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Taxonomic and ecologic aspects of zoosporic fungi in coastal and steppe soils Booth, Thomas 1971

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TAXONOMIC AND ECOLOGIC ASPECTS OF ZOOSPORIC FUNGI I N COASTAL AND STEPPE SOILS  by  TOM BOOTH B.A., E a s t e r n B a p t i s t C o l l e g e M.S., Ohio U n i v e r s i t y  A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY i n t h e Department of Botany  i We a c c e p t t h i s t h e s i s as conforming  to the  required standard  THE UNIVERSITY OF BRITISH COLUMBIA April  >  1  9  7  1  In p r e s e n t i n g t h i s  thesis  an advanced degree at the L i b r a r y I  for  freely  f u l f i l m e n t , o f the of B r i t i s h  available  for  requirements  Columbia, I agree  for  that  r e f e r e n c e and s t u d y .  t h a t p e r m i s s i o n f o r e x t e n s i v e copying o f t h i s  thesis  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 of  the U n i v e r s i t y  s h a l l make i t  f u r t h e r agree  in p a r t i a l  this  representatives. thesis  for  It  financial  i s understood that copying o r p u b l i c a t i o n gain s h a l l  written permission.  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 Vancouver 8, Canada  Apri-139,  mi  not  be allowed without my  ABSTRACT  C e r t a i n c r i t e r i a used t o d i s t i n g u i s h c h y t r i d taxa a r e h i g h l y v a r i a b l e and u n s t a b l e .  This v a r i a b i l i t y  i s evident  from a study  of t e n s i n g l e - s p o r e i s o l a t e s of an E n t o p h l y c t i s s p e c i e s c o l l e c t e d i n Western Canada, Oregon, C a l i f o r n i a and Nevada. v a r i a t i o n s place  The observed  these E n t o p h l y c t i s i s o l a t e s i n d i f f e r e n t  species,  g e n e r a , s u b f a m i l i e s , f a m i l i e s , and s e r i e s , thus demonstrating various  current  taxonomic d i s p o s i t i o n s and concepts are of  that  questionable  value. Zoosporic  f u n g i , mainly C h y t r i d i a l e s , a r e w i d e l y  c o a s t a l and steppe s o i l s . determination  distributed i n  Based on d i s t r i b u t i o n a l r e c o r d s and  o f p h y s i c a l and chemical  parameters f o r each  soil  c o l l e c t i o n , eleven  c h y t r i d and c h y t r i d i a c e o u s s p e c i e s are d i v i s i b l e  i n t o f o u r groups:  a) o b l i g a t e l y marine, b) marine, c) f a c u l t a t i v e l y  marine and d) marine o c c a s i o n a l s . problematic of  C e r t a i n aspects  o f t h i s study are  which emphasizes the n e c e s s i t y o f a more s o p h i s t i c a t e d l i n e  approach. Temperature-salinity  single-spore  growth responses o f f i f t y - s e v e n a x e n i c  cultures of zoosporic  f u n g i , m a i n t a i n e d under s i m i l a r  c o n t r o l l e d c o n d i t i o n s f o r a minimum o f s i x months, c o r r e l a t e w i t h habitat  types and c e r t a i n e n v i r o n m e n t a l parameters.  These r e s p o n s e s ,  s i m i l a r over s e v e r a l i s o l a t e s of a s p e c i e s from the same s o i l c o l l e c t i o n , are temporally  constant  and, thus, a r e e c o t y p i c .  iii  TABLE OF CONTENTS  1.  Introduction  2.  P r o b l e m a t i c a l Taxonomic C r i t e r i a i n t h e C h y t r i d i a l e s : Comparative Morphology of Ten E n t o p h l y c t i s s p . I s o l a t e s  3.  4.  5.  . . . .  O c c u r r e n c e and D i s t r i b u t i o n o f Z o o s p o r i c F u n g i Devon I s l a n d , Canadian E a s t e r n A r c t i c  1  . .  7  . . .  36  .  63  from  O c c u r r e n c e and D i s t r i b u t i o n o f Some Z o o s p o r i c F u n g i from S o i l s o f Hibben and Morseby I s l a n d s , Queen Charlotte Islands . . . . . . . O c c u r r e n c e and D i s t r i b u t i o n of Z o o s p o r i c F u n g i and Some A c t i n o m y c e t a l e s i n C o a s t a l S o i l s o f Southwestern B r i t i s h Columbia and the San J u a n I s l a n d s . . . . . . .  105  6.  O c c u r r e n c e and D i s t r i b u t i o n o f C h y t r i d s , C h y t r i d i a c e o u s F u n g i and some A c t i n o m y c e t a l e s from S o i l s o f Oregon, C a l i f o r n i a and Nevada . . . . . . . . . . . 137  7.  Occurrence of ,,Chytrids and C h y t r i d i a c e o u s Long I s l a n d , N.Y. Beaches  8.  9.  10.  171  O c c u r r e n c e of C h y t r i d s and C h y t r i d i a c e o u s F u n g i from Some Mexican Beach and D e s e r t S o i l s  175  D i s t r i b u t i o n of C e r t a i n S o i l I n h a b i t i n g C h y t r i d and C h y t r i d i a c e o u s S p e c i e s R e l a t e d to some P h y s i c a l and Chemical F a c t o r s  178  E c o t y p i c Responses of C h y t r i d and C h y t r i d i a c e o u s to V a r i o u s  11.  Fungi from Two  Species  S a l i n i t y and Temperature Combinations  Discussion  203 . . . . . .  Literature Cited  230  Appendices I. II. III. IV.  V.  226  •  253  Habitat descriptions S p e c i e s name and numbers H a b i t a t names and numbers A. Times each s p e c i e s occurs a t 0 ° / and 7 ° / i n a l l s i t e s and s t a r t s of each h a b i t a t type . B. O c c u r r e n c e and frequency o f each s p e c i e s i n t e n Base Camp Lowland h a b i t a t s . . . . . . . . Number of d i f f e r e n t s p e c i e s r e c o v e r e d a t 0 ° / and 7 ° / i n each h a b i t a t type o o  253 267 271  0 0  273 350  o o  0 0  355  VI.  VII.  Computed o c c u r r e n c e s , f r e q u e n c i e s and d i s t r i b u t i o n a l i n t e n s i t y i n d e x o f each s p e c i e s a t 0 loo and 7 /oo i n h a b i t a t types . . . . . . . H a b i t a t s and v a l u e s o f p h y s i c a l and parameters  VIII. IX.  X.  XI.  chemical  of c o l l e c t i o n 'sites  454  S t a t i s t i c s o f p h y s i c a l and c h e m i c a l f a c t o r s in habitats Ranges, means, v a r i a n c e s , s t a n d a r d d e v i a t i o n s and f i v e p e r c e n t c o n f i d e n c e i n t e r v a l s of c h e m i c a l f a c t o r s i n o c c u r r e n c e s i t e s of some species C o r r e l a t i o n o f c h e m i c a l and p h y s i c a l w i t h DII o f c e r t a i n s p e c i e s  358  491  ^5.06  factors  D i s c u s s i o n of each ecotype and s t a t i s t i c a l d a t a on growth a t , and comparisons between s a l i n i t y and temperature c o m b i n a t i o n s f o r each s p e c i e s . .  518  561  LIST OF TABLES  Table I.  Table I I .  Table I I I .  T a b l e IV.  T a b l e V.  Developmental and m o r p h o l o g i c a l d e t a i l s o f E r i t o p h l y c t i s s p . i s o l a t e s 1-10 . . . . . . . . . . . -  31  S i t e name and number, l a t i t u d e and l o n g i t u d e and h a b i t a t type o f Queen C h a r l o t t e I s l a n d s collections . . . . . •. . . . . . . . . . . . . . .  101  S i t e name and number, l a t i t u d e and l o n g i t u d e and h a b i t a t type o f Southwestern B.C. and San Juan I s l a n d s c o l l e c t i o n s . . . . . . . . . . . .  133  S i t e name, c o l l e c t i o n numbers, l a t i t u d e and l o n g i t u d e and h a b i t a t type o f Oregon, C a l i f o r n i a and Nevada c o l l e c t i o n s . . . . . . . . . . . . . . .  167  L o c a t i o n name, s i t e number and h a b i t a t type o f Long I s l a n d , N.Y. c o l l e c t i o n s  . . . . •. . . . . . .  174  Table VI.  D . I . I , o f each s p e c i e s over h a b i t a t s . . . •. . . . .  197  Table V I I .  Mean v a l u e s of environmental habitats . .  199  Table V I I I .  Ranges and means o f environmental f a c t o r s o c c u r r e n c e s o f each s p e c i e s . . . . .  T a b l e IX.  T a b l e X.  Table XI.  factors  over over . . . - 200  C o r r e l a t i o n s of each s p e c i e s w i t h environmental factors . . . . . . . . . . . . . . . . -. . . . . .  202  F i g u r e number, taxon, c o l l e c t i o n s i t e name and number and h a b i t a t type of each i s o l a t e . . . . . .  220  C e r t a i n environmental s i t e s o f each I s o l a t e  224  parameters a t r e c o v e r y . . . . . .  . .  LIST OF ILLUSTRATIONS  Map  I.  North American c o l l e c t i n g  Map  II.  Devon I s l a n d Base Camp Lowland  Map  III.  Queen C h a r l o t t e I s l a n d s c o l l e c t i n g  Map  IV.  Southwestern B.C. and Northwestern Washington collecting sites . . . . . . . . . . . . . . . . .  129  Oregon, C a l i f o r n i a and Nevada c o l l e c t i n g  . .  163  -.  15  • .  18  Map V.  sites  . . .  6  . . . . . . . . . .  61  sites';  . . . .  sites  P l a t e IPlate'II  . . . . . . . . . . . . . . . . . . . . . . . . .  Plate III P l a t e IV Plate V  . . . . .  Plate'VI  •  99  . . . . . . . . . . . . . .  21  . . . . . . . . . .  24  . . . . . . . . . . . .  27  . . . . . . . . . .  48  Plate VII  53  Plate VIII  58  P l a t e IX . Plate X  . . . . . . . . . . . . . . . . .  70  . . . . . . . . . . . . . . . . . . . . . . . . . . .  77  P l a t e XI  84  Plate XII  . . . . . . . . . . . . . . . . . . .  .  PlateXIII P l a t e XIV  90 96  . . . . . . . . . . . . . . . . . . . . .  113  P l a t e XV"... . .  125  P l a t e XVI  141  P l a t e XVII . . . . . .  . . . .  . . . -. . . . . . . . .  .  P l a t e XVIII P l a t e XIX  147 152  . . . . . . . . . . . . . . . . . . . . . .  158  vii  Figures S e c t i o n 9. F i g u r e 1.  DII o f c h y t r i d s and c h y t r i d i a c e o u s s p e c i e s over n i n e t e e n h a b i t a t s . . . . .  F i g u r e 2.  Mean h a b i t a t v a l u e s parameters  Figure'3.  F i g u r e s 4-19.  ^  of environmental \1§5^  Ranges, means and standard d e v i a t i o n s of e n v i r o n m e n t a l f a c t o r s over s i t e s where s e l e c t e d s p e c i e s occur  189  DII p l o t t e d a g a i n s t h a b i t a t means f o r c e r t a i n species . . . . . . . . . . . . .  192, "193  Growth c l o c k s o f c h y t r i d s and chytridiaceous species at various t e m p e r a t u r e - s a l i n i t y combinations . . . .  .209^213  S e c t i o n 10. F i g u r e s 1-60.  F i g u r e 61.  Environmental f a c t o r l e v e l s f o r i s o l a t e s grouped i n t o ecotypes  218-^  .viii  ACKNOWLEDGEMENTS  P a t i e n t a d v i c e , common s e n s e , w i l l i n g d i s c u s s i o n and comments of Dr.  G. C. Hughes are v e r y much a p p r e c i a t e d .  critical  D r s . W.  B.  S c h o f i e l d , V. J . K r a j i n a , R. J . Bandoni and D. J . S. B a r r k i n d l y i  r e v i e w e d p o r t i o n s of the d i s s e r t a t i o n d u r i n g i t s p r e p a r a t i o n . D r s . Hughes, S c h o f i e l d and  I . M.  Brodo c o n t r i b u t e d m a t e r i a l l y  d u r i n g t h e c o l l e c t i o n phase of t h i s work and Mr.  P. E. B a r r e t t  Dr. G. Rank r e s p e c t i v e l y s e c u r e d samples from Devon I s l a n d and S p e c i a l thanks t o Mr.  N e i l Carey whose seamanship and  and Mexico.  perseverance  made c o l l e c t i n g i n the Queen C h a r l o t t e I s l a n d s p o s s i b l e .  Access to  s i t e s i n the H u n t i n g t o n Harbor R e g i o n (N.Y.) was  provided  by  C_. G_. A. V. Voyager I I owned and  J . R. Newcomer.  o p e r a t e d by Mr.  a l s o i n d e b t e d t o the o f f i c e r s and helpful  I  am  crew of £. j>_. _S. V e c t o r f o r t h e i r  assistance.  L a b o r a t o r y f a c i l i t i e s were f r e e l y p r o v i d e d C. 0. P e r s o n , R. J . B a n d o n i , W. and V. J . K r a j i n a . e f f o r t s o f Mrs.  by D r s .  Hughes,  K. F l e t c h e r , R. F. S c a g e l , T.  Bisalputra  I am t h a n k f u l f o r the t i r e l e s s computer programming  Dolores  Lauriente.  1  1.  INTRODUCTION  Introduction Study Purposes  Taxa o f t h e C h y t r i d i a l e s , and c h y t r i d - l i k e B l a s t o c l a d i a l e s , H y p h o c h y t r i a l e s , S a p r o l e g n i a l e s , L a g e n i d l a l e s and P e r o n o s p o r a l e s a r e included i n t h i s presentation w i t h occasional reports of Actinomycetales even though they a r e n o t f u n g a l .  M o r p h o l o g i c a l f e a t u r e s used t o  d i s t i n g u i s h t h e f u n g a l o r d e r s and t h e i r s u b o r d i n a t e groups a r e p r e s e n t e d by K a r l i n g Fuller  ( 1 9 3 2 ) , Sparrow  (1966).  (1943, 1958, 1960), L i t v i n o v (1958) and  Among m o r p h o l o g i c a l f e a t u r e s used t o d i s t i n g u i s h  and c h y t r i d i a c e o u s t a x a a r e : arrangement  f l a g e l l a r t y p e and attachment  chytrid  location;  and c o m p l e x i t y o f s p o r e i n t e r n a l s t r u c t u r e s ; t h a l l u s e x t e n t  and r e p r o d u c t i v e c e n t e r arrangements; o p e r c u l a t i o n v s . i n o p e r c u l a t i o n ; eucarpism v s . holocarpism; m o n o c e n t r i c i t y v s . p o l y c e n t r i c i t y ;  relationship  of t h a l l u s t o s u b s t r a t e ( e p i b i o t i c , e n d o b i o t i c , o r i n t e r b i o t i c ) ;  zoospore  s i z e ; sporangium shape, s i z e and w a l l o r n a m e n t a t i o n s , d i s c h a r g e p a p i l l a numbers, shape, and l o c a t i o n ; a p o p h y s i s p r e s e n c e o r absence and s t r u c t u r e ; r h i z o i d a l b r a n c h i n g , c o a r s e n e s s and number o f r h i z o i d a l axes and t h a l l u s development.  Several workers, K a r l i n g  (1931, 1932, 1941, 1945, 1946a,  1947a, 1948a, 1949a, 1949b, 1953, 1954, 1967a, 1967b, 1967c), Couch (1932), Sparrow  (1932, 1933, 1943, 1958, 1960, 1968), B a r t s c h (1939), Berdan  (1941a, 1941b , .1942), W h i f f e n (1944), A n t i k a j i a n ( 1 9 4 9 ) , Emerson ( 1 9 5 0 ) , ;  H a s k i n s ( 1 9 5 0 ) , R e i t h ( 1 9 5 0 ) , Koch (1951, 1957, 1968), Johnson 1957a, 1957b, 1969), M i l l e r  (1955,  (1955, 1961, 1968), Sparrow and P a t e r s o n  2  ( 1 9 5 5 ) , P a t e r s o n (1956, 1963), W i l l o u g h b y (1956, 1959, 1961a, 1961b, 1962, 1964, 1 9 6 5 ) , L i t v i n o v  (1958), S c h o l z (1958a), B o s t i c k (1968),  Booth and M i l l e r ( 1 9 6 8 ) , Booth ( 1 9 6 9 ) , Dogma (1969a), Konno (1969) , B a r r (1970a, 1 9 7 0 c ) , H a s i j a and M i l l e r  (1970) and S a l k i n (1970) q u e s t i o n  the v a l i d i t y of one o r more of the c h a r a c t e r s l i s t e d above a t i n t e r s p e c i f i c , g e n e r i c and f a m i l i a l l e v e l s .  To d a t e few s t u d i e s , w i t h t h e  e x c e p t i o n o f K a r l i n g (1931, 1947a, 1951), Ber.dan (1942), Hanson (1945a), A n t i k a j i a n ( 1 9 4 9 ) , H a s k i n s and Weston ( 1 9 5 0 ) , Koch (1951, 1957), W i l l o u g h b y (1957, 1958), F u l l e r ( 1 9 6 2 ) , M i l l e r ( 1 9 6 8 ) , P a t e r s o n ( 1 9 6 3 ) , B a r r and Hickman (1967a, 1967b), Booth and M i l l e r ( 1 9 6 8 ) , B a r r 1970a, 1970b) and H a s i j a and M i l l e r  (1969,  (1970), have c o n s i d e r e d t h e range of  phenotypic v a r i a t i o n w i t h i n a s i n g l e s p e c i e s , i n s p i t e of urgings of Sparrow (1943) and Emerson (1950) t o s t u d y m o r p h o l o g i c ; e x p r e s s i o n i n a l a r g e number of i n d i v i d u a l s . On c o n s u l t i n g t h e works o f Sparrow (1960) and Johnson and  Sparrow  (1961) i t i s e v i d e n t t h a t a c h y t r i d and c h y t r i d i a c e o u s mycota abounds i n most, of the e a r t h ' s s o i l s .  S i n c e the appearance o f t h e s e comprehensive  t r e a t i s e s , p u b l i c a t i o n s by K a r l i n g (1960, 1965, 1966, 1967a, 1967b, 1967c, 1967d, 1967e, 1967f, 1967g, 1967h, 1968a, 1968b, 1968c, 1968d,  1968e,  1969a, 1969b), W i l l o u g h b y (1958, 1959, 1961a, 1961b, 1962, 1963, 1965, 1966, 1969), H a r d e r and Uebelmesser  ( 1 9 5 9 ) , P e r s i e l (1959-, 1960a,  1960b), Sparrow and Koch ( 1 9 5 9 ) , G o l d s t e i n ( 1 9 6 0 ) , M i l l e r 1965, 1 9 6 8 ) , W i l l o u g h b y and Townley  (1961a,  (1961, 19.62,  1961b)., Harder and P e r s i e l  (1962) , S c o t t ( 1 9 6 2 ) , G r i f f i t h s and Jones (1963), A p i n i s (1964, 1965, 1966, 1 9 7 0 ) , Sparrow e t a l .  1964,  (19641,  (1965)., G a e r t n e r and  Ulken  Sparrow  ( 1 9 6 6 ) , Sparrow (1965, 1966, 1968a, 1968b, 1969), G a e r t n e r (1966, 1967a,  3  1967b, 1968a, 1968b, 1968c), Umphlett and Olson (1967), Barr (1969, 1970a, 1970b, 1970d), Booth (1969), Dogma (1969a, 1969b), Dogma and Sparrow (1969) , Howard and Johnson (1969) , Johnson (1969), Umphlett and Koch (1969), Hasija and M i l l e r (1970), Salkin (1970) and Thornton (1970), add to this impression.  Early d i s t r i b u t i o n a l studies,  conducted by workers at Gb'ttingen, indicate:  aquatic fungi are common  i n t r o p i c a l s o i l s (Sorgel, 1941); disturbed s o i l s are greater i n species d i v e r s i t y than natural s o i l s (Sergei, 1941; Harder, 1948; Remy, 1948; Gaertner, 1954); more genera of aquatic Phycomycetes occur i n clay s o i l s than i n sand s o i l s (Remy, 1948); moist s o i l s are richer i n species d i v e r s i t y than dry s o i l s (So'rgel, 1941; Remy, 1948); lower Phycomycetes are  not tied to d e f i n i t e niches (Remy, 1948); aquatic Phycomycetes occur  i n d e f i n i t e clusters i n s o i l and numbers of individuals collected from s o i l i n summer depend upon s o i l moisture and not on s o i l temperature (Reinboldt, 1951); frequency of aquatic fungi i s either impeded or enhanced i n the rhizosphere depending on the higher plant contributing roots to the s i t e and plants at flowering stages generally impeded fungus frequency (Reinboldt, 1951); fewer species occur i n A r c t i c and Antarctic s o i l s than s o i l s at lower latitudes (Harder, 1954; Gaertner, 1954; Harder and P e r s i e l , 1962); and d i v e r s i t y of species i n s o i l s decreases from i n t e r t i d a l to dune s i t e s (Harder and Uebelmesser, 1955); and most chytrids from i n t e r t i d a l areas are only f a c u l t a t i v e l y marine (Scholz, 1958b).  Studies by HShnk (1952, 1956) support the notion that the  d i v e r s i t y of species increases from the i n t e r t i d a l , landward. Willoughby (1961a, 1962, 1964, 1965) carried out d i s t r i b u t i o n a l studies and found: c e r t a i n species are found i n wet s o i l s while others are recovered from  4  dry s o i l s ,  l a k e margins have a r i c h e r c h y t r i d mycota  than p a s t u r e l a n d  above o r submerged muds below; d i s t r i b u t i o n o f c e r t a i n s p e c i e s i s a f u n c t i o n o f s o i l pH; and o c c u r r e n c e o f R h i z o p h l y c t i s r o s e a i s h i g h e r i n n u t r i e n t r i c h s i t e s than n u t r i e n t poor  locations.  Most e c o l o g i c a l i n f o r m a t i o n on c h y t r i d s and c h y t r i d i a c e o u s f u n g i i s r e l a t e d to t h e i r d i s t r i b u t i o n .  Few attempts have been made  ( R e i n b o l d t , 1951; W i l l o u g h b y , 1961, 1965) t o c o r r e l a t e fungus w i t h s o i l p h y s i c a l and c h e m i c a l parameters.  S t u d i e s by S c h o l z  distribution (1958a)  and B a r r (1970a, 1970b, 1970d) i n d i c a t e t h a t c h y t r i d s a r e m o r p h o l o g i c a l l y and/or p h y s i o l o g i c a l l y adapted t o t h e i r  environment.  S i n c e no e x t e n s i v e d i s t r i b u t i o n a l study over s e v e r a l degrees o f l a t i t u d e has y e t been attempted i n North America, one aim o f my study i s to account f o r o c c u r r e n c e and d i s t r i b u t i o n of z o o s p o r i c f u n g i from s i t e s as f a r noth as Devon I s l a n d , Canadian E a s t e r n A r c t i c and as f a r south as M a z a t l a n , Mexico  ( f i g . 1 ) . Before reporting t h i s information, I  propose t o d i s c u s s p r o b l e m a t i c taxonomic c r i t e r i a i n t h e f u n g i and p r e s e n t examples  o f such problems i n o r d e r to s e t the p h i l o s o p h i c a l  approach taken i n s p e c i e s i d e n t i f i c a t i o n . is  Another purpose of the study  t o r e l a t e fungis d i s t r i b u t i o n t o c h e m i c a l and p h y s i c a l parameters b u t  the b a s i c t h e s i s i s t h a t these f u n g i a r e adapted t o t h e i r  environments  and, hence, a t the i n t r a s p e c i f i c l e v e l they a r e composed o f s e v e r a l ecotypes.  Map  I . — F i g u r e 1.  Approximate l o c a t i o n s ,  indicated  by s o l i d  of c o l l e c t i n g s i t e s i n N o r t h A m e r i c a .  triangles,  6  7  2.  PROBLEMATIC TAXONOMIC CRITERIA IN THE CHYTRIDIALES: COMPARATIVE MORPHOLOGY OF TEN ENTOPHLYCTIS SP. ISOLATES  Introduction Methods and m a t e r i a l s Typical  development  Developmental and m o r p h o l o g i c a l v a r i a t i o n s Comparative development Discussion  INTRODUCTION S e v e r a l c h a r a c t e r s used t o d i s t i n g u i s h c h y t r i d t a x a a r e v a r i a b l e and among t h e s e a r e zoospore s i z e , shape, p o s i t i o n of f l a g e l l a r a t t a c h V ment and numbers of s p o r e g u t t u l e s ; sporangium s i z e and shape; r . E l z o i d a l n a t u r e , e x t e n t and b r a n c h i n g ; sporangium w a l l e n a t i o n s o r p u n c t a e ; d i s c h a r g e p a p i l l a numbers, shape and l o c a t i o n ; a p o p h y s i s p r e s e n c e and shape, r h i z o i d a l axes number, t h a l l u s development, c o n f i g u r a t i o n , and s u b s t r a t e r e l a t i o n s and o p e r c u l a t i o n . Z o o s p o r e s — A number o f w o r k e r s (Koch, 1957, 1968; Dogma, 1969a; S a l k i n , 1970; H a s i j a and M i l l e r , 1970) have observed v a r i a t i o n s i n s p o r e and l i p i d body d i a m e t e r s , number of l i p i d b o d i e s , shape and p o s i t i o n of  f l a g e l l a r attachment I n p l a n o n t s from s p o r a n g i a on d i f f e r e n t s u b s t r a t e s .  C o n v e r s e l y P a t e r s o n (1963) r e p o r t s t h a t s p o r e s o f two Rhizophydium s p e c i e s (Rhizophydium sp. 1 and Rhizophydium sp. 2) have c o n s t a n t s i z e on a l l s u b s t r a t e s t e s t e d , and B a r r (1970) d e t e c t e d no s i z e d i f f e r e n c e s i n spores of P h l y c t o c h y t r i u m r e i n b o l d t a e P e r s i e l from s p o r a n g i a on d i f f e r e n t s u b s t r a t e s ( p i n e p o l l e n and YpSs medium).  8  S p o r a n g i a — K o c h (1957) and S a l k i n (1970a) found d i f f e r e n c e s i n sporangium shape on v a r i o u s s u b s t r a t e s b u t a t l e a s t some s p e c i e s have s p o r a n g i a o f h i g h l y v a r i a b l e shape on t h e same s u b s t r a t e (Koch, 1951, 1957; H a s i j a and M i l l e r , 1970). are:  Among f a c t o r s c a u s i n g s i z e v a r i a t i o n s  d i f f e r e n t s u b s t r a t e s ( B o s t i c k , 1968; B a r r , 1969, 1970a;  Salkin,  1970; H a s i j a and M i l l e r , 1970); s u b s t r a t e volume ( P a t e r s o n , 1963); b a i t c o n c e n t r a t i o n and t e m p e r a t u r e ( B o s t i c k , 1968). R h i z o i d s — V a r i a t i o n i n the e x t e n t o f t h e s e s t r u c t u r e s on d i f f e r e n t s u b s t r a t e s has been n o t e d by B o s t i c k (1968) and H a s i j a and M i l l e r (1968). I have observed ( u n p u b l i s h e d d a t a ) t h a t a t 20°C r h i z o i d s o f Rhizophydium s p h a e r o t h e c a Zopf, C h y t r i o m y c e s h y a l i n u s K a r l i n g , P h l y c t o c h y t r i u m punctatum Koch and R h i z o p h l y c t i s h a r d e r i i Uebelmesser  (see Booth and  B a r r e t t , 1971) on MHU medium a r e l e s s branched and l o n g e r a t 1 5 ° / s a l i n i t y than a t 0 " /  o o  0 0  salinity.  Sporangium w a l l m a r k i n g s — S o m e o f t h e s e s t r u c t u r e s a r e i n f l u e n c e d by e n v i r o n m e n t a l c o n d i t i o n s , f o r i n s t a n c e , o c c u r r e n c e and d i s t i n c t i v e n e s s of P h l y c t o c h y t r i u m punctatum punctae a r e i n f l u e n c e d by p o l l e n t y p e and w a t e r s o u r c e (Koch, 1957).  When f u l l y s t u d i e d , normal s p e c i e s v a r i a b i l i t y  may e s t a b l i s h a c l i n e i n c l u d i n g many p r e v i o u s l y d e s c r i b e d t a x a .  One such  c l i n e , based on t h e number o f t e e t h i n a row around t h e sporangium  apical  o r s u b a p i c a l p o r t i o n , i n v o l v e s P h l y c t o c h y t r i u m p l a n i c o r n a e A t k i n s o n sensu Sparrow ( 1 9 3 8 ) , P* - p l a n i c o r n a e sensu W i l l o u g h b y (1961a), P_. c i r c u l i d e n t a t u m Koch and P_. m u l t i d e n t a t u m U m p h l e t t .  V a r i a t i o n s i n number, l o c a t i o n and  c o m p l e x i t y o f e n a t i o n s on s p o r a n g i a o f M i l l e r ' s Booth's  (1968) " d e n t a t e " and  (1971b) Rhizophydium s p . - P h l y c t o c h y t r i u m a u r i l a e A j e l l o  suggest a n o t h e r such c l i n e .  complex  F i n a l l y , s i n c e the o r i g i n a l r e p o r t  ( W i l l o u g h b y and Townley, 1961a) o f C h y t r i o m y c e s p o c i i l a t u s W i l l o u g h b y and  9  Townley s e v e r a l w o r k e r s ( W i l l o u g h b y , 1965; Sparrow, 1968;  Dogma, 1969a  and Booth and B a r r e t t , 1971) have r e p o r t e d t h a l l i w i t h c u p u l e - f r e e sporangia.  C o n s i d e r i n g these o b s e r v a t i o n s t h e r e i s , as Dogma (1969a)  s u g g e s t s , a p o s s i b l e C_, p o c u l a t u s - C .  a n r i u l a t u s Dogma sporangium type  cline. Discharge papillae—Numbers  o f t h e s e s t r u c t u r e s a r e a f f e c t e d by  sporangium s i z e ( B a r r , 1969) and s a l i n i t y  ( S c h o l z , 1958a).  Since s e v e r a l  P h l y c t o c h y t r i u m s p e c i e s a r e d i s t i n g u i s h e d by p a p i l l a e numbers t h e above o b s e r v a t i o n s r a i s e doubt about t h e i r v a l i d i t y as s e p a r a t e t a x a . becomes more o b v i o u s when M i l l e r ' s  This  (1961) s t a t e d d i f f i c u l t y i n r e c o g n i z i n g  m u l t i p o r o u s P h l y c t o c h y t r i u m s p e c i e s and Booth's (1971c) P_. a f r i c a r i u m Gaertner-P_.  s p e c t a b i l e Uebelmesser p a p i l l a e number complex a r e c o n s i d e r e d .  A p o p h y s i s — C o u c h (1932'), Koch (1957), and M i l l e r  (1961, 1968) have  q u e s t i o n e d use o f t h i s s t r u c t u r e t o s e p a r a t e P h l y c t o c h y t r i u m from Rhizophydium.  O t h e r w o r k e r s ( K a r l i n g , 1967a; Johnson, 1969) have  r e c o g n i z e d P h l y c t o c h y t r i u m s p e c i e s w i t h o u t an apophysis  and I observed  ( u n p u b l i s h e d d a t a ) nonapophysate t h a l l i o f P h l y c t o c h y t r i u m K a r l i n g on MRU medium and P i n u s s y I v e s t r i s L. p o l l e n . of P h l y c t o c h y t r i u m , a r e s e p a r a t e d p a r t i a l l y by apophysis  chaetlferum  S e v e r a l species? shape.  However,  the u s e f u l n e s s o f t h i s c h a r a c t e r i s d o u b t f u l s i n c e each o f s e v e r a l s i n g l e spore i s o l a t e s o f a few P h l y c t o c h y t r i u m s p e c i e s were observed with?, a h i g h l y v a r i a b l e apophysis  shape (Booth, 1971b, 1971c, 197Id)..  Rhizoidal n a t u r e — A haustorium-like or tubular untapering  rhizoidal  system d i s t i n g u i s h e s c e r t a i n genera ( P h l y c t i d i u m and Septcsperma) o f t h e Phlyctidioideae. and O l s o n  Yet, Paterson  (1956), W i l l o u g h b y  (1965). and U m p h l e t t .  (1965) d e s c r i b e d P h l y c t o c h y t r i u m s p e c i e s w i t h r h i z o i d a l  s i m i l a r t o P h l y c t i d i u m and I noted  stages  (Booth, 1971b) a fungus (,?Phlyctochytriuro  10  sp.)  producing e i t h e r h a u s t o r i a  or tapering  rhizoids.  established P o l y p h l y c t i s , with Phlyctochytrium the  type, and  p a r t i a l l y distinguished  P h l y c t i d i o i d e a e by  the  catenulate  Townley (1961b) observed.  as  the  and  unispinum P a t e r s o n  (Reinboldt,  from a t a p e r i n g  1951;  r h i z o i d s t h a t he and W i l l o u g h b y  and  however, may  (1957) and  a x i s per  Paterson  to c a t e n u l a t e  Other s p e c i e s  n a t u r e have been  exactly with reported  genera a r e d i s t i n g u i s h e d by  sporangium.  However, K a r l i n g  the  presence  (1949b), Koch rhizoidal  i n t y p i c a l l y u n i - a x i a t e genera and  axis  I have a l s o  (unpublished) s e v e r a l r h i z o i d a l axes on s p o r a n g i a of Rhizdphydium  s p h a e r o t h e c a and  R.  pollinis-pini  phydium n o r m a l l y has  uniaxiate  (Braun) Zopf on MHU  medium. ' R h i z o -  sporangia.  Thallus development—Whiffen type as  reliable  f i g u r e d P_. unispinum m a t e r i a l  (1963) have observed more than one  on s p o r a n g i a o f s p e c i e s observed  not be  K a r l i n g , 1967a; S a l k i n , 1970).  Rhizoidal axes—Certain of more than one  as the  type, w i t h a h a u s t o r i a l r h i z o i d a l system.  rhizoids varying  (1967b)  i t from the o t h e r genera of  This character,  s i n c e Dogma (1969a) d e s c r i b e d  Karling  (1944) proposed t h a l l u s development  the b a s i s of c h y t r i d f a m i l i e s but  a few  subsequent  studies  i n d i c a t e v a r i a b i l i t y i n the developmental pathway of c e r t a i n s p e c i e s genera.  T h a l l i of R h i z o p h l y c t i s , Chytriomyces, R h i z i d i u m ,  Allochytridium  and  Asterophlyctis  species  can  Phlyctochytrium  develop from s w e l l i n g  e i t h e r the c y s t or germ tube ( K a r l i n g , 1947b, 1949b; A n t i k a j i a n , Koch, 1957;  (Olpidiaceae,  and  of  1949;  S a l k i n , 1970).  Thallus-substrate  epibiotic  and  relations—Monocentric  t h a l l i are  S y n c h y t r i a c e a e , E n t o p h l y c t o i d e a e and  ( P h l y c t i d i o i d e a e and  Chytridioideae).  endobiotic  Endochytrioideae) ,  C h y t r i d i o d e a e ) or i n t e r b i o t i c  (Rhizidiaceae  W h i f f e n (1944) suggests t h a t i n the case of  saprobic  11  c h y t r i d s chance a l o n e may determine whether^ t h a l l i a r e i n t r a m a t r i c a l or e x t r a m a t r i c a l . P_. c h a e t i f e r u m  F o r example, Phlyctochytrium,punctaturn (Koch, 1957),  (Booth, 1969) and R h i z i d i u m  richmondense Willoughby  ( K a r l i n g , 1967b) may be e i t h e r e p i b i o t i c or i n t e r b i o t i c and R h i z o p h l y c t i s s p i n o s a (fearling) Sparrow may even produce t h a l l i which a r e e n d o b i o t i c and  e p i b i o t i c a t the same time ( K a r l i n g , 1974b). Monocentricity  used  vs. P o l y c e n t r i c i t y — T h e s e  thalli  configurations,  to s e p a r a t e taxa a t the f a m i l y l e v e l , a r e v a r i a b l e .  Some taxa  which a r e n o r m a l l y monocentric may sometimes be p o l y c e n t r i c , f o r instance, Rhizophlyctis hyalina  ( K a r l i n g ) Sparrow and R h i z i d i u m  varians  K a r l i n g may be monocentric or p o l y c e n t r i c ( K a r l i n g , 1947b, 1949b). v a r i a t i o n r e l a t e s these taxa, n o r m a l l y , c o n s i d e r e d the  i n the R h i z i d i a c e a e to  Cladochytriaceae. Operculation—Of  a l l the c h a r a c t e r s  used  to s e p a r a t e c h y t r i d  t h i s one has p o s s i b l y come under the most d i s f a v o r and r i g o r o u s The  Such  primary argument has r e v o l v e d  this character  i s to be used.  around  Since  taxa  scrutiny.  the taxonomic l e v e l a t which  Sparrow's (1942) u s e o f o p e r c u l a t i o n  to e s t a b l i s h two s e p a r a t e s e r i e s i n the o r d e r , W h i f f e n (1944) has presented s e v e r a l reasons f o r n o t p l a c i n g much importance on o p e r c u l a t i o n and Karling  (1967d) s t a t e d  that separation  of E n t o p h l y c t i s and D i p l o p h l y c t i s  from Endochytrium and Nephrochytrium " . . . p l a c e s undue taxonomic emphasis on the presence o f an operculum above the g e n e r i c S e v e r a l o t h e r workers, Koch (1951, 1957), Willoughby Miller  (1968) and B o s t i c k  level  (1956, 1961a, 1961b),  (1968) have observed v a r i a t i o n s i n v o l v i n g  operculum presence and absence, o c c u r r e n c e o f o p e r c u l u m - l i k e ;  and/or i n c o m p l e t e l y inoperculate.  . . . ."  formed o p e r c u l a  i n species  normally  structures  considered  I n t e r p r e t a t i o n of v a r i a b i l i t y and comparison o f these  12  s t u d i e s a r e p r o b l e m a t i c because each worker seems to have h i s own concept of what c o n s t i t u t e s an operculum. Koch  F o r i n s t a n c e , Umphlett and  (1969) d e s c r i b e the q u a s i - o p e r c u l u m which they c o n s i d e r  between i n o p e r c u l a t e and o p e r c u l a t e c o n d i t i o n s .  intermediate  Johnson (1969), on the  o t h e r hand, s u g g e s t s that the q u a s i - o p e r c u l u m i s a type of f a l s e operculum.  B e s i d e s the n e c e s s i t y  o f a d e f i n i t i v e study of o p e r c u l a t i o n  and i n o p e r c u l a t i o n , r e c o g n i z e d by Koch i n 1957, perhaps common a c c o r d i s required concerning  the n a t u r e of the operculum and, f o r t h a t m a t t e r , a l l  o t h e r taxonomic c h a r a c t e r s .  METHODS AND  MATERIALS  D u r i n g a survey of c o a s t a l and i n t e r i o r s o i l s from the Queen C h a r l o t t e I s l a n d s to C a l i f o r n i a and Nevada, s e v e r a l s i n g l e spore i s o l a t e s of an E n t o p h l y c t i s s p e c i e s were o b t a i n e d by s t r e a k i n g or p l a c i n g b a i t s on m o d i f i e d Harder and Uebelmesser (1955) medium (MHU, 0.3% peptone, 0.1% d e x t r o s e , 1.1% agar i n 1 1 s a l i n i t y with G—4  x 10  0.5%'malt e x t r a c t ,  water o f 0 ° / o or O  .03% S t r e p t o m y c i n s u l p h a t e — 7 7 4 mcg/mg and  units).  7°/  O0  .03% P e n i c i l l i n  F o r t h r e e days a f t e r s t r e a k i n g each p l a t e was  surveyed  f o r s i n g l e encysted spores or i n c i p i e n t s p o r a n g i a ; these were removed p l a c e d on MHU  medium minus a n t i b i o t i c s .  several characters include:  and  The i s o l a t e s , which v a r y i n  l , " * " S t a t e l i n e Pond, C a l i f .  (41°59.0'N.,  121°54.0'W.); 30, Soda Lake F a l l o n , f e v . (39°32.0'N., 118°48.0'W.); 43, Kwuna P t . , B.C. .(53°13.1'N., 131°59.3'W.); 46, One-half Beach, (48°47.1'N., 123°04.8'W.); 47, 54, Hibben I s l a n d , B.C. 132°19.1'.W.); 52, Long Beach P r o v i n c i a l Park, B.C.  Au thor's c u l t u r e no.  B.C.  (52°58.6'N.,  (49°03.2'N., 125°43.2'W.);  13  61, A n v i l I s l a n d , B.C. (50°12.3'N.,  (49°31.4'N.,  125°05.8'W.)  119°05.0'W.).  Morphology  on p o l l e n and MHU  and 74, Mono Lake I I , C a l i f . and development  of the i s o l a t e s were s t u d i e d (1971).  MORPHOLOGY OF ALL ISOLATES  medium the s p h e r i c a l  zoospore c y s t germinates by a  tube ( f i g . 13), a broad bl'eb ( f i g s . 1, 14) or a broad  narrow  tube ( f i g . 2).  r h i z o i d a l a x i s pushes out of the expanding i n c i p i e n t sporangium 15-16).  t h i s time the zoospore c y s t may  17-19).  (figs.  ( f i g s . 3, 18, 19).  appear d e v o i d of protoplasm  (figs.  The c y s t g e n e r a l l y remains empty through the next s t a g e s i n  which s e v e r a l r h i z o i d a l axes a r i s e on the d e v e l o p i n g sporangium 4, 20). 5, 6,  As  the i n c i p i e n t sporangium  25) or i r r e g u l a r l y  dichtomously  ( f i g s . 3-6,  (figs.  (figs.  c o n t i n u e s to s w e l l , r e g u l a r l y  21, 22), and  5, 6, 21, 22).  i s s p h e r i c a l w i t h s e v e r a l r h i z o i d a l axes.  A t m a t u r i t y the  The c y s t may  R h i z o i d s a r e t u b u l a r and have b l u n t t i p s  Discharge p a p i l l a e  be v e r y e v i d e n t  (figs.  the  25, ^ 6 ) .  ( f i g . 28) or tubes ( f i g . 29) of v a r y i n g l e n g t h break  double contoured sporangium w a l l a t one  zoospore c y s t may  may  sporangium  ( f i g . 23), d i m i n u t i v e ( f i g . 26) or c o m p l e t e l y i n c o r p o r a t e d i n t o sporangium.  (figs.  the r h i z o i d s branch  21, 22) and e l o n g a t e , the zoospore c y s t  r e f i l l w i t h projioplasm ( f i g s .  the  A  Soon the a x i s branches a l o n g the c o u r s e of the r h i z o i d ( f i g .  17) or a t the j u n c t u r e w i t h the d e v e l o p i n g sporangium At  B.C.  (37°57.0'N.,  medium by the methods of Booth and B a r r e t t  TYPICAL DEVELOPMENT AND On MHU  123°17.3'W.); 66, B i r d Cove,  to s e v e r a l p l a c e s .  The  d e l i q u e s c e and f u n c t i o n as a d i s c h a r g e pore ( f i g . 27).  Zoospore d i s c h a r g e i s preceded by p a p i l l a r d i s s o l u t i o n or d e l i q u e s c e n c e of are  a large  (18-26 x 8-18y) g e l a t i n o u s p l u g a t the tube o r i f i c e .  m o t i l e i n the sporangium p r i o r  to d i s c h a r g e and escape s i n g l y  Spores through  P l a t e 1 — F i g u r e s 1-6.  Entophlyctis  sp.  F i g . 1.  Cyst germinating  2 by a broad b l e b , f h . ( f r e e h a n d ) , germ tube, f h . , xlOOO. end, x500.  F i g . 4.  an empty c y s t , xlOOO.  x200.  F i g . 2.  Germinating  c y s t and  F i g . 3.  R h i z o i d a l a x i s d e v e l o p i n g a t germ tube  Developing  t h a l l u s w i t h s e v e r a l r h i z o i d a l axes and  F i g . 5.  Developing  t h a l l u s w i t h protoplasm  c y s t and d i c h t o t o m o u s l y b r a n c h i n g r h i z o i d s , xlOOO.  F i g . 6.  w i t h sporangium almost c o m p l e t e l y expanded, xlOOO.  F i g s . 1;, 8-10,  filled  Thallus scale  A; F i g s . 2, 4-6, 11, 12, s c a l e B; F i g . 7, s c a l e C; F i g . 3, s c a l e D; F i g s . 13-16, 19, 24-26, s c a l e E; F i g s . 17, 19, 20, 27, 29-33, s c a l e F; F i g s . 21-23, 24, s c a l e G.  'Unless d e s i g n a t e d f h . , a l l drawings done w i t h camera l u c i d a .  15  1 6  pores or tubes.  M o t i l e spores a r e s p h e r i c a l , each c o n t a i n i n g a s i n g l e  large r e f r a c t i v e  globule.  On p o l l e n the c y s t germinates and produces a tube which p e n e t r a t e s the m i c r o s p o r e e x i n e . n o r m a l l y one  The germ tube s w e l l s w i t h i n the s u b s t r a t e and  to f i v e r h i z o i d a l axes a r i s e from the i n c i p i e n t  At maturity sporangia are u s u a l l y s p h e r i c a l be e l l i p s o i d a l motile prior  sporangium.  ( f i g s . 7, 30) b u t they can  ( f i g . 31), p y r i f o r m , s a c c a t e or i r r e g u l a r .  Zoospores  to d i s c h a r g e and escape the s p o r a n g i a l c o n f i n e s through  the zoospore c y s t  ( f i g s . 7, 30).  DEVELOPMENTAL AND  MORPHOLOGICAL VARIATIONS COMMON TO ALL ISOLATES  F r e q u e n t l y on MHU sporangium  medium the germ tube ceases to expand and  then develops from  remaining c y s t may  then appear  the f i r . p t r h i z o i d a l a x i s nipple-like  medium and p o l l e n , s p o r a n g i a may rather  are  (fig.  23).  (fig.  the  24).  On both  The  MHU  develop by. expansion of the c y s t  than the germ tube w i t h v a r y i n g degrees of frequency f o r each  isolate.  On p o l l e n an endo-exogenous type of development  c y s t and germ tube s w e l l ) may  occur and  b i o t i c a t the same time ( f i g . 11). a r e observed on p o l l e n . 31)-, e p i b i o t i c 12) or tubes  ( i . e . both  t h a l l i a r e e n d o b i o t i c and e p i -  Various other t h a l l u s  Sporangia can be e n d o b i o t i c  ( f i g s . 32, 33) or i n t e r b i o t i c  relationships  ( f i g s . 7, 12,  ( f i g . 34).  ( f i g s . 31-33) f r e q u e n t l y a r i s e from  the c y s t .  Spores  However, as p r e v i o u s l y  e n d o b i o t i c s p o r a n g i a i n p o l l e n g e n e r a l l y d i s c h a r g e t h e i r spores the zoospore c y s t . any p o i n t around  30,  Rhizoids ( f i g .  charge from e p i b i o t i c and i n t e r b i o t i c s p o r a n g i a on p o l l e n almost through tubes or t u b e - l i k e s t r u c t u r e s .  the  exclusively/  mentioned, through  But sometimes d i s c h a r g e tubes, which can a r i s e  the i n c i p i e n t sporangium, a r e formed^ ( f i g i  dis-  11) and  from the  P l a t e I I — F i g u r e s 7-12.  E n t o p h l y c t i s sp.  F i g . 7.  b i o t i c sporangium w i t h two r h i z o i d a l axes, f h . , x750. d i s c h a r g e tubes, f h . , x2000. x2000.  F i g . 10.  epibiotic  Branched  F i g . 9.  inter-  F i g . 8. • Simple  Complex d i s c h a r g e tubes, f h . ,  d i s c h a r g e tube, f h . , x2000.  sporangium w i t h d i s c h a r g e tubes, xlOOO.  zoosporangium w i t h i n t e r b i o t i c  Empty  rhizoids.  F i g . 11.  F i g . 12.  Endo-,  Endobiotic  18  zoospore c y s t i s f u n c t i o n l e s s  i n discharge.  D i s c h a r g e tubes may be  s t r a p - l i k e and i r r e g u l a r ( f i g . 31), s i m p l e ( f i g s . 8, 32, 34), f o r k e d (fig.  11) o r complex ( f i g s . 9, 10, 3 3 ) .  generally frequently  Sporangia and zoospores a r e  l a r g e r from t h a l l i on MHU medium than on p o l l e n .  Very i n -  zoospores escape the s p o r a n g i a l c o n f i n e s through a s p l i t  i n the sporangium w a l l  ( f i g . 32).  COMPARATIVE DEVELOPMENT AND MORPHOLOGY OF ALL ISOLATES Developmental  and m o r p h o l o g i c a l f e a t u r e s of the t e n i s o l a t e s a r e  presented i n Table I. I s o l a t e s 1-5 and 9 develop t h a l l i p r i m a r i l y by germ tube T h a l l i o f i s o l a t e 7 a l s o develop g e n e r a l l y  swelling.  by germ tube s w e l l i n g b u t  o c c a s i o n a l l y both c y s t and germ tube e n l a r g e s i m u l t a n e o u s l y .  Fifty  p e r c e n t of the t h a l l i of i s o l a t e s 6 and 10 develop from the c y s t . T h a l l i of i s o l a t e 8 on MHU medium develop p r i m a r i l y by c y s t According  to t h e i r mean d i a m e t e r , p o l l e n grown s p o r a n g i a f a l l  three s i z e classes:  into  16-18.5u, i s o l a t e s 4, 5, 8 and 9; 20-22u, i s o l a t e s  3, 6 and 10; and 29-33.5u, i s o l a t e s 1, 2 and 7. s i z e c l a s s e s can be r e c o g n i z e d :  On MHU medium three  20.5-38p, i s o l a t e s 1, 4 and 6; 50-65u,  i s o l a t e s 2, 3 and 7; and 85-100u, i s o l a t e s 8 and 10. isolates 1,2,  swelling.  Sporangia of  5, 6 and 8-10 a r e u s u a l l y s p h e r i c a l b u t s p o r a n g i a o f  i s o l a t e s 3-7 a r e s p h e r i c a l  to I r r e g u l a r .  i s o l a t e s 1-3 have one to f i v e  On p o l l e n , s p o r a n g i a of  ( r a r e l y s e v e r a l ) r h i z o i d a l axes.  of i s o l a t e s 4, 5, 8 r - and 9 have one to s e v e r a l r h i z o i d a l axes.  Sporangia Several  r h i z o i d a l axes t y p i c a l l y occur on p o l l e n grown s p o r a n g i a of i s o l a t e s 6, 7 and 10.  D i s c h a r g e tubes a r e always m i s s i n g from s p o r a n g i a of i s o l a t e s  4, 6 and 10 grown on MHU medium.  Isolate 6 also  lacks  tubes on p o l l e n .  P l a t e I I I — F i g u r e s 13-23.  E n t o p h l y c t i s sp.  g e r m i n a t i n g by a narrow tube, x l 3 2 5 . a broad b l e b , x l 3 2 5 . sporangium, x l 3 2 5 . xl325.  F i g . 17.  F i g . 15. F i g . 16\  Branching  F i g . 14.  Cyst germinating  of r h i z o i d a l a x i s , x650.  i r r e g u l a r development of sporangium, r e f i l l e d  s e v e r a l dichotomously  F i g . 22.  cyst,  F i g . 20.  F i g . 21. zoospore  tube,  F i g . 18.  empty zoospore  sporangium w i t h t h r e e r h i z o i d a l axes, x650.  by  incipient  R h i z o i d a l a x i s t e r m i n a l on germ  F u r t h e r r h i z o i d a l a x i s b r a n c h i n g , x650.  r h i z o i d a l axes, x200.  Cyst  R h i z o i d a l a x i s d e v e l o p i n g on  T h a l l u s w i t h b r a n c h i n g r h i z o i d a l a x i s and F i g . 19.  F i g . 13.  xl325.  Incipient  Thallus with c y s t and s e v e r a l  I r r e g u l a r l y d e v e l o p i n g sporangium w i t h  b r a n c h i n g r h i z o i d s , x200.  on n e a r l y mature sporangium, x200.  F i g . 23.  Cyst  remaining  21  22  Both on p o l l e n and MHU medium i s o l a t e s 1-3, 5 and 7-9 may o r may n o t have d i s c h a r g e tubes.  The zoospore c y s t o f a l l i s o l a t e s can be  p r e s e n t o r absent from s p o r a n g i a on p o l l e n o r MHU medium. g e n e r a l l y n o t the zoospore escape r o u t e on p o l l e n developed of most i s o l a t e s ,  i s o l a t e 6 excepted.  10 on MHU medium escape the sporangium  It is zoosporangia  Zoospores o f i s o l a t e s 4, 6 and primarily  c y s t i s g e n e r a l l y f u n c t i o n l e s s ,in sporangium  through the c y s t .  The  d i s c h a r g e of the remaining  i s o l a t e s on MHU medium. From p o l l e n and MHU medium, zoospores o f most i s o l a t e s have a s i n g l e , h y a l i n e , conspicuous  (1.0-1.5u i n diameter) r e f r a c t i v e  globule.  The s i n g l e g u t t u l e o f i s o l a t e 2 spores i s minute and zoospore g l o b u l e s -of i s o l a t e 8 a r e p a l e orange.  Zoospore f l a g e l l a e o f a l l i s o l a t e s on  p o l l e n o r MHU medium a r e 22-25y l o n g .  Three mean spore diameters  s p o r a n g i a on MHU medium can be r e c o g n i z e d :  from  3.5u, i s o l a t e 10; 4.5u,  i s o l a t e s 1, 7, 8 and 5.0u, i s o l a t e s 2-4, 6.  On p o l l e n , mean spore  diameters a r e 3.0u, i s o l a t e 6; 3.5u, i s o l a t e 7; 4.0y, i s o l a t e s 1-5, 8; and 4.5y, i s o l a t e s 9 and 10.  DISCUSSION T y p i c a l development to E n t o p h l y c t i s .  and morphology of i s o l a t e s 1-10 r e l a t e them  On MHU medium and i n p o l l e n , e u c a r p i c t h a l l i develop  by germ tube s w e l l i n g and s p o r a n g i a a r e i n o p e r c u l a t e .  Rhizoids arise  d i r e c t l y from the sporangium. E n t o p h l y c t i s , e s t a b l i s h e d by F i s c h e r , 1892, i n c l u d e s t e n s p e c i e s r e c o g n i z e d by Sparrow (1960) and s p e c i e s s u b s e q u e n t l y d e s c r i b e d by Willoughby and Townley  (1961a), E_. l o b a t a ; Canter (1965), E_. m o l e s t a ;  Cook (1966), _E. r e t i c u l o s p o r a and K a r l i n g  (1967d), E_. c r e n a t a .  Of  P l a t e I V — F i g u r e s 24-29. . E n t o p h l y c t i s sp. sporangium d e v e l o p i n g from  the r h i z o i d a l a x i s , x l 3 2 5 .  mature sporangium w i t h c y s t remaining, x l 3 2 5 . sporangium w i t h c y s t remaining 27.  F i g . 29.  F i g . 26.  Incipient  F i g . 25.  F i g . 28.  Nearly  Mature  (same^fehallus as f i g . 25), x l 3 2 5 .  P a p i l l a r sporangium c y s t , x650.  papillum, xl325.  F i g . 24.  Fig.  Sporangium w i t h h y a l i n e  Short d i s c h a r g e tube on sporangium, x650.  24  25  these s p e c i e s , J 3 .  texana K a r l i n g , E_. c o n f e r v a e - g l o m e r a t a e  Sparrow, E,. aurea Haskins and E_. conf ervae-glomeratae  (Cienkowski)  (Cienkowski)  Sparrow f . marina Kobayashi and Ookubo a r e c o n s i d e r e d d o u b t f u l t a x a . Sporangium s i z e and shape, p o s i t i o n and number o f r h i z o i d s and the i d e n t i t y of the h o s t a r e c h a r a c t e r s s t r e s s e d ( K a r l i n g , 1931; Sparrow, 1960). of  i n distinguishing  specifes  Other c h a r a c t e r s used i n c l u d e :  behavior  zoospores a t d i s c h a r g e ; zoospore s i z e ; c o l o r , number and s i z e of  s p o r e g u t t u l e s ; f l a g e l l u m l e n g t h ; presence o f the c y s t a t sporangium maturity;  the c y s t ' s r o l e i n sporangium  discharge; protoplasm c o l o r ;  r e s t i n g spore presence and the r e s t i n g spore ornamentation. C h a r a c t e r v a r i a t i o n i n my i s o l a t e s r e l a t e them to s e v e r a l d i f f e r e n t e s t a b l i s h e d s p e c i e s of E n t o p h l y c t i s . to E_. b u l l i g e r a  I s o l a t e s 4, 5, 8 and 9 a r e s i m i l a r  (Zopf) F i s c h e r , E_. conf ervae-glomeratae and E_. conf e r v a e -  glomeratae f . marina;  i s o l a t e s 3, 6 and 10 a r e s i m i l a r  (Schenk) Minden, E_. pygmaea  to E_. r h i z a  (Servinow) Sparrow and E_. c r e n a t a and  i s o l a t e s 1, 2 and 7 a r e s i m i l a r  to E. h e l i o f o r m i s  (Dang.) Ramsbottom,  E_. m o l e s t a , E_. r e t i c u l o s p o r a and E_. texana i n s p o r a n g i a l d i a m e t e r s . Sporangia of i s o l a t e s 2, 5, 6, 8-10, E_. a p i c u l a t a  (Braun) F i s c h e r , E_.  b u l l i g e r a , E_. v a u c h e r i a e ( F i s c h ) F i s c h e r , E_. r h i z n a , _E. r e t i c u l o s p o r a , E_. moles ta and E_. h e l i o f o r m i s a r e s p h e r i c a l or o b p y r i f o r m w h i l e s p o r a n g i a of i s o l a t e s 3, 4, 7, E_. texana, E_. pygmaea j E. conf ervae-glomeratae, E_. a u r e a , E_. conf ervae-glomeratae f . marina, E_. l o b a t a and E_. c r e n a t a a r e spherical  to i r r e g u l a r . . I s o l a t e s 1-3, w i t h one to f i v e r h i z o i d a l axes  per sporangium,  are closely related  to E_. a p i c u l a t a , E_. m o l e s t a , E_.  b u l l i g e r a , E_. v a u c h e r i a e , E_. h e l i o f o r m i s , E.  conf ervae-glomeratae, E_.  pygmaea; E_. texana and E_. conf ervae-glomeratae f . marina.  Entophlyctis  r h i z i n a , E_. aurea, E_. c r e n a t a , E_. l o b a t a , E_. r e t i c u l o s p o r a a n d  isolates  Plate V—Figures  30-34.  E n t o p h l y c t i s sp.  F i g . 30.  e n d o b i o t i c t h a l l i f u n c t i o n i n g as d i s c h a r g e tubes, x650.  C y s t s of F i g . 31.  S t r a p - l i k e i r r e g u l a r d i s c h a r g e tube of an e n d o b i o t i c t h a l l u s , x650. F i g . 32.  Epibiotic  w a l l , x650.  t h a l l u s w i t h d i s c h a r g e through  F i g . 33.  sporangium, x650.  sporangium  I r r e g u l a r , branched d i s c h a r g e tube on e p i b i o t i c  F i g . 34.  d i s c h a r g e tubes, x200.  splitting  I n t e r b i o t i c t h a l l u s w i t h long sporangium  27  28  4, 5, 8 and 9 have one  to s e v e r a l r h i z o i d a l axes per sporangium.  All  i s o l a t e s a r e s i m i l a r to E_. v a u c h e r i a e , E_. b u l l i g e r a , E_. l o b a t a , E_. r e t i c u l o s p o r a and E_. c r e n a t a w i t h r h i z o i d s a r i s i n g anywhere on the sporangium.  R h i z o i d s of i s o l a t e s 1, 3-9  s t r o n g l y dichotomously branched. v a u c h e r i a e , E. rhizoids.  and E.  pygmaea are u s u a l l y  I s o l a t e s 2, 10, E_. a p i c u l a t a , E_.  h e l i o f o r m i s and E_. confervae-glomeratae have d e l i c a t e  D i s c h a r g e tubes are m i s s i n g from s p o r a n g i a of i s o l a t e  6,  E,. a p i c u l a t a , E_. b u l l i g e r a , _E. l o b a t a , E_. m o l e s t a and E_. pygmaea w h i l e the remainder of the i s o l a t e s and o t h e r E n t o p h l y c t i s s p e c i e s g e n e r a l l y have one o r more d i s c h a r g e tubes on each sporangium. of E.  aurea and most of my  D i s c h a r g e tubes  i s o l a t e s a r e t i p p e d by l a r g e s u b s p h e r i c a l  gelatinous plugs.  The  zoospore c y s t may  of i s o l a t e s 1-10.  When p r e s e n t the c y s t f u n c t i o n s i n sporangium  charge o f i s o l a t e s 4, 6 and 10 on MHU  o r may  n o t remain on s p o r a n g i a dis-  medium and i s o l a t e 8 on p o l l e n .  Sporangium d i s c h a r g e i n the remaining i s o l a t e s g e n e r a l l y o c c u r s through a tube or pore.  Cysts of E_. a p i c u l a t a and E_. b u l l i g e r a a r e p e r s i s t e n t  and f u n c t i o n i n sporangium  d i s c h a r g e w h i l e the c y s t i s evanescent  i n o p e r a t i v e i n most o t h e r E n t o p h l y c t i s s p e c i e s . c y s t s may  or may  Entophlyctis  and  lobata  not remain but even when p r e s e n t the c y s t i s i n o p e r a t i v e  during zoosporangial discharge.  A l l i s o l a t e s f a l l w i t h i n the zoospore  s i z e range of E_. h e l i o f ormis , E_. texana, E.  conf ervae-glomeratae, E_.  aurea, E_. conf ervae-glomeratae f . m a r i n a , E_. moles t a , E_. r e t i c u l o s p o r a , E_. l o b a t a and E_. c r e n a t a .  Zoospore  longer than those of _E. texana, E.  f l a g e l l a e of i s o l a t e s c r e n a t a , E_. m o l e s t a , E.  1-10  are  l o b a t a and  r e t i c u l o s p o r a but s h o r t e r than the f l a g e l l u m o f E_. h e l i o f ormis s p o r e s . Spores of E_. aurea and E_. conf ervae-glomeratae have s e v e r a l s m a l l r e f r a c t i v e g l o b u l e s and the g u t t u l e s o f E. texana and E. l o b a t a spores  E.  29  are orange.  Other than spores of i s o l a t e 2 w i t h one minute c o l o r l e s s  g l o b u l e and  i s o l a t e 8 w i t h a l a r g e p a l e orange g l o b u l e , most of  my  i s o l a t e s have spores w i t h a s i n g l e l a r g e h y a l i n e r e f r a c t i v e body. Thus, spore c h a r a c t e r i s t i c s of most of my b u l l i g e r a , E.  conf ervae-glomeratae and  V a r i a t i o n of my  E n t o p h l y c t i s and  i s o l a t e s a r e r e l a t e d to E_. •  E_. c r e n a t a . Karling's  (1931) g i a n t t h a l l i  E n t o p h l y c t i s i n d i c a t e t h a t such c h a r a c t e r s as sporangium s i z e and number and discharge  p l a c e of o r i g i n of r h i z o i d s , r h i z o i d coarseness and tube presence, c y s t presence and  s p o r e g u t t u l e diameter and genus.  Resting  s p o r e f e a t u r e s and  taxonomic v a l u e  substrate d i f f e r e n c e s are  remaining c h a r a c t e r s d i s t i n g u i s h i n g E n t o p h l y c t i s s p e c i e s . s t r a t e d i f f e r e n c e s and species  i s questionable  a p i c u l a t a , E.  shape,  branching,  f u n c t i o n , spore s i z e ,  c o l o r a r e of l i t t l e  of  and  i n the  the  Use  of sub-  p a r a s i t i s m v s . saprophytism to d i s t i n g u i s h c h y t r i d (Emerson, 1950;  r e t i c u l o s p o r a , and  M i l l e r , 1968).  Thus, perhaps E_.  E_. c r e n a t a a r e the o n l y v a l i d  and  useful  taxa. Characters level.  of i s o l a t e s 1-10  a r e a l s o v a r i a b l e above the  Thallus-substrate p o s i t i o n v a r i a t i o n s ( i . e . , endobiotic  e p i b i o t i c v s . i n t e r b i o t i c ) may  r e l a t e my  P h l y c t i d i o i d e a e ; Olpidiaceae,(Olpidium)  Phlyctidioideae If discharge  (Rhizophydium) and  and R h i z i d i a c e a e , ( R h i z o p h l y c t i s  Inoperculatae  Scherffeliomyces  by sporangium w a l l s p l i t t i n g  E n t o p h l y c t i s may and  Considering  and  Entophlyctis Rhizidiaceae.  i s i n t e r p r e t e d as  r e l a t e to P h l y c t i d i a c e a e and  Chytridiaceae,  vs.  E n t o p h l y c t i s to Rhizophydium,  and R h i z i d i u m X ' C T h a l l u s development v a r i a t i o n s r e l a t e my  my  species  opercular,  Rhizidiaceae,  Operculatae.  such v a r i a t i o n , as W h i f f e n  (1944) s t r e s s e d ,  the  operculum warrants l e s s importance than Sparrow (1942) a s s i g n e d i t .  to  30  A l s o , as Johnson (1969) h i n t s , the operculum c h a r a c t e r careful definition.  F a m i l i a l and  c e r t a i n generic  delineations require careful re-evaluation.  concepts  This i s achieved  different conditions. conditions the s p e c i e s  fails  genetical  species  by s t u d y i n g many i s o l a t e s under s e v e r a l  Study of a s i n g l e i s o l a t e under s e v e r a l d i f f e r e n t  to account a d e q u a t e l y f o r s p e c i e s v a r i a t i o n .  concept as a p p l i e d  appearance and  and  Emerson (1950) has  expressed the u r g e n t need to study e n v i r o n m e n t a l and variability.  needs more  activities  to the organisms should  i n nature.  Finally,  r e l a t e to  their  Table I.  I s o l a t e and Collection Numbers*  Developmental and m o r p h o l o g i c a l d e t a i l s o f i s o l a t e s 1-10.  Substrate  Sporangium S i z e (H x W)  Development type  1(1) 1(1) 2(30) 2(30) 3(43) 3(43)  MHU medium pollen MHU medium pollen MHU medium pollen  p r i m a r i l y from germ as above as above as above as above as above  4(46) 4(46)  MHU medium pollen  as above as above  5(47) 6(52) 6(52) 7(54) 7(54)  pollen MHU medium pollen MHU medium pollen  as above from c y s t o r tube e q u a l l y as above as above p r i m a r i l y from germ tube but c y s t may a l s o expand  8(61) 8(61) 9(66) 10(74) 10(74)  MHU medium pollen pollen MHU medium pollen  primarily primarily primarily as above from tube  ' " C o l l e c t i o n no. b r a c k e t e d .  tube  from c y s t from tube from germ tube or cyst  equally  27- (38)-50y 16- (31)-65y 36- (65)-110u 22.J j-(33.5)-48y 38- [50)-70y 11- (22)-35y (sph) 27- (32)-42X 14- (19)-28y (asph) 17- (28)-38y 12- (16)-20y (sph) 14- (17.5)-27X 6- (15)-28y (asph) 12- (18)-30y 17- (20.5)-25y 16- [20.0)-28y 50- (65)-80y 15- C29)-75y (sph) 20- (49)-70X 17- [38)-55y (asph) 75- U 0 0 ) - 1 2 0 y 9- C18.5)-37y 10- U 7 ) - 2 2 y 65- C 8 5 ) - 1 1 0 y 14- [22)-30y  Shape spherical as above as above as above as above spherical-varied  Papillae  + .+ ,+ ,+ ,+ ,-  Spherical spherical-varied  spherical as above as above as above spherical-varied  + ,+ .+ ,+  spherical as above as above as above as above  + ,+ ,+ .+ »+ .-  Table I (continued)  I s o l a t e No. 1 1 2 2 3 3  MHU Pollen MHU Pollen MHU Pollen  4 MHU 4 Pollen 5 Pollen 6 MHU 6 Pollen 7 MHU 7 Pollen 8 8 9 10 10  MHU Pollen Pollen MHU Pollen  Presence + ,+ ,+,+,+ ,+,-  + ,+,-  + ,+,+ ,+ + ,+,-  D i s c h a r g e Tubes No./Sporangium Size  (L x W a t base)  1-2 1-2 1-2 1(3) 1-3 1-(plexus o f several)  2-(5.5)-14 x 3-(5)-6y 3.5-(15)-28 x 2-(4.5)-7y 2-(5)-12 x 2-(3)-4p 2-(6)-15 x 3-(4)-5y 4-(13)-18 x 3.5-(4.5)-6y  1 1-(plexus of several)  3-(5)-7 x (3)4y  4-(18)-24 x 3 - ( 4 ) - 5 ( l l ) u  7-(14)-9 x 3-(5)-8u  1-3 1-(plexus of several) 1-2 1 1-2  13-(21.5)-43 x 5-(7.5)-9y 3-(16.5)-40 x 3-(4.5)-5y 7-(13)-29 x 7-(8)-9(13)y 3-(6)-12 x 2.5-(4)-5y 3-(6)-10 x 2.5-(3)-5y  1-(plexus of several)  9.5-(18)-27 x 3-(5)-6p  Cyst presence at maturity  + ,+ ,+ ,+ ,+ ,+  + + ,-  + ,-  + +  Table I continued Rhizoids Isolate 1 1 2 2 3 3 4 4 5 6 6 7 7 8 8  No.  MHU Pollen MHU Pollen MHU Pollen MHU Pollen Pollen MHU Pollen MHU Pollen MHU Pollen  9 Pollen 10 MHU 10 P o l l e n  Nature coarse, blunt tipped coarse, tapering coarse, blunt tipped f i n e , bushy coarse, blunt tipped coarse, tapering coarse, blunt tipped coarse, tapering as above coarse, blunt tipped coarse, tapering coarse, blunt tipped coarse, tapering coarse, blunt tipped v e r y c o a r s e , grow from p o l l e n i n t o environment coarse, tapering f i n e , blunt tipped coarse, tapering  No. axes several l-(3)-5(9) several l-(3)-8 several 1-5 several 1-several as above several several several several several 1-5 (several) 1-5 several several  Color  i n culture  creamy w h i t e creamy w h i t e creamy w h i t e creamy w h i t e  creamy w h i t e creamy w h i t e y e l l o w brown  creamy w h i t e  Relation to  substrate  epibiotic endo-, e p i - , and i n t e r b i o t i c epibiotic endo-, e p i - , and i n t e r b i o t i c epibiotic endo-, e p i - , and i n t e r b i o t i c epibiotic endo-, e p i - , and i n t e r b i o t i c endo- or e p i b i o t i c epibiotic epibiotic epibiotic endo-, e p i - , and i n t e r b i o t i c epibiotic endobiotic p r i m a r i l y endo-, e p i - , and epibiotic endo-, e p i - , and  interbiotic interbiotic  Table I continued Zoospores I s o l a t e No.  Size  No. r e f . g l o b u l e s  1 MHU 1 Pollen 2 MHU 2 Pollen 3 MHU 3 Pollen 4 MHU 4 Pollen 5 Pollen 6 MHU 6 Pollen 7 MHU 7 Pollen 8 MHU 8 Pollen 9 Pollen 10 MHU 10 P o l l e n  4 . 0 - ( 4 . 5 ] -5.0y 3 . 0 - ( 4 . 0 ) -5. Op  1(2) 1 1-2 1-2 1(2) 1(2) 1(2) 1 1 1 1 1 1 1 1 1 1 1  3.03.03.53.03.03.03.04.0-  (5.0) -7.0y ( 4 . 0 ) -5. Oy (5.0) - 6 . O p ( 4 . 0 ) -5.0y (5.0) - 6 . Op (4.0) -4.5p (4.0) ( 5 . 0 :  2.5- ( 3 . 0 ) 3.53.04.03.54.03.03.5-  -5.Op -7.0y - 3 . 5 p  (4.5) - 6 . ( 3 . 5 ;  (4.5) (4.0; (4.5)  Op  Op -5.5y  - 4 .  -5.Op -5. Op  ( 3 . 5 ;  )-4.0p  ( 4 . 5 ;  -5. Op  Globule s i z e 1.5p l.Ou 1.5u minute 1.5p l.Op 1.5u l.Op l.Op  l.Op l.Op  1.5p l.Op 1.5p  l.Op 1.5p l.Op 1.5p  Globule  color  hyaline as above as above as above as above as above as above as above as above as above as above as above as above orange orange hyaline as above as above  Isolate  No.  Flagellum  length  1 1 2 2 3 3 4  MHU Pollen MHU Pollen MHU Pollen MHU  22-25y as above as above as above as above as above as above  4 5 6 6 7 7 8 8 9 10 10  Pollen Pollen MHU Pollen MHU Pollen MHU Pollen Pollen MHU Pollen  as as as as as as as as as as as  above above above above above above above above above above above  Remarks  D i s c h a r g e p r i m a r i l y through 1.5-(3.5)-8 x 4-(5.5)-7u  cyst,  Discharge p r i m a r i l y  through  cyst  Discharge p r i m a r i l y  through  cyst  D i s c h a r g e p r i m a r i l y through  cyst  36  3.  OCCURRENCE AND  DISTRIBUTION OF ZOOSPORIC FUNGI  FROM DEVON ISLAND, CANADIAN EASTERN ARCTIC  Introduction Methods and m a t e r i a l s Taxonomy and morphology  INTRODUCTION Devon I s l a n d , a p p r o x i m a t e l y A r c h i p e l a g o along a n o r t h - s o u t h  c e n t e r e d i n the Canadian A r c t i c  a x i s , l i e s between 74°25' to 77°30'N.  l a t . and 78°45' to 97°10'W. l o n g ( f i g . 4 0 ) . the h i g h a r c t i c b e l t ( N o r d e n s k j o l d , 1928;  Most o f i t i s l o c a t e d i n  P o l u n i n , 1951).  Major  p h y s i o g r a p h i c components i n c l u d e a low c o a s t a l f r i n g e backed by  an  a b r u p t l y r i s i n g escarpment and p l a t e a u . I n f o r m a t i o n on o c c u r r e n c e , d i s t r i b u t i o n and e c o l o g y o f z o o s p o r i c f u n g i i n the A r c t i c i s s c a n t , p a r t i c u l a r l y i f the A r c t i c i s d e f i n e d i n  i' the r e s t r i c t e d sense ( N o r d e n s k j o l d , 1928;  P o l u n i n , 1951).  Early reports  of f u n g i , summarized by L i n d (1934), were based on s t u d i e s of phanerogamic herbarium  specimens.  Consequently,  Synchytrium,  Physoderma and  P e r o n o s p o r a r e p r e s e n t e d the b u l k of the Phycomycete component of a r c t i c mycota.  L i n d e r (1947) made the f i r s t s u b s t a n t i a l c o n t r i b u t i o n t o our  understanding  o f a q u a t i c f u n g i from a r c t i c r e g i o n s .  s p e c i e s , f o u r new  He d e s c r i b e d  to s c i e n c e , from m a t e r i a l c o l l e c t e d and p r e s e r v e d  N. P o l u n i n d u r i n g the Canadian E a s t e r n A r c t i c E x p e d i t i o n s of 1934 1936.  S a v i l e (1961, 1968)  12 by and  and S a v i l e and Parmelee (1964) made e x t e n s i v e  b o t a n i c a l and m y c o l o g i c a l c o l l e c t i o n s a t p o i n t s s c a t t e r e d over  the  37  Canadian A r c t i c . on phanerogams.  T h e i r s t u d i e s were r e s t r i c t e d  to f u n g i  parasitic  As i n the c a s e of e a r l i e r s t u d i e s , r e p o r t s of z o o s p o r i c  fungi are r e s t r i c t e d  to Synchytrium, Physoderma and  A few German workers  studied  Peronospora.  z o o s p o r i c f u n g i from a r c t i c  b u t " A r c t i c " i s too b r o a d l y i n t e r p r e t e d i n t h e i r r e p o r t s . samples  labeled a r c t i c  Most  soil  ( i . e . , from Lappland and I c e l a n d ) were p r o b a b l y  s u b a r c t i c s o i l s i n the sense of Tedrow (1970). i n Lappland would b e s t be r e f e r r e d  Also high elevation  to as a l p i n e .  s i x s o i l c o l l e c t i o n s from S p i t s b e r g e n .  Harder  (1954) examined  and  (Nowak.) S c h r o e t e r i n a sample from a Poa meadow.  the s i t e s y i e l d e d f u n g i .  sites  He r e c o v e r e d O l p i d i u m pendulum  Zopf i n a sample from a Dupontia moor near Longyearbyen elegans  soils,  Nowakowskiella Only 33% of  G a e r t n e r (1954) a l s o surveyed s i x s o i l  c o l l e c t i o n s from S p i t s b e r g e n and found the same organisms as Harder. In a d d i t i o n he found s e v e r a l Pythium s p e c i e s . samples  Hohnk (1960) c o l l e c t e d  10  a c r o s s a beach a l o n g King F r e d e r i k VI C o a s t , Greenland a t  a p p r o x i m a t e l y 63°N. l a t . - F i v e of the samples  came from the emergent  p o r t i o n of the beach and f o u r from the submerged p o r t i o n . y i e l d e d a t l e a s t one s p e c i e s .  Every c o l l e c t i o n  Among the s p e c i e s found a r e :  Saprolegnia  sp., T h r a u s t o c h y t r i u m p r o l i f e r u m Sparrow, Sapromyces elongatus  (Cornu)  Coker, Apodachlya brachynema ( H i l d e b r a n d ) P r i n g s h e i m , Pythium monospermum P r i n g s h e i m , I?, graminicolum v a r . s t a g n i Hb'hnk, I?, i r r e g u l a r e . ' l Bulsman, Pythium spp., P y t h i o g e t o n ramosum Minden, O l p i d i u m l u x u r l a n s F i s c h e r , Rhizophydium R. mamillatum  polllnls-pini  (Braun) Zopf, R.  (Tomaschek)  sphaerotheca Zopf,  (Braun) F i s c h e r , P h l y c t o c h y t r i u m c h a e t i f e r u m K a r l i n g  and  R h i z o p h l y c t l s r o s e a (deBary and Woronin) F i s c h e r . Other workers have l o o k e d s p a r i n g l y f o r z o o s p o r i c f u n g i i n a r c t i c soils.  Karling  (1951) i s o l a t e d C a t e n o p h l y c t i s  (Phlyctorhlza) v a r i a b i l i s  38  ( K a r l i n g ) K a r l i n g from tundra s o i l lat.,  171°W. l o n g . ) .  Kobayasi  Alaska for a l l fungi.  Barr  gically  Pythium c a r o l i n i a n u m Matthews.  Among other  i s o l a t e was  p h y s i o l o g y of an i s o l a t e of  completely  r e g i o n s to the s o u t h .  10C, , o p t i m a l l y a t 20,  25 and  similar  a l l a t 10C.  30C,  and  B a r r suggests  not a t a l l a t 35C.  METHODS AND  Conversely,  25C  these d i f f e r e n c e s may  genetic selection i n l o c a l conditions.  Allomyces,  t r o p i c i n d i s t r i b u t i o n , was  I s l a n d s i t e by G.C.  from  However, the a r c t i c i s