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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 IN COASTAL AND STEPPE SOILS by TOM BOOTH B.A., Eastern B a p t i s t College 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 the Department of Botany i We accept t h i s t h e s i s as conforming to the req u i r e d standard THE UNIVERSITY OF BRITISH COLUMBIA A p r i l > 1 9 7 1 In present ing th i s thes is in pa r t i a l fu l f i lment , o f the requirements fo r an advanced degree at the Un ive rs i t y of B r i t i s h Columbia, I agree that the L ibrary sha l l make it f r ee l y ava i l ab le for reference and study. I fu r ther agree that permission for extensive copying of th i s thes i s fo r s cho la r l y purposes may be granted by the Head of my Department or by h is representa t i ves . It is understood that copying or pub l i c a t i on of th i s thes is fo r f i nanc i a l gain sha l l not be allowed without my wr i t ten permiss ion. Department of Botany  The Un ivers i t y of B r i t i s h Columbia Vancouver 8, Canada A p r i - 1 3 9 , mi ABSTRACT Certain c r i t e r i a used to d i s t i n g u i s h c h y t r i d taxa are highly v a r i a b l e and unstable. This v a r i a b i l i t y i s evident from a study of ten single-spore i s o l a t e s of an Entophlyctis species 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. The observed v a r i a t i o n s place these Entophlyctis i s o l a t e s i n d i f f e r e n t species, genera, subfamilies, f a m i l i e s , and s e r i e s , thus demonstrating that various current taxonomic di s p o s i t i o n s and concepts are of questionable value. Zoosporic fungi, mainly C h y t r i d i a l e s , are widely d i s t r i b u t e d i n coastal and steppe s o i l s . Based on d i s t r i b u t i o n a l records and determination of ph y s i c a l and chemical parameters f o r each s o i l c o l l e c t i o n , eleven c h y t r i d and chytridiaceous species are d i v i s i b l e into four 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 occasionals. Certain aspects of th i s study are problematic which emphasizes the necessity of a more sophisticated l i n e of approach. Temperature-salinity growth responses of f i f t y - s e v e n axenic single-spore cultures of zoosporic fungi, maintained under s i m i l a r c o n t r o l l e d conditions f o r a minimum of s i x months, cor r e l a t e with habitat types and cer t a i n environmental parameters. These responses, s i m i l a r over several i s o l a t e s of a species from the same s o i l c o l l e c t i o n , are temporally constant and, thus, are ecotypic. i i i TABLE OF CONTENTS 1. Introduction . . . . 1 2. Problematical Taxonomic C r i t e r i a i n the C h y t r i d i a l e s : Comparative Morphology of Ten Entophlyctis sp. Isolates . . 7 3. Occurrence and D i s t r i b u t i o n of Zoosporic Fungi from Devon Island, Canadian Eastern A r c t i c . . . 36 4. Occurrence and D i s t r i b u t i o n of Some Zoosporic Fungi from S o i l s of Hibben and Morseby Islands, Queen Charlotte Islands . . . . . . . . 63 5. Occurrence and D i s t r i b u t i o n of Zoosporic Fungi and Some Actinomycetales i n Coastal S o i l s of Southwestern B r i t i s h Columbia and the San Juan Islands . . . . . . . 105 6. Occurrence and D i s t r i b u t i o n of Chytrids, Chytridiaceous Fungi and some Actinomycetales from S o i l s of Oregon, C a l i f o r n i a and Nevada . . . . . . . . . . . 137 7. Occurrence of ,,Chytrids and Chytridiaceous Fungi from Two Long Island, N.Y. Beaches 171 8. Occurrence of Chytrids and Chytridiaceous Fungi from Some Mexican Beach and Desert S o i l s 175 9. D i s t r i b u t i o n of C e r t a i n S o i l Inhabiting Chytrid and Chytridiaceous Species Related to some Physical and Chemical Factors 178 10. Ecotypic Responses of Chytrid and Chytridiaceous Species to Various S a l i n i t y and Temperature Combinations 203 11. Discussion . . . . . . 226 L i t e r a t u r e C i t e d 230 Appendices • 253 I. Habitat descriptions 253 I I . Species name and numbers 267 I I I . Habitat names and numbers 271 IV. A. Times each species occurs at 0 ° / o o and 7 ° / 0 0 i n a l l s i t e s and s t a r t s of each habitat type . 273 B. Occurrence and frequency of each species i n ten Base Camp Lowland habitats . . . . . . . . 350 V. Number of d i f f e r e n t species recovered at 0 ° / o o and 7 ° / 0 0 i n each habitat type 355 VI. Computed occurrences, frequencies and d i s t r i b u t i o n a l i n t e n s i t y index of each species at 0 loo and 7 /oo i n h a b i t a t types . . . . . . . 358 VII. H a b i t a t s and values of p h y s i c a l and chemical parameters of c o l l e c t i o n 'sites 454 VIII. S t a t i s t i c s of p h y s i c a l and chemical f a c t o r s i n h a b i t a t s 491 IX. Ranges, means, v a r i a n c e s , standard d e v i a t i o n s and f i v e percent confidence i n t e r v a l s of chemical f a c t o r s i n occurrence s i t e s of some species 5^.06 X. C o r r e l a t i o n of chemical and p h y s i c a l f a c t o r s w i t h DII of c e r t a i n species 518 XI. 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 data on growth a t , and comparisons between s a l i n i t y and temperature combinations f o r each species . . 561 LIST OF TABLES Table I. Developmental and morphological d e t a i l s of Eritophlyctis sp. i s o l a t e s 1-10 . . . . . . . . . . . - 31 Table I I . S i t e name and number, l a t i t u d e and longitude and habitat type of Queen Charlotte Islands c o l l e c t i o n s . . . . . •. . . . . . . . . . . . . . . 101 Table I I I . S i t e name and number, l a t i t u d e and longitude and habitat type of Southwestern B.C. and San Juan Islands c o l l e c t i o n s . . . . . . . . . . . . 133 Table IV. 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 longitude and habitat type of 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 Table V. Location name, s i t e number and habitat type of Long Island, N.Y. c o l l e c t i o n s . . . . •. . . . . . . 174 Table VI. D.I.I, of each species over habitats . . . •. . . . . 197 Table VII. Mean values of environmental factors over habitats . . 199 Table VIII. Ranges and means of environmental factors over occurrences of each species . . . . . . . . - 200 Table IX. Correlations of each species with environmental factors . . . . . . . . . . . . . . . . -. . . . . . 202 Table X. Figure number, taxon, c o l l e c t i o n s i t e name and number and habitat type of each i s o l a t e . . . . . . 220 Table XI. Certain environmental parameters at recovery s i t e s of each Isolate . . . . . . . . 224 LIST OF ILLUSTRATIONS Map I. North American c o l l e c t i n g s i t e s . . . 6 Map I I . Devon Island Base Camp Lowland . . . . . . . . . . 61 Map I I I . Queen Charlotte Islands c o l l e c t i n g s i t e s ' ; . . . . 99 Map IV. Southwestern B.C. and Northwestern Washington c o l l e c t i n g s i t e s . . . . . . . . . . . . . . . . . 129 Map V. Oregon, C a l i f o r n i a and Nevada c o l l e c t i n g s i t e s . . 163 Plate I- -. 15 P l a t e ' I I . . . . . . . . . . . . . . . . . . . . . . . . . • . 18 Pla t e I I I . . . . . . . . . . . . . . 21 Plate IV . . . . . . . . . . 24 Pla t e V . . . . . . . . . . . . . . . . . 27 Plate'VI • . . . . . . . . . . 48 P l a t e VII 53 Pla t e VIII 58 Plate IX . . . . . . . . . . . . . . . . . . 70 P l a t e X . . . . . . . . . . . . . . . . . . . . . . . . . . . 77 Pla t e XI 84 P l a t e XII . . . . . . . . . . . . . . . . . . . . 90 P l a t e X I I I 96 Plate XIV . . . . . . . . . . . . . . . . . . . . . 113 Pla t e XV"... . . 125 Plate XVI 141 Pla t e XVII . . . . . . . . . . . . . -. . . . . . . . . . 147 Pla t e XVIII 152 Pla t e XIX . . . . . . . . . . . . . . . . . . . . . . 158 v i i Figures Section 9. Figure 1. Figure 2. Figure'3. Figures 4-19. Section 10. Figures 1-60. DII of chytrids and chytridiaceous ^ species over nineteen habitats . . . . . Mean habitat values of environmental parameters \1§5^ Ranges, means and standard deviations of environmental f a c t o r s over s i t e s where selected species occur 189 DII plotted against habitat means for c e r t a i n species . . . . . . . . . . . . . 192, "193 Growth clocks of chytrids and chytridiaceous species at various temperature-salinity combinations . . . . .209^213 Figure 61. Environmental factor l e v e l s f o r i s o l a t e s grouped into ecotypes 218-^ . v i i i ACKNOWLEDGEMENTS P a t i e n t advice, common sense, w i l l i n g d i s c u s s i o n and c r i t i c a l comments of Dr. G. C. Hughes are v e r y much appreciated. Drs. 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. Barr k i n d l y i reviewed p o r t i o n s of the d i s s e r t a t i o n during i t s p r e p a r a t i o n . Drs. 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 during the 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 and Dr. G. Rank r e s p e c t i v e l y secured samples from Devon I s l a n d and Mexico. S p e c i a l thanks to Mr. N e i l Carey whose seamanship and 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 Islands p o s s i b l e . Access to s i t e s i n the Huntington Harbor Region (N.Y.) was provided by C_. G_. A. V. Voyager I I owned and operated by Mr. J . R. Newcomer. I am a l s o indebted to the o f f i c e r s and crew of £. j>_. _S. Vector f o r t h e i r h e l p f u l a s s i s t a n c e . Laboratory f a c i l i t i e s were f r e e l y provided by Drs. Hughes, C. 0. Person, R. J . Bandoni, W. K. F l e t c h e r , R. F. Scagel, T. B i s a l p u t r a and V. J . K r a j i n a . I am t h a n k f u l f o r the t i r e l e s s computer programming e f f o r t s of Mrs. Dolores L a u r i e n t e . 1 1. INTRODUCTION I n t r o d u c t i o n Study Purposes Taxa of the 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 Peronosporales are in c l u d e d i n t h i s p r e s e n t a t i o n w i t h o c c a s i o n a l r e p o r t s of Actinomycetales even though they are not f u n g a l . Morphological features used to d i s t i n g u i s h the f u n g a l orders and t h e i r subordinate groups are presented by K a r l i n g (1932), Sparrow (1943, 1958, 1960), L i t v i n o v (1958) and F u l l e r (1966). Among morphological f e a t u r e s used to d i s t i n g u i s h c h y t r i d and c h y t r i d i a c e o u s taxa are: f l a g e l l a r type and attachment l o c a t i o n ; arrangement and complexity of spore 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 extent and r e p r o d u c t i v e center 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 ; r e l a t i o n s h i p of t h a l l u s to sub 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 , or 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 ornamentations, discharge p a p i l l a numbers, shape, and l o c a t i o n ; apophysis presence or absence and s t r u c t u r e ; r h i z o i d a l branching, coarseness and number of 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), Bartsch (1939), Berdan (1941a, 1941b;, .1942), Whiffen (1944), A n t i k a j i a n (1949), Emerson (1950), Haskins (1950), R e i t h (1950), Koch (1951, 1957, 1968), Johnson (1955, 1957a, 1957b, 1969), M i l l e r (1955, 1961, 1968), Sparrow and Paterson 2 (1955), Paterson (1956, 1963), Willoughby (1956, 1959, 1961a, 1961b, 1962, 1964, 1965), L i t v i n o v (1958), Scholz (1958a), B o s t i c k (1968), Booth and M i l l e r (1968), Booth (1969), Dogma (1969a), Konno (1969) , Barr (1970a, 1970c), H a s i j a and M i l l e r (1970) and S a l k i n (1970) question the v a l i d i t y of one or more of the characters l i s t e d above at 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 date few s t u d i e s , w i t h the exception of 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 (1949), Haskins and Weston (1950), Koch (1951, 1957), Willoughby (1957, 1958), F u l l e r (1962), M i l l e r (1968), Paterson (1963), Barr and Hickman (1967a, 1967b), Booth and M i l l e r (1968), Barr (1969, 1970a, 1970b) and H a s i j a and M i l l e r (1970), have considered the 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) to study morphologic;expression 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 the works of Sparrow (1960) and Johnson and Sparrow (1961) i t i s evident that 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 earth's s o i l s . Since the appearance of these 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), Willoughby (1958, 1959, 1961a, 1961b, 1962, 1963, 1964, 1965, 1966, 1969), Harder and Uebelmesser (1959), P e r s i e l (1959-, 1960a, 1960b), Sparrow and Koch (1959), G o l d s t e i n (1960), M i l l e r (1961, 19.62, 1965, 1968), Willoughby and Townley ( 1 9 6 1 a , 1961b)., Harder and P e r s i e l (1962) , Scott (1962), G r i f f i t h s and Jones (1963), A p i n i s ( 1 9 6 4 1 , Ulken (1964, 1965, 1966, 1970), Sparrow et a l . (1965)., Gaertner and Sparrow (1966), Sparrow (1965, 1966, 1968a, 1968b, 1969), Gaertner (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 Miller (1970), Salkin (1970) and Thornton (1970), add to this impression. Early distributional studies, conducted by workers at Gb'ttingen, indicate: aquatic fungi are common in tropical soils (Sorgel, 1941); disturbed soils are greater in species diversity than natural soils (Sergei, 1941; Harder, 1948; Remy, 1948; Gaertner, 1954); more genera of aquatic Phycomycetes occur in clay soils than i n sand soils (Remy, 1948); moist soils are richer in species diversity than dry soils (So'rgel, 1941; Remy, 1948); lower Phycomycetes are not tied to definite niches (Remy, 1948); aquatic Phycomycetes occur in definite clusters in 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 in the rhizosphere depending on the higher plant contributing roots to the site and plants at flowering stages generally impeded fungus frequency (Reinboldt, 1951); fewer species occur in Arctic and Antarctic soils than soils at lower latitudes (Harder, 1954; Gaertner, 1954; Harder and Persiel, 1962); and diversity of species i n soils decreases from intertidal to dune sites (Harder and Uebelmesser, 1955); and most chytrids from intertidal areas are only facultatively marine (Scholz, 1958b). Studies by HShnk (1952, 1956) support the notion that the diversity of species increases from the int e r t i d a l , landward. Willoughby (1961a, 1962, 1964, 1965) carried out distributional studies and found: certain species are found in wet soils while others are recovered from 4 dry s o i l s , lake margins have a r i c h e r c h y t r i d mycota than pastureland above or submerged muds below; d i s t r i b u t i o n of c e r t a i n species i s a function of s o i l pH; and occurrence of Rhizophlyctis rosea i s higher i n n u t r i e n t r i c h s i t e s than nutrient poor l o c a t i o n s . Most e c o l o g i c a l information on chytrids and chytridiaceous fungi i s re 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 (Reinboldt, 1951; Willoughby, 1961, 1965) to co r r e l a t e fungus d i s t r i b u t i o n with s o i l p h y s i c a l and chemical parameters. Studies by Scholz (1958a) and Barr (1970a, 1970b, 1970d) i n d i c a t e that chytrids are morphologically and/or p h y s i o l o g i c a l l y adapted to t h e i r environment. Since no extensive d i s t r i b u t i o n a l study over several degrees of l a t i t u d e has yet been attempted i n North America, one aim of my study i s to account f o r occurrence and d i s t r i b u t i o n of zoosporic fungi from s i t e s as f a r noth as Devon Island, Canadian Eastern A r c t i c and as f a r south as Mazatlan, Mexico ( f i g . 1). Before reporting t h i s information, I propose to discuss problematic taxonomic c r i t e r i a i n the fungi and present examples of such problems i n order to set the ph i l o s o p h i c a l approach taken i n species i d e n t i f i c a t i o n . Another purpose of the study i s to r e l a t e fungis d i s t r i b u t i o n to chemical and ph y s i c a l parameters but the basic thesis i s that these fungi are adapted to t h e i r environments and, hence, at the i n t r a s p e c i f i c l e v e l they are composed of several ecotypes. Map I . — F i g u r e 1. Approximate l o c a t i o n s , i n d i c a t e d by s o l i d t r i a n g l e s , of c o l l e c t i n g s i t e s i n North America. 6 7 2. PROBLEMATIC TAXONOMIC CRITERIA IN THE CHYTRIDIALES: COMPARATIVE MORPHOLOGY OF TEN ENTOPHLYCTIS SP. ISOLATES I n t r o d u c t i o n Methods and m a t e r i a l s T y p i c a l development Developmental and morphological v a r i a t i o n s Comparative development D i s c u s s i o n INTRODUCTION Several characters used to d i s t i n g u i s h c h y t r i d taxa are v a r i a b l e and among these are zoospore s i z e , shape, p o s i t i o n of f l a g e l l a r attachV ment and numbers of spore g u t t u l e s ; sporangium s i z e and shape; r. E l z o i d a l nature, extent and branching; sporangium w a l l enations or punctae; discharge p a p i l l a numbers, shape and l o c a t i o n ; apophysis presence 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 . Zoospores—A number of workers (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 spore and l i p i d body diameters, number of l i p i d bodies, shape and p o s i t i o n of f l a g e l l a r attachment In planonts from sporangia on d i f f e r e n t s u b s t r a t e s . Conversely Paterson (1963) r e p o r t s that spores of two Rhizophydium species (Rhizophydium sp. 1 and Rhizophydium sp. 2) have constant s i z e on a l l s u b s t rates t e s t e d , and Barr (1970) detected no s i z e d i f f e r e n c e s i n spores of Phlyctochytrium r e i n b o l d t a e P e r s i e l from sporangia on d i f f e r e n t s u b strates (pine p o l l e n and YpSs medium). 8 Sporangia—Koch (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 substrates but at l e a s t some species have sporangia of h i g h l y v a r i a b l e shape on the same su b s t r a t e (Koch, 1951, 1957; H a s i j a and M i l l e r , 1970). Among f a c t o r s causing s i z e v a r i a t i o n s are: d i f f e r e n t substrates ( B o s t i c k , 1968; B a r r , 1969, 1970a; S a l k i n , 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 (Paterson, 1963); b a i t c o n c e n t r a t i o n and temperature ( 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 extent of these s t r u c t u r e s on d i f f e r e n t s u b strates has been noted 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 (unpublished data) that a t 20°C r h i z o i d s of Rhizophydium  sphaerotheca Zopf, Chytriomyces h y a l i n u s K a r l i n g , P hlyctochytrium  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 are l e s s branched and longer at 15°/ 0 0 s a l i n i t y than at 0 " / o o s a l i n i t y . Sporangium w a l l markings—Some of these s t r u c t u r e s are i n f l u e n c e d by environmental c o n d i t i o n s , f o r i n s t a n c e , occurrence 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 are i n f l u e n c e d by p o l l e n type and water source (Koch, 1957). When f u l l y s t u d i e d , normal species 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 described taxa. One such c l i n e , based on the number of te e t h i n a row around the sporangium a p i c a l or 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 kinson sensu Sparrow (1938), P* - plani c o r n a e sensu Willoughby (1961a), P_. c i r c u l i d e n t a t u m Koch and P_. multidentatum Umphlett. V a r i a t i o n s i n number, l o c a t i o n and complexity of enations on sporangia of M i l l e r ' s (1968) "dentate" and Booth's (1971b) Rhizophydium sp.-Phlyctochytrium a u r i l a e A j e l l o complex suggest another such c l i n e . 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 (Willoughby and Townley, 1961a) of Chytriomyces pociilatus Willoughby and 9 Townley s e v e r a l workers (Willoughby, 1965; Sparrow, 1968; Dogma, 1969a and Booth and B a r r e t t , 1971) have reported t h a l l i w i t h cupule-free sporangia. Considering these observations there i s , as Dogma (1969a) suggests, a p o s s i b l e C_, poculatus-C. anriulatus Dogma sporangium type c l i n e . Discharge papillae—Numbers of these s t r u c t u r e s are a f f e c t e d by sporangium s i z e (Barr, 1969) and s a l i n i t y (Scholz, 1958a). Since s e v e r a l Phlyctochytrium species are d i s t i n g u i s h e d by p a p i l l a e numbers the above observations r a i s e doubt about t h e i r v a l i d i t y as separate taxa. This becomes more obvious when M i l l e r ' s (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 multiporous P h l y c t o c h y t r i u m species and Booth's (1971c) P_. africarium Gaertner-P_. s p e c t a b i l e Uebelmesser p a p i l l a e number complex are considered. Apophysis—Couch (1932'), Koch (1957), and M i l l e r (1961, 1968) have questioned use of t h i s s t r u c t u r e to separate P h l y c t o c h y t r i u m from Rhizophydium. Other workers ( K a r l i n g , 1967a; Johnson, 1969) have recognized P h l y c t o c h y t r i u m species without an apophysis and I observed (unpublished data) nonapophysate t h a l l i of Phlyctochytrium chaetlferum K a r l i n g on MRU medium and Pinus s y I v e s t r i s L. p o l l e n . Several species? of Phlyctochytrium, are separated p a r t i a l l y by apophysis shape. However, the usefulness of t h i s character i s d o u b t f u l s i n c e each of s e v e r a l s i n g l e spore i s o l a t e s of a few Phl y c t o c h y t r i u m species were observed with?, a h i g h l y v a r i a b l e apophysis shape (Booth, 1971b, 1971c, 197Id).. R h i z o i d a l n a t u r e — A h a u s t o r i u m - l i k e or tu b u l a r untapering r h i z o i d a l 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) of the P h l y c t i d i o i d e a e . Y e t , Paterson (1956), Willoughby (1965). and Umphlett. and Olson (1965) described P h l y c t o c h y t r i u m species w i t h r h i z o i d a l stages s i m i l a r to P h l y c t i d i u m and I noted (Booth, 1971b) a fungus (,?Phlyctochytriuro 10 sp.) producing e i t h e r haustoria or tapering r h i z o i d s . Karling (1967b) established P o l y p h l y c t i s , with Phlyctochytrium unispinum Paterson as the type, and p a r t i a l l y distinguished i t from the other genera of the P h l y c t i d i o i d e a e by the catenulate r h i z o i d s that he and Willoughby and Townley (1961b) observed. This character, however, may not be r e l i a b l e since Dogma (1969a) described and figured P_. unispinum material exactly as the type, with a h a u s t o r i a l r h i z o i d a l system. Other species with r h i z o i d s varying from a tapering to catenulate nature have been reported (Reinboldt, 1951; K a r l i n g , 1967a; Sal k i n , 1970). Rhizoidal a x e s — C e r t a i n genera are distinguished by the presence of more than one axis per sporangium. However, Kar l i n g (1949b), Koch (1957) and Paterson (1963) have observed more than one r h i z o i d a l axis on sporangia of species i n t y p i c a l l y uni-axiate genera and I have also observed (unpublished) several r h i z o i d a l axes on sporangia of Rhizdphydium  sphaerotheca and R. p o l l i n i s - p i n i (Braun) Zopf on MHU medium. ' Rhizo- phydium normally has uniaxiate sporangia. Thallus development—Whiffen (1944) proposed t h a l l u s development type as the basis 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 species and genera. T h a l l i of Rhizophlyctis, Chytriomyces, Rhizidium, Phlyctochytrium  Allochytridium and Asterophlyctis species can develop from swelling of e i t h e r the cyst or germ tube (Karling, 1947b, 1949b; A n t i k a j i a n , 1949; Koch, 1957; S a l k i n , 1970). Thallus-substrate r e l a t i o n s — M o n o c e n t r i c t h a l l i are endobiotic (Olpidiaceae, Synchytriaceae, Entophlyctoideae and Endochytrioideae) , e p i b i o t i c (Phlyctidioideae and Chytridiodeae) or i n t e r b i o t i c (Rhizidiaceae and Chytridioideae). Whiffen (1944) suggests that i n the case of saprobic 11 chytrids chance alone may determine whether^ t h a l l i are i n t r a m a t r i c a l or extramatrical. For example, Phlyctochytrium,punctaturn (Koch, 1957), P_. chaetiferum (Booth, 1969) and Rhizidium richmondense Willoughby (Karling, 1967b) may be either e p i b i o t i c or i n t e r b i o t i c and Rhizophlyctis  spinosa (fearling) Sparrow may even produce t h a l l i which are endobiotic and e p i b i o t i c at the same time (Karling, 1974b). Monocentricity vs. P o l y c e n t r i c i t y — T h e s e t h a l l i configurations, used to separate taxa at the family l e v e l , are v a r i a b l e . Some taxa which are normally monocentric may sometimes be p o l y c e n t r i c , f o r instance, Rhizophlyctis hyalina (Karling) Sparrow and Rhizidium varians Karling may be monocentric or po l y c e n t r i c (Karling, 1947b, 1949b). Such v a r i a t i o n r e l a t e s these taxa, normally,considered i n the Rhizidiaceae to the Cladochytriaceae. Operculation—Of a l l the characters used to separate c h y t r i d taxa t h i s one has possibly come under the most disfavor and rigorous s c r u t i n y . The primary argument has revolved around the taxonomic l e v e l at which this character i s to be used. Since Sparrow's (1942) use of operculation to e s t a b l i s h two separate s e r i e s i n the order, Whiffen (1944) has presented several reasons for not placing much importance on operculation and Karling (1967d) stated that separation of Entophlyctis and D i p l o p h l y c t i s from Endochytrium and Nephrochytrium " . . . places undue taxonomic emphasis on the presence of an operculum above the generic l e v e l . . . ." Several other workers, Koch (1951, 1957), Willoughby (1956, 1961a, 1961b), M i l l e r (1968) and Bostick (1968) have observed v a r i a t i o n s involving operculum presence and absence, occurrence ;of operculum-like structures and/or incompletely formed opercula i n species normally considered inoperculate. Interpretation of v a r i a b i l i t y and comparison of these 12 studies are problematic because each worker seems to have h i s own concept of what constitutes an operculum. For instance, Umphlett and Koch (1969) describe the quasi-operculum which they consider intermediate between inoperculate and operculate conditions. Johnson (1969), on the other hand, suggests that the quasi-operculum i s a type of f a l s e operculum. Besides the necessity o f a d e f i n i t i v e study of operculation and inoperculation, recognized by Koch i n 1957, perhaps common accord i s required concerning the nature of the operculum and, for that matter, a l l other taxonomic characters. METHODS AND MATERIALS During a survey of coa s t a l and i n t e r i o r s o i l s from the Queen Charlotte Islands to C a l i f o r n i a and Nevada, several s i n g l e spore i s o l a t e s of an Entophlyctis species were obtained by streaking or placing b a i t s on modified Harder and Uebelmesser (1955) medium (MHU, 0.5%'malt extract, 0.3% peptone, 0.1% dextrose, 1.1% agar i n 1 1 water of 0°/ Oo or 7°/O0 s a l i n i t y with .03% Streptomycin sulphate—774 mcg/mg and .03% P e n i c i l l i n G—4 x 10 u n i t s ) . For three days a f t e r streaking each plate was surveyed for s i n g l e encysted spores or i n c i p i e n t sporangia; these were removed and placed on MHU medium minus a n t i b i o t i c s . The i s o l a t e s , which vary i n several characters include: 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 Pt., B.C. .(53°13.1'N., 131°59.3'W.); 46, One-half Beach, B.C. (48°47.1'N., 123°04.8'W.); 47, 54, Hibben Island, B.C. (52°58.6'N., 132°19.1'.W.); 52, Long Beach P r o v i n c i a l Park, B.C. (49°03.2'N., 125°43.2'W.); Au thor's c u l t u r e no. 13 61, A n v i l Island, B.C. (49°31.4'N., 123°17.3'W.); 66, Bird Cove, B.C. (50°12.3'N., 125°05.8'W.) and 74, Mono Lake I I , C a l i f . (37°57.0'N., 119°05.0'W.). Morphology and development of the i s o l a t e s were studied on p o l l e n and MHU medium by the methods of Booth and Barrett (1971). TYPICAL DEVELOPMENT AND MORPHOLOGY OF ALL ISOLATES On MHU medium the s p h e r i c a l zoospore cyst germinates by a narrow tube ( f i g . 13), a broad bl'eb ( f i g s . 1, 14) or a broad tube ( f i g . 2). A r h i z o i d a l axis pushes out of the expanding i n c i p i e n t sporangium ( f i g s . 15-16). Soon the axis branches along the course of the r h i z o i d ( f i g . 17) or at the juncture with the developing sporangium ( f i g s . 3, 18, 19). At this time the zoospore cyst may appear devoid of protoplasm ( f i g s . 17-19). The cyst generally remains empty through the next stages i n which several r h i z o i d a l axes a r i s e on the developing sporangium ( f i g s . 4, 20). As the i n c i p i e n t sporangium continues to swell, r e g u l a r l y ( f i g s . 5, 6, 25) or i r r e g u l a r l y ( f i g s . 21, 22), and the r h i z o i d s branch dichtomously ( f i g s . 3-6, 21, 22) and elongate, the zoospore cyst may r e f i l l with projioplasm ( f i g s . 5, 6, 21, 22). At maturity the sporangium i s s p h e r i c a l with several r h i z o i d a l axes. The cyst may be very evident ( f i g . 23), diminutive ( f i g . 26) or completely incorporated into the sporangium. Rhizoids are tubular and have blunt tips ( f i g s . 25, ^ 6 ) . Discharge p a p i l l a e ( f i g . 28) or tubes ( f i g . 29) of varying length break the double contoured sporangium wall at one to several places. The zoospore cyst may deliquesce and function as a discharge pore ( f i g . 27). Zoospore discharge i s preceded by p a p i l l a r d i s s o l u t i o n or deliquescence of a large (18-26 x 8-18y) gelatinous plug at the tube o r i f i c e . Spores are motile i n the sporangium p r i o r to discharge and escape s i n g l y 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 bleb, f h . (freehand), x200. F i g . 2. Germinating cyst and germ tube, f h . , xlOOO. F i g . 3. Rhizo i d a l axis developing at germ tube end, x500. F i g . 4. Developing thallus with several r h i z o i d a l axes and an empty cyst, xlOOO. F i g . 5. Developing thallus with protoplasm f i l l e d c yst and dichtotomously branching r h i z o i d s , xlOOO. F i g . 6. Thallus with sporangium almost completely expanded, xlOOO. F i g s . 1;, 8-10, scale A; F i g s . 2, 4-6, 11, 12, scale B; F i g . 7, scale C; F i g . 3, scale D; Fi g s . 13-16, 19, 24-26, scale E; F i g s . 17, 19, 20, 27, 29-33, scale F; Fi g s . 21-23, 24, scale G. 'Unless designated f h . , a l l drawings done with camera l u c i d a . 15 1 6 pores or tubes. Mo t i l e spores are s p h e r i c a l , each containing 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 cyst germinates and produces a tube which penetrates the microspore exine. The germ tube swells within the substrate and normally one 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 sporangium. At maturity sporangia are usually s p h e r i c a l ( f i g s . 7, 30) but they can be e l l i p s o i d a l ( f i g . 31), pyriform, saccate or i r r e g u l a r . Zoospores are motile p r i o r to discharge and escape the sporangial confines through the zoospore cyst ( f i g s . 7, 30). DEVELOPMENTAL AND MORPHOLOGICAL VARIATIONS COMMON TO ALL ISOLATES Frequently on MHU medium the germ tube ceases to expand and the sporangium then develops from the fir.pt r h i z o i d a l axis ( f i g . 24). The remaining cyst may then appear n i p p l e - l i k e ( f i g . 23). On both MHU medium and pollen , sporangia may develop by. expansion of the cyst rather than the germ tube with varying degrees of frequency for each i s o l a t e . On p o l l e n an endo-exogenous type of development ( i . e . both the cyst and germ tube swell) may occur and t h a l l i are endobiotic and e p i -b i o t i c at the same time ( f i g . 11). Various other thallus r e l a t i o n s h i p s are observed on p o l l e n . Sporangia can be endobiotic ( f i g s . 7, 12, 30, 31)-, e p i b i o t i c ( f i g s . 32, 33) or i n t e r b i o t i c ( f i g . 34). Rhizoids ( f i g . 12) or tubes ( f i g s . 31-33) frequently a r i s e from the cyst. Spores d i s -charge from e p i b i o t i c and i n t e r b i o t i c sporangia on p o l l e n almost e x c l u s i v e l y / through tubes or tube-like structures. However, as previously mentioned, endobiotic sporangia i n p o l l e n generally discharge th e i r spores through the zoospore cyst. But sometimes discharge tubes, which can a r i s e from any point around the i n c i p i e n t sporangium, are formed^ ( f i g i 11) and the P l a t e I I — F i g u r e s 7-12. Entophlyctis sp. F i g . 7. Empty i n t e r -bio t i c sporangium with two r h i z o i d a l axes, f h . , x750. F i g . 8. • Simple discharge tubes, f h . , x2000. F i g . 9. Complex discharge tubes, f h . , x2000. F i g . 10. Branched discharge tube, f h . , x2000. F i g . 11. Endo-, e p i b i o t i c sporangium with discharge tubes, xlOOO. F i g . 12. Endobiotic zoosporangium with i n t e r b i o t i c r h i z o i d s . 18 zoospore cyst i s functionless i n discharge. Discharge 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), simple ( f i g s . 8, 32, 34), forked ( f i g . 11) or complex ( f i g s . 9, 10, 33). Sporangia and zoospores are generally larger from t h a l l i on MHU medium than on p o l l e n . Very i n -frequently zoospores escape the sporangial confines 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 morphological features of the ten i s o l a t e s are presented i n Table I. Isolates 1-5 and 9 develop t h a l l i p r i m a r i l y by germ tube swelling. T h a l l i of i s o l a t e 7 also develop generally by germ tube swelling but occasionally both cyst and germ tube enlarge simultaneously. F i f t y percent of the t h a l l i of i s o l a t e s 6 and 10 develop from the cyst. 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 cyst swelling. According to their mean diameter, p o l l e n grown sporangia f a l l into three s i z e classes: 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. On MHU medium three s i z e classes can be recognized: 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. Sporangia of i s o l a t e s 1,2, 5, 6 and 8-10 are usually s p h e r i c a l but sporangia of i s o l a t e s 3-7 are s p h e r i c a l to Irregular. On po l l e n , sporangia of i s o l a t e s 1-3 have one to f i v e (rarely several) r h i z o i d a l axes. Sporangia of i s o l a t e s 4, 5, 8 r - and 9 have one to several r h i z o i d a l axes. 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 sporangia of i s o l a t e s 6, 7 and 10. Discharge tubes are always missing from sporangia 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. Entophlyctis sp. F i g . 13. Cyst germinating by a narrow tube, xl325. F i g . 14. Cyst germinating by a broad bleb, xl325. F i g . 15. Rhizoidal axis developing on i n c i p i e n t sporangium, xl325. F i g . 16\ R h i z o i d a l axis terminal on germ tube, xl325. F i g . 17. Branching of r h i z o i d a l axis, x650. F i g . 18. Thallus with branching r h i z o i d a l axis and empty zoospore cyst, xl325. F i g . 19. Further r h i z o i d a l axis branching, x650. F i g . 20. Incipient sporangium with three r h i z o i d a l axes, x650. F i g . 21. Thallus with i r r e g u l a r development of sporangium, r e f i l l e d zoospore cyst and several r h i z o i d a l axes, x200. F i g . 22. I r r e g u l a r l y developing sporangium with several dichotomously branching r h i z o i d s , x200. F i g . 23. Cyst remaining on nearly mature sporangium, x200. 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 or may not have discharge tubes. The zoospore cyst of a l l i s o l a t e s can be present or absent from sporangia on pol l e n or MHU medium. I t i s generally not the zoospore escape route on p o l l e n developed zoosporangia of most i s o l a t e s , i s o l a t e 6 excepted. Zoospores of i s o l a t e s 4, 6 and 10 on MHU medium escape the sporangium p r i m a r i l y through the cyst. The cyst i s generally functionless ,in sporangium discharge 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 of most i s o l a t e s have a si n g l e , hyaline, 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 guttule of i s o l a t e 2 spores i s minute and zoospore globules -of i s o l a t e 8 are pale orange. Zoospore f l a g e l l a e of a l l i s o l a t e s on po l l e n or MHU medium are 22-25y long. Three mean spore diameters from sporangia on MHU medium can be recognized: 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 po l l e n , mean spore diameters are 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 Ty p i c a l development and morphology of i s o l a t e s 1-10 r e l a t e them to Entophlyctis. On MHU medium and i n po l l e n , eucarpic t h a l l i develop by germ tube swelling and sporangia are inoperculate. Rhizoids a r i s e d i r e c t l y from the sporangium. Entophlyctis, established by Fischer, 1892, includes ten species recognized by Sparrow (1960) and species subsequently described by Willoughby and Townley (1961a), E_. lobata; Canter (1965), E_. molesta; Cook (1966), _E. re t i c u l o s p o r a and Karling (1967d), E_. crenata. Of Plate I V — F i g u r e s 24-29. . Entophlyctis sp. F i g . 24. Inci p i e n t sporangium developing from the r h i z o i d a l a x i s , xl325. F i g . 25. Nearly mature sporangium with cyst remaining, xl325. F i g . 26. Mature sporangium with cyst remaining (same^fehallus as f i g . 25), xl325. F i g . 27. P a p i l l a r sporangium cyst, x650. F i g . 28. Sporangium with hyaline papillum, xl325. F i g . 29. Short discharge tube on sporangium, x650. 2 4 25 these species, J 3 . texana K a r l i n g , E_. confervae-glomeratae (Cienkowski) Sparrow, E,. aurea Haskins and E_. conf ervae-glomeratae (Cienkowski) Sparrow f . marina Kobayashi and Ookubo are considered doubtful taxa. Sporangium s i z e and shape, p o s i t i o n and number of r h i z o i d s and the i d e n t i t y of the host are characters stressed i n d i s t i n g u i s h i n g specifes (Karling, 1931; Sparrow, 1960). Other characters used include: behavior of zoospores at discharge; zoospore s i z e ; c o l o r , number and s i z e of spore guttules; flagellum length; presence of the cyst at sporangium maturity; the cyst's r o l e i n sporangium discharge; protoplasm co l o r ; r e s t i n g spore presence and the r e s t i n g spore ornamentation. Character 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 several d i f f e r e n t established species of Entophlyctis. Isolates 4, 5, 8 and 9 are s i m i l a r to E_. b u l l i g e r a (Zopf) Fischer, E_. conf ervae-glomeratae and E_. conf ervae- glomeratae f . marina; i s o l a t e s 3, 6 and 10 are s i m i l a r to E_. r h i z a (Schenk) Minden, E_. pygmaea (Servinow) Sparrow and E_. crenata and i s o l a t e s 1, 2 and 7 are s i m i l a r to E. helioformis (Dang.) Ramsbottom, E_. molesta, E_. r e t i c u l o s p o r a and E_. texana i n sporangial diameters. Sporangia of i s o l a t e s 2, 5, 6, 8-10, E_. apiculata (Braun) Fischer, E_. b u l l i g e r a , E_. vaucheriae (Fisch) Fischer, E_. rhizna, _E. r e t i c u l o s p o r a , E_. moles ta and E_. helioformis are s p h e r i c a l or obpyriform while sporangia of i s o l a t e s 3, 4, 7, E_. texana, E_. pygmaea j E. conf ervae-glomeratae, E_. aurea, E_. conf ervae-glomeratae f. marina, E_. lobata and E_. crenata are sp h e r i c a l to i r r e g u l a r . . Isolates 1-3, with one to f i v e r h i z o i d a l axes per sporangium, are c l o s e l y r e l a t e d to E_. ap i c u l a t a , E_. molesta, E_. b u l l i g e r a , E_. vaucheriae, E_. helioformis, 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_. crenata, E_. lobata, E_. r e t i c u l o s p o r a a n d i s o l a t e s P l a t e V — F i g u r e s 30-34. Entophlyctis sp. F i g . 30. Cysts of endobiotic t h a l l i functioning as discharge tubes, x650. F i g . 31. St r a p - l i k e i r r e g u l a r discharge tube of an endobiotic t h a l l u s , x650. F i g . 32. E p i b i o t i c t h a l l u s with discharge through s p l i t t i n g sporangium wal l , x650. F i g . 33. Irregular, branched discharge tube on e p i b i o t i c sporangium, x650. F i g . 34. I n t e r b i o t i c t h a l l u s with long sporangium discharge tubes, x200. 27 28 4, 5, 8 and 9 have one to several r h i z o i d a l axes per sporangium. A l l i s o l a t e s are s i m i l a r to E_. vaucheriae, E_. b u l l i g e r a , E_. lobata, E_. re t i c u l o s p o r a and E_. crenata with rhizoids a r i s i n g anywhere on the sporangium. Rhizoids of i s o l a t e s 1, 3-9 and E. pygmaea are usually strongly dichotomously branched. Isolates 2, 10, E_. ap i c u l a t a , E_. vaucheriae, E. helioformis and E_. confervae-glomeratae have d e l i c a t e r h i z o i d s . Discharge tubes are missing from sporangia 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. lobata, E_. molesta and E_. pygmaea while the remainder of the i s o l a t e s and other Entophlyctis species generally have one or more discharge tubes on each sporangium. Discharge tubes of E. aurea and most of my i s o l a t e s are tipped by large subspherical gelatinous plugs. The zoospore cyst may or may not remain on sporangia of i s o l a t e s 1-10. When present the cyst functions i n sporangium d i s -charge of i s o l a t e s 4, 6 and 10 on MHU medium and i s o l a t e 8 on p o l l e n . Sporangium discharge i n the remaining i s o l a t e s generally occurs 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 are pe r s i s t e n t and function i n sporangium discharge while the cyst i s evanescent and inoperative i n most other Entophlyctis species. Entophlyctis lobata cysts may or may not remain but even when present the cyst i s inoperative during zoosporangial discharge. A l l i s o l a t e s f a l l within the zoospore size range of E_. h e l i o f ormis , E_. texana, E. conf ervae-glomeratae, E_. aurea, E_. conf ervae-glomeratae f. marina, E_. moles ta, E_. r e t i c u l o s p o r a , E_. lobata and E_. crenata. Zoospore f l a g e l l a e of i s o l a t e s 1-10 are longer than those of _E. texana, E. crenata, E_. molesta, E. lobata and E. reticulospora but shorter than the flagellum of E_. h e l i o f ormis spores. Spores of E_. aurea and E_. conf ervae-glomeratae have several small r e f r a c t i v e globules and the guttules of E. texana and E. lobata spores 29 are orange. Other than spores of i s o l a t e 2 with one minute c o l o r l e s s globule and i s o l a t e 8 with a large pale orange globule, most of my i s o l a t e s have spores with a s i n g l e large hyaline 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 i s o l a t e s are r e l a t e d to E_. • b u l l i g e r a , E. conf ervae-glomeratae and E_. crenata. V a r i a t i o n of my Entophlyctis and Karling's (1931) giant t h a l l i of Entophlyctis i n d i c a t e that such characters as sporangium s i z e and shape, number and place 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 branching, discharge tube presence, cyst presence and function, spore s i z e , and spore guttule diameter and color are of l i t t l e taxonomic value i n the genus. Resting spore features and substrate differences are the remaining characters d i s t i n g u i s h i n g Entophlyctis species. Use of sub-s t r a t e differences and parasitism vs. saprophytism to d i s t i n g u i s h c h y t r i d species i s questionable (Emerson, 1950; M i l l e r , 1968). Thus, perhaps E_. a p i c u l a t a , E. r e t i c u l o s p o r a , and E_. crenata are the only v a l i d and useful taxa. Characters of i s o l a t e s 1-10 are also v a r i a b l e above the species l e v e l . Thallus-substrate p o s i t i o n v a r i a t i o n s ( i . e . , endobiotic vs. e p i b i o t i c vs. i n t e r b i o t i c ) may r e l a t e my Entophlyctis to Rhizophydium, Phl y c t i d i o i d e a e ; Olpidiaceae,(Olpidium) and Rhizidiaceae, (Rhizophlyctis and RhizidiumX 'C Thallus development v a r i a t i o n s r e l a t e my Entophlyctis to P h l y c t i d i o i d e a e (Rhizophydium) and Scherffeliomyces and Rhizidiaceae. If discharge by sporangium wall s p l i t t i n g i s interpreted as opercular, my Entophlyctis may r e l a t e to Phlyctidiaceae and Rhizidiaceae, Inoperculatae and Chytridiaceae, Operculatae. Considering such v a r i a t i o n , as Whiffen (1944) stressed, the operculum warrants le s s importance than Sparrow (1942) assigned i t . 30 Also, as Johnson (1969) h i n t s , the operculum character needs more ca r e f u l d e f i n i t i o n . F a m i l i a l and c e r t a i n generic concepts and delineations require c a r e f u l re-evaluation. Emerson (1950) has expressed the urgent need to study environmental and g e n e t i c a l species v a r i a b i l i t y . This i s achieved by studying many i s o l a t e s under several d i f f e r e n t conditions. Study of a s i n g l e i s o l a t e under several d i f f e r e n t conditions f a i l s to account adequately f o r species v a r i a t i o n . F i n a l l y , the species concept as applied to the organisms should r e l a t e to their appearance and a c t i v i t i e s i n nature. Table I. Developmental and morphological d e t a i l s of i s o l a t e s 1-10. Isol a t e and C o l l e c t i o n Numbers* Substrate Development type 1(1) MHU medium p r i m a r i l y from germ tube 1(1) p o l l e n as above 2(30) MHU medium as above 2(30) p o l l e n as above 3(43) MHU medium as above 3(43) p o l l e n as above 4(46) MHU medium as above 4(46) p o l l e n as above 5(47) p o l l e n as above 6(52) MHU medium from cyst or tube equally 6(52) po l l e n as above 7(54) MHU medium as above 7(54) p o l l e n p r i m a r i l y from germ tube but cyst may also expand 8(61) MHU medium p r i m a r i l y from cyst 8(61) p o l l e n p r i m a r i l y from tube 9(66) p o l l e n p r i m a r i l y from germ tube 10(74) MHU medium as above 10(74) p o l l e n from tube or cyst equally Sporangium Size (H x W) Shape P a p i l l a e 27- (38)-50y s p h e r i c a l + .-16- (31)-65y as above + ,-36- (65)-110u as above + ,-22.J j-(33.5)-48y as above + ,-38- [50)-70y as above + ,-11- (22)-35y (sph) sp h e r i c a l - v a r i e d 27- (32)-42X 14- (19)-28y (asph) 17- (28)-38y Spherical 12- (16)-20y (sph) sp h e r i c a l - v a r i e d 14- (17.5)-27X 6- (15)-28y (asph) 12- (18)-30y s p h e r i c a l 17- (20.5)-25y as above + ,-16- [20.0)-28y as above + .-50- (65)-80y as above + ,-15- C29)-75y (sph) sp h e r i c a l - v a r i e d + 20- (49)-70X 17- [38)-55y (asph) 75- U00)-120y s p h e r i c a l + ,-9- C18.5)-37y as above + ,-10- U7)-22y as above + .-65- C85)-110y as above + »-14- [22)-30y as above + .-'"Collection no. bracketed. Table I (continued) Is o l a t e No. Presence 1 MHU + ,-1 P o l l e n + ,-2 MHU +,-2 P o l l e n +,-3 MHU + ,-3 P o l l e n +,-4 MHU 4 P o l l e n + ,-5 P o l l e n +,-6 MHU 6 P o l l e n 7 MHU + ,-7 P o l l e n +,-8 MHU + ,-8 P o l l e n + 9 P o l l e n + ,-10 MHU 10 P o l l e n +,-Discharge Tubes No./Sporangium Size (L x W at base) 1-2 2-(5.5)-14 x 3-(5)-6y 1-2 3.5-(15)-28 x 2-(4.5)-7y 1-2 2-(5)-12 x 2-(3)-4p 1(3) 2-(6)-15 x 3-(4)-5y 1-3 4-(13)-18 x 3.5-(4.5)-6y 1-(plexus of several) 4-(18)-24 x 3-( 4 ) - 5 ( l l ) u 1 3-(5)-7 x (3)4y 1-(plexus of several) 7-(14)-9 x 3-(5)-8u 1-3 13-(21.5)-43 x 5-(7.5)-9y 1-(plexus of several) 3-(16.5)-40 x 3-(4.5)-5y 1-2 7-(13)-29 x 7-(8)-9(13)y 1 3-(6)-12 x 2.5-(4)-5y 1-2 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 No. Nature No. axes 1 MHU coarse, blunt tipped several 1 P o l l e n coarse, tapering l-(3)-5(9) 2 MHU coarse, blunt tipped several 2 Po l l e n f i n e , bushy l-(3)-8 3 MHU coarse, blunt tipped several 3 Pollen coarse, tapering 1-5 4 MHU coarse, blunt tipped several 4 Po l l e n coarse, tapering 1-several 5 P o l l e n as above as above 6 MHU coarse, blunt tipped several 6 P o l l e n coarse, tapering several 7 MHU coarse, blunt tipped several 7 P o l l e n coarse, tapering s e v e r a l 8 MHU coarse, blunt tipped s e v e r a l 8 Pollen very coarse, grow from 1-5 p o l l e n into environment (several) 9 P o l l e n coarse, tapering 1-5 10 MHU f i n e , blunt tipped several 10 Pollen coarse, tapering several Color i n culture Relation to substrate creamy white creamy white creamy white creamy white creamy white creamy white yellow brown creamy white e p i b i o t i c endo-, e p i - , and i n t e r b i o t i c e p i b i o t i c endo-, e p i - , and i n t e r b i o t i c e p i b i o t i c endo-, e p i - , and i n t e r b i o t i c e p i b i o t i c 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 e p i b i o t i c e p i b i o t i c e p i b i o t i c endo-, e p i - , and i n t e r b i o t i c e p i b i o t i c endobiotic p r i m a r i l y endo-, e p i - , and i n t e r b i o t i c e p i b i o t i c endo-, e p i - , and i n t e r b i o t i c Table I continued Zoospores Iso l a t e No. Size No. r e f . globules Globule s i z e Globule c o l o r 1 MHU 4.0- ( 4.5] -5.0y 1(2) 1.5p hyaline 1 P o l l e n 3.0- (4.0) -5. Op 1 l.Ou as above 2 MHU 3.0- (5.0) -7.0y 1-2 1.5u as above 2 P o l l e n 3.0- (4.0) -5. Oy 1-2 minute as above 3 MHU 3.5- (5.0) - 6 .Op 1(2) 1.5p as above 3 P o l l e n 3.0- (4.0) -5.0y 1(2) l.Op as above 4 MHU 3.0- (5.0) - 6 . Op 1(2) 1.5u as above 4 P o l l e n 3 . 0 - (4.0) - 4.5p 1 l.Op as above 5 P o l l e n 3.0- (4.0) -5.Op 1 l.Op as above 6 MHU 4 . 0 - ( 5 . 0 : -7.0y 1 l .Op as above 6 P o l l e n 2.5- (3.0) - 3 . 5 p 1 l .Op as above 7 MHU 3.5- (4.5) - 6 . Op 1 1.5p as above 7 P o l l e n 3.0- ( 3 . 5 ; - 4 . Op 1 l.Op as above 8 MHU 4 . 0 - (4.5) -5.5y 1 1.5p orange 8 P o l l e n 3.5- ( 4 . 0 ; -5.Op 1 l .Op orange 9 P o l l e n 4 . 0 - (4.5) -5. Op 1 1.5p hyaline 10 MHU 3.0- ( 3 . 5 ; ) - 4.0p 1 l.Op as above 10 P o l l e n 3.5- ( 4 . 5 ; -5. Op 1 1.5p as above I s o l a t e No. Flagellum length 1 MHU 1 P o l l e n 2 MHU 2 P o l l e n 3 MHU 3 P o l l e n 4 MHU 4 P o l l e n 5 P o l l e n 6 MHU 6 P o l l e n 7 MHU 7 P o l l e n 8 MHU 8 P o l l e n 9 P o l l e n 10 MHU 10 P o l l e n 22-25y as above as above as above as above as above as above as above as above as above as above as above as above as above as above as above as above as above Remarks Discharge p r i m a r i l y through c y s t , 1.5-(3.5)-8 x 4-(5.5)-7u Discharge p r i m a r i l y through 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 36 3. OCCURRENCE AND DISTRIBUTION OF ZOOSPORIC FUNGI FROM DEVON ISLAND, CANADIAN EASTERN ARCTIC I n t r o d u c t i o n Methods and m a t e r i a l s Taxonomy and morphology INTRODUCTION Devon I s l a n d , approximately centered i n the Canadian A r c t i c Archipelago along a north-south 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. long ( f i g . 40). Most of i t i s l o c a t e d i n the high a r c t i c b e l t (Nordenskjold, 1928; P o l u n i n , 1951). Major physiographic 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 abruptly r i s i n g escarpment and plateau. Information on occurrence, d i s t r i b u t i o n and ecology of zoosporic f u n g i i n the A r c t i c i s scant, p a r t i c u l a r l y i f the A r c t i c i s defined i n i' the r e s t r i c t e d sense (Nordenskjold, 1928; P o l u n i n , 1951). E a r l y r e p o r t s 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 Peronospora represented the bulk 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 to our understanding of aquatic f u n g i from a r c t i c regions. He described 12 s p e c i e s , four new 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 preserved by N. P o l u n i n during the Canadian Eastern A r c t i c Expeditions of 1934 and 1936. S a v i l e (1961, 1968) and S a v i l e and Parmelee (1964) made extensive b o t a n i c a l and mycological c o l l e c t i o n s at 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 . Their studies were r e s t r i c t e d to fungi p a r a s i t i c on phanerogams. As i n the case of e a r l i e r studies, reports of zoosporic fungi are r e s t r i c t e d to Synchytrium, Physoderma and Peronospora. A few German workers studied zoosporic fungi from a r c t i c s o i l s , but " A r c t i c " i s too broadly interpreted i n th e i r reports. Most s o i l samples labeled a r c t i c ( i . e . , from Lappland and Iceland) were probably subarctic s o i l s i n the sense of Tedrow (1970). Also high elevation s i t e s i n Lappland would best be referred to as alpine. Harder (1954) examined s i x s o i l c o l l e c t i o n s from Spitsbergen. He recovered Olpidium pendulum Zopf i n a sample from a Dupontia moor near Longyearbyen and Nowakowskiella  elegans (Nowak.) Schroeter i n a sample from a Poa meadow. Only 33% of the s i t e s yielded fungi. Gaertner (1954) also surveyed s i x s o i l c o l l e c t i o n s from Spitsbergen and found the same organisms as Harder. In a d d i t i o n he found several Pythium species. Hohnk (1960) c o l l e c t e d 10 samples across a beach along King Frederik VI Coast, Greenland at approximately 63°N. l a t . - Five of the samples came from the emergent por t i o n of the beach and four from the submerged portion. Every c o l l e c t i o n y ielded at l e a s t one species. Among the species found are: Saprolegnia sp., Thraustochytrium proliferum Sparrow, Sapromyces elongatus (Cornu) Coker, Apodachlya brachynema (Hildebrand) Pringsheim, Pythium monospermum Pringsheim, I?, graminicolum var. stagni Hb'hnk, I?, irregulare.'l Bulsman, Pythium spp., Pythiogeton ramosum Minden, Olpidium luxurlans (Tomaschek) Fischer, Rhizophydium p o l l l n l s - p i n i (Braun) Zopf, R. sphaerotheca Zopf, R. mamillatum (Braun) Fischer, Phlyctochytrium chaetiferum Karling and Rhizophlyctls rosea (deBary and Woronin) Fischer. Other workers have looked sparingly f or zoosporic fungi i n a r c t i c s o i l s . K a r l i n g (1951) i s o l a t e d Catenophlyctis (Phlyctorhlza) v a r i a b i l i s 38 (Karling) K a r l i n g from tundra s o i l of St. Lawrence Island (63°30'N.^ l a t . , 171°W. long.). Kobayasi et a l . (1967) surveyed s o i l s of A r c t i c Alaska for a l l fungi. Among their findings are Achlya conspicua Coker, Aplanes turfosus/(Minden) Coker, Aphanomyces sp. Apodachlya brachynema Hildebrand and Pythium carolinianum Matthews. Among other i s o l a t e s , Barr (1970b) studied morphology and physiology of an i s o l a t e of Hypho chy tr jum ca tenoid es from a Canadian high A r c t i c s o i l . Morpholo-g i c a l l y the a r c t i c i s o l a t e was completely s i m i l a r to other i s o l a t e s from geographic regions to the south. However, the a r c t i c i s o l a t e grew some at 10C, , optimally at 20, 25 and 30C, and not at a l l at 35C. Conversely, an Arizona i s o l a t e grew some at 35C;-; optimally at 30 and 25C and not at a l l at 10C. Growth at d i f f erenfr :pH l e v e l s showed a s i m i l a r c u l t u r e -environment coupling. Barr suggests these differences may r e f l e c t genetic s e l e c t i o n i n l o c a l conditions. Allomyces, o r i g i n a l l y thought to be p r i m a r i l y t r o p i c i n d i s t r i b u t i o n , was recently recovered from a Devon Island s i t e by G.C. Hughes (pers. comm.). METHODS AND MATERIALS During the 1967-1969 summers Paul Barrett conducted researches on phytogeocoenoses i n a 16 square m i l e ; a r e a on the base camp lowland ( f i g . 40). Several s i t e s i n teniphytogeocoenoses were considered. Several general habitats were recognized during these studies and these are discussed elsewhere (Appendix I ) . Since edaphic factors comprised a considerable portion of these studies, s o i l s were returned to la b o r a t o r i e s at the U n i v e r s i t y of B r i t i s h Columbia for chemical and p h y s i c a l analyses. Immediately upon a r r i v a l at the U n i v e r s i t y of B r i t i s h Columbia, 39 the s o i l samples were removed from large p l a s t i c bags and d r i e d . As soon as t h i s procedure was started I took subsamples of most c o l l e c t i o n s and treated them for recovery of aquatic fungi. Techniques used were i d e n t i c a l with those reported i n an e a r l i e r study (Booth, 1969). Baits included Pinus s y l v e s t r i s L. po l l e n , d i a l y z i n g membrane cellophane and snakeskin. Each subsample was started at l e a s t twice by flooding with Capilano River water at 0 ° / o o s a l i n i t y . . Baits were examined p e r i o d i c a l l y for 6 weeks. Plates remained at room temperature 20 ± 5C during this time. Baits with fungi growing on them were either streaked ( i . e . pollen) or placed ( i . e . cellophane and snakeskin) on modified (Harder and Uebelmesser, 1955) medium, MHU (0.3% malt extract, 0.3% peptone, 0.1% dextrose, 1.1% agar i n 1 1 water of 0°/oo s a l i n i t y with .03% Streptomycin sulphate—774 mcg/mg and .03% P e n i c i l l i n G—4 x 10 u n i t s ) . Each p l a t e surface was then surveyed f o r s i n g l e encysted spores or i n c i p i e n t sporangia from 1 to 3 days afterward. Usually by the t h i r d day filamentous higher fungi covered the slower growing Phycomycetes. As s i n g l e spores were seen they were removed and placed on MHU medium minus a n t i b i o t i c s . Fourteen s i n g l e spore i s o l a t e s were obtained from d i f f e r e n t locations on Devon Island. A l l subsequent manipulations were performed without a n t i b i o t i c s . Stored i s o l a t e s were kept at 6C and transferred every 8 weeks. Each i s o l a t e was grown i n 50 mis s t e r i l e l i q u i d MHU medium for two weeks at 20 + IC on a table top shaker with a 3" stroke at 60 cycles/min. Contents of each f l a s k were then homogenized for a minute i n a s t e r i l e Waring Blendor semi-microcup and poured into a 125 ml f l a s k containing a magnetic s t i r r i n g bar. The fungal material suspension was maintained 40 on magnetic s t i r r e r during the following treatments. F i r s t , 1.0 ml was removed by a s t e r i l e p i p e t t e and added to 25 ml gas s t e r i l i z e d Pinus s y I v e s t r i s p o l l e n i n a p e t r i p l a t e . Also 5 ml were placed on the surface of s o l i d MHU medium i n the bottom of a 250 ml beaker. Aft e r 3 days, fungi developing on the p o l l e n could be studied. Inoculated beakers were kept at 20 ± IC for a week. Fungi covering the medium surface were flooded with 5 ml of s t e r i l e Emerson (1958) d i l u t e s a l t s o l u t i o n (DSS). A f t e r 6 hrs 0.25 ml supernatant was removed by pipette and dispensed i n s i n g l e drops on s o l i d modified ;jT-U./ medium i n a p e t r i p l a t e . Plates were incubated at 20 IC and fungus development studied c l o s e l y . TAXONOMY' AND MORPHOLOGY C h y t r i d i a l e s , B i a s t o c l a d i a l e s and Hyphochytriales 1. Catenophlyctis v a r i a b i l i s (Karling) K a r l i n g , Amer. Jour. Bot.; 52:134, F i g s . 1-12, 1965. 2. Phlyctorhiza v a r i a b i l i s K a r l i n g , Amer. Jour. Bot., 34:27, Fi g s . 1-46, -3-947. On snakeskin: an i c e wedge polygon s i t e , 386. Sporangia, highly v a r i a b l e i n shape, from 15.5-51.5y' x 20.5-64.Oy. Two or more r h i z o i d a l axes per sporangium ( f i g . 11). The sporangial protoplast hyaline. Spores, 3-4 x'4^ -;5y, discharged through short tubes, t h a l l i generally monocentric. Taxonomic deposition of t h i s species has been problematic. 41 K a r l i n g (1946b) o r i g i n a l l y assigned the nomen nudum P e r i r h i z a  endogena. He subsequently (1947a) described i t formally as Phlyctorhiza v a r i a b i l i s . Generic assignment was tentative since sporangia are not formed from r h i z o i d a l v e s i c u l a t i o n but by s w e l l -ing of the germ tube. A f t e r f i n d i n g several p o l y c e n t r i c s t r a i n s and observing side bodies and conspicuous nuclear caps i n the zoospores, Ka r l i n g (1951) considered P_. v a r i a b i l i s somewhat re l a t e d to Catenaria. Sparrow (1957) states that Karling's (1951) material i s undoubtedly a species of Catenaria. He alsoCsuggests (1960) that P_. v a r i a b i l i s would be best accommodated i n Entophlyctis. Karling (1965) established Catenophlyctis i n the Catenariaceae. Phlyctorhiza  v a r i a b i l i s became a synonym of the new combination Catenophlyctis  v a r i a b i l i s . . ^ However, Nolan (1970) demonstrated n u t r i t i o n a l heterogeneity by the i n c l u s i o n of C_. v a r i a b i l i s i n the Catenariaceae and B l a s t o c l a d i a l e s and suggests possible recognition of a t r a n s i t i o n a l family between C h y t r i d i a l e s and B l a s t o c l a d i a l e s . K a r l i n g (1951) reported this species from the a r c t i c . 3. Chytriomyces annulatus Dogma, Nova Hedwigia, 18:349, F i g s . 1-18, 1969. On p o l l e n : i c e wedge polygon s i t e s , 383, 395 and a r a i s e d beach slope s i t e , 410. Zoosporangia sph e r i c a l ( f i g s . 1, 20) or pyriform ( f i g s . 2, 21). On pine p o l l e n mature s p h e r i c a l sporangia 11.5-(18.0)-28.5p, pyriform sporangia 13-(30)-46y high and 9-(19)-26p wide. On agar nature sporangia mostly s p h e r i c a l or s l i g h t l y angular and 18.5-(37.0)-50.Op i n diameter. R i n g - l i k e undulations usually on sporangium w a l l near 42 o r i g i n of r h i z o i d a l system ( f i g s . 2, 20, 21). Occasionally rings w e l l up on the wall ( f i g . 1). Rhizoids d i r e c t l y from the sporangium ( f i g . 21) or from an apophysis. Refractive globules commonly f i l l apophysis; On p o l l e n , sporangia often s t i p i t a t e ( f i g s . 1, 2, 20) and an apophysis may or may riot appear i n s i d e the p o l l e n grain, where s t i p e penetrates through the exine and i n t i n e . Rhizoids are f i n e and greatly branched ( f i g . 21). At discharge, upper portion of the zoosporangial w a l l cracks and the operculum pushes back with escape of spore mass. Opercula at two or even three places on the wall of a s i n g l e sporangium. Spores quiescent for a time, rock slowly and f i n a l l y swim away. Zoospores 3.5-(4.5)-6.5u on p o l l e n and 3.5~(5.0)-7.5u on agar. A lustrous globule f i l l s one-third of the spore body. The posterior flagellum V30u long. A f t e r a short swarming period spore enDysts. On agar a l l but a small portion of the cyst w a l l , where the r h i z o i d a l system a r i s e s , swells at germination. Swelling stops for a time and then resumes with the exception of a small area adjacent to the previously dominant r i n g of wall m a t e r i a l . The cycle of enlarging, stopping, and enlarging again may occur several times and r e s u l t s i n the undulations or rings on the sporangial wall described above. ! ? Sporangial dimensions of a r c t i c material on p o l l e n and MHU medium are nearly i d e n t i c a l with Dogma's figures (1969b). Presence of sporangium wall undulations, v a r i a b i l i t y of apophysis occurrence, r h i d o i d a l habit, zoospore behavior at discharge and f l a g e l l a r length are further s i m i l a r i t i e s of material i n the two studies. Zoospores from t h a l l i grown on modified fE-U medium are s l i g h t l y 43 l a r g e r than any described p r e v i o u s l y . Contrary to Dogma's observations, undulations heed not be r e s t r i c t e d to the lower h a l f of the sporangium. F i n a l l y , sporangia of a r c t i c C_. annulatus discharge from 1-3 opercula per sporangium u n l i k e the s i n g l e operculum type. This tends to r e l a t e my m a t e r i a l to Chytriomyces  poculatus Willoughby and Townley, but s p o r a n g i a l w a l l undulations were never seen to develop i n t o the conspicuous cupules of t h i s s p e c i es. This i s only the second report of C_. annulatus and the f i r s t from A r c t i c regions. 4. Chytriomyces poculatus Willoughby and Townley, Trans. B r i t . Mycol. S o c , 44:183, F i g . 3, 1961. On p o l l e n : i c e wedge polygon s i t e s , 383, 395. Sporangia on pine p o l l e n c l a v a t e ( f i g s . 22, 23) or e l l i p s o i d ( f i g . 3 ) , 16.5-(30.0)-41.0p high and 10.5-(20.0)-26.0p wide at mat u r i t y . Sporangium w a l l s wavy ( f i g s . 3, 22, 23) and p e r s i s t e n t a f t e r discharge ( f i g . 23). Rhizoids a r i s i n g d i r e c t l y from the sporangium and s p a r i n g l y branched ( f i g . 3). At discharge operculum, 2.5-3.5p high and 7.5-12.5p wide, s w e l l i n g out ( f i g . 22) and separating from sporangium w a l l along one s i d e ( f i g . 3). Zoospores passing from s p o r a n g i a l confines as a mass and remaining quiescent f o r ^2-4 minutes, rocking s l o w l y and appearing roughened i n o u t l i n e as the mass breaks up. M o t i l e spores 4-(5)-6p i n diameter w i t h one r e f r a c t i v e g l o b u l e , ^1.5p wide, surrounded by s e v e r a l s m a l l bodies i n p r o t o p l a s t ( f i g s . 3, 4 ) ; f l a g e l l a e ^23p long. Spores o f t e n f a i l i n g to escape sporangium, germinating and developing i n s i t u . 44 Zoospores, larger than those described by Willoughby and Townley (1961a) and Dogma (1969b) ,','are about the same s i z e as Sparrow (1968a) reported. Absence of overlapping cupules i s a feature shared by Willoughby's (1965), Sparrow's (1968a) and my ma t e r i a l . Multioperculate sporangia, such as Willoughby (1965) observed, were not seen. Dogma'; (1969b) suggests that cupule free sporangia without basal undulations may belong to a taxon d i f f e r e n t from Chytriomyces  poculatus. He adds further credence to t h i s point by pointing out zoospore s i z e d i f f e r e n c e s between cupuled and cupule free forms. My material would seem to support this viewpoint but, as i n Dogma's case, I consider C_. poculataus a complex of sporangial forms and highly v a r i a b l e . This i s the f i r s t report of C. poculatus from the A r c t i c . 5. Hyphochytrium catenoides K a r l i n g , Amer. Jour. Bot., 26:513, F i g s . 1-18, 1939. On snakeskin: i c e wedge polygon s i t e s , 383, 384, 395; a Carex-moss sub-association, 423 and an A r c t a g r o s t i s l a t i f o l i a meadow, 438. Sporangia s p h e r i c a l ( f i g s . 6, 24, 25, 27), 12.5-35.Op i n diameter, ovoid ( f i g s . 5, 24), 15.0-17.5y x 18.5-26.Oy or i r r e g u l a r l y elongate ( f i g . 26), 16-62y x l l - 2 1 y . T h a l l i usually p o l y c e n t r i c but occasionally monocentric forms ( f i g . 5), resembling Karling's (1967g) figures of Hyphochytrium oceanum Ka r l i n g , occurring. Septae d e l i m i t i n g sporangia along the mycelium but occasionally appearing between sporangia ( f i g s . 17, 24, 27). 45 Irregular swellings ( f i g s . 7-9) developing along the course of the tubular mycelium, l-3y i n diameter. Swellings functioning as sporangia ( f i g . 26). Sporangial protoplast extrusion accomplished through a sporangial wall opening or a tube ( f i g . 17) up to 85y long. Spores d i f f e r e n t i a t i n g and swimming from naked extruded protoplasm, a n t e r i o r l y u n i f l a g e l l a t e , f i l l e d with several small r e f r a c t i v e globules and 2-4y long and 2-3y wide. Barr (1970b) reports Hyphochytrium catenoides from s o i l s of A n t a r c t i c a and the Canadian High A r c t i c . 6. Nowakowskiella elegans (Nowak.) Schroeter, Engler and P r o n t l . , Naturlichen' Pflanzehfam.; 1:82, F i g . 35, 4-G, 1892. On cellophane and with p o l l e n : Carex hummock sub-associations, 396, 442; g r a n i t i c boulder s i t e s , 397, 405; a r a i s e d beach ridge s i t e , 420; a Saxifraga-Rhacomitrium slope s i t e , 419; and a snowbed-moss s i t e , 435. Sporangia ovoid ( f i g . 28), elongate or i r r e g u l a r ( f i g . 29), with greatest dimension ranging from 18-35y. Spindle shaped ( f i g . 30) or irregular ( f i g . 31) swellings.occurring at various points along mycelium of the p o l y c e n t r i c t h a l l u s . Spores, 5-8y i n diameter, passing from sporangium through an operculum ( f i g . 30) as a quiescent mass, soon swimming away. Occasionally a portion of the spore mass f a i l s to escape sporangial confines ( f i g . 32). Besides t h i s report of N. elegans from the A r c t i c the only other reports are Harder (1954) and Gaertner (1954). P l a t e V I — F i g u r e s 1, 2. Chytrioniyces annulatus. F i g . 1. • S t i p i t a t e s p h e r i c a l zoosporangium with widely spaced undulations, x925. F i g . 2. Basal undulations or rings on a pyriform sporangium, x925. Figures 3, 4. C_. poculatus. F i g . 3. • Cupule f r e e zoosporangium and reduced r h i z o i d a l system.- x925. F i g . 4. Granule f i l l e d zoospore, fh . (freehand),^ xlOOO. Figures 5-9. Hyphochytrium catenoides. F i g . 5. Monocentric t h a l l u s , sporangia discharge tube short, x600. F i g . 6. Sporangia with p a p i l l o i d discharge pore, x600. F i g . 7. Twice con-s t r i c t e d sporangium, x600. F i g . 8. Two c l o s e l y associated sporangia, x600. F i g . 9. Fusiform sporangium, x600. Figure 10. Nowakowskiella  macrosporum; sporangium f i l l e d with large uniguttulate spores, x925. Figure 11. Catenophlyctls v a r i a b i l i s ; three r h i z o i d a l axes on the large Irregular t h a l l u s , x925. Figure 12. Rhlzophydlum ,sp. ( u t r i c u l a r e ? ) ; large p a p i l l a e on the angular or s a c - l i k e zoosporangium, x925. Figures 13, 14. R. p o l l l n i s - p i n i . F i g . 13. Rhi z o i d a l system branched, d i s -charge papillum a p i c a l and broad; x925. F i g . 14. Empty zoosporangium and i n c i p i e n t sporangium within, x925. Figures 15, 16. Rhizophydium sp. (patellarium?). F i g . 15. Thallus with tufted r h i z o i d s and thickened basal w a l l , x925. F i g . 16. P e r s i s t e n t basal portion of sporangium w a l l a f t e r discharge, x925. F i g . 17. Hyphochytrium  catenoides;,septate intersporangial hyphae and long discharge tube on sporangium, x600. Figures 18, 19. Rhizophydium coronum. F i g . 18. Laminated halo around sporangium, x925. F i g . 19. Empty zoosporangium with s i n g l e a p i c a l pore, x925. F i g s . 1-4, 10-16, 18, 19, scale A*, "'"Unless designated as f h . , a l l drawings done with camera l u c i d a . F i g s . 5-9, 17, scale B; F i g s . 20-32, 34-39, scale C; F i g . 33, scale D. 48 49 7. ?Nowakowskiella macrospora K a r l i n g , Amer. Jour. Bot., 32:29, F i g . 1-30, 1945. On cellophane and p o l l e n : a Carex hummock s i t e , 443. M a t e r i a l on cellophane and p o l l e n very s i m i l a r to N. macrospora w i t h one exception: as i n m a t e r i a l observed by Sparrow (1960), no opercular s t r u c t u r e formed. Sporangia 18-28p high x 13-22p wide and s l i g h t l y apophysate when ter m i n a l ( f i g . 10). Spores 10-12p i n diameter w i t h a s i n g l e l a r g e r e f r a c t i v e globule ( f i g . 10). 8. ?Nowakowskiella ramosa E. J . B u t l e r , Mem. Dept. Agr. I n d i a , Bot. Ser. , 1:141, P I . 10, F i g s . 3-10, 1907. On cellophane: Carex hummock s i t e s , 396, 441 and a r a i s e d beach ri d g e s i t e , 420. Resting spores on septated s p i n d l e bodies o c c a s i o n a l l y seen containing a s i n g l e l a r g e r e f r a c t i v e body surrounded by s e v e r a l s m a l l e r ones and w i t h low verrucae over the e x t e r i o r spore w a l l . Resting spores on the same t h a l l u s may or may not be on "pseudo-parenchyma." Thus, my m a t e r i a l may be a v a r i a n t of _N. elegans. 9. Olpidium pendulum Zopf, A. Schenk, Handbuch d. Bot., . . . , 4:555, F i g . 66 (1-5), 1890. In p o l l e n : i c e wedge polygon s i t e s , 383, 386, 395; S a x i f r a g a - Rhacomitrium slope s i t e s , 413, 419; a r a i s e d beach r i d g e s i t e , 416; a Carex hummock s u b - a s s o c i a t i o n , 429; and a Carex-moss s u b - a s s o c i a t i o n , 434. Sporangia s p h e r i c a l , endobiotic and 10.0-17.5p i n diameter. 50 Usually no more than three t h a l l i per p o l l e n grain. Spores, 3.5-4.5p i n diameter, escaping sporangium through a short tube. My material d i f f e r s l i t t l e from that previously described. Linder (1947) found two species of Olpidium i n material from the Canadian A r c t i c but neither was _0. pendulum. Harder (1954) and Gaertner (1954) obtained i t from Spitzbergen. 10. Rhizidium verrucosum K a r l i n g , Amer. Jour. Bot., 31:255, F i g s . 34-63, 1944. On snakeskin: a non-sorted c i r c l e s i t e , 409; a Saxifraga- Rhacomitrium slope s i t e , 413; and a r a i s e d beach ridge s i t e , 417. Zoosporangia on snakeskin s p h e r i c a l , 18.0-55.Op, or oblong, 19.5-51.5 x 30.0-56.Op, on MHU medium s p h e r i c a l , 55-130p, oviform, 55-75 x 80-100p, or angular. High c o n i c a l discharge p a p i l l a e breaking through the sporangium w a l l ( f i g . 34), r h i z o i d s generally . a r i s i n g from one place on the w a l l , coarse on snakeskin and tufted on MHU medium. At maturity the zoospore with r e f r a c t i v e globules imparting a yellowish-orange hue to sporangial protoplast, before becoming motile spores i n a v e s i c l e , motile spores, 3.5-5.Op wide, with a s i n g l e r e f r a c t i v e body,'1.0-1.5p i n diameter. This i s the f i r s t report of R. verrucosum from the a r c t i c . 11. Rhizophydium angulosum Ka r l i n g , Arch. M i k r o b i o l . , 61:116, F i g . 1, • M-X, 1968. On p o l l e n : a raised beach ridge s i t e , 390; a Saxifraga- Rhacomitrium slope s i t e , 413 and Carex hummock s i t e s , 425, 428, 429. 51 Zoosporangia angular early i n development and at maturity two to many angled ( f i g . 39), 11.0-(18.5)-31 .0u on p o l l e n and 16.0-(20.5)-31 .0p on MHU medium. Rhizoids branched, occasionally a r i s i n g from an apophysoid structure up to 5p wide. Zoospores 2.5-3.Op i n diameter with a s i n g l e minute r e f r a c t i v e globule, discharged through sporangium wall angles. This material i s s i m i l a r to that described by K a r l i n g (1968c) and Booth (1969). Rhizophydium corohum Hanson, Torreya, 44:31, 1944. O h p o l l e n : a Carex-moss s i t e , : 4 0 7 ; Carex hummock s i t e s , 425, 426; snowbed-moss s i t e s , 435-436 and A r c t a g r o s t i s l a t i f o l i a s i t e s , 438, 439. Sporangia s p h e r i c a l ( f i g s . 18, 38), 11.0-23.5p i n diameter or e l l i p s o i d ( f i g . 19), 13.0-20.5 x 15-24.Op with a surrounding halo ( f i g s . 18, 38), 5.0-12.5p thick. Halo with laminations p a r a l l e l to sporangium surface ( f i g . 18) and disappearing at time of spore discharge. Mature zoosporangial walls l i g h t brown. Rhizoids coarse and extensive. Uniguttulate spores, 3.5-5p wide, escaping sporangium through one ( f i g . 19) or two broad p a p i l l a e en masse,.^ remaining motionless for a time and swimming away. Resting bodies l i g h t l y golden, germination not observed. My material i s s i m i l a r to Hanson's (1944) except for sporangial and zoospore sizes and the number and s i z e of discharge pores. Sporangia are smaller and spores larger than o r i g i n a l l y described by Hanson. I do not consider the halo as part of the sporangial wall which may account f o r the d i f f e r e n c e i n sporangia Plate V I I — F i g u r e s 20, 21. Chytriomyces annulatus; F i g . 20. Developing s t i p i t a t e sporangium with s i n g l e basal r i n g or undulation. F i g . 21. Thallus with bushy r h i z o i d s and several rings i n sporangium base region. Figures 22, 23. C_. poculatus. F i g . 22. E l l i p t i c a l sporangium with broad a p i c a l operculum. F i g . 23. Pseudo i n t e r n a l p r o l i f e r a t i o n i n empty sporangium. Figures 24-27. Hyphochytrium  catenoides. F i g . 24. Spherical and s l i g h t l y oval sporangia joined by septate hypha. F i g . 25. Long, i r r e g u l a r discharge tube on sporangium. F i g . 26. A p a r t i a l l y septate sporangium. F i g . 27. Broad septate hyphae around a sporangium. Figure 28. Nowakowskiella  elegans; s p h e r i c a l sporangium with a s i n g l e a p i c a l operculum. F i g s . 20-28, xl325. 53 54 s i z e s . Zoospores are larger than the type. Unlike the reports of Willoughby (1965) and Dogma (1969b), halo laminations are v i s i b l e . This i s the f i r s t report of R. coronum from the A r c t i c . 13. Rhizophydium eleyensis Sparrow, Trans. B r i t . Mycol. Soc;, 40: 525, F i g . 2, I-M, 1957. On snakeskin: a raised beach ridge s i t e , 121 and an i c e wedge polygon s i t e , 394. Th i s • c h y t r i d , with angular sporangia, 20-25p i n diameter, i s reported f or the f i r s t time from the A r c t i c . 14. Rhizophydium k a r l i n g i i Sparrow, Aquatic Phycomycetes 243, F i g . 17, A, 1960. On p o l l e n : a r a i s e d beach slope s i t e , 403 and a Carex hummock s i t e , 443. Rhizoids f i n e , sparsely branched, a r i s i n g at end of a sub-sporangial peg, 4 x 8-15p. Sporangia s p h e r i c a l , 10-26p, or ovoid, 10.0-15.5 x 12.5-18.0p?.) Minutely uniguttulate zoospores, 3p wide, escaping sporangium s i n g l y through one or two pores, 3-5p i n diameter * P a p i l l a e low and not as pronounced as figured by Kar l i n g (1938). This i s the f i r s t report of this species i n the A r c t i c and possibly the f i r s t since Sparrow's d e s c r i p t i o n . 15. Rhizophydium p o l l i n i s - p i n i (Braun) Zopf, Abhandl. Naturforsch. G e s e l l . Halle, 17:82, PI. 1, F i g s . 16-20, 1887. On polle n : i c e wedge polygon s i t e s , 383, 388, 393; a rai s e d beach ridge s i t e , 390; a Carex hummock s i t e , 428; a snowbed-moss 55 s i t e , 435 and an A r c t a g r o s t i s l a t i f o l i a meadow s i t e , 438. On p o l l e n mature sporangia 8.0-(18.5)-29.0y i n diameter and on MHU medium, 18-31y i n diameter, d e h i s c i n g through a s i n g l e a p i c a l p a p i l l u m ( f i g s . 13, 36), 5.5-13.5u hroad on p o l l e n and 7—18JJ wide on MHU medium. U n i g u t t u l a t e spores m o t i l e p r i o r to discharge, from p o l l e n grown t h a l l i 3-5y across and from MHU 3.5-5.0y. F l a g e l l a 25-30y long. Empty sporangia urniform w i t h o c c a s i o n a l pseudointernal p r o l i f e r a t i o n o c c u r r i n g ( f i g . 14). Rhizoids once ( f i g . 13) or many times branched. Hohnk (1960) found Rhizophydium p o l l i n i s - p i n i i n Greenland s o i l s . 16. Rhizophydium spherotheca Zopf, Abhandl. N a t u r f o r s c h , G e s e l l . H a l l e , 17:92, P I . 2, F i g s . 33-41, 1887. On p o l l e n : i c e wedge.polygon s i t e s , 383, 389, 393-395, 402; Carex hummock s i t e s , 396, 428-430, 441, 443; r a i s e d beach slope s i t e s , 403, 406, 410, 416; Saxifraga-Rhacomitrium slope s i t e s , 411, 413, 419; r a i s e d beach r i d g e s i t e s , 390, 418; a g r a n i t i c boulder s i t e , 405; a Carex-moss s i t e , 407; an A r c t a g r o s t i s l a t i f o l i a meadow s i t e , 422 and a snowbed-moss s i t e , 435. Sporangia on p o l l e n s p h e r i c a l , 9-26y wide, or subglobose 13-18 x 16-23y, on MHU medium s p h e r i c a l , 13.0-33.5y i n diameter. M o t i l e zoospores escaping sporangium through one to f i v e wide p a p i l l a e anywhere on sporangium w a l l ; from t h a l l i on p o l l e n spores 2.5-4.0y wide and from sporangia on MHU medium 2.0-3.5y; spore contents i n c l u d i n g a minute l u s t r o u s body and a l a r g e r , M.y i n diameter, c o l o r l e s s body; f l a g e l l u m of each spore 16-20y long. On 56 p o l l e n r h i z o i d s generally f i n e and much branched, on MHU medium coarse or f i n e and much branched. My material d i f f e r s s l i g h t l y from that o r i g i n a l l y . d e s c r i b e d . Spores are more v a r i a b l e i n s i z e , 2-4p wide as compared with 2.5-3.Op i n diameter. Sporangia are l a r g e r , to 33p i n diameter, than the type, which i s seldom more than 22p wide. These differences have been noted p r w i o u s l y ( M i l l e r , 1961; Booth, 1969). This fungus has been reported from Greenland (Hohnk, 1960). Rhizophydium sp. ( u t r i c u l a r e ? ) On p o l l e n : a Carex-moss s i t e , 407 and a Saxifraga-Rhacomitrium slope s i t e , 413. Sporangia angular with broad discharge p a p i l l a e i n the angles (figs_. L'12, 37), at maturity on p o l l e n subglobose 15-32 x 18-37p, and on MHU medium s p h e r i c a l 33-52p i n diameter; one to f i v e p a p i l l a e . Spores 3.5-5p motile i n sporangium and escaping p a p i l l a r d i s s o l u t i o n . Rhizoids coarse and branched ( f i g . 12). Angular sporangia r e l a t e this material to Rhizophydium angulosum. However, sporangia and zoospores of my material are larger than those of R. angulosum and i t i s possibly most c l o s e l y r e l a t e d to Rhizophydium utriculare^^U-ebelmesser. Spores are smaller, 3.5-5.0 vs. 8.Op, than those of R. utriculargT^; Also, spores of R. u t r i c u l a r , ^ have no r e f r a c t i v e bodies whil those of my Isolate have a s i n g l e , vLp wide, r e f r a c t i v e globule. Rhizoids of Uebelmesser's (1956) material were hardly, i f at a l l , branched but r h i z o i d s of my i s o l a t e are branched. The Rhizophydium described here i s s i m i l a r to Booth's v (1969) account of R. u t r i c u l a r e . y P l a t e V I I I — F i g u r e s 29-32. Nowakowskiella elegans. F i g . 29... I r r e g u l a r l y shaped sporangium. F i g . 30. Fusiform spindle body. F i g . 31.. Irregular spindle body. F i g . 32. Encysted spores within sporangial confines. Figures 33, 34. Rhizidium verrucosum. F i g . 33. Warty sporangial walls. F i g . 34. Discharge p a p i l l a e . Figure 35. Rhizophydium sp. (nodulosum?); thallus with a thickened r h i z o i d a l axis and crescent shaped discharge p a p i l l a e on sporangium. Figure 36. R. p o l l i n i s - p i n i ; sporangium with broad a p i c a l discharge papillum. Figure 37. Rhizophydium sp. ( u t r i c u l a r e ? ) ; somewhat angular sporangium with broad discharge p a p i l l a e . Figure 38. R. coronum; young sporangium with halo about i t . Figure 39Q R. angulosum; small angular sporangium. F i g s . 29-32, 34-39, xl325. F i g . 33, x200. 58 59 18. Rhizophydium sp. (nodulosum?) On snakeskin: a Carex hummock s i t e , 428. On p o l l e n sporangia spherical,.17-48y i n diameter, or ovoid, 20-43 x 27-55M, on MHU medium 55-90y wide. Crescent-shaped discharge p a p i l l a e ( f i g . 35), one to three per sporangium, broad, 12-22y on MHU medium, and elevated, 3-20u high, swelling s l i g h t l y and d i s s o l v i n g at spore discharge. Uniguttulate spores, 5-7p wide discharged as a non-motile mass and assume m o t i l i t y a f t e r a short time. The sporangial w a l l p e r s i s t i n g a f t e r discharge, covered by low verrucae. Rhizoids, often a r i s i n g from more than one place on sporangium, coarse and branched ( f i g . 35) with the main r h i z o i d up to 10y i n diameter at sporangium base. Save for the much larger spore of my i s o l a t e , 5-7 vs.>'!3u, there i s l i t t l e d i f f e r e n c e between i t and Karling's (1948a) Rhizophydium nodulosum. 19. Rhizophydium sp. (patellarium?) On p o l l e n : a r a i s e d beach ridge s i t e , 418 and a snowbed-moss s i t e , ' 4 3 5 . Basal portions of sporangial wall thickened (figs.115, 16) to 5y. On p o l l e n sporangia 14-20u i n diameter, on MHU medium 13-24u wide. Spores escaping sporangium by deliquescence of upper one-half to two-thirds of sporangial wall with deliquescence often incomplete and fragments of upper sporangial wall p e r s i s t i n g . No discharge p a p i l l a formed. Uniguttulate planonts, 2;5—3u wide, not motile immediately a f t e r sporangial w a l l d i s s o l u t i o n but remaining as a motionless mass before assuming m o t i l i t y and swimming away. Lower one-third to one-Map I I — F i g u r e 40. Location of the Base Camp Lowland on Devon Island. Approximate p o s i t i o n of Devon Island i n Canadian A r c t i c Archipelago shown by i n s e t . 62 ha l f of the sporangial wall remaining a f t e r discharge ( f i g . 16). Rhizoids f i n e and branched with a s i n g l e unbranched haustorium-l i k e structure sometimes occurring. On the extent of the r h i z o i d a l system and sporangial wall thickness separate my material from Rhizophydium patellarium Scholz. A s i n g l e unbranched r h i z o i d was t y p i c a l for Scholz.'js (1958a) fungus. Also, basal portions of the sporangial w a l l were always as t h i n as the r e s t of the w a l l . Perpnosporales 20. Py.thium irregulare""; Buisman, Meded. phytopath. Lab. W i l l i e Commelin Scholten, 11:38, F i g s . 9-11, 1927. (See Matthews, 1931, p. 96; Middleton, 1943, p. 121; and Waterhouse, 1968, p. 37.) With p o l l e n : r a i s e d beach ridge s i t e s , 390, 418, 420; ra i s e d beach slope s i t e s , 403, 406, 410; i c e wedge polygon s i t e s , 393, 402; a g r a n i t i c boulder s i t e , 405; a Carex hummock s i t e , 424; a Carex-moss s i t e , 432 and an Ar c t a g r o s t i s l a t i f o l i a meadow s i t e , 438. Hohnk (1960) reported this species from Greenland. Beisbrock and Hendrix (1967) demonstrate taxonomic d i f f i c u l t i e s produced by the environmental and c u l t u r a l v a r i a b i l i t y of this species. 21. Pythium spp. With p o l l e n : an i c e wedge polygon s i t e , 386; an Arc t a g r o s t i s  l a t i f o l i a meadow s i t e , 391 and a Saxifraga-Rhacomitrium slope s i t e , 412. 63 4. OCCURRENCE AND DISTRIBUTION OF SOME ZOOSPORIC FUNGI FROM SOILS OF HIBBEN AND MORSEBY ISLANDS, QUEEN CHARLOTTE ISLANDS. I n t r o d u c t i o n Methods and m a t e r i a l s Taxonomy and morphology INTRODUCTION The Queen C h a r l o t t e I s l a n d s , 51°57' to 54°16'N. and 130°56' to 133°12'W., l i e immediately southwest of the t i p of Alaska's "panhandle" and o f f B r i t i s h Columbia's northwest coast. Although there are approximately 35 to 40 l a r g e i s l a n d s i n t h i s wedged ar c h i p e l a g o , two i s l a n d s , Graham and Moresby, comprise n e a r l y 90% of the land mass ( f i g . 61). Mountains f r o n t most of the western and northwestern f j o r d coasts of both i s l a n d s . Sand dunes backed by a broad lowland form the coast of eastern and northeastern Graham I s l a n d . Although the v a s c u l a r p l a n t f l o r a i s w e l l known (Calder and Taylor, 1968; Taylor and M u l l i g a n , 1968), there i s r e l a t i v e l y l i t t l e i n f o r m a t i o n on the occurrence and d i s t r i b u t i o n of cryptogams. Recently the f i r s t study of freshwater algae was completed ( S t e i n and Gerrath, 1969) and marine macro-algae were studi e d e a r l i e r (Scagel, 1957; Druehl, 1968). Bryophytes have been studied by Persson (1958) and S c h o f i e l d (1962, 1966, 1969) and W. B. S c h o f i e l d and I . M. Brodo are preparing a comprehensive r e p o r t of bryophytes and l i c h e n s . Of a l l cryptogams on the i s l a n d s f u n g i are the l e a s t known. In Calder and Taylor (1968), S a v i l e reported Exobasidium empetri I t o and Otani and Pu c c i n i a p i m p i n e l l a e ( S t r . ) Rohl. ssp. c h a r l o t t a e S a v i l e . S a v i l e 64 (1968) also reported Peronospora grisea (Uhger) Unger and I s a r i o p s i s  veronicae ((Pass.) S a v i l e . METHODS AND MATERIALS During the summer of 1968, as part of a survey of coas t a l s o i l s for zoosporic fungi, s o i l samples were taken from one s i t e on Hibben Island and several Moresby Island l o c a t i o n s ( s o l i d t r i a n g l e s , f i g . 61). As each sample was taken i t s general habitat was noted. Table II summarizes the s i t e name, c o l l e c t i o n number, l a t i t u d e and longitude, and habitat type of each c o l l e c t i o n . Habitat types are described elsewhere (Appendix I ) . S o i l samples were baited for fungi as outlined by Couch (1939). Two subsamples per sample were treated with Capilano River water at 0 ° / o o s a l i n i t y and two subsamples of samples from marine or marine-influenced habitats were treated with sea water adjusted to 7 ° / 0 0 s a l i n i t y . Pinus s y l v e s t r l s L. p o l l e n , d i a l y z i n g tubing cellophane, snakeskin, p u r i f i e d shrimp exoskeleton and blond baby ha i r were used as b a i t s . Each " s t a r t " was surveyed for fungi over a 4-6 week period. Pure cultures of some species were prepared and studied as described i n an e a r l i e r report (Booth and Barrett, 1971). TAXONOMY AND MORPHOLOGY a. C h y t r i d i a l e s 1. Blyttlomyces aureus - Booth, Syesis, 2:142, F i g s . 15-22, 58-61, 1969. On p o l l e n : coastal western hemlock s i t e s , 195, 251, 271 and 65 a dune s i t e , 258. Chytridium megastomum Sparrow, Dansk. Bot. Ark., 8:21, p i . 4, A-D, 1934. On pol l e n : a t i d a l edge, 192. Chytriomyces appendiculatus K a r l i n g , B u l l . Torrey Bot. Club, 74: 355, f i g s . 16-37, 43-48, 1947. On snakeskin and polle n : c o a s t a l western hemlock s i t e s , 193, 286. Sporangia i r r e g u l a r l y spheroid, 39-105u wide, or elongate, 63-90u x 75-145u. At maturity sporangium wall l i g h t brown and 1-3.5u thick with a portion of the zoospore cyst wall remaining on the sporangium ( f i g . 27). Rhizoids coarse and much branched. Opercula 6-13u wide, one or two on each sporangium with a gelatinous"matrix extending below to one h a l f sporangium height. At dehiscence, matrix exuding through opercular opening p r i o r to spore escape. Zoospores 5-6u i n diameter with a 1.5-2.5u globule push into the matrix and eventually swim free . Resting spores not seen. Spores and sporangia described here are s l i g h t l y smaller than the type. Also, as Dogma (1969a) notes, the gelatinous matrix may extend'^deeper than o r i g i n a l l y described. Otherwise there i s no dif f e r e n c e between my material and that of K a r l i n g . Willoughby (1961a), M i l l e r (1965) and Dogma (1969a) report C. appendiculatus from habitats with an acid pH. Other reports, K a r l i n g (1948b, 1967c), involve C_. appendiculatus with an acid environment. 66 4. Chytriomyces hyalinus K a r l i n g . Amer. Jour. Bot.32:363, Figs. 46-61, 1945. On p o l l e n and c h i t i n : beach i n t e r t i d a l s i t e s , 165, 178, 187, 188, 190, 191, 230, 231, 250, 253, 264, 280, 281; t i d a l edge loc a t i o n s , 179, 254; flushes, 204, 208, 225, 227; marine high beach s i t e s , 211, 212, 232, 248, 255, 261 and coastal western hemlock s i t e s , 168, 262. On p o l l e n sporangia s p h e r i c a l ( f i g . 29), 7-(14.5)-25 i n diameter, or ovoid ( f i g . 28), 9-24 x 7-23p and on MHU medium (see Booth and Barrett, 1971) sporangia s p h e r i c a l , 40-(50)-85p wide. Sporangium wall t h i n and c o l o r l e s s , broken by 1-3 opercula ( f i g s . 28, 29). Opercula of sporangia on p o l l e n 5-(7)-10p wide and on MHU medium 7-(16)-21p wide. A small inverted saucer-like structure often above the operculum surface ( f i g . 29). Rhizoids from the sporangium base or an apophysate structure. Sporangial dehiscence by opening of hinged or unhinged opercula. Spores swarm i n v e s i c l e before swimming f r e e l y . Free swimming spores s p h e r i c a l , 3-(4.5)-5(6)p wide from sporangia on p o l l e n and 3.5-4p wide from sporangia on MHU medium with one to several r e f r a c t i v e globules i n protoplast. The above d e s c r i p t i o n , based on s i x very s i m i l a r axenic i s o l a t e s ( c u l . nos. 38, 53, 65, 79, 80, 83)"*" departs l i t t l e from those of Karling and Bostick (1968). Spores of my i s o l a t e s are nearly s p h e r i c a l and may have several r e f r a c t i v e globules. Both Karling and Bostick describe oval spores with a s i n g l e r e f r a c t i v e body. The small saucer-like structure on the operculum has hereto-"*"Number of culture i n author's c o l l e c t i o n -67 fore gone unreported. The o r i g i n of this structure i s not f u l l y understood. I t may be a piece of the i n c i p i e n t sporangium w a l l which has broken away during opercular development. Karling (1967c) distinguished Chytriomyces hyallnus var.' granulatus based p r i m a r i l y on zoospores containing several r e f r a c t i v e globules. Two of my i s o l a t e s form spores with one to many r e f r a c t i v e bodies per spore. On t h i s basis i t i s imprudent to recognize t h e ' v a r i e t y . Chytriomyces poculatus Willoughby and Townley. Trans. B r i t . Mycol. S o c , 44:183, F i g . 3, 1961. On p o l l e n : c o a s t a l western hemlock s i t e s , 167, 186, 195, 249, 286; flushes, 196, 209, 218, 219, 221, 225, 228, 229; blanket bog s i t e s , 201, 203, 207; a t i d a l edge, 213; a marine high beach s i t e , 246 and an alp i n e meadow l o c a t i o n , 260. ?Entophlyctis confervae-glomeratae (Cienkowski) Sparrow f . marina Kobayashi and Ookubo. B u l l . Nat. S c i . Mus. (Tokyo), (N.S.), 1(2), no. 35:67, F i g . 6, 1954. In p o l l e n : a marine high beach s i t e , 255.-' Sporangia s p h e r i c a l ( f i g s . 1, 2), pyriform ( f i g . 4) or i r r e g u l a r ( f i g . 3), 9-22p i n diameter, hyaline protoplast, one ( f i g s . 2-4) or two ( f i g . 1) r h i z o i d a l axes. Rhizoids coarse, branched, d i r e c t l y from the sporangium ( f i g s . 1-3) or an apophysoid structure ( f i g . 4). Zoospore cyst p e r s i s t e n t ( f i g s . 1-3) or evanescent ( f i g . 4). Zoospores and spore discharge not seen. Resting spores 12-20p i n diameter, large eccentric r e f r a c t i v e body i n s i d e , 3-5 spines, to 68 4y long, on sporangium, w a l l . F a i l u r e to note spores and spore discharge make my i d e n t i -f i c a t i o n t entative. . Spiny r e s t i n g spores and presence of zoospore cysts on mature sporangia make this taxonomic d i s p o s i t i o n even more uncertain. Entophlyctis confervae-glomeratae f. marina i s the only taxon i n the genus ex h i b i t i n g fechinulate r e s t i n g spores and yet the zoospore cyst i s evanescent. A l l other species have smooth walled r e s t i n g spores and the cyst i s p e r s i s t e n t i n two of them. Of these two, my fungus i s possibly c l o s e s t to Entophlyctis b u l l i g e r a (Zopf) Fischer. More pronounced spines on r e s t i n g spores as compared with those figured by Kobayasi and Ookubo (1954) add further d i f f i c u l t y . F i n a l l y , the apophysoid structure suggests that my c o l l e c t i o n may be rel a t e d to D i p h l o p h l y c t i s . Entophlyctis sp. In p o l l e n : a headland s i t e , 176; a coastal western hemlock ^ f j / s i t e , 238 and a tide edge lo c a t i o n * 266. My previous d e s c r i p t i o n of this species (Booth, 197l4)__is p a r t i a l l y based on two axenic i s o l a t e s from the Queen Charlotte Islands Karlingiomyces granulatus (Karling)^ ^ Sparrow. Mycologia, 39:57, F i g s . 1-22, 1947. On cellophane: a headland s i t e , 183, and a coastal western hemlock s i t e , 193. Other than zoospores which are s l i g h t l y l a r g e r , 6 . 0 - 8 . 0 u wide, there i s no d i f f e r e n c e between this material and Karl i n g ' s . Plate I X — F i g u r e s 1-4. Entophlyctis confervae-glomeratae. F i g . 1. Spherical endobiotic sporangium with sub-cyst swelling and two r h i z o i d a l axes. F i g . 2. Spherical zoosporangium with attached cyst and s i n g l e r h i z o i d a l a x i s . F i g . 3. Irregular sporangium with cyst and s i n g l e r h i z o i d a l a x i s . F i g . 4. Pyriform sporangium with apophysis and s i n g l e r h i z o i d a l a x i s . Figures 5-14. Rhizophydium sp.-Phlyctochytrium a u r i l a e complex. F i g s . 5-9. Side view of enations of Rhizophydium sp.-2 Phlyctochytrium a u r i l a e complex, f h . (freehand). F i g s . 10-14. Plane view of Rhizophydium sp.-Phlyctochytrium a u r i l a e complex enations, f h . Figure 15. Phlyctochytrium k n i e p i i ; sporangium with several pronounced discharge p a p i l l a e . Figures 16-18. Phlyctochytrium sp. F i g . 16. Spherical sporangium with s i n g l e discharge pore and apophysis. F i g . 17. Unbranched tubular haustorium attached to an angular sporangium. F i g . 18. Angular sporangium with a zoospore cyst segment attached and a tubular haustorium. Figures 19, 20. Rhizophlyctls harder!,--" F i g . 19. Developing type II sporangium with three apophysate r h i z o i d a l axes. F i g . 20. Type I sporangium with branched r h i z o i d s i n the r h i z o i d a l bundle. Figure 21. Rhizophydium sp. ( u t r i c u l a r e ? ) ; t h a l l u s with broad discharge p a p i l l a e and s l i g h t l y swollen r h i z o i d s . Figure 22. Rhizophydium  sphaerocarpum; empty sporangium with broad discharge pore. Figure 23. Rhizophydium sphaerotheca; i n c i p i e n t sporangium with s t r a p - l i k e r h i z o i d a l axes. F i g s . 1-4, 15-23, x460; F i g s . 5-14, x800. F i g s . 1-4, 15-23, 30-35, scale A; F i g s . 5-14, scale B; F i g s . 24, 25, 38, 41, 48, 49, 54, 55, scale C; F i g s . 26-29, 36, 37, 39, 40, 42, 44-47, 50-53, 56-60, scale D and F i g . 43, scale E. 2 -----Unless designated as f h . , all_drawings done with camera l u c i d a . 70 71 9. Nowakowsklella elegans (Nowak.) Schroeter. Engler and P r a n t l , Naturlichen Pflanzenfam., 1:82, 1892. On s o i l : c oastal western hemlock s i t e s , 168, 195 and a f l u s h , 219. Sporangia predominantly s p h e r i c a l , 22.5-40u i n diameter, with 2 or 3 r h i z o i d a l axes ( f i g . 38). Operculum, 5-8.5y wide, f l u s h on the sporangium or at t i p of tube up to 80u long. Motile spores are 6.5-8.5y i n diameter. No appreciable d i f f e r e n c e exists between my material and that o r i g i n a l l y described. 10. Olpidium gregarum (Nowak.) Schroeter. Krypogamenf1. Schlesien, 3: 182, 1885. In a protozoan: a t i d a l edge s i t e , 254. 11. Olpidium luxurians (Tomaschek) Fischer . Rabenhorst.- Kryptogamen-Fl., 1:29, 1892. In p o l l e n : flushes, 218, 220. 12. Olpidium pendulum Zbpf. A. Schenk, Handbuch d. Bot., 4:555, f i g . 66 (1-5), 1890. In p o l l e n : disturbed s i t e s , 162, 163; beach i n t e r t i d a l s i t e s , 164, 1966, 210, 245, 280; marine high beach l o c a t i o n s , 171, 172, 211, 237, 46, 248, 261, 267, 270, 275, 284, 285; headland s i t e s , 176, 177, 183-185; dune s i t e s , 189, 252, 258; flushes, 196, 200, 202, 208, 209, 220, 221, 226, 227; a blanket bog s i t e , 203; t i d a l edge s i t e s , 216, 257, 266, 274; coastal western hemlock s i t e s , 234, 238, 249, 251, 271, 286; an alpi n e meadow s i t e , 260, and tide marsh s i t e s , 72 278, 279. 13. Phlyctidium megastomum Sparrow. Aquatic Phycomycetes, p. 148, 1943. On po l l e n : a blanket bog s i t e , 207, and a coas t a l western hemlock s i t e , 224. 14. Phlyctochytrium africanum Gaertner. Arch. f . Mikrobiol., 21:120, f i g s . 3, 4, 1954. On polle n : a headland s i t e , 169; a coastal western hemlock s i t e , 238, and a beach i n t e r t i d a l s i t e , 273. Sporangia e p i b i o t i c or i n t e r b i o t i c , 21-53u i n diameter with 2-5 discharge p a p i l l a e ( f i g . 39), zoosporangium wall smooth and up to 3p thick. Rhizoids coarse, much branched, a r i s i n g from a branched apophysis. Zoospores swarming i n the sporangium p r i o r to discharge, escaping through pores formed by d i s s o l u t i o n of p a p i l l a e . Motile spores 4-5u wide with a s i n g l e r e f r a c t i v e globule, 1 . 5 n i n diameter. This d e s c r i p t i o n i s based on a s i n g l e axenic i s o l a t e ( c u l . no. 69). Sporangia may be larger than those described by Gaertner or Booth (1969). Zoospores are larger than the type species but the same s i z e I previously reported. Otherwise there i s no d i s t i n c t i o n between these i s o l a t e s and those described e a r l i e r . 15. Phlyctochytrium chaetiferum K a r l i n g . Mycologia, 29:179, f i g s . 1-3, 1937. On polle n : marine high beach s i t e s , 255, 270, 285; a f l u s h , 259, and t i d a l edge s i t e s , 269, 274. 73 16. Phlyctochytrium k n i e p i i Gaertner. Arch. f . Mikrobiol.; 21:117, F i g s . 1, 2, 1954. On p o l l e n : a t i d a l edge s i t e , 179. Only the number and length of r e s t i n g spore p a p i l l a e d i s t i n g u i s h this i s o l a t e from the type. On pine p o l l e n r e s t i n g sporangia ( f i g . 15) have 5-20 p a p i l l a e which may be 10-27p long. Gaertner figures 3-6 p a p i l l a e , 6-16p long. 17. Phlyctochytrium mucronatum Canter. Trans. B r i t . Mycol. S o c , 32:240, F i g s . 1, 2, 1949. On p o l l e n : blanket bog s i t e , 203.. 18. Phlyctochytrium palustre Gaertner. Arch. f . Mikrobiol., 21:123, Fi g s . 5, 6, 1954. On p o l l e n : a disturbed s i t e , 162; coastal western hemlock s i t e s , 238, 286; a beach i n t e r t i d a l s i t e , 247; and a marine high beach s i t e , 255. 19. Phlyctochytrium punctatum Koch. Jour; E l i s h a M i t c h e l l S c i . S o c , 73: 109, Figs. 1-16, 1957. . On p o l l e n : a disturbed s i t e , 163; and a blanket bog s i t e , 201. Sporangium s p h e r i c a l ( f i g s . 40, 41), 15-(21.5)-32p i n diameter on p o l l e n , 23-(32)-46p i n diameter on s o l i d MHU medium and 16-(30)-50p . i n diameter i n l i q u i d MHU medium; subtended by an apophysis ( f i g . 41). Sporangium wall thin, covered with minu.fee punctae. Discharge p a p i l l a e , 2-(3)-6 per sporangium on p o l l e n and 2-(4)-6 per sporangium on s o l i d and i n l i q u i d MHU medium, 2.5-3.0P wide by 2.5-4.OP high on sporangia 74 on p o l l e n and 3.0-6.Op wide by 3.0-5.Op high, on sporangia grown on H-U medium. Zoospores, 3.5-5.5p wide from sporangia on p o l l e n and 2-4p i n diameter from sporangia grown on MHU medium, escaping sporangium i n d i v i d u a l l y from one or more pores formed by d i s s o l u t i o n of p a p i l l a e . Frequently autoparasitism among t h a l l i grown on MHU medium. There i s no s u b s t a n t i a l d i f f e r e n c e between this i s o l a t e and that described by Koch. My d e s c r i p t i o n i s based on one axenic culture ( c u l . no. 50). 20. Phlyctochytrium reinboldtae P e r s i e l . Arch. f . M i k r o b i o l . , 32:414, F i g . 1, 1959. On p o l l e n : a coastal western hemlock s i t e , 238. Thallus ( f i g . 42) dimensions and morphology are s i m i l a r to P e r s i e l ' s d e s c r i p t i o n . 21. Phlyctochytrium semiglobiferum Uebelmesser. Arch. f. M i k r o b i o l . , 25:317, F i g . 5, 1956. On p o l l e n : a marine high beach s i t e , 170; a beach i n t e r t i d a l s i t e , 239; and a c o a s t a l western hemlock s i t e , 286. On p o l l e n sporangia ( f i g . 43) are smaller than those reported by Uebelmesser and Sparrow (1968a). They more nearly approximate the s i z e , <20-30p i n diameter, reported by Booth (1969). Discharge p a p i l l a e are hemispherical ( f i g . 43) and, at 4-8p wide, smaller than the type. A l l other d e t a i l s of the Queen Charlotte material agree with the o r i g i n a l d e s c r i p t i o n . 75 22. ?Phlyctochytrium sp. On po l l e n : a f l u s h , 218. Sporangia s p h e r i c a l ( f i g . 16), 11-34 wide, subangular ( f i g s . 17, 56, 57), 14-34p at greatest width, subglobose ( f i g . 18), e l l i p t i c a l or pyriform with broad p a p i l l a e at angles ( f i g s . 56, 57). Sporangium w a l l thin and c o l o r l e s s with a portion of the zoospore cyst sometimes remaining ( f i g . 18). Rhizo i d a l system, v a r i a b l e , a r i s i n g from a branched or unbranched apophysis. Rhizoids tapering ( f i g s . 16, 57) or unbranched and haustoroid ( f i g s . 17, 18). Zoospores motile p r i o r to discharge and escaping sporangium through pores formed by p a p i l l a r d i s s o l u t i o n , s p h e r i c a l , 3-5p i n diameter, with a si n g l e r e f r a c t i v e globule, 1.5p wide, f l a g e l l a e 15.5-20p long. This c o l l e c t i o n has a f f i n i t i e s with Phlyc to chy t r ium, Blyttiomyces and Phlyctidium species. The apophysis when branched, r e l a t e s i t to Phlyctochytrium africanum, J?. k n i e p i i and _P. reinboldtae. Otherwise, i t i s s i m i l a r to multiporous phlyctochytriums with unbranched apophyses. With an a p i c u l o i d structure, i t i s l i k e Blyttiomyces l a e v i s Sparrow, 13. r h i z o p h l y c t i d i s Dogma and Sparrow and B. aureus Booth. Dogma and Sparrow (1969) point out the possible danger of assigning a Phly c to chy trium species to Blyttiomyces. F i n a l l y , a subsporangial endobiotic haustoroid structure r e l a t e s my material to the smooth walled species of Phlyctidium that possess elongate tubular haustoria. 23. Phlyctochytrium sp. (dentiferum?) On p o l l e n : a f l u s h , 199 and a blanket bog s i t e , 203. Four a p i c a l bumps and one or two l a t e r a l ridges on i n c i p i e n t sporangium ( f i g . 44) developing into an a p i c a l crown of four b i p a r t i t e Plate X — F i g u r e 24. Achlya sp.; ornate oogonium. Figures 24, 25. B l a s t o c l a d i e l l a b r i t a n n i c a . F i g . 24. Portion of a thallus on shrimp exoskeleton. F i g . 25. Resting sporangium on pol l e n . Figure 27. Chytriomyces appendiculatus; portion of sporangium with cyst attached. Figures 28, 29. C_. hyalinus. F i g . 28. Inverted saucer on an a p i c a l operculum. F i g . 29. B i -operculate sporangium. Figures 30-35. Rhizophydium sp.-Phlyctochytrium a u r i l a e complex. F i g . 30. Spherical sporangium with simple teeth. F i g . 31. Pyriform sporangium with simple • teeth and bumps. F i g . 32. Irregular sporangium with broad b i p a r t i t e teeth and spher i c a l sporangium devoid of teeth. F i g . 33. Incipient sporangium and spher i c a l sporangium covered with simple teeth. F i g . 34. Spherical sporangium with short simple and b i p a r t i t e teeth. F i g . 35. Enationless sph e r i c a l sporangium, i r r e g u l a r sporangium and empty sporangium with bumps and b i f u r c a t e spines. F i g s . 24, 26-35, xl325. F i g . 25, x650. 77 78 and one or two l a t e r a l rows of b i p a r t i t e ( f i g . 46) to pentapartite ( f i g . 47) teeth. Teeth fused giving rows a flanged appearance. D e l i c a t e , branched r h i z o i d s from small apophysis subtending sporangium. Zoospores and zoosporangial discharge not seen. This .material i s s i m i l a r to Phlyctochytrium dentiferum Sparrow i n some respects. In both, there i s a set of a p i c a l and at l e a s t one subtending whorl of teeth. However, i n my material, the i n d i v i d u a l teeth of a row are fused. Sparrow's (1938) figures (15-41) i n d i c a t e that teeth of P^ : dentif erum maintain th e i r i n d i v i d u a l i t y and are only b i p a r t i t e . Also, he figures no more than one subtending row of teeth. 24. Phlyctochytrium sp. (reinboldtae?) On p o l l e n : a f l u s h , 211. Sporangia with very pronounced discharge p a p i l l a e ( f i g . 48) and subtending broad apophyses ( f i g . 48). Zoospores and zoosporangium discharge not observed. Sporangia are generally shaped l i k e those of Phlyctochytrium  reinboldtae but discharge p a p i l l a e of my material are much more pronounced. 25. Phlyctochytrium sp. (semigloblferum?) On p o l l e n : a c o a s t a l western hemlock s i t e , 194. Sporangia p r i m a r i l y e l l i p t i c a l , 10.5-27p - 11.5-34p, with two to ten broad, 7.5-10p, hemispherical p a p i l l a e which impart a gibbose appearance. Sporangium wall t h i n , smooth and c o l o r l e s s . Coarse, s l i g h t l y branched r h i z o i d s a r i s i n g from a tapering apophysis, 5-10p wide at sporangium ibase. Zoospores motile i n the sporangium p r i o r 79 to discharge, 3-4y i n diameter with a s i n g l e minute r e f r a c t i v e globule. Sporangia and zoospores of this material are smaller than those of Phlyctochytrium semiglobiferum. A greater number of discharge p a p i l l a e andgibbose sporangia also distinguishes i t from P. semiglobiferum. However, broad discharge p a p i l l a e and a turnip-shaped apophysis r e l a t e my fungus more c l o s e l y to J?.1  semiglobiferum than to any other taxon. 26. Phlyctochytrium spectabile Uebelmesser. Arch. f . Mikrobiol., 25: 315, F i g . 4, 1956. On p o l l e n : a f l u s h , 259. 27. Rhizidium l a e v i s K a r l i n g . Amer. J. Bot., 31:254, F i g s . 20-22, 1944. On shrimp exoskeleton: a blanket bog s i t e , ,207. 28. Rhizidium, sp. (nowakowskii?) On p o l l e n : a f l u s h , 225. Sporangia i n t e r b i o t i c ( f i g . 49), s p h e r i c a l to i r r e g u l a r , and 23-65u i n diameter. R h i z o i d a l axis 4-7y broad with r h i z o i d s branched ending abruptly on contacting substrate. Mature sporangium proto-plasm d i f f e r e n t i a t e d into spore anlagen which discharges into a v e s i c l e ( f i g s . 50, 51) through a s i n g l e a p i c a l opening i n the sporangium w a l l . A f t e r a short quiescent period spores swarming wit h i n v e s i c l e and eventually breaking out. Motile spores, 4.5-5.5p i n diameter c o l o r l e s s with a s i n g l e r e f r a c t i v e globule, 1.5-2.5y wide. Morphological features of this material r e l a t e i t most c l o s e l y 80 to Rhizidium nowakowskii K a r l i n g . As i n Karling's (1944) study, no r e s t i n g spores were observed. 29. Rhizidium spp. On p o l l e n and shrimp exoskeleton: coastal western hemlock s i t e s , 224, 271 and a f l u s h , 225. 30. Rhizidium varians K a r l i n g . Amer. J . Bot., 36:681, f i g s . 1-26, 1949. On polle n : a f l u s h , 218. 31. Rhizidium verrucosum K a r l i n g . Amer. J . Bot., 31:255, f i g s . 34-63, 1944. On snakeskin: a f l u s h , 221. 32. Rhizophlyctis harder!" Uebelmesser. Arch. f . Mikrobiol., 25:324, f i g . 7, 1956. On po l l e n : beach i n t e r t i d a l s i t e s , 178, 188, 244, 263, 264; marine high beach s i t e s , 180, 237, 248, 265, 267, 275; and t i d a l edge s i t e s , 243, 254, 266, 269, 274. Type II sporangia ( f i g s . 19, 52, 53) s p h e r i c a l , smooth walled and 7-(13)-23.5p i n diameter on po l l e n , 14-(22.5)-34p i n diameter on s o l i d MHU medium and 8-(14)-18.5p i n l i q u i d MHU medium. Rhizoids d e l i c a t e , branched, on MHU medium from small apophyses ( f i g s . 19, 52), on p o l l e n d i r e c t l y from the sporangium at one or several places. Zoospores discharged through a large a p i c a l opening of the sporangium, rocking slowly at the discharge opening before swimming a c t i v e l y . When motile 3.5-(4.5)-5.5p wide on a l l substrates with a s i n g l e minute 8 1 r e f r a c t i v e globule. Empty sporangia appearing urniform ( f i g . 53). Type I sporangia ( f i g s . 20, 54, 55) s p h e r i c a l , 8-(ll)-15y i n diameter, or oviform, 9-(13)-18 x 7-(10)-15y with a large c e n t r i c or eccentric r e f r a c t i v e body ( f i g s . 20, 54), formed only on poll e n . A bundle of branched ( f i g s . 20, 55) or unbranched and twisted ( f i g . 54) or r e l a t i v e l y untwisted ( f i g s . 20, 55) rh i z o i d s a p i c a l l y located. Zoospore discharge was not observed. This material d i f f e r s l i t t l e from that described by Uebelmesser, Sparrow (1968a) and Booth (1969). However, this i s the f i r s t report of branching rh i z o i d s on Type I sporangia. The above d e s c r i p t i o n i s based on two axenic i s o l a t e s ( c u l . nos. 44, 51). 33. Rhizophlyctis hyalina (Karling) Sparrow. Aquatic Phycomycetes, p. 445, 1960. On cellophane: a blanket bog s i t e , 207. 34. Rhizophlyctis rosea (deBary and Woronin) Fischer. Rabenhorst. Kryptogamen-Fl., 1:122, 1892. On cellophane: a marine high beach s i t e , 170; a t i d a l edge s i t e , 192; and coastal western hemlock, 223, 287. 35. Rhizophydium ampullaceum (Braun) Fischer. Rabenhorst. Kryptogamen-F l . , 1:101, 1892. On poll e n : a dune s i t e , 258, and a coastal western hemlock s i t e , 262. 36. Rhizophydium carpophilum (ZOpf) Fische r . Rabenhorst. Kryptogamen-82 F l . , 1:95, 1892. On p o l l e n : beach i n t e r t i d a l s i t e s , 169, 256; coastal western hemlock s i t e s , 193, 224, 271; flushes, 197, 199, 208, 228, 229; a t i d a l edge s i t e , 215; and a dune s i t e , 258. 37. Rhizophydium chitinophilum A n t i k a j i a n . Mycologia, 39:613, f i g s . 1-20, 1947. On shrimp exoskeleton: blanket bog s i t e s , 201, 207; coastal western hemlock s i t e s , 223, 224; and an alpine meadow s i t e , 260. Sporangia ( f i g . 59) 20.5-44p i n diameter with a si n g l e r h i z o i d a l axis and one or two s e s s i l e p a p i l l a e . Rhizoids coarse and. branched. Zoospores motile before escaping sporangium confines, 3.5-4.Op wide with a sing l e minute r e f r a c t i v e globule. Resting sporangia not seen. For the most part t h i s material i s s i m i l a r to that described by A n t i k a j i a n . Sporangia of my material are s l i g h t l y smaller but within the accepted l i m i t s of the species. 38. Rhizophydium chytriomycetis K a r l i n g . Mycologia, 38:115, f i g s . 1-8, 1946. On Chytriomyces hyalinus: beach i n t e r t i d a l s i t e s , 165, 190, 230, 265; a coastal western hemlock s i t e , 167; t i d a l edge s i t e s , 179, 254; a marine high beach s i t e , 261 and an alpine meadow s i t e , 260. 39. Rhizophydium coronum Hanson. Torreya, 44:31, 1944. On p o l l e n : a marine high beach s i t e and a coastal western hemlock s i t e , 203. Plate X I — F i g u r e s 36, 37. Rhizophydium sp.-Phlyctochytrium_ a u r i l a e complex. Spherical sporangium covered with p o l y p a r t i t e teeth i n side view. F i g . 37. Spherical sporangium covered with p o l y p a r t i t e teeth i n plane view. Figure 38. Nowakowskiella elegans; apophysate sporangium. Figure 39. Phlyctochytrium africanum; sporangium with several discharge p a p i l l a e . Figures 40, 41. „..}?_. punctatum. F i g . 40. Punctae i n wall of empty sporangium. F i g . 41. Developing thallus with a s l i g h t l y branched apophysis and coarse branched r h i z o i d s . Figure 42. P_. reinboldtae; pronounced p a p i l l a e on sporangium and branched apophysis. Figure 43. P. semlgloblferum; broad hemispherical discharge p a p i l l a e and lower?half of turnip shaped apophysis. Figures 44, 45. Phlyctochytrium sp. (dentiferum?). F i g . 44. Inc i p i e n t sporangium. F i g . 45. Sporangium with an a p i c a l crown and two l a t e r a l rows of b i f u r c a t e teeth. F i g s . 36, 37, 39, 40, 42, 44, 45, xl325. F i g s . 38, 41, x650. F i g . 43, xl900. 84 85 Sporangia s p h e r i c a l , 15.5-29.5p i n diameter with three to f i v e discharge p a p i l l a e , 3-4 x 3-6p, enclosed by an unlaminated gelatinous sheath, 3-4p thick, frequently borne on a r h i z o i d a l s t a l k up to 5p long. Rhizoids f i n e and much branched. Zoospores motile i n sporangium about one minute before discharge, 4-5p i n diameter with a r e f r a c t i v e body ^ l p i n width. F a i n t l y golden r e s t i n g spores ( f i g . 58), 16.5-22p i n diameter, associated with this fungus. As Booth and Barrett (1971) reported, sporangia are smaller and zoospores are s l i g h t l y larger than those.of the type. Like Willoughby (1965) and Dogma (1969b) observed, laminations are absent from the sheath surrounding the sporangium. Resting sporangia are smaller than Hanson's material. • 40. Rhizophydium gibbosum (Zopf) Fisch e r . Rabenhorst. Kryptogamen-F l . , 1:102, 1892. On p o l l e n : a c o a s t a l western hemlock s i t e , 272. 41. Rhizophydium minutum Atkinson. Bot. Gaz., 48:328, F i g . 4, 1909. On p o l l e n : a f l u s h , 208 and a co a s t a l western hemlock s i t e , 262.: 42. Rhizophydium nodulosum K a r l i n g . Mycologia, 40:328, F i g s . 1-10, 1948. On snakeskin: flushes, 198, 259 and a coastal western hemlock s i t e , 224. 43. Rhizophydium parsiticum Shen and Siang. Science Reports of 8 6 National Tsing Hua University, Ser. B: Biol, and Psych. Sciences 3:181, f i g . 2, 1948. On Rhizophydium sphaerotheca; flushes, 198, 209 and a ti d a l edge, 213. Sporangial shape (fi g . 60) and a l l other morphological features agree with the original description. 44. Rhizophydium p o l l i n i s - p i n i (Braun) Zopf. Abhandl. Naturforsch. Gesell. Halle, 17:82, p i . 1, figs. 16-20, 1887. On pollen: a beach intertidal s i t e , 166, and coastal western hemlock sites, 181, 193. 45. Rhizophydium racemosum Gaertner. Arch. f. Mikrobiol. 21:125, f i g . 7, 1954. On pollen: a headland s i t e , 176; a beach intertidal s i t e , 247; a tide marsh si t e , 277 and a marine high beach s i t e , 284. 46. Rhizophydium sp.-Phlyctochytrium aurilae Ajello complex. On pollen and shrimp exoskeleton; flushes, 197, 199, 206, 209, 218, 221 and a blanket bog'site, 203, Sporangia apophysate or non-apophysate with varying numbers and complexity of spines. (A series of increasing spine number and complexity is seen by starting with figure 30 and ending at figure 37.) In side view enations unipartite (figs. 5, 30-34), bipartite (figs. 6-8, 35) or polypartite. Polypartite enations constructed of a common base bearing any number of pointed spines. In plane view (figs. 10-14, 37) polypartite enations variable i n numbers 87 and r e g u l a r i t y of spines. Frequently, i n s o i l p l a t e cultures c e r t a i n i n d i v i d u a l s may be ascribed to Phlyctochytrlum a u r i l a e . Conversely, a dentate form re l a t e d to Rhizophydium can occur i n the same plat e s . Several character combinations between these are also found. Because of such v a r i a b l e form and a lack of knowledge about the o r i g i n a l inoculum i t i s probably wisest, following M i l l e r ' s (1968) suggestion, to consider t h i s a species complex. F i n a l l y , those forms with p o l y p a r t i t e enations may go beyond A j e l l o ' s (1945) concept of J?. a u r i l a e which has u n i p a r t i t e and b i p a r t i t e enations only. My poly-p a r t i t e forms could be considered more c l o s e l y r e l a t e d to Sparrow's (1966) Phlyctochytrium furcatum. However, u n t i l the nature and v a r i a b i l i t y of enations in. the Rhizophydium sp.-Phlcytochytrium  a u r i l a e complex i s better understood, my p o l y p a r t i t e forms would best be retained i n i t . 47. Rhizophydium sp. (bullaturn?) On p o l l e n : a blanket bog site,'207; flushes, 209, 229 and a co a s t a l western hemlock s i t e , 249. 48. Rhizophydium sp. (sphaerotheca?) On p o l l e n : a coastal western hemlock s i t e , 223 and a f l u s h , 225. At maturity sporangia smooth walled, c o l o r l e s s , s p h e r i c a l , 12-28u i n diameter. Sporangium w a l l double contoured and broken by three to ten' s e s s i l e p a p i l l a e . Rhizoids profusely branched, from a s i n g l e a x i s . Zoospores undergoing short rocking period w i t h i n 88 sporangium, prior to discharge, escaping singly through 2-3p wide pores formed by papillar dissolution, 2.5-4.5y in diameter with a single minute refractive globule and a posterior flagellum ^25-30p long. Resting sporangia, 10.5=28.Op in diameter, protoplasts with a large eccentric refractive globule. Resting spore rhizoids slender and l i t t l e branched. This fungus differs from Rhizophydium sphaerotheca in the greater number of discharge papillae, 49. Rhizophydium sp. (utriculare?) On pollen: coastal western hemlock sites, 173, 186, 223, 262, 271, 272; a headland s i t e , 183; flushes, 196, 198-200, 206, 208, 220, 222, 226, 228, 259; a blanket bog s i t e , 203; tid a l edge sites, 205, 213; and a beach intertidal s i t e , 264. On pollen sporangia epibiotic or interbiotic, 8-(17)-31p i n diameter with one to four discharge papillae, 3-7y broad, imparting an angular appearance (fi g . 21). Sporangia on MHU medium 22-(37)-55p in diameter with three to ten discharge papillae. Rhizoids fine and branched. Zoospores, motile prior to discharge, 2.5-(3.5)-4.0p in diameter with a single refractive globule ly in diameter. This material i s similar to Rhizophydium sp. (utriculare?) described by Booth and Barrett (1971). Zoospores of their collections are slightly larger than those described here. My above description i s based on three axenic isolates (cul. nos. 39, 56, 77). 50. Rhizophydium sphaerocarpum (Zopf) Fischer. Rabenhorst. Kryptogamen-F l . , 1:95, 1892. Plate X I I — F i g u r e s 46, 47. Phlyctochytrium sp. (dentiferum?). F i g . 46. Plane view of b i p a r t i t e teeth. F i g . 47. Plane view of three to four part teeth. Figure'48. Phlyc tochytrium sp. (reinboldtae?); s t a r - l i k e sporangium with broad subtending apophysis. Figures 49-51. Rhizidium sp. (nowakowskii?). F i g . 49. I n t e r b i o t i c sporangium. F i g . 50, 51. Discharging sporangium. F i g s . 52-54. Rhizophlyctis  h a r d e r i ^ V F i g . 52. Type II sporangium with three apophysate r h i z o i d a l axes. F i g . 53. Empty type II sporangium. F i g . 54. Type I sporangium with branched and twisted r h i z o i d s i n a bundle. F i g s . 46, 47, 50-53, xl325. F i g s . 48, 49, 5,4, x650. 90 91 On pollen: coastal western hemlock sites, 167, 182, 224, 262; flushes, 196, 208, 220, 259; a blanket bog s i t e , 207, and a marine high beach s i t e , 267. Thalli are composed of a spherical sporangium with a broad apical papillar pore and meagerly branched delicate rhizoids ( f i g . 22). Except for the extent of the rhizoidal system, this material agrees i n a l l morphological details with the type. Rhizophydium sphaerotheca Zopf. Abhandl. Naturforsch. Gesell. Halle, 17:92, p i . 2, fig s . 33-41, 1887. On pollen: disturbed sites, 163; beach intertidal sites, 165, 169, 245, 256, 273, 282; marine high beach sites, 170, 237, 284, 285; coastal western hemlock sites, 173, 182, 193, 224, 233, 238, 249, 262, 286, 287; headland sites, 176, 184; flushes, 197, 199, 200, 202, 204, 206, 208, 209, 219-221, 226, 228; a blanket bog s i t e , 203; t i d a l edge sites, 213, 215, 217, 243; a dune s i t e , 252 and a stream bank s i t e , 289. Sporangia 10-(18)-28p in diameter on pollen and 12-(27)-60p in diameter on MHU medium with one to five papillae, 3-l]pwide. One to five papillate pores occur in the sporangium wall distinctly double contoured. Rhizoids fine and much branched, usually arising directly from sporangium base but occasionally originating from the base of a strap-like structure ( f i g . 23). Motile zoospores 2.5-(3)-4p in diameter with one large refractive globule. This description is based on two axenic isolates (cul. nos. 14, 48). 92 52. Rhizophydium s t l p i t a t u m Sparrow. Trans. B r i t . Mycol. S o c , 40: 528, f i g . 1, A-C, 1957. On snakeskin: a c o a s t a l western hemlock s i t e , 173. Sporangia, 30-70y i n diameter, borne i n d i v i d u a l l y on a s t i p e l i k e r h i z o i d up to 200u long and 13u i n diameter w i t h up to e i g h t l a r g e p a p i l l a e . Zoospores 5-5.5u wide w i t h one or two s m a l l r e f r a c t i v e g l o b u l e s . Sporangia of t h i s m a t e r i a l are s m a l l e r than those described by Sparrow. A l l other morphological features of my fungus agree w i t h the type. 53. Rhizophydium transversum (Braun) Rabenhorst. F l o r a Euopaea algarum, 3:281, 1868. On'pollen: a c o a s t a l western hemlock s i t e , 262. 54. Rhizophydium u t r i c u l a r e Uebelmesser. Arch. f . M i k r o b i o l . , 25:314, f i g . 3, 1956. On p o l l e n : a beach i n t e r t i d a l s i t e , 264. 55. Septosperma r h i z o p h i d i i Whiffen. Mycologia, 34:552, f i g s . 28-52, 1942. On Rhizophydium sphaerotheca, Chytriomyces h y a l i n u s and C. poculatus: f l u s h e s , 204, 219, 221; a blanket bog s i t e , 207; a c o a s t a l western hemlock s i t e , 223; and a dune s i t e , 258. 93 Bias t o c l a d i a l e s 56. B l a s t o c l a d i e l l a b r i t a n n i e a Horenstein and Cantino. Trans. B r i t . Mycol. S o c , 44:197, p i . 15, f i g s , a-g, 1961. On snakeskin and p o l l e n : a c o a s t a l western hemlock s i t e , 181; a headland l o c a t i o n , 184; a f l u s h , 209; and a dune s i t e , 258. Morphological d e t a i l of t h i s m a t e r i a l agrees w i t h t h a t described by Willoughby (1959) and Horenstein and Cantino. U s u a l l y t h a l l i develop on snakeskin ( f i g . 25). O c c a s i o n a l l y , both a c t i v e and r e s t i n g sporangia ( f i g . 26) w i l l grow on p o l l e n . 57. Bias t o c l a d i e l l a spp. On snakeskin: a d i s t u r b e d s i t e , 162. 58. Catenaria a n g u i l l u l a e Sorokin. Ann. S c i . Nat. Bot., V I , 4:67, p i . 3, f i g s . 6-28, 1876. On snakeskin: headland s i t e s , 176, 184 and a f l u s h , 198. 59. Catenophlyctis v a r i a b i l i s ( K a r l i n g ) K a r l i n g . Amer. Jour. Bot., 52:134, f i g s . 1-12, 1965. On snakeskin: a marine high beach s i t e , 170; a t i d a l edge s i t e , 215; and a beach i n t e r t i d a l s i t e , 236. Hyphochytriales 60. Hypho chy t r ium catenoides K a r l i n g . Amer. Jour. Bot., 26:513, f i g s . 1-18, 1939. 94 In po l l e n : a headland s i t e , 176. Saprolegniales 61. Achlya spp. With p o l l e n : t i d a l edge s i t e s , 215, 217. Species of this genus were observed growing i n plates heavily duated with p o l l e n . Oogonia may be ornamented ( f i g . 24). 62. Schizochytrium aggregatum Goldstein and Belsky. Amer. Jour. Bot^'*« '51: 71- 7 8 F i g s / 1-16,1964. On p o l l e n : beach i n t e r t i d a l s i t e s , 166, 169, 178, 210, 236, 240, 244, 245, 253, 264, 265; a coastal western hemlock s i t e , 168; t i d a l edge s i t e s , 179, 283; a headland s i t e , 184; marine high beach s i t e s , 237, 242; tide marsh s i t e s , 2 7 7 , 278; a tide affected stream s i t e , 288; and a sirearn bank s i t e , 289. 63. Thraus to chy trium aggregatum Ulken. Verb'ffentl. Inst. Meeresf orsch. Bremerh., 9:291, F i g s . 3-6, 1965. On p o l l e n : beach i n t e r t i d a l s i t e s , 230, 247; a t i d a l edge s i t e , 274; a t i d e marsh s i t e , 278; a tide affected stream s i t e , 288. 64. Thraus to chy t r ium mo tivum-Thraus tochytrium aureum complex. On p o l l e n : beach i n t e r t i d a l s i t e s , 210, 242, 273; a tide marsh s i t e , 277; and a tide affected stream s i t e , 288. Specimens showing morphological features of either or both Thraustochytrium motivum Goldstein and Thraustochytrium aureum Plate X I I I — F i g u r e 55. Rhizophlyctis harder1; type I sporangia with branched a p i c a l r h i z o i d s . Figures 56, 57. Phlyctochytrium sp. F i g . 56. Angular sporangium with four v i s i b l e discharge p a p i l l a e . F i g . 57. Thallus with branched apophysis. Figure 58. Rhizophydium  coronum; re s t i n g spore. Figure 59. R. chitinophilum; sporangium with two broad discharge p a p i l l a e on shrimp exoskeleton. Figure 60. R. parasiticum; sporangium on R. sphaerotheca. F i g . 55, x650. Figs. 56-60. xl325. 96 97 G o l d s t e i n are considered components of the T. motivum-T. aureum complex as d e l i n e a t e d by Booth and M i l l e r (1968). 65. Thraustochytrium m u l t i r u d i m e n t a l e G o l d s t e i n . Amer. Jour. Bot., 50: 273, f i g s . 12-21, 1963. On p o l l e n : a beach i n t e r t i d a l s i t e , 256 and a stream bank s i t e , 289. 66. Thraustochytrium pachydermum Scholz. Arch. f . M i k r o b i o l . , 29:361, f i g s . 3, 4, 1958. On p o l l e n : a marine hig h beach s i t e , 237; a t i d e marsh s i t e , 279 and a t i d e a f f e c t e d stream s i t e , 288. 67. Thraustochytrium p r o l i f e r u m Sparrow. B i o l . B u l l . , 70:258, f i g s . 22-28, p i . 1, f i g . 2, 1936. On p o l l e n : a beach i n t e r t i d a l s i t e , 210 and a marine high beach s i t e , 242. 68. Thraustochytrium roseum G o l d s t e i n . Mycologia, 55:801, f i g s . 1-14, 1964. On p o l l e n : beach i n t e r t i d a l s i t e s , 164, 165, 169, 191, 210, 230, 235, 236, 240, 247, 268; c o a s t a l western hemlock s i t e s , 168, 234; marine high beach s i t e s , 237, 242, 261, 267, 270; a beach r i s e r , 241; a t i d a l edge s i t e 243; t i d e marsh s i t e s , 278, 279; a t i d e a f f e c t e d stream s i t e , 288 and a stream bank s i t e , 289. Map I I I — F i g u r e 61. Queen Charlotte Islands. Approximate c o l l e c t i n g areas indicated by dark triangles:; 99 1 0 0 Lagenidiales 69. Lagenidium humanum Ka r l i n g . Mycologia, 39:225, f i g s . 1-39, 1947. On snakeskin: a coastal western hemlock s i t e , 224 and a dune s i t e , 252. 70. Lagenidium pygmaeum Zopf. Abhandl. Naturforsch. G e s e l l . H a l l e , 17: 96, p i . 1, f i g s . 29-31, p i . 2, f i g s . 1-12, 1887. In pol l e n : a disturbed s i t e , 162; a coastal western hemlock s i t e , 175 and a headland s i t e , 185. Peronosporales 71. Pythium ir r e g u l a r e Buisman. Meded. phytopath. Lab. W i l l i e Commelin Scholten, 11:38, f i g s . 9-11, 1927. With p o l l e n and vegetable debris: flushes, 200, 209, 219, 229. 72. Pythium spp With p o l l e n and vegetable debris: flushes, 220, 222. 101 Table I I . S i t e name and number, l a t i t u d e and longitude and h a b i t a t type. Moresby I s l a n d 1. South Sandspit (53°14.0»N., 132°05.0'W.) S i t e No. 162 163 2. Tasu Creek (52°51.6'.N., 132°05.0'W.) S i t e No. 164 165 166 167 168 169 3. Blunt P t . (52°49.4'N. , 132°02.8