ) . As w i t h h i g h e r p l a n t o r d i n a t i o n s , t h e d i s t r i b u t i o n of s p e c i e s i s u t i l i z e d t o a s s o c i a t e s i t e s and s p e c i e s a s s o c i a t i o n s can a l s o be d e t e r m i n e d . In t h e p r e s e n t s t u d y , an o r d i n a t i o n was p e r f o r m e d t o summarize the samples w i t h r e s p e c t t o t h e m y c o b i o t a p r e s e n t and t h e a s s o c i a t i o n s of t h e v a r i o u s s i t e s and s u b s i t e s . In u n d e r t a k i n g t h i s s t u d y , some s p e c i f i c q u e s t i o n s c o n c e r n i n g t h e f u n g i d e c a y i n g r e d c e d a r l i t t e r , were i n v e s t i g a t e d . F i r s t , what s p e c i e s were p r e s e n t i n r e d c e d a r l i t t e r ? T h e s e were i s o l a t e d and i d e n t i f i e d . S e c ond, were t h e r e d i f f e r e n c e s i n t h e m y c o b i o t a as a f u n c t i o n of r e l a t i v e l y m i n o r v e r t i c a l or h o r i z o n t a l s p a t i a l c h a n g e s i n t h e s i t e ? I f so, s p e c i e s a s s o c i a t i o n s c o u l d ' b e d e t e c t e d w h i c h would c h a r a c t e r i z e t h e s i t e s , s u b s i t e s or l a y e r s . An o r d i n a t i o n of t h e d a t a was u t i l i z e d t o d e t e r m i n e t h i s . L a s t , do s l i g h t d i f f e r e n c e s i n e l e v a t i o n , hence d r a i n a g e , i n f l u e n c e c o m p o s i t i o n o f t h e m y c o b i o t a ? T h i s would i n v o l v e e x a m i n i n g s e a s o n a l d i f f e r e n c e s i n t h e m y c o b i o t a , as w e l l as s p a t i a l d i f f e r e n c e s i n r e l a t i o n t o t h e s t r e a m . 8 I I . M a t e r i a l s and Methods 1. D e s c r i p t i o n of the Study S i t e Two study s i t e s were s e l e c t e d a l o n g streams i n the U n i v e r s i t y Endowment Lands (UEL), U n i v e r s i t y of B r i t i s h Columbia campus, Vancouver, B.C. (49°15' N L a t . , 123°12' W Long.). The a r e a s d i f f e r e d somewhat i n topography but were both dominated by T h u j a . S i t e A was l o c a t e d i n the E c o l o g i c a l Reserve s e c t i o n of the UEL a l o n g an unnamed stream. Over a p p r o x i m a t e l y f i v e meters, the t e r r a i n went from a r e l a t i v e l y h i g h , dry a r e a near the base of a tree., down t o the water's edge, a drop of s l i g h t l y over one meter. S i t e B was l o c a t e d j u s t o f f C l i n t o n T r a i l a l o n g T i n Can Creek. I t was a more open a r e a than S i t e A, w i t h a f a l l e n t r e e a c r o s s the stream but c o v e r e d a p p r o x i m a t e l y the same a r e a . The stream c u r v e d around a mound, w i t h the l o w e s t a r e a e x t e n d i n g from the stream t h r o u g h a d e p r e s s i o n on the o t h e r s i d e of the mound. The mound i t s e l f was 50 - 75 cm above the water l e v e l . A l t h o u g h the two s i t e s were s l i g h t l y d i f f e r e n t , they both were p a r t of the Wet Subzone of the C o a s t a l Douglas F i r B i o g e o c l i m a t i c Zone ( K r a j i n a , 1965, 1969). The dominant t r e e i n the immediate area was Thuja p i i c a t a Donn, w i t h some Tsuga  h e t e r o p h y l l a ( R a f . ) S a r g . The ground cover was s p a r s e and i n c l u d e d G a u l t h e r i a s h a l l o n P u r s h . ( s a l a l ) , w i t h L y s i c h i t u m  americanum H u l t e n & S t . John (skunk cabbage) i n some a r e a s . The s i t e s were p o o r l y d r a i n e d , as i n d i c a t e d by the presence of T h u j a . 9 At each study s i t e , A and B, the area was d i v i d e d i n t o t h r e e s u b s i t e s , d e s i g n a t e d low, m i d d l e and h i g h . The names r e f l e c t a g e n e r a l i n d i c a t i o n of t h e i r p r o x i m i t y t o , and e l e v a t i o n above, the stream. At S i t e A, the low s u b s i t e was a r e c t a n g u l a r a r e a , 150 x 60 cm, l o c a t e d a l o n g the stream margin. I t was submerged much of the time and c o n s e q u e n t l y was v e r y s i l t y . W i th r e s p e c t t o the water l e v e l i n the stream, i t ranged from 0 - 19 cm above. The middl e s u b s i t e was s l i g h t l y h i g h e r and f u r t h e r from the stream, but the l i t t e r of t h i s s u b s i t e was s a t u r a t e d much of the t i m e . I t was 20 - 47 cm above the stream, depending on the water l e v e l , and c o v e r e d an area a p p r o x i m a t e l y 260 x 200 cm. The h i g h s u b s i t e was s i t u a t e d near the base of a t r e e , on a v e r y g e n t l e r i d g e (51 - 114 cm above the water l e v e l ) . I t measured 160 x 125 cm and was the f u r t h e s t from the stream. A l l t h r e e s u b s i t e s were wet d u r i n g p e r i o d s of h i g h r a i n f a l l , but d r a i n a g e r a p i d l y r e s u l t e d i n l o s s of much of the water a t the h i g h e s t s u b s i t e . S i t e B was g e o g r a p h i c a l l y s i g h t l y d i f f e r e n t from S i t e A, and was d i v i d e d i n t o t h r e e s u b s i t e s . The low a r e a was 120 x 75 cm and was not as f r e q u e n t l y f l o o d e d as the c o r r e s p o n d i n g s u b s i t e i n S i t e A. I t ranged from 0 - 35 cm above the stream l e v e l . The m i d d l e s u b s i t e was a mound a l o n g the stream margin and measured 250 x 100 cm and was 49 - 70 cm above the water l e v e l . A l t h o u g h t h i s s u b s i t e was c l o s e t o the stream, i t was d e s i g n a t e d as the middl e s u b s i t e because i t was h i g h e r i n e l e v a t i o n above the stream. The h i g h a r e a was beyond the low d e p r e s s i o n , r a i s e d 39 - 69 cm above the stream l e v e l , and measured 200 x 175 cm. 10 The i n f o r m a t i o n on p r e c i p i t a t i o n and temperature was taken from the C l i m a t o l o g i c a l S t a t i o n Report of the Atmospheric Environment S e r v i c e of Environment Canada. The data were g a t h e r e d at s t a t i o n 94238, 110 8487-267 BC, which i s l o c a t e d a t the UBC a g r i c u l t u r a l p l o t s , a p p r o x i m a t e l y 2 km N of the study s i t e s . T h i s i s p r e s e n t e d t o g i v e a g e n e r a l i n d i c a t i o n of the c l i m a t e i n the area and i s summarized f o r the sample d a t e s i n T a b l e I and Table I I . In these t a b l e s the average temperature and the t o t a l p r e c i p i t a t i o n are g i v e n f o r one and two weeks p r i o r t o the s a m p l i n g d a t e . 11 Ta b l e I . C l i m a t i c Data For Sample Dates At S i t e A Ra i n f a l l Temperature  For p r i o r For p r i o r For p r i o r For p r i o r 2 weeks week 2 weeks week Sample Date i n mm i n mm i n °C i n °C A2 Apr 5, 77 13.22 0.00 6.98 8.33 A3 J u l 25, 77 17.54 0.00 16.08 17.03 A4 Nov 7, 77 107.57 50.86 8.26 7.27 A5 Feb 13, 78 69.70 29.40 4.95 4.69 A6 May 19, 78 68.30 51.90 11.50 12.04 hi J u l 26, 78 0.00 0.00 19.11 20.31 T a b l e I I . C l i m a t i c Data For Sample Dates At S i t e B Ra i n f a l l T e m p e r a t u r e  F o r p r i o r F o r p r i o r F o r p r i o r F o r p r i o r 2 weeks week 2 weeks week Sample Date i n mm i n mm i n °C i n °C BI May 17, 77 20.83 17.29 10.24 9.76 B2 Sep 21, 77 34.55 33.31 13.32 12.98 B3 Jan 18, 78 63.10 5.80 4.76 5.59 B4 Apr 5, 78 66.00 26.00 8.71 8.15 12 2. C l a s s i f i c a t i o n of the L i t t e r L i t t e r has been d i v i d e d c l a s s i c a l l y i n t o l a y e r s s e g r e g a t e d w i t h r e s p e c t t o the degree of decay (Kubiena, 1953). The L or l i t t e r l a y e r i s the f r e s h l y f a l l e n l i t t e r , which i s j u s t b e g i n n i n g t o decay. In t h i s l a y e r , the l i t t e r i s s t i l l r e c o g n i z a b l e as t o i t s o r i g i n and has u s u a l l y t u r n e d brown. Below t h i s l i e s the F or f e r m e n t a t i o n l a y e r of decomposing l i t t e r . T h i s l a y e r sometimes can be d i v i d e d i n t o the F l and the F2, based on completeness of decay. The F l l a y e r i s c h a r a c t e r i z e d by l e a v e s which a r e b e g i n n i n g t o break down c o m p l e t e l y but are s t i l l i d e n t i f i a b l e . O f t e n they have begun t o t u r n b l a c k i s h t o gra y . In the F2 l a y e r , l e a v e s a re l e s s r e c o g n i z a b l e , f r a g m e n t a r y , gray and almost c o m p l e t e l y decayed. T h i s d i v i s i o n of the f e r m e n t a t i o n l a y e r may be observ e d e a s i l y i n many t y p e s of l i t t e r but not i n the case of the p r e s e n t s t u d y . A t h i r d l a y e r , the H or humus l a y e r , c o n s i s t s of u n r e c o g n i z a b l e o r g a n i c m a t t e r , and l i e s d i r e c t l y on the m i n e r a l s o i l . T h i s l a y e r was not i n v e s t i g a t e d i n the c u r r e n t s t u d y . Thuja l i t t e r i s not made up of d i s t i n c t l e a v e s , but of p i e c e s of b r a n c h l e t s which have f a l l e n w i t h the s c a l e - l i k e l e a v e s s t i l l a t t a c h e d . T h e r e f o r e any f u r t h e r mention of l i t t e r or l e a v e s a c t u a l l y r e f e r s t o b r a n c h l e t s . The d i v i s i o n of the l i t t e r i n t o the l a y e r s o u t l i n e d above proved t o be i m p o s s i b l e . The even d i s t r i b u t i o n of the d i f f e r e n t s t a g e s of decay was hampered by the i n c o n s i s t e n c y of the s i z e of the b r a n c h l e t s and the l a r g e amount of s i l t w i t h i n the l i t t e r l a y e r . Thus the l i t t e r was d i v i d e d i n t o o n l y two l a y e r s . The uppermost or L 13 l a y e r c o n t a i n e d the f r e s h l y f a l l e n and l e s s decomposed l i t t e r . I t was c h a r a c t e r i z e d by b r a n c h l e t s which were r a t h e r l o n g (up t o 20 cm), brown t o b l a c k and j u s t b e g i n n i n g t o show s i g n s of decay. Sometimes, as a f t e r a r e c e n t storm, the L l a y e r c o n t a i n e d b r a n c h l e t s which were s t i l l green or j u s t b e g i n n i n g t o t u r n y e l l o w . The l a y e r below t h i s was the F l a y e r . I t was made up of s h o r t e r p i e c e s of l i t t e r , brown t o b l a c k t o gray , showing d e f i n i t e s i g n s of decay. They were t y p i c a l l y f r a g i l e , f l a t t e n e d , and w i t h v e r y l i t t l e i n t e r n a l t i s s u e r e m a i n i n g . In many, the s c a l e - l i k e l e a v e s were detached from the a x i s . The low s u b s i t e of S i t e A, on the stream margin, was e x t r e m e l y s i l t y , such t h a t the F l a y e r appeared t o be m a i n l y s i l t . The L l a y e r was up t o 1 cm deep, w i t h s i l t prominent i n the lower p o r t i o n . In consequence the F l a y e r was c o m p l e t e l y i n t e r m i x e d w i t h mud and was up t o 2 cm t h i c k . The middl e s u b s i t e had a t h i c k e r L l a y e r , as much as 2 cm, w i t h l i t t l e s i l t . The F l a y e r was a p p r o x i m a t e l y 3 cm t h i c k , s a t u r a t e d and s i l t y , but l e s s than i n the low s u b s i t e . The h i g h s u b s i t e had l i t t l e or no s i l t , w i t h the L l a y e r up t o o n l y 1.5 cm t h i c k . The F l a y e r was a p p r o x i m a t e l y 4 cm t h i c k and, i n the lower p o r t i o n , i n t e r m i x e d w i t h the humus. S i t e B had a s i m i l a r low s u b s i t e and was a l s o s i l t y ; the L l a y e r was 0 - 0.5 cm deep. The F l a y e r was mixed w i t h mud and was a p p r o x i m a t e l y 2.5 cm t h i c k . The middl e s u b s i t e was on the mound, and thus was not ve r y s i l t y . The L l a y e r was up to 1 cm deep and the F l a y e r was a p p r o x i m a t e l y 2 cm deep. The h i g h s u b s i t e had a r e l a t i v e l y t h i c k L l a y e r , up t o 2.5 cm deep, and the u n d e r l y i n g F l a y e r was a p p r o x i m a t e l y 4 cm t h i c k . 14 A l t h o u g h l i t t e r can be d i v i d e d i n t o l a y e r s , i t i s impo r t a n t to r e a l i z e t h a t t h e r e a re no c l e a r d i v i s i o n s . The e n t i r e l i t t e r l a y e r i s made up of b r a n c h l e t s i n v a r y i n g s t a g e s of decay. T h e i r r e l a t i v e placement on t o p of one a n o t h e r r e f l e c t s the time they o r i g i n a l l y f e l l from the t r e e . Those t h a t have been i n the l i t t e r l o n g e s t would t h e o r e t i c a l l y a l s o be the most decayed, and would be ex p e c t e d i n the lower l a y e r s . T h e r e f o r e , the f u n g i p r e s e n t i n the two l a y e r s would r e f l e c t a sequence i n the decay; the L l a y e r s p e c i e s would be e a r l y c o l o n i z e r s , w h i l e the F l a y e r s p e c i e s would be secondary c o l o n i z e r s . But i n v e r t e b r a t e s , f u n g a l m y c e l i a and spores can p o s s i b l y move from one l e a f t o the n e x t , or from one l a y e r t o the n e x t . Thus the d i s t r i b u t i o n of the s p e c i e s i n v o l v e d i n decay does not c o m p l e t e l y i n d i c a t e t h e i r p o s i t i o n i n the time sequence of decay, but t e n d e n c i e s can be obs e r v e d . The purpose of d i v i d i n g up the l i t t e r was t o t r y t o determine what f u n g i were p r e s e n t i n the l e a v e s at t h e i r r e s p e c t i v e s t a g e s of decay and thus observe a sequence of s u c c e s s i o n . 3. Treatment of the Samples B r a n c h l e t s were c o l l e c t e d a p p r o x i m a t e l y once every t h r e e months over a p e r i o d of about a year ( S i t e A, 6 times i n 15 months, T a b l e I ; S i t e B, 4 tim e s i n 12 months, T a b l e I I ) . One sample was made i n each of the t h r e e sub-areas i n the s i t e and d i v i d e d i n t o the L and F l a y e r s . The l e a v e s were p l a c e d i n f r e s h p l a s t i c bags and brought t o the l a b o r a t o r y f o r p r o c e s s i n g ; t h i s 15 was completed w i t h i n 48 hou r s . The p o s i t i o n s of the samples i n the s u b s i t e s v a r i e d throughout the sampl i n g p e r i o d . Each sample p o s i t i o n was c h a r a c t e r i z e d by the d i s t a n c e from the stream and the e l e v a t i o n above the stream. The water l e v e l i n the stream was a l s o measured. These t h r e e v a r i a b l e s were combined i n a graph t o g i v e some i d e a of the s p a t i a l r e l a t i o n s h i p of the samples ( F i g s . 1 and 2 ) . The b r a n c h l e t s were cut i n t o p i e c e s about 4 cm l o n g , i f n e c e s s a r y , and then washed a c c o r d i n g t o the method d e s c r i b e d by H a r l e y and Waid (1955) as m o d i f i e d s l i g h t l y (Bandoni and B a r r , 1976). The l e a v e s were washed i n s t e r i l e water and j a r s f o r 2 minutes on a r o t a r y shaker a t 400 rpms. They were then t r a n s f e r r e d t o new s t e r i l e j a r s and water, and- the p r o c e s s was r e p e a t e d 30 t i m e s . The a d d i t i o n of a 30 second s o a k - i n 50% e t h a n o l between washes 25 and 26 was i n s t i t u t e d t o h e l p keep <• down the y e a s t and b a c t e r i a l growth on the o u t s i d e of the l e a v e s . The s h o r t d u r a t i o n of the soak p r e c l u d e d i t s p e n e t r a t i o n i n t o the l e a v e s and thus the i n t e r n a l f u n g i s e e m i n g l y were unaf f e c t e d . A f t e r washing, the l e a v e s were b l o t t e d dry on s t e r i l e paper t o w e l s and t r e a t e d i n t h r e e ways. The f i r s t group was p l a c e d on s i x agar p l a t e s , two l e a v e s per p l a t e , and i n c u b a t e d a t 15° C. They were t u r n e d every 24 hours f o r f o u r days and then removed. The medium employed was S o r b o s e - Y e a s t - T e t r a c y c l i n e (SYT), s l i g h t l y m o d i f i e d from t h a t used by Bandoni and B a r r (1976). I t c o n s i s t e d of 0.4% s o r b o s e , 0.05% y e a s t e x t r a c t , and 1.5% agar (Ba n d o n i , 1977). A f t e r a u t o c l a v i n g and c o o l i n g t o 45° C, 100 16 mg/1 of t e t r a c y c l i n e h y d r o c h l o r i d e was added t o the medium. Sorbose was f i r s t used t o produce c o l o n i a l paramorphs i n Neurospora s t u d i e s (Tatum et a l . , 1949). I t seems t o a f f e c t the c e l l w a l l c o m p o s i t i o n and r e s t r i c t s h y p h a l spread ( T e r r a and Tatum, 1961). T e t r a c y c l i n e , a broad spectrum a n t i b i o t i c , p r e v e n t s the growth of b a c t e r i a . The p l a t e s were examined on the second and t h i r d days a f t e r the l e a v e s were f i r s t put i n and f r e q u e n t l y t h e r e a f t e r . The f u n g i i s o l a t e d were u s u a l l y from the f i r s t two s p o t s , which r e p r e s e n t e d those t h a t were a c t i v e l y growing w i t h i n the l e a v e s . L a t e r s p o t s were u s u a l l y overgrown q u i c k l y and r e p r e s e n t e d d u p l i c a t e s of those c o l o n i e s p r e s e n t on the f i r s t two s p o t s . T h i s treatment p r o v i d e d the m a j o r i t y of the c u l t u r e s o b t a i n e d . The remainder of the l e a v e s were d i v i d e d among f o u r s t e r i l e p e t r i p l a t e s . One was f i l l e d w i t h s t e r i l e water and b a i t e d w i t h p o l l e n f o r c h y t r i d i s o l a t i o n . The o t h e r t h r e e p l a t e s c o n t a i n e d s t e r i l e wet paper t o w e l and were used as m o i s t chambers. Two were kept a t 15° C, w h i l e the t h i r d was put a t 5° C. Not a l l of the c o n i d i o p h o r e s produced i n the 15° C p l a t e s were i s o l a t e d s i n c e they were numerous. The number of ti m e s a s p e c i e s was i s o l a t e d gave a r e l a t i v e i n d i c a t i o n of i t s abundance. I s o l a t i o n s were made a f t e r one and two weeks. 17 F i g u r e I . S p a t i a l Arrangement Of The Samples At S i t e A 2 - 7 = Sample Dates (see Table I ) LOW SUBSITE ' / .0 E l e v a t i o n above t h e s t r e a m (cm) 18 F i g u r e 2. S p a t i a l Arrangement Of The Samples At S i t e B 1 - 4 = Sample Dates (see T a b l e I I ) 19 4. M u l t i v a r i a t e A n a l y s i s of the Data The s p e c i e s i s o l a t e d from the SYT p l a t e s and the m oist chambers were used t o c h a r a c t e r i z e the samples. They were i d e n t i f i e d and the number of times they were i s o l a t e d was taken as an i n d i c a t i o n of t h e i r r e l a t i v e importance, or abundance. Each sample, i . e . L or F l a y e r i n the h i g h , m i d d l e or low s u b s i t e s , was c o n s i d e r e d a d i s t i n c t u n i t . They were i n c l u d e d i n a p r i n c i p a l component a n a l y s i s (PCA), w i t h the o c c u r r e n c e of the s p e c i e s of f u n g i c h a r a c t e r i z i n g the samples. T h i s i s a m u l t i v a r i a t e a n a l y s i s based e i t h e r on a c o v a r i a n c e or c o r r e l a t i o n m a t r i x from which an a x i s i s drawn r e p r e s e n t i n g the l a r g e s t amount of v a r i a t i o n among a l l the samples. A second a x i s , at n i n e t y degrees t o the f i r s t , spreads the p o i n t s out f u r t h e r , and the a n a l y s i s c o n t i n u e s l i k e t h i s i n a s t e p w i s e f a s h i o n . The r e s u l t i s a m u l t i - d i m e n s i o n a l p i c t u r e i n which the c l o s e n e s s of p o i n t s r e f l e c t s t h e i r s i m i l a r i t i e s . By u s i n g t h i s type of a n a l y s i s the raw data c o u l d be v i s u a l l y r e p r e s e n t e d and t r e n d s of s p e c i e s a s s o c i a t i o n s or sample a s s o c i a t i o n s would be e a s i l y seen. In a d d i t i o n t o PCA, a minimum v a r i a n c e c l u s t e r i n g of samples based on E u c l i d e a n d i s t a n c e was performed. The r e s u l t s , p r e s e n t e d i n the form of a dendrogram, i l l u s t r a t e the h i e r a r c h i c a l r e l a t i o n s h i p s which e x i s t among the samples. Both of t h e s e t y p e s of a n a l y s e s p r o v i d e a way of i l l u s t r a t i n g the d a t a such t h a t t r e n d s apparent from the i s o l a t i o n s c o u l d be seen g r a p h i c a l l y and o t h e r p o s s i b l e a s s o c i a t i o n s , not so a p p a r e n t , would be d i s c o v e r e d . The computer a n a l y s i s was run on the MIDAS program 20 d i s t r i b u t e d by the S t a t i s t i c a l R esearch L a b o r a t o r y of the U n i v e r s i t y of M i c h i g a n (Fox and G u i r e , 1976), which i s a v a i l a b l e on the UBC computer (an Amdahl 470V/6 Model I I , run under the M i c h i g a n T e r m i n a l System). In t h i s program the a n a l y s i s was based on a cases v s . v a r i a b l e s m a t r i x . The d a t a were s t r u c t u r e d such t h a t the cases were the i n d i v i d u a l sample u n i t s (eg_. L l a y e r from low s u b s i t e , F l a y e r from low s u b s i t e , L l a y e r from middle s u b s i t e , e t c . ) . The v a r i a b l e s were the v a r i o u s s p e c i e s of f u n g i t h a t were i s o l a t e d i n the s u b s i t e s and l a y e r s . The d a t a f o r S i t e A and S i t e B were a n a l y z e d s e p a r a t e l y . Only the more f r e q u e n t l y i s o l a t e d s p e c i e s were u t i l i z e d i n the a n a l y s i s , as they were thought t o r e p r e s e n t the f u n g i most a c t i v e l y d e c a y i n g the l i t t e r a t the samp l i n g t i m e . The PCA was performed w i t h both the u n s e a l e d and s c a l e d o p t i o n s . The u n s e a l e d o p t i o n means t h a t the d a t a are c e n t e r e d by s p e c i e s , but a r e l e f t u n s t a n d a r d i z e d . T h i s ensures t h a t the more f r e q u e n t l y i s o l a t e d s p e c i e s a re more i m p o r t a n t i n d e t e r m i n i n g the f i n a l r e l a t i o n s h i p s . The b a s i c premise i s t h a t these s p e c i e s are the f u n g i most a c t i v e i n the decay p r o c e s s , and t h e i r i nvolvement i s r e l a t e d d i r e c t l y t o the number of i s o l a t i o n s made. The importance of the l e s s f r e q u e n t l y i s o l a t e d s p e c i e s c o u l d t h e r e f o r e be o v e r l o o k e d . The s c a l i n g o p t i o n means t h a t i n a d d i t i o n t o c e n t e r i n g by s p e c i e s , the data are s t a n d a r d i z e d by d i v i d i n g t h r ough by s p e c i e s s t a n d a r d d e v i a t i o n s . T h i s s t e p e q u a l i z e s the importance of s p e c i e s i n the a n a l y s i s , w i t h the e f f e c t of g i v i n g weight t o l e s s common s p e c i e s . From t h i s combined approach, a b e t t e r v i s u a l i z a t i o n of the s p e c i e s a s s o c i a t i o n s and sample a s s o c i a t i o n s c o u l d be o b t a i n e d . 21 I I I . R e s u l t s And O b s e r v a t i o n s In the c o u r s e of t h i s i n v e s t i g a t i o n , 1641 c u l t u r e s , r e p r e s e n t i n g 68 s p e c i e s , were i s o l a t e d u s i n g the t e c h n i q u e s o u t l i n e d above. A l l of the s p e c i e s i s o l a t e d from S i t e A a r e l i s t e d i n T a b l e s I I I - V, a l o n g w i t h t h e i r d i s t r i b u t i o n throughout the s a m p l i n g t i m e . Each of the t a b l e s g i v e s the d i s t r i b u t i o n of s p e c i e s i n one of the t h r e e s u b s i t e s ( h i g h , m i d d l e , or l o w ) . The same i n f o r m a t i o n i s a l s o g i v e n f o r S i t e B i n T a b l e s V I I - IX. The i n f o r m a t i o n c o n c e r n i n g the s p e c i e s i s o l a t e d from the l i v i n g l e a v e s i s g i v e n i n Table V I , f o r both s i t e s . Two of the t e c h n i q u e s employed i n t h i s work e i t h e r worked p o o r l y or p r o v i d e d no u s e f u l i n f o r m a t i o n , e.g., the m o i s t chambers i n c u b a t e d a t 5° C. Most of .the f u n g i which o c c u r r e d on the l e a v e s i n c u b a t e d a t 15° C a l s o appeared on the 5° C p l a t e s but a t a much slower r a t e . Because of the l e n g t h of time i t took f u n g i t o d e v e l o p , these 5° C p l a t e s o f t e n became i n f e s t e d w i t h m i t e s . For t h i s reason and s i n c e no new i n f o r m a t i o n was o b t a i n e d from them, the i s o l a t e s from the 5° C p l a t e s were o m i t t e d from the s t u d y . Another t e c h n i q u e which proved r e l a t i v e l y u n s u c c e s s f u l was the c h y t r i d i s o l a t i o n method. Only one s p e c i e s was i s o l a t e d and i t was t e n t a t i v e l y i d e n t i f i e d as Rhizophydium s p h a e r o t h e c a Zopf. In most ca s e s t h a l l i d e v e l o p e d on the p o l l e n g r a i n s but o t h e r f i l a m e n t o u s f u n g i a l s o appeared. I s o l a t i o n of the c h y t r i d was d i f f i c u l t because of these o t h e r f u n g i and a l s o because of b a c t e r i a l growth. The c h y t r i d a l s o proved somewhat s e n s i t i v e t o 22 t e t r a c y c l i n e , so e l i m i n a t i n g the b a c t e r i a o f t e n e l i m i n a t e d the c h y t r i d as w e l l . Any tre a t m e n t of i t s s e a s o n a l i t y was f e l t t o be somewhat b i a s e d because of the s e t e c h n i c a l d i f f i c u l t i e s , t h e r e f o r e i t s d i s t r i b u t i o n was not i n c l u d e d i n the d i s c u s s i o n s . O b s e r v a t i o n s made d u r i n g the cour s e of t h i s study i n d i c a t e t h a t t h i s c h y t r i d was p r e s e n t on almost a l l sample d a t e s a t a l l s i t e s and s u b s i t e s . The o c c u r r e n c e of c h y t r i d s on t e r r e s t r i a l l i t t e r has been r e p o r t e d p r e v i o u s l y by Bandoni and B a r r (1976), but t h i s i s the f i r s t r e p o r t from washed c o n i f e r l i t t e r . S t e r i l e m y c e l i a t y p i c a l l y a r e i s o l a t e d i n l i t t e r s t u d i e s . These i s o l a t e s u s u a l l y a re d i v i d e d i n t o two groups, s t e r i l e w h i t e m y c e l i a and s t e r i l e dark m y c e l i a . O f t e n s t u d i e s on l i t t e r and s o i l f u n g i i n c l u d e these groups i n t h e i r d i s c u s s i o n s on s e a s o n a l i t y and s p a t i a l d i s t r i b u t i o n . T h i s i s an u n s a t i s f a c t o r y p r a c t i c e s i n c e n e i t h e r group r e p r e s e n t s known t a x a . The s t e r i l e dark m y c e l i a i s o l a t e d i n t h i s study grew v e r y s l o w l y i n c u l t u r e . T h e i r v e g e t a t i v e morphology was s i m i l a r t o t h a t found i n i s o l a t e s of Endophragmia a l t e r n a t a , which a l s o were o f t e n slow i n "forming s p o r e s . In f a c t many of the i s o l a t e s of E. a l t e r n a t a were f i r s t i s o l a t e d as s t e r i l e dark m y c e l i a . Because of the a r t i f i c i a l , and p o s s i b l y h e terogenous, n a t u r e of the s e g r o u p i n g s n e i t h e r was i n c l u d e d i n the a n a l y s e s a l t h o u g h they a r e r e c o r d e d i n the s p e c i e s l i s t s p r o v i d e d . D e c o m p o s i t i o n of the l i t t e r i s a complex p r o c e s s i n v o l v i n g many v a r i a b l e s and organisms. In the p r e s e n t s t u d y , the f i l a m e n t o u s f u n g i ( p r e d o m i n a n t l y Fungi I m p e r f e c t i ) i n v o l v e d i n decay are examined. In s e l e c t i n g f o r i m p e r f e c t f u n g i , b a c t e r i a and y e a s t s have been s e l e c t e d a g a i n s t . The invol v e m e n t of these 23 two groups of organisms i n the d e c o m p o s i t i o n p r o c e s s has been reviewed elsewhere (Anderson and Domsch, 1973; Faure-Raynaud and G o u r b i e r e , 1976; L a s t and D e i g h t o n , 1965; L a s t and Warren, 1972; Remacle, 1970; Suberkropp and K l u g , 1976). I n v e r t e b r a t e s are a l s o i n v o l v e d i n the breakdown of l i t t e r and are i m p o r t a n t i n the p r e p a r a t i o n of the l i t t e r f o r subsequent c o l o n i z a t i o n by f u n g i . T h e i r involvement has been i g n o r e d here s i n c e o t h e r workers have reviewed t h e i r e f f e c t on decay (Cummins et a l . , 1973; van der D r i f t and Witkamp, 1960; Hayes, 1963; S e d e l l et a l . , 1975; T r i s k a and B u c k l e y , 1978; Witkamp and C r o s s l e y , 1966) . No B a s i d i o m y c e t e s and few Ascomycetes were i s o l a t e d , p o s s i b l y a r e s u l t of t h e i r slow growth r a t e . T h i s , p l u s the d i f f i c u l t i e s i n v o l v e d i n i d e n t i f i c a t i o n , has u s u a l l y e l i m i n a t e d them from most r e s e a r c h e s on d e c o m p o s i t i o n of l i t t e r . N e v e r t h e l e s s a few workers have examined t h e i r i nvolvement ( C h a s t u k h i n , 1946, 1962; Cromack et a l . , 1975; Egorava, 1968; L i n d b e r g , 1946; M i k o l a , 1956, 1960; S a i t o , 1965). Only the Fungi I m p e r f e c t i were examined i n t h i s s tudy s i n c e they r e p r e s e n t the m a j o r i t y of f u n g i i n v o l v e d i n the decay of l i t t e r , and a r e r e a d i l y i s o l a t e d and i d e n t i f i e d . 24 Tab l e I I I . S p e c i e s I s o l a t e d In The High S u b s i t e At S i t e A Acremonium s t r i c t u m  A crodontium c r a t e r i f o r . A r t i c u l o s p o r a t e t r a c l a . Chaetomium sp. C h a l a r a c o n s t r i c t a C. c y l i n d r o s p e r m a C. l o n g i p e s C l a d o s p o r i u m c l a d o s p o r . C y l i n d r o c a r p o n didymum C. tenue Endophragmia a l t e r n a t a  F l a g e l l o s p o r a s t r i c t a  F u s a r i u m l a t e r i t i u m  G l i o c l a d i u m roseum  G l i o m a s t i x sp. H e l i c o d e n d r o n t r i g l i t z . K r i e g e r i a s e a v e r i  L i b e r t e l l a sp. Monoci11ium s t a t e M o r t i e r e l l a i s a b e l l i n a M. ramanniana v a r . ram. Mucor h i e m a l i s f. hiem. P a e c i l o m y c e s sp. HIGH SUBSITE L LAYER AP JU NO FE MY JU 11 1 2 1 3 1 2 1 2 3 1 2 1 2 1 7 5 12 1 1 1 3 1 4 1 F LAYER AP JU NO FE MY JU 1 2 2 1 1 1 9 2 4 4 5 25 Tab l e I I I . ( c o n t i n u e d ) P e n i c i l l i u m b r e v i - compactum P. c i t r i n u m P. decumbens P. f r e q u e n t a n s P. n i g r i c a n s P. notatum P. r a i s t r i c k i i P. verrucosum v a r . eye. P e s t a l o t i a monochaet. P o l y s c y t a l u m f e c u n d i s s . R a m i c h l o r i d i u m s u b u l a t . Selenophoma sp. Septonema.chaetospira S. c h a e t o s p . v a r . p i n i Thysanophora p e n i c i 1 1 i . Tr ichoderma k o n i n q i i T. polysporum T. v i r i d e V a r i c o s p o r i u m elodeae  V e r t i c i l l i u m b u l b i l l o s . V. l e c a n i i S t e r i l e Dark M y c e l i a HIGH SUBSITE L LAYER AP JU NO FE MY JU 3 1 2 3 1 2 3 1 1 1 1 1 1 2 2 2 1 1 1 1 1 1 2 1 1 1 F LAYER AP JU NO FE MY JU 1 3 2 3 3 3 4 4 1 1 2 1 2 1 1 3 1 3 5 1 1 1 1 1 26 Table IV. S p e c i e s I s o l a t e d In The M i d d l e S u b s i t e At S i t e A Acremonium s t r i c t u m  A crodontium c r a t e r i f o r . A r t i c u l o s p o r a t e t r a c l a . C h a l a r a c o n s t r i c t a C. l o n g i p e s C l a d o s p o r i u m c l a d o s p o r . C y l i n d r o c a r p o n didymum C. tenue Endophragmia a l t e r n a t a Epicoccum sp. F l a g e l l o s p o r a s t r i c t a H e l i c o d e n d r o n t r i g l i t z . H e l i c o o n fuscosporum K r i e g e r i a s e a v e r i L i b e r t e l l a sp. Monoci11ium s t a t e Mucor h i e m a l i s f . hiem. Oedocephalum sp. P a e c i l o m y c e s sp. Pen i c i 1 1 i um brev i - compactum P. c i t r i n u m P. e c h i n u l a t u m MIDDLE SUBSITE L LAYER AP JU NO FE MY JU 1 1 1 4 1 2 1 1 1 2 1 2 2 1 2 9 5 1 9 2 1 2 4 7 1 2 2 1 3 2 1 1 3 2 1 F LAYER AP JU NO FE MY JU 8 2 3 1 5 1 1 9 2 2 5 2 7 4 6 9 1 2 1 1 1 27 Tab l e IV. ( c o n t i n u e d ) MIDDLE SUBSITE L LAYER F LAYER AP JU NO FE MY JU AP JU NO FE MY JU P. f r e q u e n t a n s 1 P. n i g r i c a n s 2 1 1 1 5 P. notatum 1 P. r a i s t r i c k i i 1 2 P. verrucosum v a r . eye. 1 1 1 1 3 P e r i c o n i e l l a sp. 1 P e s t a l o t i a monochaet. 3 2 4 P o l y s c y t a l u m f e c u n d i s s . 8 1 1 2 3 1 1 R a m i c h l o r i d i u m s u b u l a t . 1 Selenophoma sp. 1 5 1 Septonema c h a e t o s p i r a 2 1 1 S. c h a e t o s p . v a r . p i n i 1 2 Sporidesmium flexum 1 5 1 1 5 1 Trichoderma k o n i n g i i 1 1 T. v i r i d e 5 2 8 4 V a r i c o s p o r i u m elodeae 2 V e r t i c i l l i u m b u l b i l l o s . 1 S t e r i l e Dark M y c e l i a 1 1 1 2 1 1 S t e r i l e White M y c e l i a 1 2 1 1 1 1 2 28 Ta b l e V. S p e c i e s I s o l a t e d In The Low S u b s i t e At S i t e A A r t i c u l o s p o r a t e t r a c l a . C h a l a r a c o n s t r i c t a C. l o n q i p e s C. s t i p i t a t a C l a d o s p o r i u m c l a d o s p o r . C y l i n d r o c a r p o n didymum C. tenue Endophraqmia a l t e r n a t a F l a q e l l o s p o r a s t r i c t a F u s a r i u m l a t e r i t i u m G l i o m a s t i x sp. Gnomonia sp. H e l i c o d e n d r o n giqanteum H. t r i q l i t z i e n s i s H yalodendron l i g n i c o l a I d r i e l l a sp. K r i e g e r i a s e a v e r i L i b e r t e l l a sp. Monoci11ium s t a t e M o r t i e r e l l a i s a b e l l i n a M. ramanniana v a r . ram. Mucor h i e m a l i s f. hiem. P e n i c i l l i u m b r e v i - compactum LOW SUBSITE L LAYER AP JU NO FE MY JU 5 2 1 1 4 4 1 2 2 1 1 1 1 2 3 10 1 3 2 1 1 1 5 1 2 5 1 F LAYER AP JU NO FE MY JU 3 6 1 2 2 1 2 3 2 1 1 1 2 3 2 1 1 4 3 3 3 5 1 2 1 1 2 Table V. ( c o n t i n u e d ) P. c i t r i n u m P. decumbens P. n i g r i c a n s P. notatum P. r a i s t r i c k i i P. verrucosum v a r . eye. P e s t a l o t i a monochaet. P o l y s c y t a l u m f e c u n d i ss . R a m i c h l o r i d i u m s u b u l a t . Selenophoma sp. Septogloeum sp. Septonema c h a e t o s p i r a S. c h a e t o s p . v a r . p i n i S e s q u i c i l l i u m sp. Sporidesmium flexum Tr ichoderma k o n i n g i i T. polysporum T. v i r i d e Tripospermum m y r t i  V a r i c o s p o r i u m elodeae  V e r t i c i l l i u m b u l b i l l o s . V. c e p h a l o s p o r i u m S t e r i l e White M y c e l i a LOW SUI 3SITE L LAYER F LAYER AP JU NO FE MY JU AP JU NO FE MY JU 1 1 1 2 1 2 1 1 3 - 1 . 1 2 2 1 1 1 3 3 1 2 6 1 2 1 1 1 1 1 1 1 3 1 2 2 5 5 5 5 2 4 1 1 1 1 1 1 1 1 2 1 1 1 4 3 1 1 2 6 2 1 1 1 2 2 2 5 1 1 1 2 1 1 1 1 3 2 2 6 2 1 2 3 4 Table V I . S p e c i e s I s o l a t e d From L i v i n g Leaves At Both S i t e s Acrodont ium c r a t e r i forme  C l a d o s p o r i u m c l a d o s p o r i o i d e s  Endophragmia a l t e r n a t a  Hyalodendron 1 i g n i c o l a  K r i e q e r i a s e a v e r i  Oedocephalum sp. P e n i c i 1 1 i u m brevi-compactum P. notatum P. verrucosum v a r . c y c l o p i u m P e s t a l o t i a m o n o c h a e t i o i d e s Selenophoma sp. Septogloeum sp. Tr ichoderma k o n i n g i i T. v i r i d e Tripospermum m y r t i SITE A SITE B AP MY JU SE AP 77 78 78 77 78 6 1 4 1 1 2 2 1 1 4 2 1 1 2 4 5 2 2 2 3 1 2 1 5 31 T a b l e V I I . S p e c i e s I s o l a t e d In The High S u b s i t e At S i t e B Acremonium s t r i c t u r n  A c rodontium c r a t e r i f o r m e  C h a l a r a c o n s t r i c t a C. c y l i n d r o s p e r m a C. l o n q i p e s C. s t i p i t a t a C h l o r i d i u m v i r e s c e n s v a r . chlamyd, C l a d o s p o r i u m c l a d o s p o r i o i d e s C y l i n d r o c a r p o n tenue Endophragmia a l t e r n a t a Epicoccum sp. F l a g e l l o s p o r a s t r i c t a F u sarium l a t e r i t i u m G l i o c l a d i u m roseum H e l i c o d e n d r o n t r i g l i t z i e n s i s H yalodendron 1 i q n i c o l a M o r t i e r e l l a i s a b e l l i n a M. ramanniana v a r . ramanniana Mucor h i e m a l i s f. h i e m a l i s Pen i c i 1 1 i um brevi-compactum P. decumbens P. f r e q u e n t a n s P. n i g r i c a n s HIGH SUBSITE L LAYER MY SE JA AP 1 3 1 2 1 2 1 1 8 8 1 1 1 2 2 3 1 1 2 2 1 3 F LAYER MY SE JA AP 1 1 1 6 1 1 1 2 1 2 2 2 2 9 11 32 Table V I I . ( c o n t i n u e d ) P. r a i s t r i c k i i P. verrucosum v a r . c y c l o p i u m  P e s t a l o t i a m o n o c h a e t i o i d e s  P o l y s c y t a l u m fecundissimum  Selenophoma sp. Septogloeum sp. Septonema c h a e t o s p i ra S. c h a e t o s p i r a v a r . p i n i  T o r u l a sp. Tr ichoderma kon i n g i i T. polysporum-T. v i r i d e T r i p o s p o r i u m e l e g a n s S t e r i l e Dark M y c e l i a S t e r i l e White M y c e l i a HIGH SUBSITE L LAYER MY SE JA AP 1 1 1 2 1 2 1 1 2 9 1 1 4 F LAYER MY SE JA AP 1 1 1 6 2 1 1 1 33 T a b l e V I I I . S p e c i e s I s o l a t e d In The M i d d l e S u b s i t e At S i t e B MIDDLE £ 5UBSITE L LAYER F LAYER MY SE JA AP MY SE JA AP Acremonium s t r i c t u m 1 3 1 B o t r y t i s sp. 2 C h a l a r a c y l i n d r o s p e r m a 1 3 1 1 C l a d o s p o r i u m c l a d o s p o r i o i d e s 3 2 1 2 1 C y l i n d r o c a r p o n tenue 3 4 5 1 6 4 3 Endophragmia a l t e r n a t a 1 F l a g e l l o s p o r a s t r i c t a 2 1 1 F u s a r i u m l a t e r i t i u m 1 G l i o c l a d i u m roseum 1 1 2 1 2 H e l i c o d e n d r o n t r i g l i t z i e n s i s 1 Hyalodendron l i g n i c o l a 5 1 L i b e r t e l l a sp. 1 1 M o r t i e r e l l a ramanniana v a r . raman. 2 1 Mucor h i e m a l i s f. h i e m a l i s 1 1 P e n i c i l l i u m brevi-compactum 6 1 5 P. c i t r i n u m 1 P. f r e q u e n t a n s 1 P. n i g r i c a n s 2 1 P. verrucosum v a r . c y c l o p i u m 1 2 1 1 3 1 P e s t a l o t i a m o n o c h a e t i o i d e s 3 P o l y s c y t a l u m fecundissimum 1 1 3 1 Selenophoma sp. 4 1 2 6 2 Septogloeum sp. 1 34 T a b l e V I I I . ( c o n t i n u e d ) MIDDLE SUBSITE L LAYER F LAYER MY SE JA AP MY SE JA AP Septonema c h a e t o s p i r a v a r . p i n i 1 T h y s a n o p h o r a p e n i c i 1 1 i o i d e s 1 T o r u l a s p . 1 T r i c h o d e r m a k o n i n g i i 1 3 1 7 2 T. p o l y s p o r u m 1 2 1 4 T. v i r i d e 1 T r i p o s p o r i u m e l e g a n s 3 1 S t e r i l e Dark M y c e l i a 2 3 1 1 1 35 T a b l e I X . S p e c i e s I s o l a t e d I n The Low S u b s i t e A t S i t e B A l t e r n a r i a s p . A r t i c u l o s p o r a t e t r a c l a d i a C h a l a r a c o n s t r i c t a C. c y l i n d r o s p e r m a C. s t i p i t a t a C l a d o s p o r ium c l a d o s p o r i o i d e s  C l a t h r o s p h a e r i n a s p . C y l i n d r o c a r p o n t e n u e  C y l i n d r c c a r p o n didymum  G l i o c l a d i u m roseum  H a r p o s p o r i u m s p . H e l i c o d e n d r o n t r i g l i t z i e n s i s  H y a l o d e n d r o n 1 i q n i c o l a  L i b e r t e l l a s p . M o n o c i 1 1 i u m s t a t e o f N i e s s l i a M o r t i e r e l l a i s a b e l l i n a M. ramann i a n a v a r . ramann i a n a Mucor h i e m a l i s f . h i e m a l i s P e n i c i l l i u m b r e v i - c o m p a c t u m P. f r e q u e n t a n s P. n i g r i c a n s P. n o t a t u m P. r a i s t r i c k i i LOW SUBSITE L LAYER MY SE JA AP 1 1 2 1 2 8 4 8 1 1 1 2 1 1 1 2 F LAYER MY SE JA AP 1 2 8 4 4 1 1 1 1 1 36 T a b l e IX. ( c o n t i n u e d ) P. verrucosum v a r . c y c l o p i u m P e r i c o n i e l l a sp. P e s t a l o t i a m o n o c h a e t i o i d e s P o l y s c y t a l u m fecundissimum Selenophoma sp. Septogloeum sp. Septonema c h a e t o s p i r a S e s q u i c i 1 1 i u m sp. Tr ichoderma k o n i n g i i T. polysporum T. v i r i d e Tripospermum m y r t i  V e r t i c i l l i u m b u l b i l l o s u m V. l e c a n i i S t e r i l e Dark M y c e l i a S t e r i l e White M y c e l i a LOW SUBSITE L LAYER MY SE JA AP 2 5 2 3 3 2 6 1 1 1 3 7 2 1 2 F LAYER MY SE JA AP 1 2 1 5 10 6 9 1 1 1 1 J 37 I n e x a m i n i n g t h e d e c o m p o s i t i o n o f r e d c e d a r l i t t e r a n d s e a s o n a l i t y o f t h e f u n g a l s p e c i e s i n v o l v e d , t h e d a t a c a n be p r e s e n t e d i n a number o f ways. Some o f t h e s e t e n d t o c o n f u s e t h e r e a d e r , so p r e l i m i n a r y g u i d e l i n e s a r e p r e s e n t e d . N ot a l l s p e c i e s l i s t e d i n T a b l e s I I I - I X were deemed i m p o r t a n t i n t h e d e c a y p r o c e s s . I n f a c t , many o f t h e s e were i s o l a t e d o n l y o n c e o r t w i c e d u r i n g t h e e n t i r e s a m p l i n g p e r i o d . O n l y t h e more f r e q u e n t l y i s o l a t e d s p e c i e s a r e d i s c u s s e d . I n t h e f o l l o w i n g d i s c u s s i o n s e m p h a s i s i s p l a c e d on t h e l i t t e r l a y e r s i n t h e d i f f e r e n t s i t e s and s u b s i t e s . The f i r s t p o r t i o n c o n c e r n s t h e d o m i n a n t f u n g i i n t h e L a n d F l a y e r s a n d t h e i r d i s t r i b u t i o n i n t h e s u b s i t e s o f e a c h s i t e . The s e a s o n a l v a r i a t i o n i n e a c h l a y e r i s t h e n d i s c u s s e d , f o l l o w e d by a l i s t o f t h e i m p o r t a n t s p e c i e s ' a n d t h e i r c h a r a c t e r i s t i c s . 1. P r e d o m i n a n t S p e c i e s i n t h e L L a y e r The s p e c i e s i s o l a t e d f r o m t h e L l a y e r i n S i t e A v a r i e d i n s e a s o n a l o c c u r r e n c e a n d between s u b s i t e s . G e n e r a l l y t h e r e were s p e c i e s w h i c h o c c u r r e d more f r e q u e n t l y i n one o r two o f t h e d i f f e r e n t s u b s i t e s . O t h e r s p e c i e s t e n d e d t o o c c u r a t a l l t h r e e s u b s i t e s , a l t h o u g h t h e number o f i s o l a t i o n s a t e a c h m i g h t h a v e been d i f f e r e n t . T a b l e s X and XI s u m m a r i z e t h e d i s t r i b u t i o n o f s p e c i e s i n t h e L l a y e r a t S i t e s A a n d B, r e s p e c t i v e l y . The s p e c i e s h a v e been d i v i d e d i n t o g r o u p s b a s e d on t h e i r o c c u r r e n c e i n t h e s u b s i t e s ; G r o u p I - s p e c i e s w h i c h o c c u r r e d e q u a l l y i n a l l t h r e e s u b s i t e s ; G r o u p I I - s p e c i e s f o u n d i n t h e h i g h a n d low 38 s u b s i t e s ; Group I I I - s p e c i e s found i n the m i d d l e and low s u b s i t e s ; Group IV - s p e c i e s found i n the h i g h and m i d d l e s u b s i t e s ; Group V - s p e c i e s found i n the h i g h s u b s i t e ; Group VI - s p e c i e s found i n the m i d d l e s u b s i t e ; Group V I I - s p e c i e s found i n the low s u b s i t e . 39 T a b l e X. D i s t r i b u t i o n Of S p e c i e s In The L Layer At S i t e A TOTAL # OF I S O I J A T E S HIGH MIDDLE LOW SUBSITE SUBSITE SUBSITE GROUP I ( a l l s i t e s ) C y l i n d r o c a r p o n tenue 26 28 19 H e l i c o d e n d r o n t r i q l i t z i e n s i s 11 16 15 C l a d o s p o r i u m c l a d o s p o r i o i d e s 6 5 7 Trichoderma v i r i d e 5 5 7 C h a l a r a c o n s t r i c t a 6 4 4 GROUP I I ( h i g h and low) P e n i c i l l i u m verrucosum v a r . c y c l o p i u m 6 2 6 Trichoderma k o n i n g i i 4 1 2 C y l i n d r o c a r p o n didymum 3 3 GROUP I I I (middle and low) P o l y s c y t a l u m fecundissimum 3 12 10 A r t i c u l o s p o r a t e t r a c l a d i a 4 9 9 GROUP IV ( h i g h and mid d l e ) F l a g e l l o s p o r a s t r i c t a 3 2 1 P e n i c i l l i u m c i t r i n u m 3 2 1 Acremonium s t r i c t u m 3 2 GROUP V ( h i g h ) Mucor h i e m a l i s f. h i e m a l i s 9 3 1 P e n i c i l l i u m n i g r i c a n s 9 2 1 P e n i c i l l i u m r a i s t r i c k i i 6 1 L i b e r t e l l a sp. 5 2 R a m i c h l o r i d i u m subulatum 3 1 Endophraqmia a l t e r n a t a 2 1 GROUP VI (mi d d l e ) Selenophoma sp. 3 7 2 P e s t a l o t i a m o n o c h a e t i o i d e s 1 9 1 Sporidesmium flexum 7 2 H e l i c o o n fuscosporum 4 P e n i c i l l i u m brevi-compactum 1 3 GROUP V I I (low) Septonema c h a e t o s p i r a 2 2 9 V a r i c o s p o r i u m elodeae 1 2 6 C h a l a r a l o n g i p e s 2 5 Septonema c h a e t o s p i r a v a r . p i n i 1 1 4 40 Ta b l e X I . D i s t r i b u t i o n Of S p e c i e s In The L Layer At S i t e B GROUP I ( a l l s i t e s ) C y l i n d r o c a r p o n tenue Selenophoma sp. Pen i c i11ium brevi-compactum P o l y s c y t a l u m f e c u n d i ssimum Septogloeum sp. GROUP I I ( h i g h and low) Tr ichoderma k o n i n g i i H e l i c o d e n d r o n t r i g l i t z i e n s i s C h a l a r a c o n s t r i c t a GROUP I I I (middle and low) Cla d o s p o r ium c l a d o s p o r i o i d e s P e s t a l o t i a m o n o c h a e t i o i d e s GROUP IV ( h i g h and mi d d l e ) Hyalodendron l i q n i c o l a  Pen i c i11ium verrucosum . v a r . c y c l o p i u m M o r t i e r e l l a ramanniana var ramanniana F l a g e l l o s p o r a s t r i c t a F u sarium l a t e r i t i u m GROUP V (hig h ) C h a l a r a l o n q i p e s  Septonema c h a e t o s p i r a  G l i o c l a d i u m roseum  C h l o r i d i u m v i r e s c e n s v a r . chlamydosporum Mucor h i e m a l i s f. h i e m a l i s GROUP VI (middle) T r i p o s p o r i u m e l e q a n s GROUP V I I (low) L i b e r t e l l a sp. Septonema c h a e t o s p i r a  Tr ichoderma polysporum v a r . p i n I T O T A L # O F I S O I J A T E S H I G H M I D D L E L O W S U B S I T E S U B S I T E S U B S I T E 27 12 22 17 13 14 5 6 5 4 5 7 1 1 1 8 5 13 3 1 4 2 2 5 3 1 3 2 5 6 1 3 3 1 2 2 1 2 2 1 1 8 6 1 4 1 1 3 3 * 1 3 1 3 1 2 1 2 41 From an e x a m i n a t i o n of the p r e c e d i n g t a b l e s , i t i s e v i d e n t t h a t d i f f e r e n t s p e c i e s were o f t e n found a t the two s i t e s , and those t h a t were common between the s i t e s , were o f t e n not found i n the same s u b s i t e s . The s p e c i e s i s o l a t e d i n the L l a y e r a t S i t e A, but not S i t e B, were: Acremonium s t r i c t u m  A r t i c u l o s p o r a t e t r a c l a d i a  C y l i n d r o c a r p o n didymum  Endophragmia a l t e r n a t a  H e l i c o o n fuscosporum  Pen i c i 1 1 i urn c i t r inum P. n i g r i c a n s P. r a i s t r i c k i i  R a m i c h l o r i d i u m subulatum  Sporidesmium flexum  Tr ichoderma v i r i d e  V a r i c o s p o r i u m elodeae Some of these s p e c i e s were t o t a l l y absent from the o t h e r s i t e ; o t h e r s d i d o c c u r , but o n l y once or- t w i c e . The s p e c i e s found a t S i t e B, but not S i t e A, i n c l u d e d : C h l o r i d i u m v i r e s c e n s v a r . chlamydosporum  Fusarium l a t e r i t i u m  G l i o c l a d i u m roseum  Hyalodendron l i g n i c o l a M o r t i e r e l l a ramanniana v a r . ramanniana Septogloeum sp. Tr ichoderma polysporurn T r i p o s p o r i u m e l e g a n s D e s p i t e the above d i f f e r e n c e s , a few s p e c i e s had s i m i l a r d i s t r i b u t i o n s i n both S i t e A and S i t e B. Only one s p e c i e s was p r e s e n t i n Group I i n bo t h s i t e s , and t h a t was C y l i n d r o c a r p o n  tenue. T h i s was not s u r p r i s i n g , s i n c e i t was the fungus most f r e q u e n t l y i s o l a t e d i n t h i s s t u d y . Tr ichoderma k o n i n g i i was 42 found i n Group I I ( h i g h and low s u b s i t e s ) ; F l a g e l l o s p o r a s t r i c t a was p r e s e n t i n Group IV ( h i g h and middle s u b s i t e s ) ; Mucor  h i e m a l i s f. h i e m a l i s was a member of Group V ( h i g h s u b s i t e ) ; Septonema c h a e t o s p i r a was found i n Group V I I (low s u b s i t e ) , i n both S i t e A and S i t e B. 2. Predominant S p e c i e s i n the F Lay e r The F, or f e r m e n t a t i o n l a y e r , r e p r e s e n t s l e a v e s which have undergone decay f o r a l o n g e r p e r i o d of ti m e . Some of the s p e c i e s i s o l a t e d were the same as i n the L l a y e r , but some were new. Many of the same s p e c i e s were d i s t r i b u t e d d i f f e r e n t l y among the s u b s i t e s . The s p e c i e s found i n t h i s l a y e r a r e summarized i n T a b l e s X I I 'and X I I I , f o r S i t e A and S i t e B, r e s p e c t i v e l y . They are d i v i d e d i n t o the same groups as f o r the L l a y e r . 43 Ta b l e X I I . D i s t r i b u t i o n Of S p e c i e s In The F Layer At S i t e A GROUP I ( a l l s i t e s ) P e n i c i 1 1 i u m verrucosum • v a r . e y e l o p i u m  Septonema c h a e t o s p i r a v a r C h a l a r a l o n g i p e s p i r n GROUP I I ( h i g h and low) Cl a d o s p o r i u m c l a d o s p o r i o i d e s GROUP I I I (middle and low) H e l i c o d e n d r o n t r i g l i t z i e n s i s  A r t i c u l o s p o r a t e t r a c l a d i a Trichoderma v i r i d e GROUP IV ( h i g h and midd l e ) C y l i n d r o c a r p o n tenue  C h a l a r a c o n s t r i c t a Mucor h i e m a l i s f. h i e m a l i s GROUP V (h i g h ) Pen i c i11ium n i g r i c a n s  P e n i c i l l i u m r a i s t r i c k i i  Tr ichoderma kon i n g i i  P e n i c i l l i u m brevi-compacturn  V e r t i c i l l i u m b u l b i l l o s u m G l i o c l a d i u m roseum GROUP VI (middle) C y l i n d r o c a r p o n didymum Sporidesmiurn flexum GROUP V I I (low) Septonema c h a e t o s p i r a P o l y s c y t a l u m fecundissimum  V a r i c o s p o r i u m elodeae L i b e r t e l l a sp. Tr ichoderma P e n i c i l l i u m polysporum notatum Hyalodendron 1 i g n i c o l a T O T A L # O F I S O I J A T E S H I G H M I D D L E L O W S U B S I T E S U B S I T E S U B S I T E 4 5 4 3 2 3 1 1 1 4 1 6 2 9 16 14 12 4 14 8 24 33 9 5 5 2 5 3 1 19 8 3 10 2 1 6 1 4 1 3 1 1 4 3 13 2 7 3 1 2. 22 1 5 9 12 2 5 1 4 1 1 2 3 44 Tab l e X I I I . D i s t r i b u t i o n Of S p e c i e s In The F Laye r At S i t e B • TOTAL #• OF I SOI J A T E S HIGH MIDDLE LOW SUBSITE SUBSITE SUBSITE GROUP I ( a l l s i t e s ) C y l i n d r o c a r p o n tenue 26 14 15 P e n i c i l l i u m verrucosum v a r . c y c l o p i u m 11 6 7 P e n i c i l l i u m brevi-compactum 3 6 4 P e n i c i l l i u m f r e q u e n t a n s 1 1 1 GROUP IV ( h i g h and m i d d l e ) G l i o c l a d i u m roseum 8 •6 2 Cl a d o s p o r i u m c l a d o s p o r i o i d e s 5 4 1 P e n i c i l l i u m n i g r i c a n s 3 3 F l a g e l l o s p o r a s t r i c t a 2 2 GROUP V (h i g h ) Mucor h i e m a l i s f. h i e m a l i s 4 2 1 P e n i c i l l i u m r a i s t r i c k i i 4 1 C h l o r i d i u m v i r e s c e n s v a r . chlamydosporum 3 GROUP VI (middle) Trichoderma polysporum 1 7 C h a l a r a c y l i n d r o s p e r m a 1 6 Acremonium s t r i c t u m 4 Selenophoma sp. 2 1 GROUP V I I (low) Trichoderma k o n i n q i i 4 9 26 Trichoderma v i r i d e 1 • 3 Septonema c h a e t o s p i r a 2 45 As can be obse r v e d , t h e r e were a g a i n some s p e c i e s i n the F l a y e r which were p r e s e n t o n l y a t S i t e A. These i n c l u d e d : A r t i c u l o s p o r a t e t r a c l a d i a  C h a l a r a c o n s t r i c t a  C h a l a r a l o n g i p e s  C y l i n d r o c a r p o n didymum  H e l i c o d e n d r o n t r i g l i t z i e n s i s  Hyalodendron 1ign i c o l a  L i b e r t e l l a sp. P e n i c i l l i u m notatum  P o l y s c y t a l u m f e c u n d i ss imum  Septonema c h a e t o s p i ra v a r . p i n i  Sporidesmium flexum  V a r i c o s p o r i u m elodeae V e r t i c i l l i u m b u l b i l l o s u m S p e c i e s p r e s e n t at S i t e B, but not S i t e A i n c l u d e d : Acremonium s t r i c t u m  C h a l a r a c y l i n d r o s p e r m a C h l o r idium v i r e s c e n s v a r . chlamydosporurn  F l a g e l l o s p o r a s t r i c t a  P e n i c i l l i u m f r e q u e n t a n s  Selenophoma sp. In the F l a y e r , t h e r e were fewer s p e c i e s common t o the same groups a t the two s i t e s , , than i n the L l a y e r . The o n l y s p e c i e s found c o n s i s t e n t l y i n Group I was P e n i c i l l i u m verrucosum v a r . c y c l o p i u m . C y l i n d r o c a r p o n tenue was the most f r e q u e n t l y i s o l a t e d s p e c i e s , but a t S i t e B i t o c c u r r e d more o f t e n i n the h i g h and mi d d l e s u b s i t e s , than i t d i d i n the low- s u b s i t e . P e n i c i l l i u m r a i s t r i c k i i was p r e s e n t i n Group V ( h i g h s u b s i t e ) ; Septonema c h a e t o s p i ra was found i n Group V I I (low s u b s i t e ) . In an o v e r a l l view of the d i s t r i b u t i o n of the s p e c i e s i s o l a t e d from the v a r i o u s s u b s i t e s , r e g a r d l e s s of which l a y e r 46 they were i s o l a t e d from, c e r t a i n s p e c i e s were always r e p r e s e n t a t i v e of p a r t i c u l a r groups, when they o c c u r r e d . The s p e c i e s a re as f o l l o w s : Group I ( a l l s u b s i t e s ) C y l i n d r o c a r p o n tenue Group I I I (middle and low) A r t i c u l o s p o r a t e t r a c l a d i a Group IV ( h i g h and middle) F l a g e l l o s p o r a s t r i c t a Mort i e r e l l a ramann iana v a r . ramann i a n a  P e n i c i l l i u m c i t r inum  Fusarium l a t e r i t i u m Group V (hi g h ) P e n i c i l l i u m n i g r i c a n s  P e n i c i l l i u m r a i s t r i c k i i  Mucor h i e m a l i s f. h i e m a l i s  G l i o c l a d i u m roseum C h l o r i d i u m v i r e s c e n s v a r . chlamydosporum  V e r t i c i l l i u m b u l b i 1 l o s u m  R a m i c h l o r i d i u m subulatum  Endophraqmia a l t e r n a t a . Group VI (middle) Spor idesmi um flexum  C h a l a r a c y l i n d r o s p e r m a  H e l i c o o n fuscosporum  T r i p o s p o r i u m e l e g a n s Group V I I (low) Septonema c h a e t o s p i r a V a r i c o s p o r i u m elodeae P e n i c i l l i u m notatum In examining the d i s t r i b u t i o n of the f u n g i i s o l a t e d a t S i t e A and S i t e B, i n c l u d i n g a l l s p e c i e s i s o l a t e d from the L l a y e r , F l a y e r and l i v i n g l e a v e s , some s p e c i e s tend t o be found e x c l u s i v e l y i n one or the o t h e r s i t e . These are summarized i n Tab l e XIV, and are l i s t e d i n d e c r e a s i n g o r d e r of freq u e n c y of o c c u r r e n c e . 47 T a b l e XIV. S p e c i e s R e s t r i c t e d To One Or The Other Main S i t e (see T a b l e s XXI and XXII f o r number of i s o l a t e s ) SITE A A r t i c u l o s p o r a t e t r a c l a d i a  C y l i n d r o c a r p o n didymum  V a r i c o s p o r i u m elodeae  Sporidesmium flexum  Pen i c i 1 1 i urn c i t r inum  H e l i c o o n fuscosporum  Tripospermum m y r t i  V e r t i c i l l i u m b u l b i l l o s u m  Pen i c i 1 1 i urn notatum  R a m i c h l o r i d i u m subulatum SITE B C h a l a r a c y 1 i n d r o s p e r m a  Septogloeum sp. C h l o r i d i u m v i r e s c e n s v a r . chlamydosporurn  M o r t i e r e l l a ramanniana v a r . ramanniana  Tr i p o s p o r ium e l e g a n s Fusarium l a t e r i t i u m 48 3. Comparison of S p e c i e s Found i n the L and F L a y e r s In the f o r e g o i n g d i s c u s s i o n s , the d i s t r i b u t i o n of s p e c i e s i n the L and F l a y e r s , between the d i f f e r e n t s u b s i t e s , have been t r e a t e d . These s u b s i t e s were s u b j e c t i v e l y d e f i n e d a r e a s i n the s i t e s t h a t were i n t e n d e d t o show what v a r i a t i o n s might e x i s t i n s p e c i e s c o m p o s i t i o n , w i t h s l i g h t changes i n d i s t a n c e from a stream. The L and F l a y e r s , as mentioned e a r l i e r , r e p r e s e n t d i f f e r e n t s t a g e s i n the decay of the l i t t e r , and thus a r e a more o b j e c t i v e l y d e f i n e d group. These two l a y e r s have many f u n g a l s p e c i e s i n common, but a l s o have some s p e c i e s which are found p r e d o m i n a n t l y i n one or the o t h e r l a y e r . T a b l e s XV and XVI l i s t the s p e c i e s , common t o both l a y e r s , found p r e d o m i n a n t l y i n the L l a y e r , and found p r e d o m i n a n t l y i n the F l a y e r , f o r S i t e A and S i t e B r e s p e c t i v e l y . In these T a b l e s , the s p e c i e s a r e l i s t e d i n d e c r e a s i n g f r e q u e n c y of o c c u r r e n c e . 49 T a b l e XV. Comparison Of S p e c i e s In The L And F L a y e r s At S i t e A S p e c i e s common t o both L and F l a y e r s C y l i n d r o c a r p o n tenue A r t i c u l o s p o r a t e t r a c l a d i a P e n i c i l l i u m verrucosum v a r . c y c l o p i u m C h a l a r a c o n s t r i c t a Sporidesmium flexum Septonema c h a e t o s p i ra v a r . p i n i L i b e r t e l l a sp. Tr ichoderma kon i ng i i P e n i c i l l i u m brevi-compactum Endophraqmia a l t e r n a t a S p e c i e s found p r e d o m i n a n t l y i n the L l a y e r H e l i c o d e n d r o n t r i g l i t z i e n s i s  P o l y s c y t a l u m fecundissimum  C l a d o s p o r ium c l a d o s p o r i o i d e s  Mucor h i e m a l i s f. h i e m a l i s  Selenophoma sp. P e s t a l o t i a monochaet i o i d e s  C h a l a r a l o n q i p e s  F l a g e l l o s . p o r a s t r i c t a  P e n i c i l l i u m c i t r inum  Acremonium s t r i c t u m  H e l i c o o n fuscosporum R a m i c h l o r i d i u m subulatum S p e c i e s found p r e d o m i n a n t l y i n the F l a y e r Pen i c i11ium n i g r i c a n s  Tr ichoderma v i r i d e  Septonema c h a e t o s p i ra  C y l i n d r o c a r p o n didymum  V a r i c o s p o r i u m elodeae  P e n i c i l l i u m r a i s t r i c k i i  Tr ichoderma polysporum  V e r t i c i l l i u m b u l b i l l o s u m  G l i o c l a d i u m roseum  P e n i c i l l i u m notatum  Hyalodendron l i q n i c o l a 50 Table XVI. Comparison Of S p e c i e s In The L And F L a y e r s At S i t e B S p e c i e s common t o both L and F l a y e r s C y l i n d r o c a r p o n tenue  Pen i c i 1 1 i um brevi-compactum  C l a d o s p o r i um c l a d o s p o r i o i d e s  F l a g e l l o s p o r a s t r i c t a C h l o r i d i u m v i r e s c e n s v a r . chlamydosporum  Septonema c h a e t o s p i ra Fusarium l a t e r i t i u m S p e c i e s found p r e d o m i n a n t l y i n the L l a y e r Selenophoma sp. P o l y s c y t a l u m fecundissimum Hyalodendron l i g n i c o l a C h a l a r a l o n g i p e s H e l i c o d e n d r o n t r i g l i t z i e n s i s Septonema c h a e t o s p i ra v a r . p i n i P e s t a l o t i a m o n o c h a e t i o i d e s M o r t i e r e l l a ramann i a n a v a r . ramann i a n a C h a l a r a c o n s t r i c t a L i b e r t e l l a sp. T r i p o s p o r i u m e l e g a n s Septoqloeum sp. S p e c i e s found p r e d o m i n a n t l y i n the F l a y e r Tr ichoderma k o n i n g i i P e n i c i l l i u m verrucosum v a r . c y c l o p i u m G l i o c l a d i u m roseum Tr ichoderma polysporum C h a l a r a c y 1 i n d r o s p e r m a Mucor h i e m a l i s f. h i e m a l i s P e n i c i l l i u m n i g r i c a n s P e n i c i l l i u m r a i s t r i c k i i Acremonium s t r i c t u r n Tr ichoderma v i r i d e Pen i c i 1 1 i um f requentans 51 4. S e a s o n a l i t y of Fungi i n the L and F L a y e r s In the c o u r s e of t h i s s t u d y , samples were taken a t v a r i o u s i n t e r v a l s t hroughout the y e a r , and the f u n g i p r e s e n t were i s o l a t e d and i d e n t i f i e d . The o c c u r r e n c e of these s p e c i e s a t the sample d a t e s ( i . e . , number of times i s o l a t e d ) , r e f l e c t e d a p a t t e r n of d i s t r i b u t i o n t h roughout the y e a r . Of c o u r s e , the s e a s o n a l d i s t r i b u t i o n of s p e c i e s i s not a b s o l u t e l y d e f i n e d s i n c e c o n d i t i o n s v a r y from year t o y e a r , but n e v e r t h e l e s s g e n e r a l t r e n d s of o c c u r r e n c e can be d i s c e r n e d . The times of the year when samples were taken and the c o r r e s p o n d i n g c l i m a t i c c o n d i t i o n s a r e summarized f o r S i t e A i n Table I and f o r S i t e B i n T a ble I I . In the f o l l o w i n g d i s c u s s i o n s on s e a s o n a l i t y , the mention of a s p e c i e s as o c c u r r i n g d u r i n g a c e r t a i n time of year s i m p l y i n d i c a t e s t h a t i t s r e l a t i v e f r e q u e n c y i s h i g h e r a t t h a t time of the y e a r . I t may have o c c u r r e d a t o t h e r t i m e s , but a t a much lower f r e q u e n c y . Some s p e c i e s showed l i t t l e s e a s o n a l f l u c t u a t i o n and were thus l a b e l l e d n o n s e a s o n a l . V a r i o u s s e a s o n a l p a t t e r n s were e x h i b i t e d by the r e m a i n i n g s p e c i e s , depending on the s i t e or l a y e r examined. The most c l e a r l y d i s t i n g u i s h e d groups were those o c c u r r i n g i n the s p r i n g and summer, summer, and f a l l and w i n t e r . The s e a s o n a l d i s t r i b u t i o n of s p e c i e s throughout the sam p l i n g time i s shown i n T a b l e s XVII and X V I I I , f o r i s o l a t e s from the L l a y e r a t S i t e A and S i t e B, r e s p e c t i v e l y . I t s h o u l d be noted t h a t the samples from S i t e B were not made a t the same time as those i n S i t e A, so comparison between the two s i t e s i s 52 s l i g h t l y d i f f e r e n t . The same i n f o r m a t i o n i s g i v e n i n T a b l e s XIX and XX f o r the i s o l a t e s from the F l a y e r . 53 Table XVII S e a s o n a l D i s t r i b u t i o n Of S i t e A Sp e c i e s In The L Layer At NONSEASONAL C y l i n d r o c a r p o n tenue  P o l y s c y t a l u m fecundissimum  V a r i c o s p o r i u m elodeae SPRING & SUMMER H e l i c o d e n d r o n t r i g l i t z i e n s i s  A r t i c u l o s p o r a t e t r a c l a d i a  C l a d o s p o r i u m c l a d o s p o r i o i d e s  Septonema c h a e t o s p i r a  Sporidesmium flexum  C h a l a r a l o n g i p e s L i b e r t e l l a sp. F l a g e l l o s p o r a s t r i c t a  Septonema c h a e t o s p i r a v a r . p i n i Acremonium s t r i c t u m SUMMER Tr ichoderma  P e n i c i11ium P e n i c i11ium v i r i d e  n i g r i c a n s r a i s t r i c k i i C y l i n d r o c a r p o n didymum FALL & WINTER Pen i c i 1 1 i um verrucosum v a r . e y e l o p i u m  Mucor h i e m a l i s f. h i e m a l i s  Selenophoma sp. P e s t a l o t i a m o n o c h a e t i o i d e s  Tr ichoderma k o n i n q i i  Pen i c i 1 1 ium c i t r i n u m  H e l i c o o n fuscosporum  Pen i c i 1 1 i um brevi-compactum SPRING, SUMMER & WINTER C h a l a r a c o n s t r i c t a  R a m i c h l o r i d i u m subulatum  Endophragmia a l t e r n a t a TOTAI J # OI " ISOLATES i APR JUL NOV FEB MAY JUL 6 19 13 9 23 3 10 5 5 1 4 1 1 3 2 2 16 12 2 4 7 1 4 9 3 3 2 1 4 7 3 1 2 1 3 5 1 2 2 2 5 2 4 1 2 5 1 1 3 3 4 1 1 1 2 2 9 2 1 5 1 6 2 3 3 1 3 1 2 1 2 2 7 3 2 1 1 5 1 5 2 3 6 1 4 1 2 4 1 2 2 2 3 3 1 3 4 4 4 1 4 1 2 2 1 1 1 54 T a b l e X V I I I . S e a s o n a l D i s t r i b u t i o n Of S p e c i e s In The L Layer At S i t e B NONSEASONAL C y l i n d r o c a r p o n tenue C l a d o s p o r i u m c l a d o s p o r i o i d e s Pen i c i11ium verrucosum v a r . c y c l o p i u m Septonema c h a e t o s p i r a v a r . p i n i L i b e r t e l l a sp. SPRING C h l o r i d i u m v i r e s c e n s v a r . chlamydosporum SPRING & FALL Tr ichoderma k o n i n g i i P o l y s c y t a l u m f e c u n d i ssimum G l i o c l a d i u m roseum F l a g e l l o s p o r a s t r i c t a Septonema c h a e t o s p i r a Tr ichoderma polysporum Fus a r i u m l a t e r i t i u m FALL P e n i c i l l i u m b r e v i - compactum Hyalodendron l i g n i c o l a  C h a l a r a l o n g i p e s  H e l i c o d e n d r o n t r i g l i t z i e n s i s  Mort i e r e l l a ramanniana v a r . ramann i a n a  C h a l a r a c o n s t r i c t a  T r i p o s p o r i u m e l e g a n s Mucor h i e m a l i s f. h i e m a l i s WINTER P e s t a l o t i a m o n o c h a e t i o i d e s SPRING & WINTER Selenophoma sp. Septogloeum sp. TOTAL # OF ISOLATES MAY 3 2 1 10 2 3 2 1 2 1 2 1 1 1 8 1 SEP 19 1 2 3 1 9 6 2 1 1 1 1 13 8 8 5 4 2 3 2 JAN 17 3 1 2 2 5 2 1 1 1 13 1 APR 22 2 3 2 2 6 1 1 1 1 16 1 55 T a b l e XIX. Se a s o n a l D i s t r i b u t i o n Of S p e c i e s In The F Layer At S i t e A NONSEASONAL C y l i n d r o c a r p o n tenue  Mucor h i e m a l i s f. h i e m a l i s L i b e r t e l l a sp. SPRING H e l i c o d e n d r o n t r i g l i t z i e n s i s  P o l y s c y t a l u m fecundissimum  C h a l a r a c o n s t r i c t a  Sporidesmium flexum  C h a l a r a l o n q i p e s  Hyalodendron 1 i g n i c o l a SUMMER Pen i c i11ium n i g r i c a n s  A r t i c u l o s p o r a t e t r a c l a d i a  Trichoderma v i r i d e  C y l i n d r o c a r p o n didymum  P e n i c i l l i u m r a i s t r i c k i i  V a r i c o s p o r i u m elodeae  Tr ichoderma polysporum  V e r t i c i l l i u m b u l b i 1 l o s u m G l i o c l a d i u m roseum FALL & WINTER Pen i c i11ium verrucosum v a r . c y c l o p i u m  Tr ichoderma k o n i n g i i P e n i c i l l i u m notatum SPRING, SUMMER & FALL Septonema c h a e t o s p i r a C l a d o s p o r ium c l a d o s p o r i o i d e s S. c h a e t o s p i ra v a r . p i n i  Pen i c i 1 1 i um brevi-compactum TOTAI ; # OE " I SOL; VTES APR J U L NOV FEB MAY JUL 16 7 13 9 12 9 2 2 2 2 1 1 1 2 1 2 11 3 2 4 5 2 10 2 3 11 1 1 7 2 2 1 3 3 6 3 4 5 9 3 14 2 3 1 3 10 2 8 6 9 3 3 2 1 5 3 5 2 5 1 1 1 2 5 2 3 2 1 1 2 2 4 4 1 1 6 1 2 1 5 7 5 6 2 1 3 4 3 2 1 3 1 1 2 3 56 Table XX. Se a s o n a l D i s t r i b u t i o n Of S p e c i e s In The F Layer At S i t e B NONSEASONAL C y l i n d r o c a r p o n tenue Tr ichoderma k o n i n g i i G l i o c l a d i u m roseum Cladosporiurn c l a d o s p o r i o i d e s FALL & SPRING Tr ichoderma polysporum F l a g e l l o s p o r a s t r i c t a WINTER & SPRING P e n i c i l l i u m verrucosum v a r . c y c l o p i u m SPRING Mucor h i e m a l i s f. h i e m a l i s Ac remoni urn s t r i c t urn  Selenophoma sp. FALL Pen i c i 1 1 i urn  C h a l a r a c y l i n d r o s p e r m a P e n i c i l l i u m r a i s t r i c k i i brevi-compacturn FALL & WINTER Pen i c i 1 1 i urn n i g r i c a n s  Tr ichoderma v i r i d e  P e n i c i l l i u m f r e q u e n t a n s WINTER C h l o r i d i u m v i r e s c e n s v a r . chlamydosporum  Septonema c h a e t o s p i r a TOTAL # OF ISOLATES MAY 2 18 4 1 2 1 SEP 14 10 3 2 2 2 11 4 4 2 1 2 JAN 21 10 7 4 10 4 2 1 2 2 57 In the p r e c e d i n g t r e a t m e n t of the s e a s o n a l i t y of the s p e c i e s found i n the L and F l a y e r s , o b s e r v a t i o n s were made a t two major s i t e s , d e s i g n a t e d A and B. These s i t e s were not i d e n t i c a l w i t h r e s p e c t t o the s p e c i e s p r e s e n t or t h e i r d i s t r i b u t i o n . F u r t h e r m o r e , the number of samples a t each s i t e was not the same. D e s p i t e these d i f f e r e n c e s , some statements can be made on the s e a s o n a l i t y of the s p e c i e s found i n the two l a y e r s , based on t h e i r o c c u r r e n c e s i n both s i t e s . Of a l l the s p e c i e s o c c u r r i n g i n the L l a y e r , o n l y one showed l i t t l e or no s e a s o n a l v a r i a t i o n and t h a t was C y l i n d r o c a r p o n tenue. The r e s t of the s p e c i e s c o u l d be p l a c e d i n t h r e e groups, s p r i n g and summer, summer, and f a l l and w i n t e r . The s p r i n g and summer group i n c l u d e d ( i n d e c r e a s i n g o r d e r of f r e q u e n c y ) : A r t i c u l o s p o r a t e t r a c l a d i a  Septonema c h a e t o s p i r a  F l a g e l l o s p o r a s t r i c t a  Acremoniurn s t r i c t u m  Sporidesmium flexum C h l o r i d i u m v i r e s c e n s v a r . chlamydosporum The summer group c o m p r i s e d : Tr ichoderma v i r i d e  Pen i c i 1 1 i urn n i g r i c a n s P. r a i s t r i c k i i  C y l i n d r o c a r p o n didymum 58 The f a l l and w i n t e r group was c h a r a c t e r i z e d by: Mucor h i e m a l i s f. h i e m a l i s  P e s t a l o t i a m o n o c h a e t i o i d e s  P e n i c i l l i u m brevi-compactum  Hyalodendron 1 i g n i c o l a Mort i e r e l l a ramann i a n a v a r . ramann i a n a  T r i p o s p o r i u m e l e q a n s  C h a l a r a c o n s t r i c t a P e n i c i l l i u m c i t r i n u m The F l a y e r c o n t a i n e d a l s o C y l i n d r o c a r p o n tenue as the o n l y s p e c i e s which showed l i t t l e s e a s o n a l v a r i a t i o n . The groups a t t h i s l a y e r were s p r i n g , summer, and f a l l and w i n t e r . The s p e c i e s found i n the s p r i n g were: H e l i c o d e n d r o n t r i g l i t z i e n s i s  P o l y s c y t a l u m f e c u n d i ssimum  C h a l a r a c o n s t r i c t a  Sporidesmium f l e x u m  Hyalodendron l i g n i c o l a The summer group i n c l u d e d : A r t i c u l o s p o r a t e t r a c l a d i a  C y l i n d r o c a r p o n didymum  V a r i c o s p o r i u m elodeae V e r t i c i 1 1 i u m b u l b i l l o s u m The s p e c i e s p r e s e n t i n the f a l l and w i n t e r i n c l u d e d : P e n i c i l l i u m verrucosum v a r . c y c l o p i u m P. notatum P. f requentans C h a l a r a c y l i n d r o s p e r m a The c o m p l e x i t y of the i n t e r a c t i o n between s p e c i e s i n v o l v e d i n the decay p r o c e s s i s such t h a t d e f i n i t e r o l e s or a s s o c i a t i o n s among s p e c i e s a r e hard t o e l u c i d a t e . In the f o r e g o i n g s e c t i o n s , an attempt has been made t o p o i n t out g e n e r a l i z a t i o n s of s p e c i e s 59 o c c u r r e n c e w i t h r e s p e c t t o p o s i t i o n w i t h i n the s i t e s , sequence i n decay and s e a s o n a l i t y . The s p e c i e s i s o l a t e d have been grouped i n each of these i n s t a n c e s , but an o v e r a l l i n d i c a t i o n of t h e i r i n v o l v m e n t i n the decay has been hard t o i l l u s t r a t e . The f o l l o w i n g s e c t i o n d e a l s w i t h the s p e c i e s as i n d i v i d u a l s , d i s c u s s i n g t h e i r d i s t r i b u t i o n s and s e a s o n a l i t y . 60 5. Fu n g a l S p e c i e s I s o l a t e d T h e . p r e c e d i n g d i s c u s s i o n s have been concerned w i t h the d i s t r i b u t i o n and s e a s o n a l i t y of the s p e c i e s i s o l a t e d , w i t h r e s p e c t t o the s i t e s and s u b s i t e s . The r e s u l t has been a t e n t a t i v e g r o u p i n g of c e r t a i n s p e c i e s t o g e t h e r , r e p r e s e n t i n g d i f f e r e n t s i t e s , s u b s i t e s or seasons. In the f o l l o w i n g s e c t i o n the i n d i v i d u a l s p e c i e s a re t r e a t e d i n o r d e r t o p o i n t out d i s t i n c t i v e f e a t u r e s , and comment on t h e i r d i s t r i b u t i o n . Ac remon i urn s t r icturn Gams. C e p h a l o s p o r i u m - a r t i g e S c h i m m e l p i l z e 42. 1971. C o l o n i e s w h i t e t o p i n k i s h , u s u a l l y w i t h l i t t l e a e r i a l mycelium; p h i a l i d e s o f t e n i n groups of t h r e e or produced l a t e r a l l y on f u n i c u l o s e s t r a n d s , 27.0 - 46.8 x 1.8 urn; spores i r r e g u l a r t o somewhat c u r v e d , 3.6 - 7.2 x 1.2 - 3.0 urn. A. s t r i c t u r n was i s o l a t e d most o f t e n from l e a v e s i n m o i s t chambers; i t was found r e l a t i v e l y i n f r e q u e n t l y ( F i g . 3 ) . I t was absent from the low s u b s i t e i n both S i t e A and B, and was found o n l y s p o r a d i c a l l y i n the L and F l a y e r s . T h i s i s a common s o i l fungus t h a t i s a l s o found on o r g a n i c d e b r i s (Gams, 1971; Domsch and Gams, 1972). 61 F i g u r e 3. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of Acremonium s t r i c t u m = L L a y e r = F L a y e r One I s o l a t e = 2.5mm V e r t i c a l D i s t a n c e X i I? AP -1—I h JU -I—I h ND -\—I—I h ±977 FE MY 197B -\ 1 1 JU SAMPLE DATE M X i CD UJ CD 1977 137B 5AMPLE DATE 62 Acrodontium c r a t e r i f o r m e (Van Beyma)de Hoog. S t u d . M y c o l . 1:26. 1972. C o l o n i e s c o t t o n y t o f l o c c o s e , g r a y ; c o n i d i o p h o r e s u s u a l l y f u n i c u l o s e , c o n i d i o g e n o u s c e l l s s y m p o d i a l , v e r y t h i n w i t h minute d e n t i c l e s , c o n i d i a 3.6 x 1.8 urn. The c u l t u r e s f i t the d e s c r i p t i o n g i v e n by de Hoog (1972). A. c r a t e r i f o r m e was i s o l a t e d i n f r e q u e n t l y from e i t h e r s i t e , i t s l a r g e s t o c c u r r e n c e b e i n g on l i v i n g l e a v e s a t the s p r i n g sample from S i t e A. I t was found i n the L l a y e r a t the summer sample, p r o b a b l y from l e a v e s which were s t i l l on the t r e e a t the p r e v i o u s sample. T h i s s p e c i e s was prominent o n l y d u r i n g the s p r i n g and summer ( F i g . 4 ) . De Hoog (1972) r e p o r t e d A. c r a t e r i forme from v a r i o u s s u b s t r a t a i n c l u d i n g l e a v e s of numerous p l a n t s ; one r e p o r t was from l i v i n g l e a v e s of F r a x i n u s  e x c e l s i o r . Yokoyama et a l . (1977) a l s o r e p o r t e d t h i s s p e c i e s on s t e r i l i z e d l e a v e s embedded i n l i t t e r . F i g u r e 4. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of Acrodontium c r a t e r i forme For Legend See F i g . 3 I < 5 I < I—I—I—I—I—I—I—I—h A P J U N O 1 9 7 7 H—I 1 1 1 1 1 F E M Y J U 1 9 7 B SAMPLE DATE 5 n I ! I CD ' 1 ' I ' ' I 1 1 I I 1 I M Y S E J A A P 1 9 7 7 1 9 7 B SAMPLE DATE 64 A r t i c u l o s p o r a t e t r a c l a d i a Ing. Trans. B r i t . M y c o l . Soc. 25: 376. 1942. In c u l t u r e , c o l o n i e s are o f t e n p i n k i s h , and appear wet on the s u r f a c e , sometimes p r o d u c i n g s m a l l rounded s c l e r o t i u m - 1 i k e s t r u c t u r e s of unknown f u n c t i o n . T h i s s p e c i e s i s e a s i l y r e c o g n i z a b l e by i t s d i s t i n c t t e t r a r a d i a t e s p o r e s , which are as l o n g as 75 urn. A. t e t r a c l a d i a was found p r e d o m i n a n t l y at S i t e A. I t o c c u r r e d o n l y once i n S i t e B. I t was i s o l a t e d m o s t l y from the lower and m i d d l e s u b s i t e s , i n both the L and F l a y e r s , but p r e d o m i n a n t l y the F l a y e r . I t o c c u r r e d i n the g r e a t e s t numbers at the f i r s t summer sample ( F i g . 5 ) . Temperature may be more im p o r t a n t than p r e c i p i t a t i o n i n i t s s e a s o n a l i t y , s i n c e l a t e r samples had more r a i n but lower t e m p e r a t u r e s , and the frequency was down. In c u l t u r e , c o l o n y growth and s p o r u l a t i o n d e c rease a t lower (5° C) and h i g h e r (25° C) t e m p e r a t u r e s (Koske and Duncan, 1974). M o i s t u r e must s t i l l be i m p o r t a n t s i n c e i t s o c c u r r e n c e d i d not i n c r e a s e a g a i n a t the second summer sample when the temperature r o s e , p r o b a b l y because t h e r e had been no r a i n f o r two weeks. A. t e t r a c l a d i a i s a member of the a q u a t i c hyphomycetes, an a r t i f i c i a l group u s u a l l y found growing on d e b r i s i n streams. I t has o f t e n been r e p o r t e d from t e r r e s t r i a l s i t u a t i o n s (Bandoni, 1972, 1977; Sanders and Webster, 1978; Webster, 1977). I t s o c c u r r e n c e i n the F l a y e r i n d i c a t e s i t s a c t i v e r o l e as a secondary c o l o n i z e r i n the decay of the t e r r e s t r i a l l i t t e r . 65 F i g u r e 5. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of A r t i c u i o s p o r a t e t r a c l a d i a For Legend See F i g . 3 5 M ! I < 2 I < A P J U -I—| 1 1 1—|—j-ND F E 1 9 7 7 M Y 1 9 7 8 -\ 1 1 J U SAMPLE DATE U i CD I—I—I—I—I—I—I—I 1—I—I—I—I M Y S E J A A P 1 9 7 7 1 9 7 B SAMPLE DATE 66 C h a l a r a c o n s t r i c t a Nag Raj and K e n d r i c k . A Monograph of C h a l a r a and A l l i e d Genera. 103. 1975. C u l t u r e s v e r y slow growing, w h i t e t o g r a y , sometimes w i t h b l a c k p a t c h e s ; c o n i d i o p h o r e s u s u a l l y f u n i c u l o s e ; p h i a l i d e s o f t e n p r o l i f e r a t i n g , sometimes s l i g h t l y c u r v e d ; spores h y a l i n e , c l a v a t e , s m a l l , n o n - s e p t a t e , 3.0 - 7.2 (10.8) x 1.0 - 1.8 urn. T h i s s p e c i e s was d e s c r i b e d by Nag Raj and K e n d r i c k (1975) from l e a v e s of A g a t h i s a u s t r a l i s from New Zealand. The p r e s e n t i s o l a t e s d i f f e r from the o r i g i n a l d e s c r i p t i o n i n t h a t the upper range of the spores i s s l i g h t l y h i g h e r i n some c u l t u r e s . F u r t h e r m o r e , i n some i s o l a t e s the c o n i d i o p h o r e s a re more than 4 c e l l e d . These d i f f e r e n c e s do not warrant e x c l u s i o n from t h i s spec i e s . In S i t e A, C. c o n s t r i c t a o c c u r r e d most f r e q u e n t l y i n the f i r s t s p r i n g sample, where i t o c c u r r e d i n a l l t h r e e s u b s i t e s , m o s t l y i n the F l a y e r . The frequ e n c y d i m i n i s h e d i n subsequent samples when i t o c c u r r e d o n l y i n the m i d d l e and h i g h s u b s i t e s , i n the L l a y e r . I t o c c u r r e d i n f r e q u e n t l y a t S i t e B ( F i g . 6 ) . T h i s s p e c i e s was i s o l a t e d almost e n t i r e l y from l e a v e s i n mo i s t chambers. I t o c c u r r e d i n the L l a y e r d u r i n g summer through w i n t e r and i n the F l a y e r d u r i n g the s p r i n g . I t d i d not occur a t the f i n a l summer sample when the temp e r a t u r e was the h i g h e s t and the r a i n f a l l the l o w e s t . Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of C h a l a r a c o n s t r i c t a For Legend See F i g . 3 1 9 7 7 1 9 7 8 SAMPLE DATE ' I I—I—I—I—I—I—I—I—I—I—l MY SE JA AP 1 9 7 7 1 9 7 8 SAMPLE DATE 6 8 C h a l a r a c y l indrosperma (Cda. )Hughes. Can. J . Bot. 3_6:747. 1958. C o l o n i e s dark gray t o b l a c k , r e s t r i c t e d ; c o n i d i o p h o r e s f u n i c u l o s e ; p h i a l i d e s w i t h a l o n g c o l l a r e t t e , 18.0 - 23.4 um; c o n i d i a c y l i n d r i c a l , h y a l i n e , 7.2 - 14.4 x 1.8 um. Nag Raj and K e n d r i c k (1975) have r e p o r t e d t h i s s p e c i e s from v a r i o u s angiosperm l e a v e s and wood. C. c y l i n d r o s p e r m a was r e s t r i c t e d m a i n l y t o S i t e B, o c c u r r i n g most f r e q u e n t l y d u r i n g e a r l y f a l l i n the F l a y e r ( F i g . 7 ) . C h a l a r a l o n g i p e s (Pr.)Cooke. G r e v i l l e a 10:50. 1881. C o l o n i e s r e s t r i c t e d , dark gray t o b l a c k ; c o n i d i o p h o r e s f u n i c u l o s e , dark; c o n i d i a c y l i n d r i c a l , 3.0 - 7.2 x 1.0 - 1.8 um. The type i s on p i n e n e e d l e s from Germany. The p r e s e n t i s o l a t e s d i f f e r from the o r i g i n a l d e s c r i p t i o n i n t h a t the spores are s l i g h t l y l a r g e r . In S i t e A, C. l o n g i p e s o c c u r r e d s p o r a d i c a l l y i n most l a y e r s and s u b s i t e s , w i t h the h i g h e s t o c c u r r e n c e i n the low s u b s i t e , i n the f i r s t s p r i n g sample ( F i g . 8 ) . T h i s s p e c i e s was most prominent i n the e a r l y f a l l a t S i t e B, i n the L l a y e r . O v e r a l l , C. l o n g i p e s was n o t i c e a b l y absent d u r i n g the summers and t h e r e f o r e may not be a c t i v e a t ti m e s of h i g h t emperature and low p r e c i p i t a t i o n . None of the p r e c e d i n g s p e c i e s of C h a l a r a have been r e p o r t e d i n s t u d i e s on f u n g i from washed c o n i f e r l i t t e r . 69 F i g u r e 7. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of C h a l a r a c y l i n d r o s p e r m a For Legend See F i g . 3 5 M I I < I 1 1 h AP JU H 1—+• 1977 I— I 1—I—1 1—1 I—I—r ND FE MY JU 197B SAMPLE: DATE M I I CD i CD i CD MY -I—I—I 1—I—I—I 1—I—I—I—I SE 1977 JA AP 1978 SAMPLE DATE 70 F i g u r e 8. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of C h a l a r a l o n g i p e s For Legend See F i g . 3 5 M X I < IS ! i i < l—l—I—I—I—I—I—I—I—I—I—I—I—I—i—i—| AP JU M3 FE MY 1977 1978 SAJVFLE DATE I i CD I I »—I—'—I—I—I 1—I—I—I—I MY SE JA AP 1977 1378 SAM=LE DATE 71 C h l o r i d i u m v i r e s c e n s ( P e r s . ex Pers.)W. Gams & H o l . - J e c h . v a r . chlamydosporum (Van Beyma)W. Gams & H o l . - J e c h . S t u d . M y c o l . 13:21. 1976. Colony growth r e s t r i c t e d , p r o d u c i n g abundant, brown chlamydospores, c o n i d i a 3.6 - 5.4 x 1.8 - 2.2 urn. T h i s s p e c i e s was i s o l a t e d r a r e l y , and o n l y from the h i g h s u b s i t e a t S i t e B. I t was prominent i n the L l a y e r d u r i n g the s p r i n g , and the F l a y e r d u r i n g the w i n t e r ( F i g . 9 ) . C. v i r e s c e n s v a r . chlamydosporum has been r e p o r t e d p r e v i o u s l y from v a r i o u s s o i l s and l i t t e r s (Domsch and Gams, 1972; Gams and Holubova-Jechova, 1976). C l a d o s p o r i u m c l a d o s p o r i o i d e s ( F r e s e n . ) d e V r i e s . C o n t r i b u t i o n t o the Knowledge of the Genus C l a d o s p o r ium L i n k ex F r . 57. 1952. T h i s s p e c i e s i s e a s i l y i d e n t i f i e d by the smooth, non-s e p t a t e , lemon shaped b l a s t o s p o r e s . I t was i s o l a t e d f r e q u e n t l y from both S i t e A and S i t e B. I t o c c u r r e d i n a l l s u b s i t e s , i n c l u d i n g the l i v i n g l e a v e s , i n both L and F l a y e r s . B e s i d e s b e i n g one of the most common s o i l and l i t t e r f u n g i , t h i s s p e c i e s has a l s o been found r e p e a t e d l y on the s u r f a c e s of l i v i n g l e a v e s (Baker et a l . , 1979; G o u r b i e r e , 1975; C o l l i n s and Hayes, 1976). C. c l a d o s p o r i o i d e s o c c u r r e d l a r g e l y d u r i n g the s p r i n g and summer at S i t e A, and d u r i n g the s p r i n g and w i n t e r a t S i t e B ( F i g . 1 0 ) . 72 F i g u r e 9. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of C h l o r i d i u m v i r e s c e n s v a r . chlamydosporum For Legend See F i g . 3 CD I—I—I—I—I—I—I—I 1—I—I—I—I M V SE J A A P 1 9 7 7 1 9 7 B SAMPLE DATE F i g u r e 10. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of Cladosporium c l a d o s p o r i o i d e s For Legend See F i g . 3 S > SAMPLE DATE 74 C y l i n d r o c a r p o n didymum ( H o r t i g ) W o l l e n w . Fus. A u t o g r . D e l . ed. 2. 650. 1926. C o l o n i e s s p r e a d i n g r e l a t i v e l y r a p i d l y , w h i t e to o f f - w h i t e , sparse a e r i a l hyphae, u s u a l l y somewhat s l i m y ; c o n i d i a produced i n s l i m y masses, 0 - 3 s e p t a t e , but u s u a l l y 1 - 2, 18.0 - 35.0 x 3.6 - 6.0 um; chlamydospores p r e s e n t , brown. T h i s s p e c i e s i s e a s i l y r e c o g n i z e d by the wide spores w i t h somewhat non-s y m m e t r i c a l s e p t a . Sometimes t h e r e were v e r y s m a l l spores p r e s e n t , 5.4 - 7.2 x 1.0 - 1.8 um. These may be m i c r o c o n i d i a , or what Booth (1966) c a l l e d p r i m a r y c o n i d i a . C. didymum was i s o l a t e d o n l y from S i t e A, m a i n l y from the F l a y e r . I t o c c u r r e d i n a l l t h r e e s u b s i t e s , most f r e q u e n t l y o c c u r r i n g d u r i n g the summer i n the m i d d l e s u b s i t e ( F i g . 1 1 ) . 75 F i g u r e 11. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of C y l i n d r o c a r p o n didymum For Legend See F i g . 3 76 C y l i n d r o c a r p o n tenue Bugn. E n c y c l . M y c o l . 1_1:175-178. 1939. C o l o n i e s r a p i d l y growing, a e r i a l hyphae c o t t o n y , w h i t e t o orange-brown, the l a t t e r c o l o r predominant i n o l d e r c u l t u r e s ; c o n i d i a 0 - 1 s e p t a t e , sometimes 2 - 3 , 10.8 - 32.4 x 1.8 -3.6 um; chlamydospores p r e s e n t , brown, u s u a l l y i n i n t e r c a l a r y c h a i n s . These i s o l a t e s f i t the d e s c r i p t i o n of the s p e c i e s g i v e n by Booth (1966), but d i f f e r i n t h a t the spore s i z e range i s l a r g e r than t h a t d e s c r i b e d . There i s no o t h e r s p e c i e s w i t h t h i s c o m b i n a t i o n of narrow spores and chlamydospores p r e s e n t . Sometimes s m a l l e r c o n i d i a are p r e s e n t , but the s e may j u s t be immature t y p e s . C. tenue was the most f r e q u e n t l y i s o l a t e d of a l l the f u n g i p r e s e n t . In S i t e A i t was p r e s e n t a t a l l sample d a t e s , more o f t e n i n the L l a y e r , but a l s o i n the F l a y e r . I t seemed t o occur under d r i e r c o n d i t i o n s , but not e x t r e m e l y d r y . T h i s i s shown by the f a c t t h a t i t was p r e s e n t i n the middl e and low s u b s i t e s , but not the h i g h , when temperature was h i g h and r a i n f a l l low. As the temperature f e l l and r a i n f a l l i n c r e a s e d , i t d e c l i n e d i n the low and mi d d l e s u b s i t e s , and rose i n abundance i n the h i g h s u b s i t e . C o n v e r s e l y , when r a i n f a l l dropped d r a s t i c a l l y d u r i n g the l a s t summer sample at S i t e A, i t was much l e s s abundant ( F i g . 1 2 ) v In S i t e B i t had much the same p a t t e r n except t h a t i t seemed t o be more e v e n l y d i s t r i b u t e d among the samples ( F i g . 1 3 ) . F i g u r e 12. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of C y l i n d r o c a r p o n tenue - - S i t e A For Legend See F i g . 3 1 9 7 7 1 9 7 8 SAMPLE DATE 78 F i g u r e 13. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of C y l i n d r o c a r p o n tenue - - S i t e B For Legend See F i g . 3 5 n I I 1—I 1 i — I 1—I 1 — I — I 1—I MY SE J A AP 1977 197B SAMPLE DATE 79 Endophragmia a l t e r n a t a Tubaki and S a i t o . T r a n s . B r i t . M y c o l . Soc. 52:477. 1969. C o l o n i e s slow growing, mycelium b l a c k ; c o n i d i o p h o r e s d a r k , s h o r t , p r o d u c i n g o n l y one or two d a r k , s e p t a t e c o n i d i a . These i s o l a t e s resemble t h a t d e s c r i b e d by Tubaki and S a i t o (1969). E. a l t e r n a t a was an e x t r e m e l y slow growing s p e c i e s i n c u l t u r e , and s u b s e q u e n t l y produced spores i n f r e q u e n t l y . Matsushima (1975) i n d i c a t e d t h i s s p e c i e s t o be i n f e r t i l e i n c u l t u r e . Some of the c u l t u r e s d e s i g n a t e d as s t e r i l e dark m y c e l i a , i n t h i s s t u d y , c o u l d have been i s o l a t e s of E. a l t e r n a t a t h a t had not s p o r u l a t e d . T h i s s p e c i e s was i s o l a t e d r a r e l y , but o c c u r r e d i n a l l s u b s i t e s a t S i t e A and i n the h i g h and m i d d l e s u b s i t e s a t S i t e B ( F i g . 1 4 ) . Tubaki and S a i t o (1969) c o n s i d e r e d t h i s s p e c i e s an i m p o r t a n t e x t e r n a l c o l o n i z e r of P i n u s d e n s i f l o r a l i t t e r . Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of Endophraqmia a l t e r n a t a For Legend See F i g . 3 I 1 1—I 1 1—I 1 1 1—I—I 1 1 1 1 1 AP JU tO FE MV JU 1977 1976 SAMPLE DATE h—I—I—I—I—I—I—I—I—I—I—I—I MY SE JA AP 1977 1978 SAMPLE DATE 81 F l a q e l l o s p o r a s t r i c t a S. N i l s s . Bot. Not. 115:82. 1962. C o l o n i e s w i t h l i m i t e d a e r i a l mycelium, p r o d u c i n g masses of s l i m y s p o r e s ; c o n i d i a s t r a i g h t , 0 - 1 s e p t a t e , t a p e r e d a t both ends, 36.0 - 43.2 x 1.3 - 2.0 urn. These i s o l a t e s d i f f e r from the o r i g i n a l d e s c r i p t i o n i n sometimes h a v i n g a s e p t a t e s p o r e . F. s t r i c t a was found i n b o t h S i t e A and S i t e B, at the e a r l i e r s a m p l i n g d a t e s , s p r i n g and summer. I t o c c u r r e d more o f t e n i n the h i g h and middl e s u b s i t e s ; c o n f i n e d t o the L l a y e r i n S i t e A, i n both l a y e r s a t S i t e B ( F i g . 1 5 ) . I t was not i s o l a t e d f r e q u e n t l y , and was r a r e l y found on l e a v e s i n moist chambers. F i g u r e 15. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s . O f F l a g e l l o s p o r a s t r i c t a For Legend See F i g . 3 .5 X i < LI i < t < 5 X i CD h-H—I—I—I—I—I—I—I 1—I—I—I—|—|—|—| AP JU NO FE MV JU 1977 197B SAMPLE DATE 2 i CD I—I—I—I 1—I—I—I 1—I—I—|—| MY SE JA AP 1977 1978 SAMPLE DATE 83 F usarium l a t e r i t i u m Nees. S y s t . P i l z e Schwamme. 31. 1817. C u l t u r e s ranged from s l i m y and orange, t o f e l t y and v i n a c e o u s ; the l a t t e r u s u a l l y produced p e r i t h e c i a of i t s p e r f e c t s t a g e , G i b b e r e l l a b a c c a t a ( W a l l r . ) S a c c . ; c o n i d i a s t r a i g h t t o f a l c a t e , 27.0 - 41.6 x 3.6 - 5.4 um. These i s o l a t e s f i t the d e s c r i p t i o n g i v e n by Booth (1971), who a l s o r e p o r t s p e r i t h e c i a of t h i s s p e c i e s on T h u j a , among o t h e r h o s t s . F. l a t e r i t i u m was i s o l a t e d i n f r e q u e n t l y , o c c u r r i n g at both s i t e s , but more f r e q u e n t l y a t S i t e B, where i t was found d u r i n g the f a l l ( F i g . 16) . G l i o c l a d i u m roseum ( L i n k ) B a n i e r . B u l l . Soc. M y c o l . F r . 23:111 -112". 1907. C o l o n i e s s p r e a d i n g , w h i t e ; c o n i d i o p h o r e s t a l l ; c o n i d i a l mass i n l a r g e d r o p l e t s , 5.4 - 9.0 x 2.5 um. T h i s s p e c i e s was e a s i l y r e c o g n i z a b l e by i t s s l i m y spore drops on p e n i c i l l i a t e c o n i d i o p h o r e s . I t o c c u r s o f t e n as a p a r a s i t e on o t h e r f u n g i ( B a r n e t t and L i l l y , 1962). I t was i s o l a t e d from the l i t t e r i n t h i s s t u d y , but whether i t o c c u r r e d as a m y c o p a r a s i t e was not d e t e r m i n e d . G. roseum o c c u r r e d i n f r e q u e n t l y a t S i t e A, but was more abundant at S i t e B. I t was found most o f t e n i n the F l a y e r , e s p e c i a l l y d u r i n g the w i n t e r and s p r i n g samples ( F i g . 1 7 ) . A l l i s o l a t e s of t h i s s p e c i e s were from l e a v e s i n m o i s t chambers. 84 F i g u r e 16. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of Fusarium l a t e r i t i u m For Legend See F i g . 3 5 X < B i < 5 r-i iX i 1 CD CD I-—»—\—I—f—I—I—I—I 1—I—I—I—I—I—j—| AP JU NO FE MY JU 1977 1978 S A f c n P L E D A T E I—I—I—I—I—I—I—I 1—J—I—I—I MY SE JA AP 1977 1978 SAMPLE D A T E 85 F i g u r e . 1 7 . S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of G l i o c l a d i u m roseum For Legend See F i g . 3 5 M I I < I 1 1 1 1 1 1 r-AP JU NO H 1 — I — I — I — I 1977 FE MY 197B SAMPLE DATE JU X i CO 2 I CD i CD I — I — I — I — I — h MY SE 1977 -\ 1 — I — I — I — I JA AP 1978 SAMPLE DATE 86 H e l i c o d e n d r o n t r i q l i t z i e n s i s ( J a a p ) L i n d e r . Ann. Mo. Bot. Gdn. 16:330. 1929. C o l o n i e s s p r e a d i n g , w h i t e w i t h abundant a e r i a l mycelium; s p o r u l a t i n g p r o f u s e l y , c o n i d i a 10.8 - 18.0 x 12.6 urn, w i t h 4-6 c o i l s . T h i s s p e c i e s i s d i s t i n c t i n i t s p r o d u c t i o n of c h a i n s of s h o r t - c o i l e d s p o r e s . H. t r i g l i t z i e n s i s was i s o l a t e d m o s t l y from S i t e A but d i d occur a t S i t e B. I t was p r e s e n t i n a l l s u b s i t e s but most o f t e n from the m i d d l e and low ones, i n both L and F l a y e r s . I t was found more commonly d u r i n g the s p r i n g and summer samples ( F i g . 1 8 ) . T h i s i s one of the " a e r o - a q u a t i c " f u n g i of F i s h e r (1977) and o f t e n has been r e p o r t e d on p a r t i a l l y submerged l i t t e r . I t i s a l s o known from angiosperm l i t t e r of t h i s r e g i o n (Goos and Bandoni, 1977). F i g u r e 18. Seaso n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of He l i c o d e n d r o n t r i q l i t z i e n s i s For Legend See F i g . 3 5 I t < 3 i < Q - J i < I—I—I—I—I—I—I—I—I 1—I—I—I—I—I—I—I AP JU NO FE MY JU 1977 197B SAMPLE DATE IT CD i CD i CD I— I—J—I— I— I—J— I 1—I—I—I—I MY SE JA AP 1977 1976 SAMPLE DATE 88 H e l i c o o n fuscosporum L i n d e r . Ann. Mo. Bot. Gard. 16:326. 1929. C o l o n i e s m o d e r a t e l y s p r e a d i n g ; mycelium dark; c o n i d i o p h o r e s unbranched, 18.0 - 70.2 x 3.6 urn; c o n i d i a f u s c o u s , 30.6 - 66.6 x 21.6 - 32.4 um, c o i l e d 9 - 15 t i m e s . T h i s i s o l a t e resembles the d e s c r i p t i o n g i v e n by van B e v e r w i j k (1953), except the c o n i d i o p h o r e s here a r e s h o r t e r and the c o n i d i a a r e l a r g e r , sometimes w i t h more c o i l s . These d i f f e r e n c e s do not warrant e x c l u s i o n from t h i s s p e c i e s . H. fuscosporum i s another member of the " a e r o - a q u a t i c " hyphomycetes ( F i s h e r , 1977). I t a l s o has a r e l a t i v e l y h i g h c e l l u l o l y t i c a b i l i t y ( F i s h e r et a_l. , 1977). I t was i s o l a t e d o n l y from the m i d d l e s u b s i t e a t S i t e A, i n both l a y e r s , m o s t l y d u r i n g f a l l and w i n t e r ( F i g . 1 9 ) . 89 F i g u r e 19. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of H e l i c o o n fuscosporum For Legend See F i g . 3 3t ! I I—j—I—(—I—I—\—I—I 1—I—I—I—I—5—I—I AP JU NO FE MY JU 1 9 7 7 1 9 7 8 SAM=LE DATE 90 Hyalodendron 1 i g n i c o l a Diddens. Z e n t r a l b l . B a k t e r i o l . , 2. Abt. 90:317-318. 1934. C o l o n i e s m o d erately s p r e a d i n g , submerged mycelium d a r k , a e r i a l mycelium w h i t e , composed m o s t l y of c o n i d i o p h o r e s and c o n i d i a ; c o n i d i o p h o r e s o f t e n g a t h e r e d i n t u f t s ; c o n i d i a i n branched c h a i n s , h y a l i n e , n o n - s e p t a t e , lemon-shaped, 5.4 - 7.2 x 1.8 - 2.7 urn. These i s o l a t e s d i f f e r s l i g h t l y from the o r i g i n a l d e s c r i p t i o n i n h a v i n g a dark submerged mycelium. The method of spore p r o d u c t i o n i s the same, as i s spore s i z e . Hyalodendron i s a c o n f u s i n g taxon i n the l i t e r a t u r e ; the most common i n t e r p r e t a t i o n i s t h a t i t i s a h y a l i n e C l a d o s p o r i u m L i n k ( B a r r o n , 1968). There i s a l s o an a f f i n i t y t o P o l y s c y t a l u m Ri'ess, which has s i m i l a r c o n i d i a l p r o d u c t i o n , but l a r g e r s p o r e s . A major - . i n v e s t i g a t i o n of t h e s e two genera would be h e l p f u l . Diddens (1934) i s o l a t e d t h i s s p e c i e s from d e c a y i n g c o n i f e r woodpulp. H. 1 i g n i c o l a was i s o l a t e d m a i n l y from S i t e B but was found a l s o a t S i t e A. I t was e n c o u n t e r e d more o f t e n i n the L l a y e r , i n the m i d d l e and h i g h s u b s i t e s , and was a l s o i s o l a t e d from l i v i n g l e a v e s once ( F i g . 20). T h i s s p e c i e s was found i n the s p r i n g and e a r l y f a l l samples, e s p e c i a l l y the l a t t e r , where i t had the h i g h e s t o c c u r r e n c e . H. l i g n i c o l a has not been r e p o r t e d o f t e n i n the l i t e r a t u r e ; when found, i t has u s u a l l y been i d e n t i f i e d o n l y t o genus ( B h a t t , 1970; K a t z and L i e t h , 1974; Sherwood and C a r r o l l , 1974; Wicklow and Whittingham, 1974, 1978). F i g u r e 20. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of Hyalodendron l i q n i c o l a For Legend See F i g . 3 2 8 — 1 > _ J I < < - i , 1 1 j — | — | j i 1 } 1 NO FE MY JU 1978 SAMPLE DATE 5 M II I m 2S CD CD MY H—l—I—\—\—I—I ¥—f SE 1977 -}—I JA AP 1978 SAMPLE DATE 92 K r i e q e r i a s e a v e r i (Rehm)Seaver . M y c o l . 3_5:493. 1943. T h i s s p e c i e s was never a c t u a l l y i s o l a t e d i n c u l t u r e , but r e a d i l y formed a p o t h e c i a on the l e a v e s i n moist chambers. The a p o t h e c i a were s m a l l , g e l a t i n o u s , g r e e n i s h t o y e l l o w - g r e e n and f i t the d e s c r i p t i o n p r o v i d e d by Seaver (1951). K. s e a v e r i i s known o n l y from Thuja p i i c a t a f o i l a g e ( S eaver, 1951; Shaw, 1973). I t was found m a i n l y a t S i t e A, on l i v i n g l e a v e s and i n the L l a y e r , d u r i n g the f a l l ( F i g . 2 1 ) . L i b e r t e l l a sp. C o l o n i e s moderately s p r e a d i n g , at f i r s t w h i t e , then t u r n i n g l i g h t t o d a r k e r brown, l i t t l e a e r i a l mycelium p r e s e n t ; c o n i d i o p h o r e s branched, p r o d u c i n g p h i a l i d e s , 4.5 - 12.6 x 1.0 -2.7 um; c o n i d i a h y a l i n e t o p a l e brown i n mass, U-shaped t o s i g m o i d , produced i n s l i m e , 3 s e p t a t e , 59.4 - 75.6 x 1.8 -2.0 um. L i b e r t e l l a sp. o c c u r r e d i n both S i t e A and S i t e B, m a i n l y i n the low and m i d d l e s u b s i t e s , but was found i n the h i g h s u b s i t e i n S i t e A d u r i n g the s p r i n g . I t o c c u r r e d i n both the L and F l a y e r s , w i t h l i t t l e change of f r e q u e n c y throughout the year ( F i g . 2 2 ) . F i g u r e 21. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of K r i e q e r i a s e a v e r i For Legend See F i g . 3 > M -J I < s M I I < M 3E I < < I—I— I— I—I—»—1—I—I—J—1—\—\—I—I—I— I AP JU NO FE 1377 MY 1978 JU SAMPLE DATE > M - J 1 CD MY A—\—I—I—I—I—» 1 ( t 1 - J SE 1977 JA AP 1978 SAMPLE DATE F i g u r e 22. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of L i b e r t e l l a sp. For Legend See F i g . 3 5 X » < 2 » < t < i S i GQ I— I—*— I— f—H-AP JU 1977 H—I 1—J—I—»—8—I—I—I NO F E MY JU 1976 SAMPLE DATE -J i GQ _ H 1 1—I—I—» !—I—J—I—I MY £E JA AP 1977 197S SAM=LE DATE 95 M o r t i e r e l l a ramanniana ( M o l l e r ) L i n n e m . v a r . ramanniana Linnem. P f l a n z e n f o r s c h u n g , Jena. 2_3:19. 1941. C o l o n i e s s p r e a d i n g - r a p i d l y ; s p o r a n g i o p h o r e s r e l a t i v e l y s h o r t ; s p o r a n g i a r e d d i s h . T h i s s p e c i e s was e a s i l y r e c o g n i z e d t h r o u g h the above f e a t u r e s . M. ramann iana v a r . ramann i a n a o c c u r r e d t w i c e a t S i t e A, and s i x times at S i t e B, m o s t l y i n the L l a y e r . I t was u s u a l l y i s o l a t e d d u r i n g the summer and e a r l y f a l l ( F i g . 23). M o r t i e r e l l a s p e c i e s are h e a v i l y s p o r i n g and r e l a t i v e l y slow growing, and a r e commonly found i n the s o i l ( S e w e l l , 1959). 96 F i g u r e 23. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of M o r t i e r e l l a ramann i a n a v a r . ramann i a n a F o r Legend See F i g . 3 • f i X I < < I—»—I—f—I—I—I—I—f—f—I—I—I—I—f—f—I J U N D F E M Y J U 1 9 7 7 1 9 7 8 S/^PLE DATE 5 M I b-4 » CD i C D \—I—f—I—h MY —I—f I—f—J—i—\ S E J A A P 1 9 7 7 1 9 7 8 SAfcPLE DATE 97 Mucor h i e m a l i s Nehmer f . h i e m a l i s S c h i p p e r . Stud. M y c o l . 4:26. 1973 . C o l o n i e s f a s t g rowing; s p o r a n g i o p h o r e s t a l l , w i t h dark b l a c k s p o r a n g i a ; l a r g e c o l u m e l l a p r e s e n t , 23.4 - 54.0 um i n d i a m e t e r , sometimes up t o 100 um i n d i a m e t e r ; spores i r r e g u l a r , g l o b o s e t o e l o n g a t e 3.6 - 9.0 x 3.6 - 4.5 um. These i s o l a t e s sometimes had a c o l u m e l l a l a r g e r than t h a t r e p o r t e d by S c h i p p e r (1973), but o t h e r w i s e f i t the d e s c r i p t i o n . M. h i e m a l i s f. h i e m a l i s i s widesp r e a d i n s o i l s and l i t t e r (Domsch and Gams, 1972), and was one of the most common f u n g i i s o l a t e d from the SYT p l a t e s . I t o c c u r r e d a t a l l sample t i m e s i n both s i t e s , i n the L and F l a y e r s ( F i g . 24). A l t h o u g h i t was p r e s e n t i n a l l t h r e e s u b s i t e s , i t was found more f r e q u e n t l y i n the h i g h s u b s i t e . 98 Figure 24. Seasonal Distribution - Number Of Isolates Of Mucor hiemalis f. hiemalis For Legend See Fi g . 3 I I < M i < < | 1—|—f—| 1—I 1 f 1—I—f 1 f—f— I—» AP JO NO FE MV JU 1377 1978 S A M P L E D A T E 5 I* CQ i m i CD I — 9 — 1 — I — » — I — \ — \ » » I—J—I MV S£ J A AP 1377 1378 SM/PLE DATE 99 P e n i c i 1 1 i u m brevi-compactum D i e r c k x . Soc. S c i e n . Brux. 25:88. 1901. T h i s s p e c i e s i s a member of the V e l u t i n a s u b s e c t i o n , a c c o r d i n g t o Raper and Thorn (1949). I t i s r e c o g n i z e d by i t s s h o r t , t i g h t p e n i c i l l u s , w i t h somewhat s w o l l e n metulae. P. brevi-compactum was found more f r e q u e n t l y i n the mi d d l e and h i g h s u b s i t e s a t S i t e A, and i n the mi d d l e and low s u b s i t e s a t S i t e B. I t was i s o l a t e d from l i v i n g l e a v e s , but o t h e r w i s e o c c u r r e d m a i n l y d u r i n g the f a l l and w i n t e r ( F i g . 25). I t has been r e p o r t e d from a v a r i e t y of s o i l t y p e s (Domsch and Gams, 1972) . P e n i c i l l i u m c i t r i n um Thorn. U.S.D.A. Bur. Anim. Ind. B u l l . 118. 61-63. 1910. T h i s s p e c i e s i s a l s o a member of the s u b - s e c t i o n V e l u t i n a (Raper and Thorn, 1949). I t i s c h a r a c t e r i z e d by no b r a n c h i n g below the l e v e l of metulae, and a b l u i s h t i n t t o the spore mass; c o n i d i a 1.5 - 2.0 um. P. c i t r i n u m was found m a i n l y a t S i t e A d u r i n g the f a l l , but i t o c c u r r e d d u r i n g the summer i n the h i g h s u b s i t e ( F i g . 26). I t s d i s t r i b u t i o n i s worldwide and common i n s o i l s (Domsch and Gams, 1972). 100 F i g u r e 25. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of P e n i c i l l i u m brevi-compactum | J — | — | 1 1 — » 1 \ 1—|—| 1 1 1 1 1 3 AP JU ND FE MV JU i 1977 1970 SAMPLE DATE DD I—|—}—|—f—|—f—| 1—|—|—|—j MY 9E JA AP 1977 1978 S A M P L E D A T E 1 0 1 F i g u r e 26. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of P e n i c i l l i u m c i t r i n u m F o r L e g e n d See F i g . 3 5 W X I < M 2£ i < < AP -f—F—I-J U H 1—f 1—f—I f — f — F — f — I 1977 NO F E SAMPLE DATE MY 1978 J U 2 » CD I » J 1—I !• M Y S£ 1977 - I — | 1 — 5 — I — J — I JA AP 1978 SAMPLE GATE 102 P e n i c i l l i u m f r e q u e n t a n s W e s t l i n g . Ark. Bot. 11:58. 133-134. 1911. T h i s s p e c i e s i s a member of the M o n o v e r t i c i l l a t a s e c t i o n , a c c o r d i n g t o Raper and Thorn (1949). I t i s r e c o g n i z e d by i t s v e l v e t y c o l o n y , m o n o v e r t i c i H a t e c o n i d i o p h o r e and globose t o subglobose c o n i d i a , 1.8 - 3.6 urn i n d i a m e t e r , s l i g h t l y roughened. P. f r e q u e n t a n s was i s o l a t e d m a i n l y from S i t e B, i n the F l a y e r , d u r i n g the f a l l and w i n t e r ( F i g . 27). T h i s i s a common s o i l fungus, e s p e c i a l l y i n f o r e s t s o i l s and l i t t e r (Domsch and Gams, 1972). P e n i c i I l i u m n i g r i c a n s ( B a i n i e r ) T h o m . The P e n i c i l l i a . 351-353. . 1930. P. n i g r i c a n s belongs t o the D i v a r i c a t a s u b s e c t i o n of Raper and Thorn (1949). I t i s r e c o g n i z e d by i t s somewhat roughened s p o r e s , 1.5 - 3.0 urn i n d i a m e t e r , which impart a d u l l gray c o l o r to the c o l o n y , and a y e l l o w r e v e r s e . I t was i s o l a t e d throughout the year a t S i t e A, and o n l y d u r i n g the f a l l and w i n t e r a t S i t e B ( F i g . 2 8 ) . In S i t e A the h i g h e s t o c c u r r e n c e s were d u r i n g t h e summer samples, both of which were c h a r a c t e r i z e d by low p r e c i p i t a t i o n and h i g h t e m p e r a t u r e . T h i s c o r r e s p o n d s w i t h the i n f o r m a t i o n r e p o r t e d by Domsch and Gams (1972) f o r t h i s common s o i l fungus. Throughout the s a m p l i n g , i t was i s o l a t e d more f r e q u e n t l y from the h i g h s u b s i t e and the F l a y e r . 103 F i g u r e 27. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of P e n i c i l l i u m f r e q u e n t a n s F o r L e g e n d See F i g . 3 5 X \ < 2 i < AP -f—f—f—f—f—f 1—I 1—I—f f—f—I—I—I JU NO FE 1977 SAMPLE DATE MV 197B JU S i f CD 2 CD i CO I — f — H MV SE 1977 H f—h H—f—f JA AP 1978 SAMPLE DATE 104 F i g u r e 28. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of P e n i c i l l i u m n i g r i c a n s For Legend See F i g . 3 5 M I I < I < I < I I CO I CO I—I—I—I—I—I—I—I—h AP J U ; NO 1977 H—I 1 1 1 1 1 FE MY JU 1970 SAMPLE DATE i CO I—I—I—I—I—I—I—h MY SE 1977 -I—I—I 1—I JA AP 1979 SAMPLE DATE 105 P e n i c i l l i u m notatum W e s t l i n g . Ark. Bot. 11_:55, 95-97. 1911. T h i s s p e c i e s i s another member of the V e l u t i n a s u b s e c t i o n of P e n i c i l l i u m (Raper and Thorn, 1949). I t i s r e c o g n i z e d by i t s b l u e - g r e e n c o l o r and subglobose t o glo b o s e c o n i d i a , 2.0 - 3.6 urn i n d i a m e t e r . P. notatum was found o n l y at S i t e A, m a i n l y i n the F l a y e r ; i t u s u a l l y o c c u r r e d d u r i n g the f a l l and w i n t e r , but o n l y i n low numbers ( F i g . 29). T h i s i s a common and widesp r e a d s o i l fungus (Domsch and Gams, 1972). P e n i c i l l i u m r a i s t r i c k i i Smith. T r a n s . B r i t . M y c o l . Soc. 18:90. 1933. T h i s s p e c i e s i s a l s o a member of the s u b s e c t i o n ' D i v a r i c a t a (Raper and Thorn, 1949). I t i s one of the e a s i e s t r e c o g n i z e d P e n i c i l l i a , i n t h a t i t u s u a l l y produces abundant s c l e r o t i a and r e l a t i v e l y s p a r s e c o n i d i o p h o r e s . The c o n i d i o p h o r e s a r e s l i g h t l y roughened, and the a e r i a l hyphae and the c o l o n y r e v e r s e a r e o f t e n y e l l o w . P. r a i s t r i c k i i was found m a i n l y d u r i n g the summer samples a t S i t e A; the e a r l y f a l l and s p r i n g samples a t S i t e B ( F i g . 3 0 ) . I t i s s i m i l a r t o P. n i g r i c a n s i n o c c u r r i n g a t tim e s of low p r e c i p i t a t i o n and h i g h t e m p e r a t u r e . 1 0 6 F i g u r e 29. S e a s o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of P e n i c i l l i u m n o t a t u m F o r L e g e n d See F i g . 3 _J I < 5 w I i < I H < < AP 4—f—|—|—f-JU -f—I h F t 1977 SAMPLE DATE H—f—I—I—I MY 1978 JU i « CD MY 9£ 1977 4—4—I—f—| JA AP 1978 SAMPLE DATE Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of P e n i c i l l i u m r a i s t r i c k i i For Legend See F i g . 3 I \ 1 1 1 » \ 1 1—\—I 1 \ MY SE *IA AP 1977 1976 SM&LE DATE 108 P e n i c i l l i u m verrucosum D i e r c k x . v a r . c y c l o p i u m ( W e s t l i n g ) Sampson, S t o l k & Hadlok. Stud. M y c o l . 11:37-41. 1976. A c c o r d i n g to Sampson e_t a l . (1976), t h i s s p e c i e s i n c l u d e s P. c y c l o p i u m and P. m a r t e n s i i , among o t h e r d e s c r i b e d s p e c i e s . These two s p e c i e s d i f f e r o n l y s l i g h t l y i n the r e l a t i v e roughness of the c o n i d i o p h o r e , t h e r e f o r e t h e i r c o m b i n a t i o n under one s p e c i e s i s w a r r a n t e d . T h i s s p e c i e s i s a member of the s u b s e c t i o n F a s c i c u l a t a of Raper and Thorn (Sampson et a l . , 1976). I t i s c h a r a c t e r i z e d by somewhat f a s i c u l a t e t o mononematous c o n i d i o p h o r e s ; u s u a l l y w i t h t h e ' c o n i d i a g i v i n g the c u l t u r e a b l u i s h shade; c o n i d i a 1.5 - 4.0 urn i n d i a m e t e r ; c o n i d i o p h o r e s f i n e t o c o a r s e l y roughened. P. verrucosum v a r . c y c l o p i u m was the most commonly o c c u r r i n g s p e c i e s of P e n i c i l l i u m i n t h i s s t u d y . I t was p r e s e n t a t a l l d a t e s i n both S i t e A and S i t e B, except f o r the l a s t summer sample, and was more p r e v a l e n t i n the F l a y e r when i t was found ( F i g . 31). I t o c c u r r e d m a i n l y d u r i n g the f a l l and w i n t e r samples, times c h a r a c t e r i z e d by low tempe r a t u r e s and h i g h p r e c i p i t a t i o n . G r i f f i n (1963) has r e p o r t e d 81 - 84% r e l a t i v e h u m i d i t y as the lower l i m i t f o r growth f o r t h i s s p e c i e s . 109 F i g u r e 31. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of Pen i c i11ium verrucosum v a r . c y c l o p i u m I 1 f 1 1 1 1 1 1—| 1 1—j MY SE JA AP 1977 1978 SAMPLE DATE 110 P e s t a l o t i a m o n o c h a e t i o i d e s Doyer. Med. P h y t o p a t h . Lab. " W i l l . Comm. S c h o t t . " 9:24. 1925. C o l o n i e s s p r e a d i n g r a p i d l y ; p r o d u c i n g r a i s e d a c e r v u l i ; c o n i d i a 5 - c e l l e d , the m i d d l e t h r e e c e l l s d a rk, the upper two c e l l s s l i g h t l y d a r k e r , the lower two c e l l s o f t e n w arty, 27.0 x 9.0 - 9.9 urn; c o n i d i a w i t h one s e t a , which i s sometimes branched, up t o 50.4 um l o n g . Guba (1961) r e p o r t s i t from Chamaecyparis. P. m o n o c h a e t i o i d e s was i s o l a t e d m o s t l y from the L l a y e r a t both s i t e s , but a l s o o c c u r r e d on l i v i n g l e a v e s a t S i t e A. I t tended t o be found d u r i n g the f a l l and w i n t e r samples ( F i g . 32). gure 32. 2 > M t < i X I < UJ I Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of o P e s t a l o t i a m o n o c h a e t i o i d e s For Legend S e e ' F i g . 3 AP JU NO 1977 FE MY JU 1378 SAMPLE DATE I—I—I—I—I—I—I—I 1—I—I—I—I MY SE JA AP 1377 1978 SAMPLE DATE 112 P o l y s c y t a l u m fecundissimum R i e s s . Bot. Z e i t u n g . 11:138-139. 1853. C o l o n i e s m o d erately s p r e a d i n g , w h i t e t o b l a c k i n some a r e a s ; c o n i d i a produced i n branched c h a i n s , sometimes g a t h e r e d i n t o t u f t s , 0-1 s e p t a t e , 7.2 - 23.4 x 1.8 - 3.6 urn. T h i s s p e c i e s shows a f f i n i t i e s w i t h Hyalodendron 1 i g n i c o l a D iddens, which i t r e s e m b l e s . These two genera are b a f f l i n g i n the l i t e r a t u r e and are i n need of major c u l t u r a l and taxonomic a n a l y s i s . P. fecundissimum was r e p o r t e d s p e c i f i c a l l y by H e r i n g (1965), but i n o t h e r r e p o r t s on l i t t e r f u n g i , i t has been i d e n t i f i e d o n l y t o genus ( A p i n i s <st a l . , 1972; Hogg and Hudson, 1966; Katz and L i e t h , 1974; P a r k i n s o n and B a l a s o o r i y a , 1969; Tubaki and Yokoyama, 1971, 1973b; Yokoyama and T u b a k i , 1973; Yokoyama e_t a l . , 1977). P. f e c u n d i ss imum was i s o l a t e d f r e q u e n t l y from both s i t e s , i n a l l s u b s i t e s , but m a i n l y the m i d d l e and low ones. I t was more prominent i n the L l a y e r and o c c u r r e d more f r e q u e n t l y d u r i n g the s p r i n g and f a l l . In S i t e A, as i t s o c c u r r e n c e t a p e r e d o f f i n the m i d d l e s u b s i t e , i t i n c r e a s e d i n the low s u b s i t e ( F i g 33). F i g u r e 33. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of P o l y s c y t a l u m fecundissimum For Legend See F i g . 3 X i < M I < I < AP JU NO 1977 FE MY JU 197B SAMPLE DATE 5 m I CD i CD I—J—I—I 1—I—I—I I-H—I—I-H MY SE JA AP 1977 1978 SAKFLE DATE 114 R a m i c h l o r i d i u m subulatum de Hoog. Stud . M y c o l . 15:83. 1977. C o l o n i e s m o d e r a t e l y s p r e a d i n g ; a e r i a l mycelium g r a y i s h ; c o n i d i o p h o r e s dark, 22.5 - 40.5 x 3.6 - 4.5 um, e l o n g a t i n g s y m p o d i a l l y ; c o n i d i a born on d e n t i c l e s , t r u n c a t e , p a l e brown, 4.5 - 6.3 x 2.0 - 2.5 um. These i s o l a t e s f i t the d e s c r i p t i o n g i v e n by de Hoog (1977). R. subulatum was i s o l a t e d o n l y from S i t e A, m o s t l y from the L l a y e r . I t was found i n f r e q u e n t l y , u s u a l l y i n the summer or w i n t e r ( F i g . 34). T h i s s p e c i e s has been r e p o r t e d from v a r i o u s s o i l s (de Hoog, 1977). Selenophoma sp. C o l o n i e s m o d erately s p r e a d i n g ; mycelium da r k , p r o d u c i n g dark p y c n i d i a o f t e n grouped t o g e t h e r , somewhat s t r o m a t i c ; c o n i d i a h y a l i n e , a l l a n t o i d , 9.0 - 10.8 x 2.7 um. Selenophoma i s a genus commonly found on g r a s s e s (Park and Sprague, 1953), but i n c l u d e s one s p e c i e s , S. drabae which i s found on d i c o t y l e d o n s . No r e p o r t s have been made of t h i s genus on c o n i f e r s , but Tubaki and Yokoyama (1973b) i s o l a t e d i t from s t e r i l i z e d l e a v e s i n angiosperm l i t t e r . Selenophoma sp. o c c u r r e d a t both s i t e s , m o s t l y i n the L l a y e r , but was i s o l a t e d i n l a r g e r numbers a t S i t e B. At S i t e A i t was prominent d u r i n g the w i n t e r , w h i l e a t S i t e B i t o c c u r r e d on l i v i n g l e a v e s i n l a t e summer, and i n the o t h e r s u b s i t e s i n s p r i n g and w i n t e r ( F i g . 35). F i g u r e 34. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of R a m i c h l o r i d i u m subulatum For Legend See F i g . 3 1977 1978 SAMPLE DATE 116 F i g u r e 35. 5 M X I < E M t < i CO Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of Selenophoma sp. For Legend See F i g . 3 I 1—i 1 1 1 1 1 1—I—I 1—| M Y S E J A A P 1977 1378 SAMPLE DATE 117 Septogloeum sp. Colony growth moderate; mycelium w h i t e t o g r a y : c o n i d i o p h o r e s i n p u s t u l e s , o f t e n b r o w n i s h ; c o n i d i a h y a l i n e , a t f i r s t c o n t i n u o u s , but becoming 1-5 s e p t a t e , m o s t l y 3, base of c o n i d i u m a p i c u l a t e , 21.0 - 34.0 x 7.2 - 11.5 um. Sometimes the o l d e r c o n i d i a appear m u r i f o r m , but t h i s seems t o be a r e s u l t of p l a s m o l y s i s . Most of the s p e c i e s d e s c r i b e d i n t h i s genus occur on d i c o t y l e d o n o u s p l a n t s . Septogloeum sp. was i s o l a t e d at both s i t e s , but more f r e q u e n t l y a t S i t e B. I t was most f r e q u e n t l y i s o l a t e d from l i v i n g l e a v e s i n l a t e summer and f a l l ( F i g . 36). O t h e r w i s e , i t o c c u r r e d .mainly i n the L l a y e r . 118 F i g u r e 36. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of Septogloeum sp. For Legend See F i g . 3 2 M > M I I < I < I \ h AP JU H 1 h NO FE H 1 1—I 1—I 1977 S A N F L E D A T E MY 1978 JU M > M i_ l i I CD ^2 M X I CD M I CD i CD 1 1 — H MY H 1 1 1 1 — I — I 1—H SE 1977 JA AP 1978 SAMPLE DATE 119 Septonema c h a e t o s p i ra (Grove)Hughes. N a t u r a l i s t . Jan.-Mar. 9-11. 1952. C o l o n i e s m o d erately s p r e a d i n g , dark brown t o b l a c k w i t h abundant a e r i a l mycelium; c o n i d i a produced i n l o n g , branched c h a i n s , p a l e brown, 0-3 s e p t a t e , 12.6 - 36.9 x 2.0 - 5.4 urn. I t i s e a s i l y r e c o g n i z e d by i t s c h a r a c t e r i s t i c s e p t a t e b l a s t o s p o r e . E l l i s (1976) has p l a c e d t h i s s p e c i e s i n the genus Heteroconium P e t r a k . S. c h a e t o s p i r a was r e l a t i v e l y abundant, o c c u r r i n g i n both S i t e A and S i t e B, more p r o m i n e n t l y i n the low s u b s i t e , i n both the L and F l a y e r s , but more o f t e n from the l a t t e r . I t o c c u r r e d i n t i m e s of warmer t e m p e r a t u r e s , but not too d r y , t h u s * i t i s found more f r e q u e n t l y i n s p r i n g and summer ( F i g . 3 7 ) . The genus has been r e p o r t e d from s o i l ( I s h i i , 1970), l i v i n g n e e d l e s ( G o u r b i e r e , 1975) and nee d l e s i n a stream ( B a r l o c h e r and K e n d r i c k , 1974). T h i s p a r t i c u l a r s p e c i e s has been r e p o r t e d once from f o r e s t s o i l (Gochenaur and Woodwell, 1974) and a g r i c u l t u r a l s o i l (Domsch and Gams, 1972). 120 F i g u r e 37. S e a s o n a l D i s t r i b u t i o n - N u m b e r O f I s o l a t e s O f S e p t o n e m a c h a e t o s p i r a F o r L e g e n d S e e F i g . 3 5 M X I < 2 i < i < A P H—I V JU H 1—I—h FE 1977 NO SAMPLE DATE MY 197Q H 1 1 JU .5 M .1 I CD i CD MY *—I—I 1—I 1—I 1—I—I \—| SE 1977 JA AP 1978 SAMPLE DATE 121 Septonema c h a e t o s p i r a (Groves)Hughes v a r . p i n i B o u r c h i e r . Can. J . Bot. 39:1782-1784. 1961. C o l o n i e s r e s t r i c t e d , dark brown t o b l a c k , l i t t l e a e r i a l mycelium p r e s e n t ; c o n i d i a produced i n branched c h a i n s , p a l e brown, 0-1 s e p t a t e , 6.0 - 14.5 x 1.8 - 3.6 urn. B o u r c h i e r (1961) d e s c r i b e d t h i s s p e c i e s from the wood of P i nus c o n t o r t a . S. c h a e t o s p i r a v a r . p i n i was i s o l a t e d i n low numbers from almost a l l samples i n S i t e A and S i t e B. At S i t e A, i t o c c u r r e d i n the low and m i d d l e s u b s i t e s i n the s p r i n g and summer, and i n the h i g h s u b s i t e i n the w i n t e r . I t was p r e s e n t i n c o n s i s t e n t l y i n both L and F l a y e r s ( F i g . 3 8 ) . Sporodesmium flexum Matsushima. Icones Microfungorum Matsushima Lectorum. 137-138. 1975. C o l o n i e s m o d e r a t e l y s p r e a d i n g , a e r i a l hyphae brown, b e a r i n g numerous s p o r e s ; c o n i d i a f l e x u o u s , brown, 6 - 8 s e p t a t e , t a p e r i n g toward the apex. T h i s s p e c i e s i s i d e n t i f i e d e a s i l y by i t s c h a r a c t e r i s t i c s p o r e . S. flexum was i s o l a t e d o n l y from l e a v e s i n moist chambers and from o n l y S i t e A. I t was found s l i g h t l y more f r e q u e n t l y i n the F l a y e r , and o n l y i n the m i d d l e and low s u b s i t e s ( F i g . 3 9 ) . I t o c c u r r e d i n s p r i n g and summer samples, e s p e c i a l l y the l a s t two samples of the s t u d y , p o s s i b l y i n d i c a t i n g t h a t i t i s more p r e v a l e n t a t t i m e s of h i g h temperature but r e q u i r i n g a moderate amount of m o i s t u r e . 122 F i g u r e 38. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of Septonema c h a e t o s p i ra v a r . p i n i For Legend See F i g . 3 5 I < I 1 1 1 1 1 1 1 1 1 — I — I — H 1 1 1 1 AP JU NO FE MY JU 1977 1979 SAMPLE DATE 2 i I—\—1—\—l—\—I—\ 1 — 1 — i — \ — J MY SE JA AP 1377 1976 SAM=LE DATE 123 F i g u r e 39. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of Sporidesmiurn flexum For Legend See F i g . 3 E M 3 1377 1978 S M v P L E D A T E 124 Tr ichoderma kon i n g i i Oud. apud Oud. et Koning. A r c h s . N e e r l . S c i . I I . 7:291. 1902. C o l o n i e s s p r e a d i n g r a p i d l y , p r o d u c i n g l o o s e spore masses w i t h abundant a e r i a l hyphae, spore c l u s t e r s y e l l o w - g r e e n ; c o n i d i a e l l i p s o i d , smooth, 3.6 - 5.4 x 2.0 - 3.0 um. T. k o n i n q i i was i s o l a t e d a t both s i t e s , but much more f r e q u e n t l y a t S i t e B. I t was found i n a l l t h r e e s u b s i t e s , i n both the L and F l a y e r s . I t tended t o occur more o f t e n i n w i n t e r and s p r i n g ( F i g . 4 0 ) . Trichoderma polysporum ( L i n k ex P e r s . ) R i f a i . Commonw. M y c o l . I n s t . M y c o l . Pap. 116:19-22. 1969. C o l o n i e s s p r e a d i n g r a p i d l y , l i t t l e or no a e r i a l mycelium except f o r the spore c l u s t e r s ; c o n i d i a and c o n i d i o p h o r e s h y a l i n e , c o n i d i o p h o r e s w i t h a p i c a l s t e r i l e appendage, c o n i d i a s m a l l , 2.8 - 3.7 x 1.8 - 2.0 um. T h i s common s o i l fungus i s r e c o g n i z e d e a s i l y by the whi t e spores and s t e r i l e appendages. T. polysporum o c c u r r e d m a i n l y i n the mi d d l e and low s u b s i t e s , m o s t l y i n the F l a y e r . I t was more common d u r i n g the s p r i n g and summer samples ( F i g . 4 1 ) . 12.5 F i g u r e 40. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of Trichoderma k o n i n q i i For Legend See F i g . 3 I 1 \ 1 1 \ 1 1 1—}—I \ f MY SE JA AP 1977 1970 SAMPLE DATE F i g u r e 41. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of Trichoderma polysporum For Legend See F i g . 3 1977 1976 SAMPLE DATE i CO I I 1 i 1 ! 1 1 \ 1 1 ) I MY SE JA AP 1977 1378 SAMPLE DATE 127 Trichoderma v i r i d e P e r s . ex S.F. Gray. Nat. A r r a n g . Br. P i . 1:560. 1821. C o l o n i e s s p r e a d i n g r a p i d l y , a e r i a l mycelium s p a r s e except i n the spore c l u s t e r s , c o n i d i a and c o n i d i o p h o r e s green, c o n i d i a v e r y m i n u t e l y to c o a r s e l y roughened, o v o i d , 3.6 - 5.0 x 2.7 -4.2 um. T h i s s p e c i e s i s i d e n t i f i e d e a s i l y by i t s c h a r a c t e r i s t i c rough s p o r e . T. v i r i d e was found i n a l l t h r e e s u b s i t e s and on l i v i n g l e a v e s , at both s i t e s , but much more f r e q u e n t l y a t S i t e A ( F i g . 4 2 ) . I t o c c u r r e d s l i g h t l y more o f t e n i n the F l a y e r . The h i g h e s t o c c u r r e n c e s were d u r i n g the summer samples, which were c h a r a c t e r i z e d by h i g h t e m p e r a t u r e s and low p r e c i p i t a t i o n . G r i f f i n (1963) r e p o r t e d optimum growth at h u m i d i t y of 91 - 96 %, which does not seem t o r e f l e c t t h i s p a t t e r n of o c c u r r e n c e . B e f o r e R i f a i ' s t r e atment of the genus (1969), a l l . green Tr ichodermas were r e f e r r e d commonly t o T. v i r i d e . G r i f f i n p o s s i b l y c o u l d have been d e a l i n g w i t h another s p e c i e s , which might e x p l a i n t h i s d i s c r e p a n c y . P o s s i b l y temperature i s more i m p o r t a n t , as S e w e l l (1959) m a i n t a i n s t h a t the m y c e l i a l a c t i v i t y i s slowed i n w i n t e r . F i g u r e 42. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of Tr ichoderma v i r i d e For Legend See F i g . 3 S > _ l I < 5 M I < M I !< I < 2 > H I _ i I CO 5 £ i CQ •HI i CO 4 CD AP H—f—I—I—t-JU H 1 I 1 1 9 7 ? ra SAM=L£ DATE -I—I—H MV 197B JU h ~ » — J — J — I — ] — \ — 1 1—|—\—I—| fcfr SE JA AP 1977 1978 SAMPLE DATE 129 Tripospermum m y r t i (Lind)Hughes. Commonw. M y c o l . I n s t . M y c o l . Pap. 46:17-18. 1951. T h i s s p e c i e s i s r e c o g n i z e d by i t s d a r k , somewhat r e s t r i c t e d c o l o n y , p r o d u c i n g i r r e g u l a r l y t e t r a r a d i a t e c o n i d i a . T. m y r t i was i s o l a t e d o n l y a t S i t e A, m o s t l y from l i v i n g l e a v e s , but once i n the L l a y e r ( F i g . 4 3 ) . G o u r b i e r e (1974a) has r e p o r t e d t h i s s p e c i e s from l i v i n g l e a v e s of A b i e s . T r i p o s p o r i u m e l e g a n s Corda. I c o n . Fung. 1:16. 1837. C o l o n i e s somewhat r e s t r i c t e d , w h i t e ; a e r i a l hyphae s p a r s e ; t h i s s p e c i e s i s r e c o g n i z e d e a s i l y by i t s d i s t i n c t dark t r i - or t e t r a r a d i a t e s p o r e , produced on e r e c t , dark c o n i d i o p h o r e s . T. e l e g a n s was i s o l a t e d o n l y from S i t e B, m a i n l y i n the L l a y e r . I t o c c u r r e d d u r i n g the l a t e summer, but was i s o l a t e d i n low numbers(Fig. 4 4 ) . 130 F i g u r e 43. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of Tripospermum m y r t i For Legend See F i g . 3 2 > I < 2: i < AP HI—I—I 1 t r JU NO \ I I—I—I—J—i—i FE 1977 MV 1978 JU SA&FLE DATE CO I—I—I—I—I—I—1—I—I—I—I—I—I MY SE JA AP 1977 1978 SAMPLE DATE gure 44. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s T r i p o s p o r i u m e l e g a n s For Legend See F i g . 3 j | 1 1 1 1 1 1 1— | 1 1—-} MY SE JA AP 1977 1970 SAMPLE DATE 132 V a r i c o s p o r i u m elodeae K e g e l . Ber. Deutsch. Bot. Ges. 24:213-216. 1906. .Colonies m o d erately s p r e a d i n g , w h i t e , t u r n i n g green i n p l a c e s ; c o n i d i a c h a r a c t e r i s t i c a l l y branched and e a s i l y r e c o g n i z a b l e . T h i s s p e c i e s was i s o l a t e d o n l y from S i t e A. V. elodeae o c c u r r e d m o s t l y i n the low and middl e s u b s i t e s , and was more p r e v a l e n t as p r e c i p i t a t i o n d e c r e a s e d . I t was most common i n the low s u b s i t e where i t was found more o f t e n i n the F l a y e r ( F i g . 4 5 ) . The o c c u r r e n c e of t h i s s p e c i e s i n both a q u a t i c and t e r r e s t r i a l h a b i t a t s has been w e l l documented (Bandoni, 1972, 1977; I n g o l d , 1975; Park, 1974; Sanders and Webster, 1978; Webster, 1977). V e r t i c i l l i u m b u l b i l l o s u m W. Gams and M a l l a . Cephalosporium-a r t i g e S c h i m m e l p i l z e . 189-190. 1971. C o l o n i e s m o d e r a t e l y s p r e a d i n g , w h i t e w i t h abundant a e r i a l mycelium. T h i s s p e c i e s was c h a r a c t e r i z e d by the presence of i r r e g u l a r shaped, u n s t a l k e d chlamydospores, and c o n i d i a , 2.7 -4.5 x 1.8 urn. V. b u l b i l l o s u m was i n f r e q u e n t l y i s o l a t e d from S i t e A and S i t e B, but c o n s i s t e n t l y from the F l a y e r . I t was more p r e v a l e n t d u r i n g the s p r i n g and summer ( F i g . 4 6 ) . 133 F i g u r e 45. Seas o n a l D i s t r i b u t i o n - Number Of I s o l a t e s Of V a r i c o s p o r i u m elodeae For Legend See F i g . 3 I 1 1 1 1 1 » 1 1 1—f 1 1 1 1 1 » AP JU NO FE MY JU 1377 1378 5 M £ DATE 134 F i g u r e 46'. Seasonal D i s t r i b u t i o n - Number Of I s o l a t e s Of V e r t i c i l l i u m b u l b i l l o s u m For Legend See F i g . 3 5 M 2 i i < i < I—I—I—I 1 1—I 1—I 1—I—I 1—I—I—I—I AP JU NO FE MY JU 1977 1378 SAMPLE DATE i I CD I—I—I—I 1 1 . 1 1 ' — H — « — I - H MY SE JA AP 1977 1978 SAMPLE DATE 135 6. M u l t i v a r i a t e A n a l y s i s of the Data The p r e c e d i n g s e c t i o n s have d i s c u s s e d s e p a r a t e l y the d i f f e r e n c e s found between the s e p a r a t e l a y e r s , s i t e s , s u b s i t e s and seasons, w i t h r e s p e c t t o s p e c i e s d i s t r i b u t i o n . The da t a were complex and a s s o c i a t i o n s have been d i f f i c u l t t o v i s u a l i z e . C e r t a i n t r e n d s c o u l d be shown when examining each a s p e c t of the study s e p a r a t e l y but an o v e r a l l i n t e g r a t i o n of a l l the i n f o r m a t i o n was d i f f i c u l t . T h e r e f o r e , an o r d i n a t i o n of the samples was undertaken t o p r o v i d e a r e p r e s e n t a t i o n of t h e i r a s s o c i a t i o n s and d i f f e r e n c e s , which c o u l d be used as a guide i n i n t e r p r e t i n g the o r i g i n a l d a t a . The s p e c i e s used t o c h a r a c t e r i z e the s i t e s a r e g i v e n i n T a b l e s XXI and X X I I , a l o n g w i t h t h e i r f r e q u e n c i e s of o c c u r r e n c e , means and v a r i a n c e s . The complete d i s t r i b u t i o n ' i n f o r m a t i o n on these s p e c i e s can be e x t r a c t e d from T a b l e s I I I - IX or from the F i g u r e s f o r the c o r r e s p o n d i n g s p e c i e s i n the p r e v i o u s s e c t i o n . 136 T a b l e XXI. Frequency And V a r i a n c e Of The Most Common S p e c i e s At S i t e A TOTAL # ISOLATES FREQUENCY % MEAN VARIANCE C y l i n d r o c a r p o n tenue 139 14 57 3 46 9 .52 H e l i c o d e n d r o n t r i q l i t z i e n s i s 69 7 .23 1 72 5 05 A r t i c u l o s p o r a t e t r a c l a d i a 48 5 03 1 23 3 70 Trichoderma v i r i d e 45 4 72 1 15 3 55 P e n i c i l l i u m n i g r i c a n s 42 4 40 1 08 2 .23 P o l y s c y t a l u m fecundissimum 40 4 19 1 03 2 87 Septonema c h a e t o s p i r a 38 '3 98 0 97 2 55 C l a d o s p o r i u m c l a d o s p o r i o i d e s 34 3 56 0 87 1 17 P e n i c i l l i u m verrucosum v a r . c y c l o p i u m 28 2 94 0 72 0 79 C h a l a r a c o n s t r i c t a 26 2 73 0 67 1 70 C y l i n d r o c a r p o n didymum 24 2 41 0 62 2 35 Mucor h i e m a l i s f. h i e m a l i s 22 2 31 0 56 0 99 V a r i c o s p o r i u m elodeae 21 2 20 0 54 1 04 P e n i c i l l i u m r a i s t r i c k i i 20 2 10 0 51 1 26 P e s t a l o t i a m o n o c h a e t i o i d e s 19 ' 1 99 0 49 1 15 Sporidesmium flexum 19 1 99 0 49 1 36 Trichoderma k o n i n g i i 15 1 57 0 38 1 08 Selenophoma sp. Septonema c h a e t o s p i r a 14 '1 47 0 36 0 82 v a r . p i n i 14 1 47 0 36 0 76 L i b e r t e l l a sp. 14 1 47 • 0 36 0 82 P e n i c i l l i u m brevi-compactum 13 1 36 0 33 0 86 C h a l a r a l o n g i p e s 10 1 05 0 26 0 56 P e n i c i l l i u m c i t r i n u m 9 0 94 0 23 0 45 Acrodontium c r a t e r i f o r m e 8 0 84 0 21 0 96 K r i e q e r i a s e a v e r i 8 0 84 0 21 0 33 Endophragmia a l t e r n a t a 6 0 63 0 15 0 13 Acremonium s t r i c t u m 6 0 63 0 15 0 19 F l a g e l l o s p o r a s t r i c t a 6 0 63 0 15 0 34 H e l i c o o n fuscosporum 6 0 63 0 15 0 29 Trichoderma polysporum 6 0 63 0 15 0 45 Tripospermum m y r t i 6 0 63 0 15 0 66 V e r t i c i l l i u m b u l b i l l o s u m 6 0 63 0 15 0 19 Hyalodendron l i g n i c o l a 5 0 52 0 13 0 33 P e n i c i l l i u m notatum 5 0 52 0 13 0 17 R a m i c h l o r i d i u m subulatum 5 0 52 0 13 0 .17 137 T able X X I I . Frequency And V a r i a n c e Of The Most Common S p e c i e s At S i t e B TOTAL # ISOLATES FREQUENCY % MEAN VARIANCE C y l i n d r o c a r p o n tenue 116 16 .89 4 .38 10 .81 Trichoderma k o n i n g i i 65 9 46 2 46 8 .58 Selenophoma sp. 54 7 .86 2 08 5 .75 P e n i c i l l i u m brevi-compactum 31 4 .51 1 19 3 .52 P e n i c i l l i u m verrucosum v a r . c y c l o p i u m 31 4 .51 1 19 2 .40 G l i o c l a d i u m roseum 22 3 20 0 85 1 .26 C l a d o s p o r i u m c l a d o s p o r i o i d e s 19 2 .77 0 73 0 .84 P o l y s c y t a l u m fecundissimum 18 2 .62 0 .69 1 42 Hyalodendron l i q n i c o l a 12 1 75 0 46 1 .38 Trichoderma polysporum 11 1 60 0 42 0 89 ' H e l i c o d e n d r o n t r i g l i t z i e n s i s 10 1 46 0 38 0 41 Mucor h i e m a l i s f. h i e m a l i s 10 1 46 0 38 0 .49 C h a l a r a c y l i n d r o s p e r m a • 9 1 31 0 35 0 .48 C. l o n g i p e s 9 1 31 0 35 2 48 F l a q e l l o s p o r a s t r i c t a 8 1 16 0 31 0 30 Septogloeum sp. 8 1 16 0 31 0 54 Septonema c h a e t o s p i r a v a r . p i n i 8 1 16 0 31 0 46 P e n i c i l l i u m n i g r i c a n s 7 1 02 0 27 0 52 Acremonium s t r i c t u m 6 0 87 0 23 0 42 C h l o r i d i u m v i r e s c e n s v a r . chlamydosporum 6 0 87 0 23 0 50 M o r t i e r e l l a ramanniana v a r . ramanniana 6 0 87 0 23 0 26 P e s t a l o t i a m o n o c h a e t i o i d e s 6 0 87 0 23 0 50 L i b e r t e l l a sp. 6 0 87 0 23 0 26 Trichoderma v i r i d e 6 0 87 0 23 0 18 P e n i c i l l i u m r a i s t r i c k i i 5 0 73 0 19 0 40 Septonema c h a e t o s p i r a 5 0 73 0 19 0 24 T r i p o s p o r i u m e l e q a n s 5 0 73 0 19 0 40 C h a l a r a c o n s t r i c t a 4 0 58 0 15 0 14 Fusarium l a t e r i t i u m 3 0 44 0 12 0 11 P e n i c i l l i u m f r e q u e n t a n s 3 0 44 0 12 0 11 138 As mentioned p r e v i o u s l y , PCA was performed w i t h both the u n s e a l e d and s c a l e d o p t i o n s . S i n c e the u n s e a l e d d a t a emphasized m a i n l y the most f r e q u e n t l y i s o l a t e d s p e c i e s , the o r d i n a t i o n s tended t o show some of the obvi o u s g r o u p i n g s of s p e c i e s and d i d not show d i s t i n c t i v e g r o u p i n g s of l a y e r s , s u b s i t e s or samples. W i t h the S i t e A d a t a , C y l i n d r o c a r p o n tenue dominated the samples and had the h i g h e s t v a r i a n c e . T h i s meant t h a t the o r d i n a t i o n of samples showed v e r y l i t t l e i n the way of g r o u p i n g s of samples p o s s i b l y because t h i s s p e c i e s was found everywhere. The c l u s t e r a n a l y s i s was s i m i l a r i n i t s l a c k of d i s t i n c t i o n . At S i t e B, the t h r e e most f r e q u e n t l y i s o l a t e d s p e c i e s , C y l i n d r o c a r p o n tenue, Tr ichoderma k o n i n g i i and Selenophoma sp., combined t o i n f l u e n c e the g r o u p i n g of samples. T h i s was because each of the t h r e e were found e x c l u s i v e of the o t h e r s . They r e p r e s e n t e d the h i g h s u b s i t e , F l a y e r of the low s u b s i t e and the L l a y e r , r e s p e c t i v e l y . Because the s p e c i e s were m u t u a l l y e x c l u s i v e , t h e s e groups were r e l a t i v e l y d i s t i n c t , but were a l s o o b v i o u s from the o r i g i n a l d a t a . T h e r e f o r e , u s i n g u n s e a l e d data was h e l p f u l i n v e r i f y i n g some of the major t r e n d s i n a s s o c i a t i o n s but because they were dominated by the s p e c i e s w i t h the h i g h e s t f r e q u e n c i e s , p o s s i b l y the more s u b t l e i n f l u e n c e of the r e s t of the s p e c i e s i s o l a t e d was o v e r l o o k e d . When the data were s c a l e d , a l l of the s p e c i e s were g i v e n e s s e n t i a l l y the same importa n c e . T h i s has the e f f e c t of r e d u c i n g the i n f l u e n c e of the most f r e q u e n t l y i s o l a t e d s p e c i e s , and i n c r e a s i n g t h a t of the r a r e r s p e c i e s . Thus, a f t e r f i r s t o b s e r v i n g how the dominant s p e c i e s determined the o r d i n a t i o n s w i t h the u n s e a l e d d a t a , i t was i n s t r u c t i v e w i t h the s c a l e d data 139 to see what a s s o c i a t i o n s were apparent as a r e s u l t of the l e s s f r e q u e n t l y o c c u r r i n g s p e c i e s . In the S i t e A a n a l y s i s u s i n g s c a l e d d a t a , a m e a n i n g f u l o r d i n a t i o n of samples was o b t a i n e d i n the p l o t of p r i n c i p a l component a x i s I v s . I l l ( F i g . 4 7 ) . The f i r s t p r i n c i p a l component a c c o u n t s f o r 12% of the t o t a l v a r i a n c e , and the t h i r d , 8%. Even though t h i s i s o n l y 20% of the t o t a l v a r i a n c e , t r e n d s c o u l d be seen which shed l i g h t on the o r i g i n a l d a t a . There was a p o l a r i z a t i o n between the f i r s t two s p r i n g and summer samples (2 and 3) and the l a s t two s p r i n g and summer samples (6 and 7 ) . These two groups o v e r l a p p e d s l i g h t l y i n the lower s u b s i t e s , e s p e c i a l l y the F l a y e r . A l t h o u g h t h e s e r e p r e s e n t e d s i m i l a r times of the y e a r , they d i f f e r e d w i t h r e s p e c t t o temperature and p r e c i p i t a t i o n , and somewhat by s p e c i e s i s o l a t e d (see T a b l e s I , XVII and X I X ) . There was a g r o u p i n g of the f a l l and w i n t e r samples (4 and 5 ) , which were s i t u a t e d between the two s e t s of s p r i n g and summer samples, showing a tem p o r a l change from the i n i t i a l s a m p l i n g through t o the l a s t . An e x c e p t i o n t o t h i s were the w i n t e r samples (5) i n the h i g h and m i d d l e s u b s i t e s , L l a y e r . These were a s s o c i a t e d more w i t h the low s u b s i t e of the f i r s t s p r i n g sample (2) than the r e s t of the f a l l and w i n t e r samples. T h i s i l l u s t r a t e s the s i m i l a r i t y of the low s u b s i t e i n the s p r i n g to the h i g h s u b s i t e i n the w i n t e r , p o s s i b l y because of s i m i l a r p r e c i p i t a t i o n and d r a i n a g e c o n d i t i o n s at these t i m e s of the y e a r . A s p e c i e s o r d i n a t i o n a t S i t e A was produced by p l o t t i n g e i g e n v e c t o r I v s . e i g e n v e c t o r I I I ( F i g . 4 8 ) . There was a group toward the l e f t of the p l o t r e p r e s e n t e d by s p e c i e s 2, 28, 20, 3, 140 7, 13 and 26. These c o r r e s p o n d e d t o H e l i c o d e n d r o n  t r i q l i t z i e n s i s , F l a g e l l o s p o r a s t r i c t a , L i b e r t e l l a sp., A r t i c u l o s p o r a t e t r a c l a d i a , Septonema c h a e t o s p i r a , Var i c o s p o r ium  elodeae and Endophraqmia a l t e r n a t a . These s p e c i e s r e p r e s e n t the groups known as " a q u a t i c " and " a e r o - a q u a t i c " hyphomycetes, except f o r L i b e r t e l l a sp., S. c h a e t o s p i ra and E. a l t e r n a t a . The main c h a r a c t e r i s t i c of the " a q u a t i c " and " a e r o - a q u a t i c " hyphomycetes i s the p r o d u c t i o n of spores which are branched or o t h e r w i s e adapted f o r d i s p e r s a l i n water. L i b e r t e l l a sp. produced c u r v e d s c o l e c o s p o r e s i n s l i m y masses, which might be advantageous f o r d i s p e r s a l i n a moist environment. The o t h e r two s p e c i e s p o s s e s s e d q u i t e d i f f e r e n t s p o r e s . S. c h a e t o s p i r a produced l o n g c h a i n s of r e l a t i v e l y d r y s p o r e s . E. a l t e r n a t a a l s o produced d r y s p o r e s , but not i n c h a i n s ; t h i s s p e c i e s was i s o l a t e d i n low numbers. A l l of these s p e c i e s were prominent i n the lower s u b s i t e s , and thus i n c l o s e p r o x i m i t y t o the stream. Another g r o u p i n g of s p e c i e s was i n the lower r i g h t c o r n e r of the graph, s p e c i e s 14 and 5. These were Pen i c i11ium  r a i s t r i c k i i and P. n i g r i c a n s . They m a i n l y r e p r e s e n t e d the h i g h s u b s i t e , o c c u r r i n g i n the summer months. P o s s i b l y a s s o c i a t e d w i t h t h i s group were s p e c i e s 30, 16 and 4; Tr ichoderma  polysporum, Sporidesmium flexum and T. v i r i d e r e s p e c t i v e l y . These a l s o tended t o occur d u r i n g the summer, but not d u r i n g the e x t r e m e l y d r y times when P. r a i s t r i c k i i and P. n i g r i c a n s o c c u r r e d . These two groups were s e p a r a t e d from the a q u a t i c and a e r o - a q u a t i c group. T h i s p a t t e r n c o n t r i b u t e d t o the s e p a r a t i o n of the two s e t s of summer and s p r i n g samples i n the o r d i n a t i o n 141 of the samples p r e v i o u s l y mentioned. The t h i r d group of s p e c i e s was r e p r e s e n t e d by Selenophoma sp., C h a l a r a l o n q i p e s and C. c o n s t r i c t a (10, 22 and 18). A l l of these tended to occur d u r i n g the f a l l and w i n t e r samples, m o s t l y i n the L l a y e r , and were c h a r a c t e r i s t i c of the w i n t e r samples s e g r e g a t e d out above. In d e s c r i b i n g the o r d i n a t i o n i t appears t h a t e i g e n v e c t o r I I I p o l a r i z e d the w i n t e r and f a l l s p e c i e s from the summer and s p r i n g ones, and e i g e n v e c t o r I seemed t o s e p a r a t e the s p e c i e s o c c u r r i n g i n m o i s t e r s u b s i t e s from those o c c u r r i n g i n d r i e r ones. In comparing t h i s o r d i n a t i o n t o t h a t produced w i t h the u n s e a l e d d a t a , the same b a s i c groups composed of the more f r e q u e n t l y i s o l a t e d s p e c i e s c o u l d be seen, but added t o these groups w i t h the s c a l e d d a t a were some of the l e s s f r e q u e n t l y i s o l a t e d spec i e s . nr 142 F i g u r e 47. O r d i n a t i o n Of Samples At S i t e A. A x i s I v s . I l l ( S c a l e d ) 2 - 7 = Sample Dates (see T a b l e I ) 0 0 = High S u b s i t e O A • = L Layer A A = M i d d l e S u b s i t e © A B = F Layer • B = Low S u b s i t e " x = L i v i n g Leaves 2 • 5 0 3 0 2 A 2 4 2 0 3D 34 i 4 * 51 4 D 6D 3A ' + \ \ 7B 2 * 54 4 * 4 0 5 * 6« 64 7D 74 ' 7X 6 0 3 * 6 * I 70 / / 6» 2« I 143 F i g u r e 48. O r d i n a t i o n Of S p e c i e s At S i t e A. A x i s I v s . I I ( S c a l e d ) S p e c i e s U t i l i z e d In The A n a l y s i s 1 C y l i n d r o c a r p o n tenue 2 H e l i c o d e n d r o n t r i g l i t z i e n s i s 3 A r t i c u l o s p o r a t e t r a c l a d i a 4 Tr ichoderma v i r i d e 5 P e n i c i l l i u m n i g r i c a n s 6 P o l y s c y t a l u m fecundissimum 7 Septonema c h a e t o s p i ra 8 C l a d o s p o r ium c l a d o s p o r i o i d e s 9 P e n i c i l l i u m verrucosum v a r . c y c l o p i u m 10 C h a l a r a c o n s t r i c t a 11 C y 1 i n d r o c a r p o n didymum 12 Mucor h i e m a l i s f. h i e m a l i s 13 V a r i c o s p o r i u m elodeae 14 Pen i c i 1 1 i urn r a i s t r i c k i i 15 P e s t a l o t i a m o n o c h a e t i o i d e s 16 Sporidesmium flexum 17 Trichoderma k o n i n g i i 18 Selenophoma sp. 19 Septonema c h a e t o s p i r a v a r . p i n i 20 L i b e r t e l l a sp. 21 Pen i c i 1 1 i urn brev i-compacturn 22 C h a l a r a l o n g i p e s 23 P e n i c i l l i u m c i t r i n u r n 24 Acrodontium c r a t e r i forme 25 K r i e g e r i a s e a v e r i 26 Endophragmia a l t e r n a t a 27 A.cremonium s t r i c t u r n 28 F l a g e l l o s p o r a s t r i c t a 29 H e l i c o o n fuscosporum 30 Tr ichoderma polysporum 31 Tripospermum m y r t i 32 V e r t i c i l l i u m b u l b i l l o s u m 33 Hyalodendron l i g n i c o l a 34 Pen i c i 1 1 i u m notatum 35 R a m i c h l o r i d i u m subulatum 144 F i g u r e 48 , ' 18 . 10 1 22 35 27 33 19 29 28 26 23 20 \ 13 / 30 32 11 -f-16 ' 34 25 15 ' 14 31 24 17 21 12 v 4 I 145 S i t e B was c h a r a c t e r i z e d by many of the same f u n g i as were p r e s e n t a t S i t e A, but i n d i f f e r e n t f r e q u e n c i e s . The s p e c i e s u t i l i z e d i n the PCA a t S i t e B are l i s t e d i n Table X X I I . The p l o t of a x i s I I v s . a x i s I I I ( F i g . 49) was c o n s i d e r e d t o g i v e the most m e a n i n g f u l o r d i n a t i o n of samples u s i n g s c a l e d d a t a . These components account f o r 12% and 10% of the t o t a l v a r i a n c e , r e s p e c t i v e l y . A l t h o u g h t h i s p l o t o n l y a c c o u n t s f o r a s m a l l amount of the t o t a l v a r i a n c e ( 2 2 % ) , i t does i n d i c a t e t r e n d s and a s s o c i a t i o n s which a re h e l p f u l w i t h r e s p e c t t o the o r i g i n a l i s o l a t i o n d a t a . W i t h i n the F l a y e r , the h i g h and mi d d l e s u b s i t e s appear s i m i l a r , e s p e c i a l l y among the f a l l and w i n t e r samples (2 and 3 ) ; however, i n the L l a y e r , the h i g h s u b s i t e i s s e p a r a t e d from the low and middl e s u b s i t e s . A l s o , the samples of l i v i n g l e a v e s t e n d t o a s s o c i a t e w i t h those of the L l a y e r . T h i s i s exp e c t e d s i n c e the l e a v e s on the t r e e , a f t e r f a l l i n g , would i n i t i a l l y be found i n the L l a y e r . The p l o t of e i g e n v e c t o r s I I v s . I l l ( F i g . 50) p r o v i d e d an i n t e r p r e t a b l e s p e c i e s o r d i n a t i o n . There was a f o u r c o r n e r e d p o l a r i z a t i o n of the s p e c i e s found a t t h i s s i t e . In the upper r i g h t hand c o r n e r , s p e c i e s 6, 13, 18 and 5 tended t o group t o g e t h e r . These p o i n t s r e p r e s e n t e d G l i o c l a d i u m roseum, C h a l a r a  c y l i n d r o s p e r m a , P e n i c i l l i u m n i g r i c a n s and P. verrucosum v a r . c y c l o p i u n T r e s p e c t i v e l y . These s p e c i e s were r e p r e s e n t a t i v e of the F l a y e r , d u r i n g the f a l l and w i n t e r , thus d i s t i n g u i s h i n g these samples i n the above o r d i n a t i o n . O p p o s i t e t h i s group, i n the lower l e f t of the diagram, was a group of s p e c i e s i n c l u d i n g Selenophoma sp., P o l y s c y t a l u m fecundissimum, P e s t a l o t i a  m o n o c h a e t i o i d e s and Septogloeum sp. (3, 8, 22 and 16, 146 r e s p e c t i v e l y ) . These s p e c i e s seemed t o c h a r a c t e r i z e the L l a y e r and a l s o the l i v i n g l e a v e s , and were found m a i n l y d u r i n g the f a l l and w i n t e r . T h i s group c o r r e s p o n d s w i t h the samples from l i v i n g l e a v e s and the L l a y e r , as shown i n the o r d i n a t i o n of the samples. In the lower r i g h t c o r n e r , t h e r e were t h r e e s p e c i e s f o r m i n g a group, 23, 29 and 27; they were L i b e r t e l l a sp., Fusar i u m l a t e r i t i u m and T r i p o s p o r i u m e l e g a n s . They r e p r e s e n t e d the f i r s t s p r i n g and e a r l y f a l l samples i n the h i g h and middle s u b s i t e s , L l a y e r . In f a c t these samples were found i n the same p o s i t i o n i n F i g . 49 as the s p e c i e s i n F i g . 50. O p p o s i t e to these were a l s o s p e c i e s which o c c u r r e d i n the e a r l y f a l l ; Mucor  h i e m a l i s f. h i e m a l i s , C y l i n d r o c a r p o n tenue, C h a l a r a l o n g i p e s and F l a g e l l o s p o r a s t r i c t a (12, 1, 14 and 15) . They were more r e p r e s e n t a t i v e of the h i g h s u b s i t e , i n the L l a y e r . They a l s o seemed t o r e p r e s e n t the f a l l samples and thus c o n t r i b u t e d t o t h e i r g r o u p i n g t o g e t h e r . In comparison w i t h the u n s e a l e d d a t a , t h i s o r d i n a t i o n shows the a s s o c i a t i o n of many o t h e r s p e c i e s w i t h two of the t h r e e most f r e q u e n t l y i s o l a t e d s p e c i e s , C y l i n d r o c a r p o n tenue and Selenophoma sp.. Tr ichoderma k o n i n g i i was c e n t e r e d i n the p l o t and thus a s s o c i a t i o n s were not o b v i o u s . In summing up the r e l a t i v e worth of PCA as used i n t h i s s t u d y , I found t h a t i t - was u s e f u l as a d e s c r i p t i v e means of v i e w i n g the d a t a and thus p o s s i b l y p o i n t i n g out a s s o c i a t i o n s which l e n d themselves t o i n t e r p r e t a t i o n t h rough the o r i g i n a l d a t a . 147 F i g u r e 49. O r d i n a t i o n Of Samples At S i t e B. A x i s I I v s . I l l ( S c a l e d ) 1 - 4 = Sample Dates (see T a b l e I ) O © = High S u b s i t e O A • = L Layer A A = M i d d l e S u b s i t e • A H = F Layer • • = Low Subsite- x = L i v i n g Leaves 20 3* \ 4* 1* 3 0 4 0 / 4D V 1* 1 A 4-2* 3* 20 10 4 A X 3 ' 3D 10 2x 4x 2m 2 4 JT 148 F i g u r e 50. O r d i n a t i o n Of S p e c i e s At S i t e B. A x i s I v s . I I ( S c a l e d ) S p e c i e s U t i l i z e d In The A n a l y s i s 1 C y l i n d r o c a r p o n tenue 2 Tr ichoderma k o n i n q i i 3 Selenophoma sp. 4 P e n i c i l l i u m brevi-compactum 5 P e n i c i l l i u m verrucosum v a r . c y c l o p i u m 6 G l i o c l a d i u m roseum 7 C l a d o s p o r ium c l a d o s p o r i o i d e s 8 P o l y s c y t a l u m f e c u n d i ssimum 9 Hyalodendron l i q n i c o l a 10 Tr ichoderma polysporum 11 H e l i c o d e n d r o n t r i g l i t z i e n s i s 12 Mucor h i e m a l i s f. h i e m a l i s 13 C h a l a r a c y l i n d r o s p e r m a 14 C h a l a r a l o n g i p e s 15 F l a g e l l o s p o r a s t r i c t a 16 Septogloeum sp. 17 Septonema c h a e t o s p i ra v a r . p i n i e 18 Pen i c i 1 1 i um n i g r i c a n s 19 Acremonium s t r i c t u m 20 C h l o r i d i u m v i r e s c e n s v a r . chlamydospor'um 21 Mort i e r e l l a ramann i a n a v a r . ramanniana 22 P e s t a l o t i a monochaet i o i d e s 23 L i b e r t e l l a sp. 24 Tr ichoderma v i r i d e 25 P e n i c i l l i u m r a i s t r i c k i i 26 Septonema c h a e t o s p i r a 27 T r i p o s p o r i u m e l e g a n s 28 C h a l a r a c o n s t r i c t a 29 Fusarium l a t e r i t i u m 30 P e n i c i l l i u m f r e q u e n t a n s 149 F i g u r e 50 / 12 14 \ 17 1 5 / 13 18 5 ' 28 26 19 11 + 10 20 25 22 24 9 30 21 150 IV. D i s c u s s i o n and C o n c l u s i o n s In the p r e s e n t s t u d y , f u n g i i n v o l v e d i n the decay of r e d cedar l i t t e r were examined. The fr e q u e n c y of i s o l a t i o n of f u n g i from the b r a n c h l e t s i n the l i t t e r l a y e r s has been taken as an i n d i c a t i o n of t h e i r r e l a t i v e i nvolvement i n decay. Granted t h a t t h i s i n d i r e c t measure of a c t i v i t y i s s u b j e c t t o a number of v a r i a b l e s , such as i s o l a t i o n t e c h n i q u e and media employed, i t n o n e t h e l e s s p r o v i d e s a b a s i s f o r some g e n e r a l i n d i c a t i o n s of the a s s o c i a t i o n s of these f u n g i . The i n f l u e n c e of m o i s t u r e upon the d i s t r i b u t i o n of s p e c i e s was o n l y i n d i r e c t l y examined. That i s , p r e c i p i t a t i o n d a t a was o b t a i n e d r e l a t i v e t o c o l l e c t i o n d a t e s . 1. D i s c u s s i o n of the S u b s i t e s As mentioned p r e v i o u s l y , the study s i t e s were d i v i d e d i n t o t h r e e s u b s i t e s , i n d i c a t i n g t h e i r r e l a t i v e p o s i t i o n i n r e l a t i o n t o a nearby stream. The d i v i s i o n s were a r t i f i c i a l and s u b j e c t i v e , but were employed to see i i t h e r e were d i f f e r e n c e s i n the mycobiota w i t h s l i g h t p o s i t i o n a l changes i n r e l a t i o n t o the stream. The s p e c i e s i s o l a t e d were used t o c h a r a c t e r i z e each sample, and the samples were grouped t h r o u g h a PCA o r d i n a t i o n . The r e s u l t s of t h i s a n a l y s i s have been p r e s e n t e d i n the f o r e g o i n g s e c t i o n . In the a n a l y s i s of the d a t a , the s u b s i t e s were not a s s o c i a t e d i n the same e x a c t groups i n each of the main s i t e s . 151 The s u b s i t e s r e p r e s e n t e d a g r a d a t i o n away from the stream and thus toward somewhat d r i e r c o n d i t i o n s . I t seemed r e a s o n a b l e t o expect the two extreme s u b s i t e s , low and h i g h , t o be more d i s t i n c t , w i t h the m i d d l e s u b s i t e a s s o c i a t e d a t t i m e s w i t h one or the o t h e r . In some r e s p e c t s t h i s was the c a s e , as shown by the c l u s t e r a n a l y s i s and PCA of the d a t a from both s i t e s . Samples from the h i g h and low s u b s i t e s were more or l e s s d i s t i n g u i s h e d as s e p a r a t e groups, w i t h samples from the m i d d l e s u b s i t e d i s t r i b u t e d between these two. At S i t e A, t h e m i d d l e s u b s i t e was more f r e q u e n t l y grouped w i t h the h i g h . At S i t e B, the m i d d l e s u b s i t e was a s s o c i a t e d e q u a l l y between the low and h i g h . 2. D i s c u s s i o n of the L i t t e r L a y e r s In an attempt t o d e t e r m i n e what e f f e c t s p a t i a l d i f f e r e n c e s had on the f u n g a l p o p u l a t i o n , the s p e c i e s c o m p o s i t i o n i n the L and F l a y e r s was a l s o examined. The f u n g i p r e s e n t on l i t t e r i n these l a y e r s were r e p r e s e n t a t i v e of t h e i r p o s i t i o n i n the time sequence of decay; the L l a y e r s p e c i e s b e i n g m o s t l y e a r l y c o l o n i z e r s , and the F l a y e r s p e c i e s r e p r e s e n t i n g those c o l o n i z i n g a t l a t e r s t a g e s of decay. In examining the o r d i n a t i o n s d i s c u s s e d p r e v i o u s l y , the r e l a t i v e s e p a r a t i o n of the samples i n t o t h e s e two l a y e r s was d e t e r m i n e d . At S i t e A, s e p a r a t i o n of l a y e r s was not v e r y s t r o n g ; t h e r e was a g r o u p i n g of the F l a y e r i n the s p r i n g and summer samples. There were o t h e r s c a t t e r e d g r o u p i n g s of samples from 152 the same l a y e r , but n o t h i n g v e r y d i s t i n c t i v e . The in c o m p l e t e s e p a r a t i o n of the two l a y e r s was even more e v i d e n t i n the c l u s t e r a n a l y s i s . On the o t h e r hand, a t S i t e B, both the u n s e a l e d and s c a l e d d a t a , and a l s o the c l u s t e r a n a l y s i s , d e p i c t e d a r e a s o n a b l e s e p a r a t i o n of the two l a y e r s . When i n v e s t i g a t i n g l i t t e r decay f u n g i , most workers have found the d i v i s i o n i n t o the L and F l a y e r s r e p r e s e n t e d u n i t s d i s t i n g u i s h a b l e by the f u n g a l s p e c i e s p r e s e n t (Borowska, 1966; Bran d s b e r g , 1969; K e n d r i c k , 1963; K e n d r i c k and Burges, 1962; Verona and R a m b e l l i , 1972; V i s s e r and P a r k i n s o n , 1975; Watson et. a l . , 1974; Wicklow and Whittingham, 1974, 1978; Widden and P a r k i n s o n , 1973). In the p r e s e n t s t u d y , the s p e c i e s c h a r a c t e r i s t i c of the d i f f e r e n t l a y e r s a re l i s t e d i n T a b l e s XVII and X V I I I . Many of these s p e c i e s have not been r e p o r t e d p r e v i o u s l y from l i t t e r s t u d i e s ; of those t h a t have, o c c u r r e n c e i n the r e s p e c t i v e l a y e r s i s c o n s i s t e n t w i t h what has been r e p o r t e d . Some of the s p e c i e s found i n the L l a y e r a t one s i t e , were found i n the F l a y e r at the o t h e r s i t e . T h i s was t r u e of Hyalodendron l i g n i c o l a , Mucor h i e m a l i s f. h i e m a l i s and Acremonium s t r i c t u m . I t i s p r o b a b l e t h a t these r e p r e s e n t s p e c i e s which can be c o n s i d e r e d t o occur e q u a l l y i n both l a y e r s . The d i f f e r e n c e s may r e f l e c t the b a s i c d i f f e r e n c e s between the s i t e s . For the most p a r t , t h i s d i v i s i o n of s p e c i e s between the l a y e r s r e p r e s e n t s a d e f i n i t e e c o l o g i c a l tendency; some s p e c i e s a re found p r e d o m i n a n t l y i n the L l a y e r , w h i l e o t h e r s a re found p r e d o m i n a n t l y i n the F l a y e r . S i n c e many of the s p e c i e s have not .been r e p o r t e d p r e v i o u s l y from l i t t e r , t h e i r i n c l u s i o n here adds 153 more i n f o r m a t i o n t o the e c o l o g y of the s e s p e c i e s . , 3. D i s c u s s i o n of S e a s o n a l i t y The samples were a n a l y z e d r e g u l a r l y throughout the study i n ord e r t o determine the s e a s o n a l d i f f e r e n c e s among them. In the o r d i n a t i o n s a l r e a d y p r e s e n t e d , some of the samples .were o f t e n c l u s t e r e d s e a s o n a l l y , r e g a r d l e s s of t h e i r l a y e r or s u b s i t e . T h i s was e s p e c i a l l y t r u e w i t h the f a l l and w i n t e r samples, and the s p r i n g and summer samples. The g r o u p i n g t o g e t h e r of these seasons seemed r e a s o n a b l e as they r e p r e s e n t major c l i m a t i c d i f f e r e n c e s d u r i n g the y e a r . Many of the s p e c i e s i s o l a t e d can a l s o be grouped t o g e t h e r on the b a s i s of t h e i r s e a s o n a l o c c u r r e n c e s . The g r o u p i n g s have been l i s t e d i n the p r e v i o u s s e c t i o n on s e a s o n a l i t y i n the L and F l a y e r s . Other workers have r e p o r t e d some of the s p e c i e s t o show the same s e a s o n a l d i s t r i b u t i o n . S p e c i e s o c c u r r i n g d u r i n g s p r i n g were Acremonium s t r i c t u r n , Tr ichoderma v i r i d e and P o l y s c y t a l u m f e c u n d i ssimum (Katz and L i e t h , 1974). The summer group i n c l u d e d C y l i n d r o c a r p o n didymum (Badurowa and Badura, 1967), Tr ichoderma v i r i d e ( G o u r b i e r e , 1979; Ruscoe, 1971; V i s s e r and P a r k i n s o n , 1975), Pen i c i 1 1 i u m n i g r i c a n s ( M a r t i n e z and Ramirez, 1972 ) and P. r a i s t r i c k i i ( G o u r b i e r e , 1979). The s p e c i e s found d u r i n g the f a l l and w i n t e r were Mucor h i e m a l i s f. h i e m a l i s ( H e r i n g , 1965; G o u r b i e r e , 1979; M a r t i n e z and Ramirez, 1972), P e n i c i 1 1 i u m brevi-compactum and P. notatum (Badura and Badurowa, 1964; Badurowa and Badura, 1967; M a r t i n e z and Ramirez, 1972), 154 P. c i t r i n u m (Yokoyama et a l . , 1977), P. f r e q u e n t a n s and M o r t i e r e l l a ramanniana v a r . ramanniana (Badurowa and Badura, 1967; G o u r b i e r e , 1979), C h a l a r a c y l i n d r o s p e r m a (Hogg and Hudson, 1966) and Hyalodendron 1 i g n i c o l a (Katz and L i e t h , 1974; Tubaki and Yokoyama, 1971). Some of these workers have r e p o r t e d d i f f e r e n t s e a s o n a l i t i e s f o r a few of the s p e c i e s , but t h i s may r e f l e c t o n l y d i f f e r e n c e s i n c l i m a t e s and s u b s t r a t a . S i n c e most of the s p e c i e s i s o l a t e d i n t h i s study have not been r e p o r t e d p r e v i o u s l y from l i t t e r , t h e i r o c c u r r e n c e s w i t h r e s p e c t t o seasons has not been d i s c u s s e d p r e v i o u s l y . Of these s p e c i e s , those i s o l a t e d d u r i n g the s p r i n g and summer were A r t i c u l o s p o r a t e t r a c l a d i a , Septonema c h a e t o s p i r a , F l a g e l l o s p o r a  s t r i c t a , Sporidesmium fle x u m , H e l i c o d e n d r o n t r i g l i t z i e n s i s , V a r i c o s p o r i u m elodeae and V e r t i c i l l i u m b u l b i l l o s u m . S p e c i e s p r e s e n t i n the f a l l and w i n t e r were P e s t a l o t i a m o n o c h a e t i o i d e s and T r i p o s p o r i u m e l e g a n s . One s p e c i e s , C h a l a r a c o n s t r i c t a , was found i n the f a l l and w i n t e r i n the L l a y e r , and i n the -spring i n the F l a y e r . 4. S p e c i e s A s s o c i a t i o n s In the p r o c e s s of a n a l y z i n g the da t a t o determine the a s s o c i a t i o n s of the samples, an o r d i n a t i o n was a l s o made of the s p e c i e s used t o c h a r a c t e r i z e the samples. The groups of s p e c i e s , mentioned i n the p r e v i o u s s e c t i o n , were based on s i m i l a r d i s t r i b u t i o n s of the s p e c i e s i n the s u b s i t e s and l a y e r s . At S i t e A, u n s e a l e d d a t a , the group of C y l i n d r o c a r p o n didymum, 155 P e n i c i l l i u m n i g r i c a n s , P. r a i s t r i c k i i and A r t i c u l o s p o r a  t e t r a c l a d i a , r e p r e s e n t e d the summer s p e c i e s group t h a t were a l s o found p r e d o m i n a n t l y i n the F l a y e r . These s p e c i e s form a b i o l o g i c a l u n i t i n d i c a t i v e o f , and f a v o r e d by, dry c o n d i t i o n s . P. n i g r i c a n s , P. r a i s t r i c k i i and C. didymum have o f t e n been found under low m o i s t u r e c o n d i t i o n s ( B i s s e t t and P a r k i n s o n , 1979a; Gochenaur and Backus, 1967). A. t e t r a c l a d i a i s u s u a l l y thought of as an i n h a b i t o r of a q u a t i c e n v i r o n m e n t s , but i t has a l s o been found t o s u r v i v e w e l l i n t e r r e s t r i a l s i t u a t i o n s (Bandoni, 1972, 1977; Sanders and Webster, 1978; Webster, 1977). Another group of s p e c i e s , H e l i c o d e n d r o n t r i g l i t z i e n s i s , P o l y s c y t a l u m f e c u n d i ss imum and C h a l a r a c o n s t r i c t a , were s p r i n g and summer s p e c i e s i n the L l a y e r . Thus a c o m b i n a t i o n of c h a r a c t e r i s t i c s has been u t i l i z e d t o p r o v i d e s p e c i e s a s s o c i a t i o n s , p a r t i c u l a r l y , s e a s o n a l i t y and o c c u r r e n c e i n one or the o t h e r l a y e r . The t h i r d major group at t h i s s i t e was s t r i c t l y the p r o d u c t of p o s i t i o n , o c c u r r i n g i n the F l a y e r , i n the low s u b s i t e . The s p e c i e s i n v o l v e d were Septonema c h a e t o s p i r a , V a r i c o s p o r i u m elodeae and Tr ichoderma polysporum. V. e l o d e a e , l i k e A. t e t r a c l a d i a , i s c o n s i d e r e d a member of the a q u a t i c hyphomycetes, but i t too has been shown t o e x i s t i n dry t e r r e s t r i a l c o n d i t i o n s (Bandoni, 1972, 1977; Bessey, 1939; G o u r l e y , 1969; Sanders and Webster, 1978; Waid, 1954; Webster, 1961, 1977; W i l l i a m s and S c h m i t t h e n n e r , 1956). When' the o r d i n a t i o n was performed u s i n g s c a l e d d a t a , the groups of s p e c i e s were based a g a i n on d i f f e r e n t p arameters. One group was s e p a r a t e d because of a h i g h f r e q u e n c y of o c c u r r e n c e i n the low s u b s i t e . Another r e p r e s e n t e d summer s p e c i e s , and t h i s 156 group was d i v i d e d i n t o the extreme dry t i m e s (sample 7; P e n i c i l l i u m n i g r i c a n s and P. r a i s t r i c k i i ) and the l e s s extreme (sample 3; Trichoderma polysporum, T. v i r i d e and Sporidesmium  f l e x u m ) . T h i s group was s i m i l a r l y d i s t i n g u i s h e d as such i n the u n s e a l e d o r d i n a t i o n . The l a s t group i n t h i s o r d i n a t i o n was of f a l l and w i n t e r o c c u r r i n g s p e c i e s , i n the L l a y e r , r e p r e s e n t e d by • C h a l a r a c o n s t r i c t a , C. l o n g i p e s and Selenophoma sp. I t f o l l o w s t h a t Selenophoma sp. would be found more f r e q u e n t l y i n the f a l l and w i n t e r s i n c e i t i s p y e n i d i a l , thus an e a r l y c o l o n i z e r , and much of the l e a f f a l l i s d u r i n g the f a l l of the year (Dimock, 1958; K e n d r i c k , 1959). As a l r e a d y s t a t e d , S i t e B was s l i g h t l y d i f f e r e n t from S i t e A i n s p e c i e s i s o l a t e d . T h e r e f o r e the o r d i n a t i o n of s p e c i e s a t S i t e B d i d not p r o v i d e the same a s s o c i a t i o n s of s p e c i e s as a t S i t e A. With the d a t a u n s e a l e d , two of the groups were s e a s o n a l l y d i s t i n c t . These were P e n i c i l l i u m brevi-compactum and P o l y s c y t a l u m fecundissimum, i n the low s u b s i t e s d u r i n g the f a l l , and P e n i c i l l i u m verrucosum v a r . c y c l o p i u m and G l i o c l a d i u m  roseum, i n the F l a y e r d u r i n g the w i n t e r . A l t h o u g h these s p e c i e s are not r e p o r t e d f r e q u e n t l y i n l i t t e r decay s t u d i e s , the i n f o r m a t i o n a v a i l a b l e tends t o support t h e s e d i s t r i b u t i o n s (Badura and Badurowa, 1964; Hogg and Hudson, 1966; Widden and P a r k i n s o n , 1973). The o t h e r s p e c i e s were s e p a r a t e d on p o s i t i o n a l grounds; Trichoderma k o n i n g i i , an F l a y e r s p e c i e s i n the low s u b s i t e , and Selenophoma sp., an L l a y e r s p e c i e s i n the h i g h s u b s i t e . When the d a t a were s c a l e d , the groups remained much the same, but more s p e c i e s were added. The w i n t e r group above g a i n e d 157 s p e c i e s ( C h a l a r a cy1indrosperma and P e n i c i l l i u m n i g r i c a n s ) , and were r e p r e s e n t a t i v e of f a l l and w i n t e r s p e c i e s i n the F l a y e r . I t i s i n t e r e s t i n g t h a t P. n i g r i c a n s was i s o l a t e d more f r e q u e n t l y i n the f a l l and w i n t e r at t h i s s i t e and not i n the summer, as was the case a t S i t e A. T h i s anomaly may be s i m p l y the r e s u l t of o v e r e m p h a s i z i n g the importance of t h i s s p e c i e s a t S i t e B, s i n c e i t o c c u r r e d i n r e l a t i v e l y low numbers t h e r e . The f a l l group l o s t one s p e c i e s (Pen i c i 1 1 i um brevi-compactum) and combined w i t h Selenophoma sp. and o t h e r s p e c i e s • (Septogloeum sp. and P e s t a l o t i a m o n o c h a e t i o i d e s ) t o r e p r e s e n t f a l l and w i n t e r s p e c i e s i n the L l a y e r . These s p e c i e s a l l r e p r e s e n t p r i m a r y c o l o n i z e r s of l e a v e s , and thus t h i s g r o u p i n g i s not unexpected. The o t h e r two a s s o c i a t i o n s were not shown i n the PCA of the u n s e a l e d d a t a . These were s p e c i e s from the f i r s t s p r i n g and f a l l samples i n the L l a y e r of the h i g h and m i d d l e s u b s i t e s ( L i b e r t e l l a sp., F u s a r i u m l a t e r i t i u m and T r i p o s p o r i u m e l e g a n s ) , and f a l l s p e c i e s i n the L l a y e r of the h i g h s u b s i t e (Mucor h i e m a l i s f. hiemalis-, C y l i n d r o c a r p o n tenue, C h a l a r a l o n g i p e s and F l a g e l l o s p o r a  s t r i c t a ) . The s p e c i e s i n the f i r s t group were found i n low f r e q u e n c i e s and t h e i r emphasis may have been a r e s u l t of the i n c r e a s e d importance g i v e n them by s t a n d a r d i z i n g the d a t a . The l a t t e r group r e p r e s e n t s s p e c i e s more f r e q u e n t l y encountered i n the h i g h s u b s i t e , and may be good i n d i c a t o r s of t h a t a r e a . In an o v e r a l l view of s p e c i e s a s s o c i a t i o n s based on s i m i l a r i t i e s of d i s t r i b u t i o n i n the v a r i o u s l a y e r s , s u b s i t e s and seasons, the o r d i n a t i o n method (PCA) p r o v i d e s i n d i c a t i o n s of groups t h a t can be pursued t h r o u g h the raw d a t a . By combining the i n f o r m a t i o n from both major s i t e s , s e v e r a l groups of s p e c i e s 158 emerge which appear t o r e p r e s e n t p o s s i b l e e c o l o g i c a l u n i t s . The f i r s t group of s p e c i e s a r e r e p r e s e n t a t i v e of low s u b s i t e s and thus moist c o n d i t i o n s . These i n c l u d e H e l i c o d e n d r o n  t r i q l i t z i e n s i s , A r t i c u l o s p o r a t e t r a c l a d i a , V a r i c o s p o r i u m  e l o d e a e , F l a q e l l o s p o r a s t r i c t a , P o l y s c y t a l u m f e c u n d i ssimum, Septonema c h a e t o s p i ra and Endophragmia a l t e r n a t a . As mentioned p r e v i o u s l y , the f i r s t f o u r s p e c i e s r e p r e s e n t a q u a t i c - t y p e hyphomycetes, a l t h o u g h they a re not unknown from t e r r e s t r i a l s i t u a t i o n s . The l a s t t h r e e s p e c i e s have not been r e p o r t e d f r e q u e n t l y i n l i t t e r s t u d i e s , and t h e i r i n c l u s i o n here i s new. A second group i n c l u d e s Tr ichoderma v i r i d e , Pen i c i11ium  n i g r i c a n s , P. r a i s t r i c k i i , T. polysporum and Sporidesmium  f1exurn. These s p e c i e s were i n d i c a t i v e of summer c o n d i t i o n s , and P. n i g r i c a n s and P. r a i s t r i c k i i were both prominant d u r i n g e x t r e m e l y d r y p e r i o d s . Another a s s o c i a t i o n of s p e c i e s i n c l u d e d Selenophoma sp., P e s t a l o t i a monochaet i o i d e s , Septogloeum sp., Pen i c i 1 1 i u m b r e v i - compactum, Mucor h i e m a l i s f. h i e m a l i s, C h a l a r a c o n s t r i c t a and C. l o n g i p e s . These were r e p r e s e n t a t i v e of the L l a y e r s . The f i r s t t h r e e a r e p y c n i d i a l and thus i m p o r t a n t p r i m a r y c o l o n i z e r s . P. brevi-compactum and M. h i e m a l i s f. h i e m a l i s a re common l i t t e r f u n g i . C. c o n s t r i c t a and C,. l o n g i p e s have not p r e v i o u s l y been r e p o r t e d i n l i t t e r s t u d i e s . A f i n a l group d i d not i n c l u d e many s p e c i e s , but they d i d tend t o occur under s i m i l a r c i r c u m s t a n c e s , u s u a l l y i n the F l a y e r . They were P e n i c i l l i u m verrucosum v a r . c y c l o p i u m , G I i o c l a d i u m roseum and C h a l a r a c y 1 i n d r o s p e r m a . The l a t t e r s p e c i e s was not r e p o r t e d from l i t t e r b e f o r e , and the former two 159 a r e common s o i l and l i t t e r f u n g i . 5. Summary In c o n c l u s i o n , t h i s study has p r o v i d e d f u r t h e r i n s i g h t i n t o the complex p r o c e s s of l i t t e r decay and the f u n g i i n v o l v e d , i n a unique environment dominated by Thuja p i i c a t a Donn. The s t a n d a r d t e c h n i q u e f o r d e t e r m i n i n g t h i s has been t o examine the o c c u r r e n c e and d i s t r i b u t i o n of s p e c i e s , u s i n g v a r i o u s i s o l a t i o n t e c h n i q u e s ( L i n d s e y , 1973; P a r k i n s o n , 1973). The f u n g i i n v o l v e d i n the decay were i s o l a t e d and i d e n t i f i e d . Many of these s p e c i e s have not been r e p o r t e d p r e v i o u s l y i n s o i l and l i t t e r s t u d i e s . T h e i r o c c u r r e n c e here may be a r e s u l t of the t e c h n i q u e s employed. S t u d i e s on l i t t e r f u n g i u s i n g the m o d i f i e d d i l u t i o n p l a t e t e c h n i q u e , have r e s u l t e d i n a predominance of s p e c i e s of P e n i c i l l i u m , Mucor and Tr ichoderma ( B r a n d s b e r g , 1969; Verona and R a m b e l l i , 1972). A c o m b i n a t i o n of washing l i t t e r and then u s i n g the d i l u t i o n p l a t e method, as used by V i s s e r and P a r k i n s o n (1975), r e s u l t e d i n a predominance of P e n i c i l l i u m spp., Tr ichoderma spp. and Phoma spp. Other workers who u t i l i z e d washing t e c h n i q u e s and c u l t u r e d the washed l i t t e r on s e l e c t i v e media or i n moist chambers, r e p o r t e d a reduced involvement of P e n i c i l l i u m spp., w i t h Tr ichoderma spp., Mort i e r e l l a spp., Mucor spp. and p r i m a r y c o l o n i z e r s such as p y c n i d i a l s p e c i e s , p r e d o m i n a n t i n g ( G o u r b i e r e , 1979; K e n d r i c k and Burges, 1962; Watson e t a l . , 1974). In the p r e s e n t study the predominant s p e c i e s was C y l i n d r o c a r p o n tenue, which has not been r e p o r t e d 160 o f t e n from l i t t e r s t u d i e s . Tr ichoderma spp. (T. v i r ide and T. k o n i n g i i ) and P e n i c i l l i u m spp. (P. n i g r i c a n s , P. verrucosum v a r . c y c l o p i u m and P. brevi-compacturn) were a l s o found t o occur w i t h h i g h f r e q u e n c i e s . A p y c n i d i a l s p e c i e s , Selenophoma sp., was a l s o i s o l a t e d as a p r i m a r y c o l o n i z e r . Two s p e c i e s , H e l i c o d e n d r o n  t r i g l i t z i e n s i s and A r t i c u l o s p o r a t e t r a c l a d i a , were i s o l a t e d w i t h h i g h f r e q u e n c i e s . These s p e c i e s have been r e p o r t e d o c c a s i o n a l l y as o c c u r r i n g on l i t t e r , but have never been found i n s t u d i e s on l i t t e r decay f u n g i . T h e i r o c c u r r e n c e here i n d i c a t e s t h e i r p r o b a b l e a c t i v e involvement i n the decay p r o c e s s . Many of the r e m a i n i n g s p e c i e s have not been i s o l a t e d b e f o r e i n l i t t e r decay s t u d i e s . The t e c h n i q u e s used i n t h i s s tudy ( i . e . washing l i t t e r and then p u t t i n g i t on SYT p l a t e s o r i n moist chambers) p r o b a b l y g i v e a good i n d i c a t i o n of the f u n g i i n v o l v e d i n the decay. S i n c e i s o l a t e s were taken from the f i r s t two sp o t s i n the SYT p l a t e s and m a i n l y a f t e r the f i r s t week of i n c u b a t i o n of the moist chambers, the f u n g i c o u l d be c o n s i d e r e d a c t i v e l y growing i n the l i t t e r . The d i f f e r e n c e s between the v a r i o u s s u b s i t e s were s l i g h t , but d i d tend t o r e p r e s e n t a g r a d i e n t between the low and h i g h s u b s i t e s . The L and F l a y e r s were o f t e n more d i s t i n c t i n the s p e c i e s i s o l a t e d , and r e p r e s e n t d i f f e r e n c e s i n the involvement of f u n g i i n the d i f f e r e n t s t a g e s of decay. As w i t h s p a t i a l d i f f e r e n c e s , s e a s o n a l changes i n the o c c u r r e n c e of s p e c i e s were s l i g h t but r e p r e s e n t e d a g r a d i e n t between f a l l and w i n t e r , and s p r i n g and summer. The d i f f e r e n c e s i n the summer were o f t e n extreme. The common denominator among a l l these d i f f e r e n c e s 161 seems t o be m o i s t u r e a l t h o u g h s u p p o r t a t i v e d a t a a re not a v a i l a b l e . The s l i g h t d i f f e r e n c e s i n d i s t a n c e from, and e l e v a t i o n above a stream, or s e a s o n a l d i f f e r e n c e s i n p r e c i p i t a t i o n , a r e deemed t o be r e s p o n s i b l e f o r s l i g h t but d i s c e r n i b l e changes i n the o c c u r r e n c e of s p e c i e s . The a s s o c i a t i o n s of s p e c i e s , shown through PCA,suggest a number of s t a r t i n g p o i n t s f o r e x p e r i m e n t a l r e s e a r c h i n t o the n a t u r e of i n t e r a c t i o n s among s p e c i e s i n v o l v e d i n the decay p r o c e s s . W h i l e these a s s o c i a t i o n s do not c h a r a c t e r i z e s p e c i f i c d i f f e r e n c e s i n s u b s i t e s , seasons or l a y e r s , they do r e p r e s e n t a c o m b i n a t i o n of these f a c t o r s . As a r e s u l t , the s p e c i e s a s s o c i a t i o n s r e p r e s e n t groups which p r o b a b l y f u n c t i o n t o g e t h e r i n the environment. 162 V. B i b l i o g r a p h y Anderson, J.M, and A. Macfadyn. 1976. The r o l e of t e r r e s t r i a l  and aquat i c organ i sms i n decomposit i o n p r o c e s s e s . B l a c k w e l l S c i e n t i f i c . O x f o r d . 474p. Anderson, J.P.E., and K.H. Domsch. 1973. 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