N o v a Hedwigia
56
1—2
1—33
Stuttgart, Februar 1993
The freshwater Ascomycetes
by
CA.
Shearer
Department of Plant Biology, University of Illindi:
U r b a n a , Illinois 61801, U . S . A .
/ '
W i t h 7 figures and 3 tables
;
Shearer, C A . (1993): The freshwater Ascomycetes. - N o v a Hedwigia 56: 1-33.
Abstract: The freshwater Ascomycetes (exclusive of yeasts and lichenized forms) are a taxonomically diverse assemblage of species with representatives i n the Plectomycetes, Discomycetes, Pyrenomycetes, and
Loculoascomycetes. Over 200 species have been reported from freshwater habitats in association with
aquatic and wetland macrophytes and/or as saprophytes of autochthonous and allochthonous plant
debris. Various degrees of morphological adaptation to aquatic habitats are represented, ranging from
adaptation of both teleomorph and anamorph, teleomorph alone, or anamorph alone. Numerous species
show no obvious morphological adaptations at all. Definition, study techniques, systematics, geographical distribution, ecology, and evolution of the freshwater Ascomycetes are discussed.
Introduction
A l t h o u g h over 200 ascomycete t a x a (exclusive o f yeasts a n d l i c h e n i z e d f o r m s ) have
been reported f r o m freshwater habitats ( T a b l e 1), this g r o u p o f f u n g i has received
m u c h less a t t e n t i o n t h a n the w e l l - k n o w n m a r i n e A s c o m y c e t e s ( K o h l m e y e r a n d K o h l meyer 1979, K o h l m e y e r a n d V o l k m a n n - K o h l m e y e r 1991) a n d the freshwater H y phomycetes (Webster a n d Descals 1981). Several reasons m a y a c c o u n t f o r this l a c k
o f a t t e n t i o n . T h e freshwater A s c o m y c e t e s ( F W A ) c o m p r i s e a t a x o n o m i c a l l y diverse
g r o u p w i t h representatives i n a w i d e variety o f families a n d orders (See Systematics
Section). T h i s diversity a n d the absence o f t a x o n o m i c m o n o g r a p h s a n d keys f o r the
i d e n t i f i c a t i o n o f F W A present a f o r m i d a b l e barrier to the r a p i d i d e n t i f i c a t i o n o f
species for even the m o s t s k i l l e d ascomycete systematist. Since m a n y F W A also o c c u r i n terrestrial habitats o r are i n genera w i t h terrestrial representatives, k n o w l e d g e
o f terrestrial t a x a is i m p o r t a n t to the successful i d e n t i f i c a t i o n o f freshwater t a x a .
A d d i t i o n a l l y , i n recent years, freshwater m y c o l o g y has been d o m i n a t e d b y the study
o f aquatic H y p h o m y c e t e s . G e n e r a l l y such studies have c o n c e r n e d f u n g i o n d e c a y i n g
deciduous leaves f r o m w h i c h ascomycete fructifications are largely absent (Shearer
1992). T h u s F W A m a y have escaped detection i n the m a j o r i t y o f freshwater f u n g a l
studies. I n spite o f these p r o b l e m s , a considerable b o d y o f k n o w l e d g e about F W A
exists ( D u d k a 1985). T h i s paper presents the current state o f o u r k n o w l e d g e o f F W A
w i t h the i n t e n t i o n o f s t i m u l a t i n g further research o n their life histories, e c o l o g y , geo g r a p h i c a l d i s t r i b u t i o n a n d systematics.
0029-5035/93/0056-0001
©
$8.25
1993 J . Cramer in der CebrUder Borntraeger
Verlagsbuchhandlung, D-1000 Berlin • D-7000 Stuttgart
1
Definition
S o m e o f the same p r o b l e m s that o c c u r i n d e f i n i n g v a s c u l a r plants as aquatic o r terrestrial ( C o r r e l l a n d C o r r e l l 1975) a p p l y to f u n g i . These p r o b l e m s stem f r o m the difficulty i n d e t e r m i n i n g w h a t habitats s h o u l d be c o n s i d e r e d aquatic a n d h o w far i n t o
the interface zones between aquatic a n d terrestrial habitats s h o u l d be considered
a q u a t i c . T h e o u t l i n e o f types o f a q u a t i c habitats presented b y C o r r e l l a n d C o r r e l l
(1975) w h i c h is a m o d i f i c a t i o n o f a n earlier one b y M a s o n (1957) has been used for
the purpose o f this r e v i e w . U n l i k e vascular h y d r o p h y t e s , h o w e v e r , presence i n these
aquatic habitats, a l o n e , m a y n o t be a p p r o p r i a t e to define a n ascomycete as a " f r e s h water A s c o m y c e t e " . M a n y A s c o m y c e t e s r e p o r t e d f r o m freshwater habitats ( T a b l e
1) have also been r e p o r t e d f r o m terrestrial habitats, a n d it is not k n o w n whether
their occurrence i n freshwater is s i m p l y f o r t u i t o u s . S o m e A s c o m y c e t e s f o u n d i n
freshwater m a y be there n o t because they g r o w a n d r e p r o d u c e w h i l e s u b m e r g e d , but
because they enter water o n a l l o c h t h o n o u s substrata. C o n s t a n t recruitment o f such
species w o u l d be necessary to m a i n t a i n their p o p u l a t i o n s i n water. A s c o m y c e t e s
f o u n d by p l a t i n g o r i n c u b a t i o n o f substrata m a y be present i n water as spores a n d
g r o w a n d develop o n l y w h e n they are r e m o v e d f r o m water. F u r t h e r , substrata o n
w h i c h F W A o c c u r m a y be alternately s u b m e r g e d a n d exposed. L i v i n g a n d s t a n d i n g
dead emergent m a c r o p h y t e s m a y be m o s t l y submerged d u r i n g wet times o f the year
a n d entirely exposed d u r i n g d r y p e r i o d s . S u b m e r g e d a l l o c h t h o n o u s substrata m a y
be deposited o n l a n d d u r i n g f l o o d s a n d r e m a i n there u n t i l they are a g a i n washed
i n t o water. I n m a n y emergent h y d r o p h y t e s w h i c h serve as substrata for F W A , the
upper p l a n t parts are never submerged u n t i l w e l l after the death o f the p l a n t . W o o d y
debris i n streams is often o n l y p a r t i a l l y s u b m e r g e d . W h e t h e r A s c o m y c e t e s s h o u l d
be considered a q u a t i c w h e n their substrata are submerged a n d terrestrial w h e n they
are n o t is p r o b l e m a t i c .
P a r k (1972) cogently discussed the p r o b l e m o f d e f i n i n g aquatic h e t e r o t r o p h i c m i c r o o r g a n i s m s a n d d i v i d e d t h e m i n t o t w o categories: indwellers a n d i m m i g r a n t s . I n d w e l lers are f u l l y adapted to freshwater, i.e. they c a n m a i n t a i n their b i o m a s s at a given
site f r o m year to year u s i n g substrata a n d nutrients at that site. A c o n s i d e r a b l e n u m ber o f F W A f a l l i n t o this category, e.g., Pseudohalonectria
lignicola,
Aniptodera
rosea, Nais inornata. I m m i g r a n t s s h o w v a r y i n g degrees o f a d a p t a t i o n to a n d e c o l o gical activity i n aquatic habitats, but they a l l require i m m i g r a t i o n to m a i n t a i n their
presence i n water. I m m i g r a n t s m a y range f r o m those that are very active e c o l o g i c a l l y to those that are n o t active at a l l . T h e r e m a y be a t h i r d category, a m p h i b i o u s species. A m p h i b i o u s species m a y o c c u r i n the interfaces between l a n d a n d water, e.g.,
edges o f p o n d s , lakes, f l o o d p l a i n s , b a c k w a t e r s o f rivers o r i n ephemeral a q u a t i c
habitats such as t e m p o r a r y streams, seasonal p o n d s , a n d drainage ditches. A m p h i b i o u s A s c o m y c e t e s w o u l d be a d a p t e d to f l u c t u a t i n g water levels. T h e r e have been
so few e c o l o g i c a l studies o f F W A that it is i m p o s s i b l e at this t i m e to classify a d e q u a tely most o f the F W A as i n d w e l l e r s , i m m i g r a n t s or a m p h i b i a n s , let alone speculate
o n their e c o l o g i c a l a c t i v i t y .
D e f i n i t i o n b y t a x o n o m y is not useful because the F W A are a t a x o n o m i c a l l y diverse
g r o u p w i t h representatives i n 15 different orders. F o r the purposes o f this r e v i e w ,
2
Table I. Ascomycetes reported from submerged and/or partially submerged substrata.
Species
Substratum
Locality
sewage sludge
Japan
PLECTOMYCETES
Corynascella
& Siichi
inquinata Udagawa
Udagawa and Siichi
1979
organic debris
Ireland
Eurotium sp.
beech wood
England
Eaton and Jones
1971b
Eurotiaceae gen. n. 4084
vegetation
USA,
Christiansen and
Whittingham 1965
Emericellopsis
terricola
van Beyma
Pseudeurotium multisporum
(Saito & Hinoura) Stolk
WI
beech wood
England
Eaton
organic debris
Ireland
Park
Subbaromyces aquaticus
Manoch. & Ramarao
maize baits
India
Manocharachary and
Ramarao 1974
Talaromyces flavus (Klocher)
Stolk & Samson var. flavus
leaf l i t t e r
USA,
creek sand
Australia
alder, willow and
Corylus twigs
England, France,
Sweden
Pseudogymnoascus roseus R a i l l o
(sub Pseudogymnoascus vinaveus
1972
1972
Raillo)
IL
Shearer and Crane
1986
DISCOMYCETES
Apostemidium fiscellum
Karst.
(Karst.)
Graddon 1965
alder twigs
Hungary
Revay and Gbnczol
Apostemidium fiscellum
(Karst.) Karst.
var. submersum Graddon
willow twigs
England, Germany,
Sweden
Graddon 1965
Apostemidium sp.
birch twig
Aquadiscula
appendiculata
Shearer & Crane
Acer rubrum L,
leaves
Belonopsis
Phragmites
excelsior
(Karst.) Rehm
leaves
Phragmites
Belonopsis
mediel la (Karst.) Aebi
retincola
(Rbh.) LeGal. & Hang.
Lamore and Goos
1978
USA,
IL
Shearer and Crane
1985, 1986
Engl and
Phragmites australis
(Cav.)
T r i n . ex Steud. stalks
Phragmites australis
Belonopsis
RI
Finland
Phragmites
Belonopsis hydrophila (Karst.) Nannf.
[sub Mollisi a arundinaceae (DC.) Phi 11.]
USA,
Phragmites,
Schoenoplectus
lacustris
L.
Phragmites, Carex,
Typha
Bisporella
citrina (Batsch:Fr.)
Korf & Carpenter
1990
Corner 1935 (not seen,
c i t e d in Nannfeldt 1985)
Ingold 1954,
USSR
Oudka 1963
1955
England
Apinis et a l . 1972a
Czechoslovakia,
Denmark, England,
Finland, France,
Germany, Poland
Scotland, Sweden
Switzerland
Nannfeldt
1985
Engl and, Finland,
France, Germany,
Ireland, Sweden,
Switzerland
Nannfeldt
1985
England, Finland,
France, Germany
Ireland, Sweden,
Switzerland
Nannfeldt
1985
USA,
Shearer and Crane
1986
IL
(Rehm.}
Eriophorum vaginatum I .
Juncus filiformis
L.
Austria
Magnes and Hafel1ner
1991
Cistella
graminicola
Raitv.
(Raitv.)
Carex rostrata Stokes
Equisetum fluviatile
L.
em. Ehrh.
Austria
Magnes and Hafellner
1991
Coleosperma lacustre
Ingold
Schoenoplectus
stalks
Engl and
Ingold 1954
Austria
Magnes and Hafellner
1991
Brunnipila cat yciaides
Baral
Coronellaria
Karst.
caricinella
(Karst.)
lacustris
Carex nigra (L.) Reichard,
Carex echinata Hurr.
3
Crocicreas megalosporum (Rea.)
Carpent. var. megalosporum
Carpent.
Carex
Cudoniella clavus (Alb. & Schw.:Fr.)
Dennis
Dasyscyphus controversus
(Cooke) Rehm
Dasyscyphus nudipes (Fuckel) Sacc.
Dasyscyphus c f . virgineus
S. F. Gray
rostrata
Austria
Magnes and Hafellner
1991
ash, alder, oak
twigs
Engl and
Willoughby and
Archer 1973
Phragmites
australis
stems
England
Taligoola et a l .
1972
Phragmites
stems
England
Apinis et a l .
1972a
leaves, rush stems,
twigs
Germany
Eckel and
Hirsch 1979
Phragmites
australis
stems
England
Taligoola et a l .
1972
USA, RI
Lamore and Goos 1978
australis
(Batsch)
Diplonaevia
emergens (Karst.) Hein
Juncus
filiformis
Eriophorum
angustifolium
Honck.
Austria
Magnes and Hafellner
1991
Diplonaevia
seriata
Carex
Austria
Magnes and Hafellner
1991
(Libert) Hein
Gorgoniceps
boltonii
(Phi11.) Dennis
(sub Pocillum boltonii
Graddonia coracina
Phill.)
rostrata
Equisetum
stems
fluviatile
England
Ingold 1954
Equisetum
fluviatile
Austria
Magnes and Hafellner
1991
England
Dennis 1978, Webster
pers. comm.
Dennis
Helotium
citrinulum
Karst.
Phragmites
stems
australis
England
Taligoola et a l .
1972
Phragmites
leaves
australis
England
Apinis et a l .
1972a
Phragmites
stems
australis
USSR
Dudka 1963
England
Taligoola et a l .
1972
USA, RI
Lamore and Goos
1978
England
Abdullah and
Webster 1983
catkins of Alnus,
spine of Castanea
sativa M i l l e r
England
Descals and
Webster 1976
twigs
USA, RI
Lamore and Goos 1978
oak twigs
England
Webster et a l . 1990
alder twigs
England
Shearer and
Webster 1991
Hymenoscyphus africanus
Descals, Fisher & Webster
alder twigs
England
Descals et a l . 1984
Hymenoscyphus equisetinus
(Velenovsky) Dennis
Equisetum
Austria
Magnes and Hafellner
1991
Hymenoscyphus foliicola
Abdullah, Descals & Webster
Acer pseudoplatanus I.
twigs
England
Abdullah et a l . 1981
Hungary
Revay and
Gonczol 1990
Malawi
Fisher & Spooner 1987
Switzerland
Fisher and
Webster 1983
USSR
Dudka 1963
England
Abdullah et a l . 1981
Helotium sp.
Hyalinia
sp
maple twig
Hyaloscypha
lignicola
Abdullah & Webster
twigs
Hyaloscypha zalewskii
Descal s & Webster
Hyalotricha
sp.
Hydrocina chaetocladia
Scheuer
fluviatile
alder twigs
Hymenoscyphus malawiensis
Fisher & Spooner
plant debris
Hymenoscyphus paradoxus
Fisher and Webster
willow twigs
Hymenoscyphus phyllophilus
(Desm.) 0. Kuntze
skeletal leaves
Hymenoscyphus splendens
Abdullah, Descals & Webster
beech cupules
4
Hymenoscyphus tetracladius
Abdullah, Descals & Webster
beech and willow
leaves
England
Abdullah et a l . 1981
Hymenoscyphus
Tubaki
twigs
Japan
Tubaki 1966
alder twigs
Hungary
Revay and
Gonczol 1990
Carex nigra
Austri a
Magnes and Hafellner
1991
Austria
Magnes and Hafellner
1991
Eriphorum vaginatum,
Carex rostrata,
C. paupercula Michx.
subsp. irrigua (Wahl.)
A. & D. Love, C. nigra,
Juncus
filiforme
Austria
Magnes and Hafellner
1991
Carex
Austria
Magnes and Hafellner
1991
Austria
Magnes and Hafellner
1991
varicosporoides
Hymenoscyphus sp.
Hypoderma alpinum Spooner
Hysteronaevia
advena (Karst.) Nannf.
Hysteronaevia
olivacea
Hysteropeziza
pusilla
(Mout.) Nannf.
Hein & Scheuer
Hysteropezizella
Nannf.
diminuens (Karst.)
Hysteropezizella
Nannf.
fuscella
(Karst.)
Eriophorum
angustifolium
rostrata
Carex rostrata
C. paupercula Michx.
subsp. irrigua (Wahl.)
A. & D. Love
Austria
Magnes and Hafellner
1991
Acer pseudoplatanus
twig
Engl and
Abdullah and
Webster 1981
Eriophorum
Honck.
Austria
Magnes and Hafellner
1991
Hysteropezizella
phragmitina
(Karst. & Starb.) Nannf.
Carex rostrata
Lambertella
tubulosa
Abdullah & Webster
Lachnum imbecille
Karst.
Loramyces juncicola
Magnes and Hafellner
1991
Carex nigra, C. rostrata
angustifolium
USA, MA
Weston 1929
dead stalks of
Scirpus,
Eleocharis,
England
Ingold 1955
Equisetum, Juncus
USA, RI
Digby and Goos
1987
England
Ingold and
Chapman 1952
Weston
Loramyces macrospora
Ingold & Chapman
internodes of
Equisetum fluviatile
Miladina
lechithina
(Cooke) Svrcek
bark o f Crategus,
Alnus wood
Engl and
Descals and
Webster 1978
Mitrula
alba W.G.Sm.
leaves
Engl and
Cannon et a l . 1985
Mitrula
paludosa Fr.
leaves, wood
Engl and
Dennis 1978
Mollisia
arundinacea
Engl and
Apinis et a l . 1972a
USSR
Dudka 1963
Finland
Fisher and
Webster 1983
(D.C.) P h i l l .
Phragmites
leaves
L.
australis
Mollisia
cinerea
(Batsch) Karst.
Mollisia
cinerea (Batsch:Fr.) Karst.
var. minutella Sacc.
[sub Mollisia minutella (Sacc.) Rehm]
Mollisia
gigantea
Fisher & Webster
Typha latifolia
Picea
I.
twigs
Mollisia melaleuca
(Fr.: Fr.) Sacc.
wood
USSR
Mollisia
uda (Pers.:Fr.) G i l l .
wood
England
Shearer and
Webster unpublished
Mollisia
sp.
twigs
Engl and
Webster 1961
Mollisia
sp.
skeletonized
oak leaves
Engl and
Webster and
Descals 1979
Mollisia
spp.
England
Willoughby and
Archer 1973
USA, RI
Lamore and Goos
1978
Mollisia
spp. 1, 2, and 3
oak, ash, alder twigs
twigs
5
alder, ash, beech
and oak twigs
Hungary
Revay and Gbnczol
1990
Carex rostrata, Eriophorum
vaginatum, Juncus
filiformis
Austria
Magnes and Hafellner
1991
Nimbomollisia macrospora
(Karst.) Nannf.
Carex rostrata
Austria
Magnes and Hafellner
1991
Nimbomollisia
melatephroides
(Rehm) Nannf.
Carex rostrata
Austria
Magnes and Hafellner
1991
Niptera excelsior
(Karst.) Dennis
[sub Belonium excelsium (Karst.) Boud.]
Phragmites stalks
England
Ingold 1954
Phragmites culms
England
Dennis 1978
alder and oak twigs
England
Shearer and
Webster 1991
Schoenoplectus
Phragmites
Scandinavia
Fries 1849 (as c i t e d in
Nannfeldt 1985)
Mollisia
sp.
Nimbomollisia
Nannf.
Niptera
eriophori
lacustris
(Opiz)
(Fr.) Fr.
Scirpus,
Niptera melanophaea Rehm,
Niptera pilosa
(Crossland)
Phragmites
Schoenoplectus
Boudier
pulla
( P h i l l . & Keith) Boud.
lacustris
Carex
Schoenoplectus
Niptera
lacustris,
lacustris
Carex, Eleocharis,
Juncus
Phragmites,
Phalaris
England
Dennis 1978
USSR
Dudka 1963
U.K.
Dennis 1978
USSR
Dudka 1963
England
Dennis 1978
Typha
Italy
Nannfeldt 1985
Carex rostrata
Switzerland
Muller et a l . 1979
Carex rostrata
Austria
Magnes and Hafellner
1991
Eriophorum sp
USA, AK
Shearer unpublished
twigs
USA, RI
Lamore and Goos 1978
stream foam
Engl and
Webster and
Descals 1979
armeniaca Kirschst.
twigs
USA, IL
Shearer and
von Bodman 1983
Pachyella babingtonii (Berk.) Boud.
[sub Pachyella depressa ( P h i l l . ) Boud.]
twigs
England
Ingold 1954
Peziza sp.
beech, Scots pine
wood
England
Eaton and Jones
1971b
Pezizella
Phragmites
australis stems
England
Taligoola et al
1972
Pezoloma rhodocarpa Fisher & Spooner
plant debris
Malawi
Fisher and Spooner 1987
Pezoloma sp.
stream foam
England
Webster and
Descals 1979
Phaeohelotium sp
twigs
USA, IL
Shearer and
von Bodman 1983
wood
USSR
Dudka 1963
twigs
France
G i l l e t 1879
Carex rostrata
Austria
Magnes and Hafellner
1991
Scutellinia
scutellata
(L.:Fr.) Labotte
[sub Lachnea scutellata
(L.) G i l l . ]
wood
USSR
Dudka 1963
Scutomollisia
morvernensis Grad.
Carex rostrata, Equisetum
fluviatile,
Eriophorum
angustifolium,
Glyceria
fluitans
(L.) R. Br.,
Juncus filiformis
I.
Austria
Magnes and Hafellner
1991
Scutomollisia
punctum (Rehm) Nannf.
Carex rostrata
Austria
Magnes and Hafellner
1991
Equisetum fluviatile
Austria
Magnes and Hafellner
1991
Niptera
tephromelas (Pass.) Nannf.
Obtectodiscus
aquaticus
Muller, Petrini & Samuels
Orbilaster
Orbilia
sp.
sp.
Orbiliella
Plicaria
sp
violaceo-nigra
Psilopezia
aurantiaca
Pyrenopeziza huetteri
& Hafellner
Stamnaria persoonii
Rehm
Gill.
Magnes
(Pers.: Fr.) Fckl.
6
Tapesia knieffii
(Wallr.) J. Kunze
(sub Belonidium rhenopalaticum Rehm)
Phragmites stalks
England
Ingold 1954
Tapesia sp.
Carex
Austria
Magnes and Hafellner
1991
Bohemi a
Vacek 1949
Schoenoplectus
lacustris
and Typha leaf bases
Engl and
Ingold 1954
Carex
Austria
Magnes and Hafellner
1991
wood
Australia
Beaton and Weste 1977
twigs
Sweden, Norway
Schumacher 1976
rostrata
Thecotheus
rivicola
(Vacek) Kimbrough & P f i s t e r
Trichobelonium
Trichopezizella
hystricula
Vibrissea
guestphalicum
Rehm
nidulus var.
(Karst.) Haines
bicolor
Beaton & Weste
Vibrissea decolorans
Sanchez & Korf
(Saut.)
rostrata
eucalyptus logs
Australia
Beaton and Weste 1976
Vibrissea filisporia
(Bon.)
Korf & Sanchez f.bouderi
willow and alder
twigs
Norway
Schumacher 1976
Vibrissea filisporia
(Bon.)
Korf & Sanchez f .
filisporia
willow twigs
Norway
Schumacher 1976
oak wood
Scotland
Graddon 1965
oak wood
Engl and
Hamad and Webster 1987
twigs
Engl and
Ingold 1954
willow twigs
Belgium, England,
France, Holland,
Sweden
Graddon 1965
Vibrissea
leptospora
(Berk. & Br.) P h i l l .
[sub Apostemidium leptospora
(Berk. & Br.) Boud.]
Prunus, Rosa, Ribes,
Sa 1 ix
England, Scotland
Sweden, Wales
Graddon 1965
Vibrissea
sporogyra
(Ingold) Sanchez
(sub Apostemidium sporogyrum Ingold)
dead stalks of
Equisetum, Carex,
Eleocharis
Engl and
Vibrissea
melanochlora
Beaton & Weste
eucalyptus logs
Australia
Beaton and Weste
1976
Vibrissea
norvegica
(Gremmen) Sanchez
Phragmites
australis
Norway
Sanchez 1967
Vibrissea
obconica
(Kanouse) Sanchez f . boudieri
alder twigs
Scandinavia
Schumacher 1976
Vibrissea
tasmanica
Rodway
eucalyptus wood
Australia
Beaton and Weste
1976
Vibrissea truncorum
(Alb. & Schwein.) Fr.
maple, alder, birch
willow, juniper branches
Scandinavia
K j u l l e r 1960
Scandinavia
K j u l l e r 1960
USA,
Shearer and von Bodman
1983
Vibrissea
dura Weste
Vibrissea
flavovirens
(Pers.:Fr.) Korf & Dixon
(sub Apostemidium torrenticola
Graddon)
Vibrissea
guernisacii
Crouan & H. Crouan
[sub Apostemidium guernisaci
(Cr.) Boud.]
twigs
Engl and
Vibrissea
vibrisseoides
(Peck) K j u l l e r
Willoughby and
Archer 1973
Ingold 1954
PYRENOMYCETES
IL
Aniptodera
chesapeakensis
Shearer & M i l l e r
wood
wood
USA, IL
Shearer and Crane 1986
Aniptodera
fusiformis
wood
USA, IL
Shearer 1989b
Aniptodera
limnetica
Aniptodera
lignatilis
Shearer
Shearer
Hyde
Aniptodera margarition
Arnium apiculatum
( G r i f f i t h s ) Lundq.
Shearer
wood
USA, IL
Shearer 1989b
balsa wood
Japan
Minoura and Muroi
1972
Hyde unpublished
wood
wood
USA, IL,
England
Shearer 1989b
Equisetum
fluviatile
USA, IL
Shearer unpubli shed
Equisetum
fluviatile
USA, IL
Shearer et a l . 1980
7
Bombardia arachnoidea
(Niessl) Cain
balsa wood
USA, MO
Shearer 1972
Calonectria
wood
USA, RI
Lamore and Goos 1978
beech wood
England
Eaton and Jones 1971a
beech wood
England
Eaton 1972
wood
USA, IL
Shearer and Crane
1986
alder and beech twigs
Hungary
Revay and Gonczdl 1990
alder, ash, oak
and willow twigs
England
Willoughby and Archer
1973
twigs
USA, IL
Shearer and von Bodman
1983
balsa wood
USA, IL
Shearer and Crane 1986
alder & beech twigs
Hungary
Revay and Gonczdl 1990
ash twigs
England
Ingold 1951
England
Apinis et a l . 1972b
sp.
Ceratosphaeria lampadophora
(Berk. & Br.) Niessl
Ceratosphaeria
pusilla
(Fuckel) Sacc.
Ceratostomella sp.
Cercophora sp.
Ceriospora
caudae-suis
Ceriosporopsis
Ingold
sp.
Phragmites
culms
Chaetomium aureum Chivers
wood
Chaetomium cochlioides
Phragmites
Pall.
australis
australis
USA, IL
Shearer and Crane 1986
England
Apinis et a l . 1972a
balsa wood
USA, IL
Shearer and Crane 1986
wood
USA, IL
Shearer and Crane 1986
Chaetomium elatum
Kunze & Schmidt:Fr.
beech wood
England
Eaton 1972
Chaetomium globosum
Kunze:Fr.
beech and Scots
pine wood
England
Eaton and Jones 1971a
Chaetomium funicolum
Cooke
beech wood
England
Eaton 1972
balsa wood
USA, IL
Shearer and Crane 1986
beech wood
England
Davis and Jones
unpublished
Chaetomium seminudum Ames
leaf l i t t e r
USA, IL
Shearer and Crane 1986
Chaetomium trilaterale
balsa wood
USA, IL
Shearer and Crane 1986
Engl and
Apinis et a l . 1972a
Chivers
Chaetomium spp.
Phragmites
leaves
australis
wood
USA, RI
Lamore and Goos 1978
balsa wood
USA, MD
Shearer 1972
wood
USA, IL
Shearer and Crane 1986
Coniochaeta leucoplaca
(Berk. & Rav.) Cain
wood
USA, IL
Shearer and Crane 1986
Coniochaeta
lignaria
(Grev.) Cooke
[sub Coniochaeta discospora
(Auers. ex Niessl) Cain]
beech wood
Engl and
Eaton 1972
Coniochaeta
velutina
(Fuckel) Munk
wood
USA, IL
Shearer and von Bodman
wood
USA, IL
Shearer and Crane 1986
bog vegetation
USA, WI
Christensen and
Whittingham 1965
wood
Sudan, England
Fisher and Petrini 1983
Scots pine wood
England
Eaton and Jones 1971b
balsa wood
Japan
Minoura and Muroi 1978
twigs
USA, IL
Shearer and von Bodman
1983
Coniochaeta kellermani
( E l l . & Ev.) Munk
Coniochaeta
sp. 4022
Chaetosphaeria anglica
Fisher & Petrini
Clypeosphaeria mamillana
(Fr.) Lambotte
Debaryella
gracilis
Debaryella sp.
1983
Munk
8
Ditopella
sp.
Scots pine wood
England
Eaton and Jones 1971b
Gaeumannomyces graminis
(Sacc.) v. Arx & O l i v i e r
Carex
Austria
Magnes and Hafellner
1991
Gnomonia papuana
Sivanesan & Shaw
1 eaves
Papua, New Guinea
Sivanesan and Shaw
1977
Gnomonia sp.
wood
USA, IL
Shearer and Crane 1986
Engl and
Eaton and Jones 1971a
Griphiosphaeria
corticola
(Fuckel) v. Hohnel
Halosarpheia
lotica
Halosarpheia
parva Shearer
Shearer
Halosarpheia
retorquens
Shearer & Crane
Halosarpheia viscosa (Schmidt)
(Schmidt) Shearer & Crane
ex Kohlm. & Vol km.-Kohlm.
Hypoderma scirpinum DC.
rostrata
Scots pine wood
USA, WI
Shearer 1984
USA, IL
Shearer unpublished
USA, IL
Shearer and Crane 1980
wood
USA, IL
Shearer and von Bodman
1983
wood
USA, IL
Shearer and Crane 1986
twigs
USA, IL
Shearer and von Bodman
1983
twigs
monocot stem
wood
Schoenoplectus
stems
lacustris
Engl and
Ingold 1954
Schoenoplectus
lacustris
USSR
Dudka 1963
Phragmites
culms
australis
England
Apinis et a l . 1972b
Phragmites
stems
australis
England
Taligoola et a l . 1972
twigs
USA, IL
Shearer and von Bodman
1983
Lasiosordaria
lignicola
(Fuckel) Chen.
beech and pine
wood
England
Davis and Jones
unpublished
Lejosphaerella
leonensis
Fisher & Petrini
wood
Africa
Fisher and Petrini 1983
Lopbiotrema
Schoenoplectus
Lasiosphaeria
Luttrellia
spp.
culmifraga Speg.
estuarina
Shearer
USSR
Dudka 1963
balsa wood
lacustris
USA, MD
Shearer 1978
twigs
USA, IL
Shearer and von Bodman
1983
balsa wood
USA, IL
Shearer and Crane 1986
Melogramma sp.
beech, Scots
pine wood
Engl and
Apinis et a l . 1972b
Micronectriella
Phragmites
culms
australis
England
Eaton and Jones 1971a
Phragmites
stems
australis
England
Taligoola et a l . 1972
Phragmites
stems
australis
England
Apinis et a l . 1972
beech, Scots
pine wood
England
Eaton and Jones 1971a
Shearer 1972
sp.
Nais inornata Kohlm.
Nectria
coccinea
balsa wood
USA, MD
twigs, balsa wood
USA, IL
Shearer and Crane 1978
balsa wood
Japan
Minoura and Muroi 1978
twigs
USA, IL
Shearer and von Bodman
1983
balsa wood
USA, IL
Shearer and Crane 1986
beech twigs
Hungary
Revay and Gbnczbl 1990
oak twigs
England
Shearer and Webster 1991
twigs
USA, IL
Shearer and von Bodman
1983
(Pers.:Fr.) Fr.
Nectria
Nectria
discophora
(Hont.) Mont.
haematococca Berk. & Br.
9
Nectria
lucidum Hohnel
twigs
Nectria
lugdunensis Webster
twigs
England
Webster 1959
oak, willow twigs
Engl and
Willoughby and Archer
1973
Nectria
penicillioides
Ranzoni
Nectria spp.
alder, oak twigs
England
Shearer and Webster 1991
maple leaves
USA, CA
Ranzoni 1956
beech wood
England
Eaton and Jones 1971b
twigs
USA, RI
Lamore and Goos 1978
alder and
beech twigs
Hungary
Revay and Gonczdl 1990
Austri a
Magnes and Hafellner
1991
USA, FL
Conway and Barr 1977
Nectriella
lacustris
(Kirsch.)
Magnes & Hafellner
Carex
Ophioceras dolichostomum
(Berk. & C u r t i s ) Sacc.
wood
Ophioceras
leptosporum
(Iqbal) Walker
(sub Gaumannomyces leptosporus
Iqbal)
Shearer and von Bodman
1983
rostrata
plant stalks
Engl and
Iqbal 1972
1 eaves
Papua, New Guinea
Shaw 1977
Shearer unpublished
Ophioceras
sp. 652
twigs
Panama
Ophioceras
sp. 837
twigs
USA,
IN
Shearer unpublished
Ophioceras
sp. 408
twigs
USA, IL
Shearer unpublished
beech and Scots
pine wood
England
Eaton and Jones 1971a
beech and Scots
pine wood
Engl and
Eaton 1972
Japan
Minoura and Muroi 1978
Phaeonectriella
lignicola
Eaton & Jones
Phomatospora aquatica
Minoura & Muroi
Phomatospora berkleyi
Sacc.
Phragmites
Carex
Phyllachora therophila
v. Arx & Muller
Physalospora
aquatica
(Desm.)
Ingold
australis
rostrata
Juncus
filiformis
Schoenoplectus
lacustris,
Typha angustifoh'a
Equisetum
fluviatile
USSR
Dudka 1963
Austria
Magnes and Hafellner
1991
Austria
Magnes and Hafellner
1991
England
Ingold 1955
Austria
Magnes and Hafellner
1991
Africa
Monod and Fisher 1983
Plagiosphaeria
nilotica
Monod & Fisher
twig
Platyspora
permunda (Cooke) Wehm.
Scirpus, Typha
USA, MN
Cavaliere 1975
Platyspora
planispora
Scirpus, Typha
USA, MN
Cavaliere 1975
Eaton 1971b
( E l l . ) Wehm.
Podospora setosa
(Winter) Niessl
beech wood
Engl and
beech wood
England
Eaton 1972
Pseudohalonectria
Shearer
adversaria
wood
USA, IL
Shearer 1989a
Pseudohalonectria
Shearer
falcata
wood
USA,
Shearer 1989a
Pseudohalonectria
lignicola
Minoura & Muroi
Pseudohalonectria
Shearer
longirostrum
IL
balsa wood
Japan
Minoura and Muroi 1972
wood
USA, MD
Shearer 1972
(as CS-182-1)
cottonwood,
cherry and
sycamore twigs
USA, IL
Shearer and von Bodman
1983
balsa wood
USA, IL
Shearer and Crane 1986
alder and beech twigs
Hungary
Revay and Gonczol 1990
wood
Panama
Shearer 1989a
10
Chile
Shearer 1989a
USA, IL
Shearer 1989a
Engl and
Webster and Hawksworth
1986
beech, Scots pine
and greenhart wood
England, Wales
Jones and Eaton 1969
sycamore twigs
USA, IL
Shearer and von Bodman
1983
Savoryella
verrucosa
Minoura & Muroi
balsa wood
Japan
Minoura and Muroi 1972
Schizothecium sp.
beech twig
Hungary
Revay and Gonczol 1990
Sillia
wood
Engl and
Eaton and Jones 1971a,b;
Eaton 1976
Pseudohalonectria
Shearer
lutea
Pseudohalonectria
Shearer
phialidica
Pyxidiophora
spinulo-rostrata
Webster & Hawksworth
Savoryella
lignicola
Jones & Eaton
ferruginea (Pers.:Fr.) Karst.
(probably an Ophioceras sp.)
Sordaria
sp. I
wood
USA, IL
Shearer and Crane 1986
Sordaria
sp. II
wood
USA, IL
Shearer and Crane 1986
Thielavia
sp.
Phragmites
stems
Engl and
Taligoola et a l . 1972
twigs
USA, IL
Shearer and von Bodman
1983
Zopfiella
latipes
(Lundq.) Halloch & Cain
balsa wood
USA, MD
Shearer 1972
balsa wood
USA, IL
Shearer and Crane 1986
Zopfiella
leucotricha
(Speg.) Lundq.
balsa wood
USA, MD
Shearer 1972
Zopfiella
lundqvistii
Shearer & Crane
balsa wood and
woody debris
USA, IL
Shearer and Crane 1978
balsa wood and
woody debris
USA, IL
Shearer and Crane 1986
twigs
USA, IL
Shearer and von Bodman
1983
wood
Australia
Zopfiella
sp.
australis
LOCULOASCOMYCETES
Ascagilis
bipolaris
Hyde
Bricookea sepalorum (Vleugel)
Barr
Juncus
Buergenerula biseptata
(E. Rostr.) H. Sydow
Carex
Byssothecium flumineum
Crane, Shearer & Huhndorf
wood
filiformis
rostrata
Hyde in press
Austria
Magnes and Hafellner
. 1991
Austria
Magnes and Hafellner
1991
USA, IL
Shearer and von Bodman
1983
(as Leptosphaeria
wood
USA,
Caryospora
callicarpa
(Curry) Nitschke ex Fuckel
wood
USA, IL
Clathrospora tirolensis
ex 0. Eriksson
Carex
Rehm
Delitschia
bispora
Eaton & Jones
Didymella equisetina
Petrak
Didymella glacialis
Didymella spp.
(H. Sydow)
Rehm
rostrata
IL
sp. 1)
Crane et a l . in press
Shearer unpublished
Austria
Magnes and Hafellner
1991
beech wood
Engl and
Eaton and Jones 1970
beech wood
England
Eaton and Jones 1971b
beech wood
England
Eaton 1972
Austria
Magnes and Hafellner
1991
Austria
Magnes and Hafellner
1991
Equisetum
Carex
fluviatile
rostrata
Phragmites
leaves
australis
Engl and
Phragmites
stems
australis
Engl and
11
Apinis et a l . 1972a
Taligoola et a l . 1972
Didymosphaeria sp.
beech wood
England
Eaton and Jones 1971b
Scots pine wood
Engl and
Davis and Jones
unpublished
Diplonaevia
Hein
emergens (Karst.)
Juncus
filiformis
Eriophorum angustifolium
Austria
Magnes and Hafellner
1991
Diplonaevia
seriata
Carex
Austria
Magnes and Hafellner
1991
(Libert) Hein
rostrata
Hadrospora fallax (Houton) Boise
(sub Trematosphaeria fallax Mouton)
balsa wood
USA, MD
Shearer and Crane 1971
Kirschsteiniothelia
sp. CS-825-1,2
woody debris
USA, IL, Chile
Shearer unpublished
Lepidopterella
palustris
Shearer & Crane
balsa wood
USA, IL
Shearer and Crane 1980a
Leptosphaeria acuta
(Hoffm.:Fr.) Karsten
Scirpus, Typha
Leptosphaeria
clavicarpa
E l l . & Ev.
Schoenoplectus
Leptosphaeria
culmifraga
(Fr.:Fr.) Ces. & deNot.
Phragmites
culms
australis
England
Apinis et a l . 1972b
Phragmites
stems
australis
Engl and
Taligoola et a l . 1972
Cavaliere 1975
lacustris
USSR
Dudka 1963
Leptosphaeria lemaneae
(Cohn) Sacc.
P a r a s i t i c on
Lemanea sp.
England
Ingold 1955
Leptosphaeria
sparganii
(Fautrey) Munk
Typha leaves
Engl and
Pugh and Mulder 1971
Leptosphaeria spp.
organic debris
Ireland
Park 1972
birch and maple
twigs
USA, RI
Lophiostoma
Fuckel
appendiculata
Salix
Lophiostoma arundinis
(Fr.) Ces. & De Not.
wood
Dudka 1963
Phragmites
Lophiostoma sp.
Lamore and Goos 1978
USSR
australis
twigs
USSR
Dudka 1963
USA, IL
Shearer unpublished
Austria
Magnes and Hafellner
1991
Massarina amphibia Magnes
& Hafellner
Equisetum
fluviatile,
Carex rostrata, C. nigra
Massarina
Engl and
twigs and roots of
Alnus glutinosa (L.) Gaertn.
Webster 1965
Massarina
arundinacea
(Sow.:Fr.) Leuchtman
(sub Leptosphaeria
arundinacea
Sow.:Fr.)
Phragmites
culms
Apinis et al 1972b
Massarina
australiensis
wood
Massarina
tetraploa
aquatica Webster
Hyde
Scheuer
australis
Australia
Scheuer 1991
alder, ash, oak and
willow twigs
England
Willoughby and Archer
1973
twigs
Engl and
Webster and Descals 1979
England
Webster and Descals 1979
USA, IL
Shearer and von Bodman
1983
alder twigs
England
Shearer and Webster 1991
alder and beech twigs
Hungary
Revay and Gonczol 1990
Carex
Austria
Magnes and Hafellner
1991
Carex nigra
Austria
Magnes and HafelIner
1991
Typha leaves
USA, MN
Cavaliere 1975
Fagus sylvaticus
twigs
Ehrh.
L.
twigs
Massariosphaeria
Criveili
rubicunda
(Muller)
Massariosphaeria
scirpina
(Wint.) Leuchtman
(sub Leptosphaeria scirpina
Hyde in press
England
Carex acutiformis
Massarina spp.
Engl and.
rostrata
Wint.)
12
Magnes and Hafellner
1991
Micropeltopsis
nigro-annulata
(Webster) Spooner & Kirk var.
papillosa
(Scheuer) Magnes
& Hafellner
Carex nigra, C. rostrata,
Equisetum
fluviatile,
Eriophorum
angustifolium,
Juncus.
filiformis
Mycosphaerella aspidii
Holm & Holm
Equisetum
fluviatile
Austria
Magnes and Hafellner
1991
Mycosphaerella
equiseticola
Bond.-Monte.
Equisetum
fluviatile
Austria
Magnes and Hafellner
1991
Mycosphaerella perexigua (Karst.)
Johanson var. minima Johanson
Carex rostrata,
Eriophorum
angustifolium
Austria
Magnes and Hafellner
1991
Mycosphaerella c f . perexigua
(Karst.) Johanson
Carex
Austria
Magnes and Hafellner
1991
Nodulosphaeria modesta
(Desm.) Munk ex Holm
[sub Leptosphaeria modesta
(Desm.) Auersw.]
Typha leaves
Engl and
Pugh and Mulder 1971
Denmark
Munk 1957
Ophiobolus gracilis
(v. Hohnel)
(Niessl) E. Muller
rostrata
Carex paniculata
Ophiobolus typhae Feltgen
Typha
Ophiobolus sp.
Phragmites
Paraphaeosphaeria michotii
(West.) Eriksson
[sub Leptosphaeria michotii
(West.) S a c c ]
Passeriniella
obiones (Crouan & Crouan)
Hyde & Mouzouras
[sub Passeriniella
discors
(Sacc. & E l l . ) Apinis
& Chesters]
Phaeosphaeria albopunctata
(West.) Shoem.
[sub Leptosphaeria albopunctata
(Westend.) S a c c ]
L.
latifolia
Engl and
Ingold 1951
Engl and
Apinis et a l . 1972b
Typha leaves
England
Pugh and Mulder 1971
Phragmites
culms
australis
England
Apinis et a l . 1972b
Phragmites
stems
australis
England
T a l i g o o l a et a l . 1972
wood
USA, IL
Shearer and Crane 1986
Typha leaves
England
Pugh and Mulder 1971
Phragmites
australis
USSR
Dudka 1963
Phragmites
stems
australis
England
Taligoola et a l . 1972
australis
USA, RI
Lamore and Goos 1978
England
Apinis et a l . 1972a
Carex rostrata,
Equisetum
fluviatile
Austria
Magnes and Hafellner
1991
Phaeosphaeria
caricinella
(Karst.) 0. Eriksson
Carex rostrata,
C. paupercula subspec.
irrigua, Juncus
filiformis
Austria
Magnes and Hafellner
1991
Phaeosphaeria culmorum (Auersw.
in Rehm) Leuchtm.
Carex rostrata,
Equisetum
fluviatile,
Glyceria
fluitans
Austria
Magnes and Hafellner
1991
Phaeosphaeria eustoma
(Fuckel) Holm
[sub Leptosphaeria eustoma
(Fuckel) S a c c ]
beech wood
England
Eaton 1972
beech wood
England
Eaton and Jones 1971a
Scirpus,
USA, MN
Cavaliere 1975
Carex rostrata,
Equisetum
fluviatile
Austria
Magnes and Hafellner
1991
Phragmites australis
leaves
England
Apinis et a l . 1972a
Phaeosphaeria halima
(Johnson) Shoem.
(sub Leptosphaeria halima Johnson)
Phragmites
stems
England
Taligoola et a l . 1972
Phaeosphaeria herpotricha (Fries) Holm
[sub Ophiobolus herpotrichus (Fries)
Sacc. ]
Typha sheaths
Engl and
Pugh and Mulder 1971
Phaeosphaeria
herpotrichoides
(de Not.) Holm
beech wood
Engl and
Eaton and Jones 1971b
Carex rostrata
Austria
Magnes and Hafellner
1991
twigs
Phragmites
leaves
Phaeosphaeria
alpina Leuchtm.
Phaeosphaeria eustomoides
( S a c c ) Shoem.
(sub Leptosphaeria eustomoides
australis
Typha
Sacc)
australis
13
USA, MN
Cavaliere 1975
Typha leaves
England
Pugh and Mulder 1971
Phaeosphaeria microscopica
(Karst.) Eriksson
(sub Leptosphaeria microscopica
Karst.)
Phragmites australis
leaves
England
Apinis et a l . 1972a
Phragmites australis
stems
England
Taligoola et a l . 1972
Phaeosphaeria sowerby
(Fuckel) Holm
[sub Leptosphaeria sowerby
(Fuckel) S a c c ]
Typha leaves
USA, MN
Cavaliere 1975
Phaeosphaeria typhae
(Karst.) Shoem.
[sub Leptosphaeria typhae
(Auersw.) Karst.]
Typha leaves
England
Pugh and Mulder 1971
Scirpus,
USA, MN
Cavaliere 1975
Phaeosphaeria typharum
(Desm.) Holm
[sub Leptosphaeria typharum
(Desm.) Karst.]
Typha leaves
England
Webster 1955
Typha leaves
USA, MN
Cavaliere 1975
Typha leaves
England
Pugh and Mulder 1971
reed leaf bases
England
Ingold 1955
Pleospora gaudefroyi Pat.
(sub Pleospora
lignicola
Webster & Lucas)
wood
England
Webster and Lucas 1961
Pleospora
Carex rostrata
Austria
Magnes and Hafellner
1991
England
Webster and Lucas 1961
Equisetum fluviatile
Austria
Magnes and Hafellner
1991
Schoenoplectus
stalks
lacustris
England
Schoenoplectus
lacustris
USSR
Scirpus,
Phaeosphaeria
juncicola
(Rehm ex Winter) Holm
(sub Leptosphaeria
juncicola
Rehm apud Wint.)
Phaeosphaeria
licatensis
(Sacc.) Shoem.
(sub Leptosphaeria licatensis
Physalospora
Ingold
Sacc.)
aquatica
incerta
Criv.
Pleospora
palustris
Berlese
Pleospora phaeospora
Ces. & de Not.
Typha
Typha
Scirpus maritimus
(Duby)
Pleospora
scirpicola
(DC.) Karst.
L.
Ingold 1955
Dudka 1963
Pleospora submersa
Webster & Lucas
Scirpus maritimus
England
Pyrenophora typhaecola
(Cooke) E. Muller
[sub Pleospora typhaecola
(Cke.) S a c c ]
Typha leaves
England
Rebentischia unicaudata
(Berk. & Br.) S a c c
deciduous leaves
USA, IL
Shearer and Crane 1986
Rebentischia sp.
Phragmites
stalks
England
Ingold 1955
Sphaerulina sp.
Phragmites
leaves
australis
England
Sporormia minima Auersw.
beech, Scots
pine wood
beech wood
England
Sporormiella sp.
twigs
USA, IL
Trematosphaeria
britzelmayriana
(Rehm) Sacc.
twigs
Hungary
Trematosphaeria circinans Fuckel
[sub Trematosphaeria vindelicorum
(Rehm) S a c c ]
alder and beech wood
Hungary
Trematosphaeria hydrela
(Rehm) S a c c
immersed tree
trunks
USSR
Petrak 1925
Trematosphaeria hydrophyla
(Karst.) Sacc.
Salix
USSR
Dudka 1963
Webster and Lucas 1961
Pugh and Mulder 1971
wood
14
England
Apinis et a l . 1972a
Eaton and Jones 1971a
Eaton 1972
Shearer and von Bodman
1983
Revay and Gonczdl 1990
Revay and Gdnczol 1990
beech, Scots
pine wood
England
Eaton and Jones 1972
beech, Scots
pine wood
England
Eaton and Jones 1971b
alder and beech
wood
Hungary
Revay and Gonczol 1990
Trematosphaeria sp.
beech twigs
Hungary
Revay and Gonczol 1990
Tubeufia paludosa
(Crouan & H. Crouan) Rossman
(sub Tubeufia helicomyces Hohnel)
Arrhenatherum,
Phragmites stems
England
Webster 1951
Wettsteinina
Muller
Phragmites
stalks
australis
England
Ingold 1955
Phragmites
stalks
australis
USSR
Dudka 1963
Phragmites
stems
australis
Engl and
Taligoola et a l . 1972
Phragmites
culms
australis
England
Apinis et a l . 1972b
USA, MD
Shearer 1972
fluviatile
Austria
Magnes and Hafellner
1991
Trematosphaeria pertusa
(Pers.:Fr.) Fuckel
niesslii
balsa wood
Wettsteinina sp.
Equisetum
all A s c o m y c e t e s that o c c u r o n s u b m e r g e d o r p a r t i a l l y s u b m e r g e d substrata i n the
aquatic habitats o u t l i n e d b y C o r r e l l a n d C o r e l l (1975) w i l l be c o n s i d e r e d " f r e s h w a ter A s c o m y c e t e s " . M u c h a d d i t i o n a l study is necessary before a m o r e precise d e f i n i t i o n o f the g r o u p c a n be m a d e .
S t u d y techniques
C O L L E C T I O N : F W A c a n be f o u n d b y e x a m i n i n g l i v i n g a n d dead m a c r o p h y t e s i n
p o n d s , lakes, bogs, s w a m p s , wetlands, drainage ditches a n d s m a l l t e m p o r a r y water
bodies. S u b m e r g e d w o o d y debris is also a g o o d source o f F W A , b u t o n l y a few species, eg. Aquadiscula
aquatica (Fig. 1), Mitrula alba a n d M. paludosa, o c c u r o n submerged d e c i d u o u s leaves. S u b s t r a t a c a n be e x a m i n e d f o r a s c o m a t a i n the field u s i n g
a h a n d lens a n d p r o m i s i n g substrata c a n be r e t u r n e d i n a plastic b a g t o the l a b o r a t o ry f o r further study. T h e absence o f a s c o m a t a o n substrata i n the field is n o t necessarily a g o o d i n d i c a t i o n that F W A are n o t present because some species o n l y appear
after i n c u b a t i o n i n m o i s t c h a m b e r s . T h u s it is a g o o d practice t o i n c u b a t e field materials as w e l l as e x a m i n e t h e m directly. M o i s t chambers f o r i n c u b a t i o n c a n be c o n structed o f glass o r plastic P e t r i dishes c o n t a i n i n g m o i s t e n e d filter paper. P l a s t i c
boxes o r bags c o n t a i n i n g m o i s t e n e d paper t o w e l i n g c a n also be u s e d . T h e choice o f
paper t o w e l i n g is i m p o r t a n t because some types c o n t a i n substances that stimulate
the g r o w t h o f o p p o r t u n i s t i c fungi such as Trichoderma a n d Penicillium
w h i c h overg r o w the F W A . I n c u b a t e d samples s h o u l d be kept at a temperature s i m i l a r to that
o f the n a t u r a l habitat a n d i n alternating light a n d d a r k w h i c h a p p a r e n t l y enhances
sexual r e p r o d u c t i o n (Shearer a n d v o n B o d m a n 1983). I f i n c u b a t i o n is used, it is i m p o r t a n t t o e x a m i n e samples i m m e d i a t e l y after c o l l e c t i o n , p r i o r t o i n c u b a t i o n , because some F W A deteriorate once they are r e m o v e d f r o m water.
15
Since it is often d i f f i c u l t to determine the source a n d length o f s u b m e r s i o n p e r i o d
o f r a n d o m l y c o l l e c t e d substrata, baits have been e m p l o y e d to f o l l o w the c o l o n i z a t i o n a n d d e v e l o p m e n t o f F W A (Shearer, 1972; Shearer a n d v o n B o d m a n 1983).
B a i t s s u c h as twigs o r w o o d b l o c k s c a n be attached to m a s o n r y b r i c k s o r cement
b l o c k s a n d p l a c e d i n the h a b i t a t . T h e y also c a n be assembled i n t o p a c k s w i t h n y l o n
t w i n e a n d attached to a l i n e tethered to a r e l a t i v e l y p e r m a n e n t structure such as a
pier o r a tree (Shearer 1972). S u b s t r a t a s u c h as d e a d m a c r o p h y t e leaves a n d stems
c a n be p l a c e d i n n y l o n mesh bags a n d p l a c e d i n the habitat as a b o v e . G i v e n the fluct u a t i n g water levels a n d often m u d d y c o n d i t i o n s c o m m o n i n freshwater habitats, a
g o o d deal o f t h o u g h t a n d p l a n n i n g s h o u l d go i n t o the selection o f s a m p l i n g sites a n d
the m e t h o d o f p l a c i n g a n d r e t r i e v i n g o f samples. A l l o w a n c e s s h o u l d be m a d e f o r
flood conditions.
Indirect c o l l e c t i o n techniques s u c h as p l a t i n g water, m u d s , h o m o g e n i z e d substrata
etc. are n o t v e r y efficient. A s c o s p o r e s o f some F W A (e.g. Aniptodera,
Lepidopterella) are d i f f i c u l t to g e r m i n a t e . A l t h o u g h ascospores o f m a n y F W A germinate
r e a d i l y , a l o n g t i m e p e r i o d a n d special c u l t u r a l procedures are necessary to i n d u c e
r e p r o d u c t i o n . S o m e species d o n o t r e p r o d u c e i n c u l t u r e . W i t h o u t r e p r o d u c t i v e
structures F W A c a n n o t be i d e n t i f i e d . T h u s it is quite l a b o r i o u s a n d often futile to
test the n u m e r o u s isolates that result f r o m p l a t i n g techniques.
T h e techniques o f f o a m e x a m i n a t i o n a n d m e m b r a n e f i l t r a t i o n o f water w h i c h are
so useful for the i d e n t i f i c a t i o n a n d q u a n t i f i c a t i o n o f a q u a t i c H y p h o m y c e t e s ( I q b a l
a n d W e b s t e r 1973, Shearer a n d W e b s t e r 1985) have little v a l u e for F W A . T h e ascospores o f F W A o c c u r at m u c h l o w e r densities t h a n the c o n i d i a o f a q u a t i c H y p h o m y cetes a n d thus are r a r e l y e n c o u n t e r e d o n m e m b r a n e filters. I n a d d i t i o n , the ascospores o f m a n y F W A are n o t very m o r p h o l o g i c a l l y d i s t i n c t i v e a n d unless the i n v e s t i gator is already v e r y k n o w l e d g e a b l e a b o u t the F W A present, they c a n n o t be i d e n t i f i e d . These l i m i t a t i o n s m a k e it very d i f f i c u l t to d i r e c t l y q u a n t i f y the s p o r a o f F W A
i n water.
I S O L A T I O N : T h e ascospores o f most F W A germinate relatively easily. F W A w i t h
closed ascomata can be isolated b y crushing mature ascomata i n 2-4 m l sterile distilled
water to f o r m a spore s u s p e n s i o n . D i s c h a r g e d ascospores w h i c h have a c c u m u l a t e d
a r o u n d ostioles c a n be r e m o v e d a f l a m e d , c o o l e d needle a n d stirred i n t o sterile water.
V i g o r o u s a g i t a t i o n o f spore suspensions w i t h a tube agitator is useful to separate
ascospores a n d d i s l o d g e attached b a c t e r i a . S p o r e suspensions are p o u r e d o n t o a n t i b i o t i c d i s t i l l e d water agar ( A W A , agar 18 g, s t r e p t o m y c i n sulfate 250 m g / 1 , p e n i c i l l i n G 250 m g / 1 , d i s t i l l e d water 11). A n t i b i o t i c s are a d d e d t o the m o l t e n agar i m m e diately after a u t o c l a v i n g . S p o r e suspensions o n A W A plates are a l l o w e d to stand
for 1-2 h r to p e r m i t settling o f the ascospores. Excess water is gently decanted f r o m
the plates w h i c h are i n c u b a t e d at a slight angle f r o m the u p r i g h t so excess water c a n
d r a i n o f f . T o isolate D i s c o m y c e t e s , a s m a l l piece o f filter paper is attached w i t h a
d r o p o f water to the l i d o f a n A W A plate a b o u t 1 c m f r o m the edge o f the l i d . A n
a p o t h e c i u m is r e m o v e d f r o m the s u b s t r a t u m w i t h a sterile needle a n d p l a c e d o n the
filter paper so that its surface is facing d o w n t o w a r d the agar surface. T h e l i d is placed
o n the b o t t o m d i s h a n d rotated about 4 5 ° every 30 minutes so that ascospores are
16
d i s c h a r g e d i n a c i r c u l a r p a t h a r o u n d the plate. T h i s p r o c e d u r e c a n be reversed for
p y r e n o m y c e t o u s f o r m s w i t h f o r c i b l e ascospore discharge. A s m a l l piece o f substratum
w i t h p e r i t h e c i a directed u p w a r d is m o u n t e d o n the filter paper a n d the P e t r i
d i s h is i n v e r t e d so the agar that is o n t o p receives u p w a r d l y d i s c h a r g e d ascospores.
A f t e r 24 h o u r s i n c u b a t i o n at r o o m temperature, plates s h o u l d be e x a m i n e d m i c r o s c o p i c a l l y w i t h t r a n s m i t t e d light a n d the p o s i t i o n o f g e r m i n a t e d ascospores m a r k e d .
It is most i m p o r t a n t that single, r e c o g n i z a b l e , u n c o n t a m i n a t e d ascospores be i s o l a t e d . T h u s e x a m i n a t i o n a n d transfer o f g e r m i n a t e d ascospores w i t h i n 24-48 hours
is a c r i t i c a l step. I f the plates are left for longer periods o f t i m e , the ascospores m a y
be o v e r g r o w n b y contaminants o r unrecognizable due to their g e r m i n a t i o n a n d g r o w t h .
Because o f v a r i a t i o n i n f r u i t i n g a b i l i t y a m o n g strains a n d the p o s s i b i l i t y that r e p r o d u c t i o n is h e t e r o t h a l l i c , at least 6-8 isolates o f a given species s h o u l d be m a d e . S i n g l e
g e r m i n a t e d ascospores s h o u l d be transferred f r o m A W A to s t a n d a r d nutrient m e d i a
such as c o r n m e a l agar ( D i f c o ) , m a l t extract agar ( D i f c o ) , peptone, yeast extract, g l u cose agar ( F u l l e r a n d J a w o r s k i 1987) or yeast extract s o l u b l e s t a r c h agar ( F u l l e r a n d
J a w o r s k i 1987). F o r the l o n g - t e r m storage o f cultures, strips o f b a l s a o r b i r c h w o o d
can be a d d e d to c u l t u r e tubes o f the a b o v e m e d i a before a u t o c l a v i n g . C u l t u r e s o f
new o r u n u s u a l species s h o u l d be deposited w i t h one o r m o r e i n t e r n a t i o n a l l y r e c o g n i z e d c u l t u r e c o l l e c t i o n s s u c h as the A m e r i c a n T y p e C u l t u r e C o l l e c t i o n o r the Intern a t i o n a l M y c o l o g i c a l Institute C o l l e c t i o n .
T o i n d u c e a s c o m a t a f o r m a t i o n , cultures are g r o w n o n a l f a l f a stems o r w o o d strips.
Six to eight pieces (4 c m l o n g ) are a d d e d to 50 m l sterile d i s t i l l e d water (filtered water
f r o m the n a t u r a l habitat m a y also be used) i n 125 m l E r l e n m e y e r flasks a n d a u t o c l a v e d for one h o u r o n three successive d a y s . F l a s k s are i n o c u l a t e d w i t h m y c e l i u m
f r o m stock tubes a n d p l a c e d o n a r o t a r y shaker (100 r p m ) at 2 2 C u n t i l substrata are
v i s i b l y w e l l - c o l o n i z e d . S u b s t r a t a are transferred aseptically to sterile m o i s t chambers
(see above), sealed w i t h p a r a f i l m , a n d i n c u b a t e d at 18-22C i n alternating l i g h t / d a r k
c o n d i t i o n s . D a y l i g h t fluorescent light is acceptable a n d an a d d i t i o n a l b l a c k light m a y
be useful i n s t i m u l a t i n g f r u i t i n g i n some species (Webster pers. c o m m . ) . S u b s t r a t a
s h o u l d be e x a m i n e d w e e k l y for the presence o f a s c o m a t a .
A q u a t i c h y p h o m y c e t e a n a m o r p h s have been r e p o r t e d for several F W A ( T a b l e 2),
thus a l l cultures s h o u l d be tested for the p r o d u c t i o n o f s u c h states (Webster a n d D e s cals 1979, W e b s t e r 1992). T h i s c a n be done b y s u b m e r g i n g slivers o f c o l o n i z e d agar
i n sterile d i s t i l l e d water at 15-18C for c o l d water species a n d 2 2 - 2 5 C for w a r m water
species. U s i n g t r a n s m i t t e d l i g h t , submerged cultures s h o u l d be e x a m i n e d for c o n i d i a
p e r i o d i c a l l y over several weeks. I f n o c o n i d i a are p r o d u c e d i n static water c u l t u r e ,
isolates m u s t t h e n be tested i n aerated c u l t u r e . T h i s c a n be d o n e b y passing air f o r c i b l y t h r o u g h a e r a t i o n stones or Pasteur pipettes d r a w n o u t to a fine p o i n t i n flasks
or c u l t u r e tubes. N u m e r o u s F W A p r o d u c e a n a m o r p h s other t h a n a q u a t i c h y p h o m y cetes, but these are u s u a l l y p r o d u c e d o n s t a n d a r d l a b o r a t o r y m e d i a so e x a m i n a t i o n
o f stock cultures is sufficient for their detection.
EXAMINATION: For known F W A ,
squash m o u n t s o f a s c o m a t a are u s u a l l y sufficient
for i d e n t i f i c a t i o n . I n the case o f u n k n o w n or p r o b l e m a t i c specimens, it is necessary
to section a s c o m a t a to determine a n a t o m i c a l details a n d c e n t r u m structure. R o u g h
17
Table II. Teleomorph-anamorph connections for freshwater Ascomycetes.
TELEOMORPH
Vibrissea flavovirens
(Pers: Fr) Korf & Dixon
(sub Apostemidium torrenticola
Graddon)
Anavirga dendromorpha
Descal5 & Sutton
Hamad and Webster 1987
Gnomonia papuana
Sivanesan & Shaw
Sesquicillium sp.
Sivanesan and Shaw 1977
Hyaloscypha lignicola
Abdullah & Webster
Pseudaegerita sp. (corticalis?)
Abdullah and Webster 1983
Hyaloscypha zalewskii
Descals & Webster
Clathrosphaerina zalewski
v. Beverwijk
Descals and Webster 1976
Hydrocina chaetocladia
Scheuer
Tricladium chaetocladium Ingold
Webster et a l . 1991
Hymenoscyphus africanus
Descals, Fisher & Webster
Geniculospora grandis
(Greathead ex Nolan) Nilsson
Descals et a l . 1984
Hymenoscyphus foliicola
Abdullah, Descals & Webster
Dimorphospora
Tubaki
Abdullah et a l . 1981
Hymenoscyphus inberbis
Anguillospora
foliicola
fustiformis
Descals and Harvanova
unpublished
Hymenoscyphus malawienis
Fisher & Spooner
Unnamed hyphomycete
Fisher and Spooner 1987
Hymenoscyphus paradoxus
Fisher & Webster
Helicodendron paradoxum
Peyronel
Fisher and Webster 1983
Hymenoscyphus splendens
Abdullah, Descals & Webster
Tricladium splendens Ingold
Hymenoscyphus tetracladius
Abdullah, Descals & Webster
Articulospora
Ingold
Hymenoscyphus varicosporoides
Tubaki
Varicosporium sp.
Lambertella tubulosa
Abdullah & Webster
Heliocodendron tubulosum
(Reiss) Linder
Loramyces juncicola
Weston
Anguillospora sp.
Massarina aquatica
Webster
Tumularia aquatica
Massarina sp.
Miladina lechithina
(Cooke) Svrcek
Abdullah et a l . 1981
Tubaki 1966
Abdullah and Webster 1981
Digby and Goos 1987
Webster 1965
(Ingold) Descals & Marv.
Clavariopsis
Massarina sp.
tetracladia
Abdullah et a l . 1981
aquatica de Wild.
Anguillospora longissima
(Sacc. & Syd.) Ingold
Actinospora megalospora Ingold
Webster and Descals 1979
Willoughby and Archer 1973
Descals and Webster 1978
Mollisia gigantea
Fisher & Webster
Helicondendron giganteum
Glenn-Bott
Fisher and Webster 1983
Mollisia uda
(Pers.: Fr.) G i l l .
Anguillospora crassa Ingold
Webster 1961
Mollisia
Filosporella
Webster and Descals 1979
sp. I
sp.
Nectria discophora
(Mont.) Mont.
Cylindrocarpon ianthothele
Wollenw.
Booth 1966
Nectria haematococca
Berk. & 8r.
Fusarium solani (Mart.)
Sacc.
Booth 1960, 1971
Nectria lucidum
Hohnel
Cylindrocarpon lucidum
Booth
Booth 1966
Nectria lugdunensis
Webster
Heliscus lugdunensis
Sacc. & Therry
Webster 1959
Nectria
penicillioides
Ranzoni
Flagellospora
Ingold
Ranzoni 1956
penicillioides
Nectria sp.
Flagellospora
curta
Webster unpublished
Orbilia
Anguillospora
rosea sp. ined.
Webster and Oescals 1979
sp.
Pezoloma sp.
Pyxidiophora
Webster
spinulo-rostrata
Tubeufia paludosa (Crouan &
H. Crouan) Rossman
(sub Tubeufia helicomyces Hohnel)
Talaromyces flavus (Klocher)
Stolk & Samson var. flavus
Anguillospora furtiva
Descals
Webster and Descals 1979
denticulate anamorphic
state
Webster and Hawksworth 1986
Webster 1951
Helicosporium phragmites Hdhn.
Pitt 1979
Penicillium
dangeardii Pitt
sections o f a s c o m a t a m a y be m a d e w i t h a r a z o r b l a d e , but m o r e d e f i n i t i v e w o r k requires t h i n sections. T h e freezing m i c r o t o m e is useful for sectioning fresh specimens,
especially D i s c o m y c e t e s a n d a s c o m a t a e m b e d d e d i n host o r substrata. E m b e d d i n g
o f specimens i n l o w v i s c o s i t y resin a n d s e c t i o n i n g w i t h the u l t r a - m i c r o t o m e ( H u h n d o r f 1991) has p r o v e d excellent f o r b o t h fresh a n d d r i e d specimens. I n a d d i t i o n to
sectioning m a t e r i a l , it is i m p o r t a n t to m a k e squash m o u n t s o f p o r t i o n s o f the cent r u m i n water, f o l l o w e d b y I n d i a i n k . T h i s permits the h y d r a t i o n a n d r e v e l a t i o n o f
gelatinous coatings o n physes a n d ascospore sheaths a n d appendages. W h e n n o sheaths
or appendages are revealed b y I n d i a i n k , it is necessary to r e m o v e the i n k b y p u l l i n g
d i s t i l l e d water u n d e r the c o v e r s l i p . T h e appendages o f some F W A stain w i t h i n k a n d
are o n l y visible after excess i n k is r e m o v e d . It is also necessary to s t a i n ascus tips
w i t h M e l t z e r ' s reagent a n d aqueous c o t t o n b l u e to determine the n a t u r e o f the a p i c a l
a p p a r a t u s a n d / o r apex.
It is c r i t i c a l that b o t h type a n d v o u c h e r specimens be deposited i n r e c o g n i z e d herbaria ( D e n n i s 1978, K o h l m e y e r a n d K o h l m e y e r 1979). A p u b l i s h e d r e c o r d o f species
occurrences is useless unless it c a n be v a l i d a t e d b y later investigators. D e p o s i t e d specimens also s h o u l d p r o v e useful i n p o p u l a t i o n studies. I n a d d i t i o n , d r i e d a s c o m a t a
can serve as a source o f genetic i n f o r m a t i o n i n the f o r m o f preserved D N A for t a x o n o m i c a n d p o p u l a t i o n studies. A s s o o n as cultures, sections a n d s q u a s h m o u n t s have
been o b t a i n e d , specimens s h o u l d be d r i e d d o w n w h i l e they are still i n g o o d c o n d i t i o n . I n d r y climates, this c a n be done at r o o m temperature w i t h n o special p r o c e dures. I n h u m i d areas, a desiccant m a y be needed a l o n g w i t h gentle heat. H i g h d r y ing temperatures s h o u l d be a v o i d e d as this c o u l d degrade D N A . It is extremely i m portant to i n c l u d e w i t h h e r b a r i u m specimens details o f habitat (e.g. standing o r f l o w ing water, water c h e m i s t r y , temperature) a n d s u b s t r a t u m (type a n d c o n d i t i o n ) . T h i s
i n f o r m a t i o n , w h e n c o m p i l e d i n the future, w i l l be i n v a l u a b l e i n i n t e r p r e t i n g the e c o l ogy a n d d i s t r i b u t i o n patterns o f F W A .
I D E N T I F I C A T I O N : W i t h the e x c e p t i o n o f a k e y to the A s c o m y c e t e s o n emergent vegetation o f a l p i n e lakes ( M a g n e s a n d H a f e l l n e r 1991), there are n o c o m p r e h e n s i v e
keys to the F W A . T h u s it is necessary to collect the o r i g i n a l descriptions o f the species k n o w n f r o m freshwater. C i t a t i o n s i n T a b l e 1 c a n be used to o b t a i n a large part
o f this literature. T h e r e are several general w o r k s that are useful: M i i l l e r (1950), H o l m
(1957), M u n k (1957), E r i k s s o n (1967), D e n n i s (1978), H e d j a r o u d e (1969), L u n d q vist (1972), K o r f (1973), L u t t r e l l (1973), M i i l l e r a n d v o n A r x (1973), B a r r (1978, 1987,
1990), K o h l m e y e r a n d K o h l m e y e r (1979), K o h l m e y e r a n d V o l k m a n n - K o h l m e y e r
(1991), L e u c h t m a n n (1984), Sivanesan (1984), a n d S h o e m a k e r a n d B a b c o c k (1989).
Systematics
The systematics o f A s c o m y c e t e s , i n general, is at present c o n t r o v e r s i a l a n d t a x o n o m ic schemes a b o u n d ( H a w k s w o r t h , et a l . 1983). T h e classification system o f B a r r (1987,
1990) is used for the L o c u l o a s c o m y c e t e s a n d p y r e n o m y c e t o u s m e m b e r s o f the class
H y m e n o a s c o m y c e t e s a n d that o f H a w k s w o r t h et a l . (1983) f o r a l l other t a x a . B a s e d
o n these systems, 275 ascomycete species i n 15 orders ( E u r o t i a l e s , O n y g e n a l e s , H e -
19
lotiales, P e z i z a l e s , R h y t i s m a t a l e s , C a l o s p h a e r i a l e s , D i a p o r t h a l e s , H y p o c r e a l e s ,
Halosphaeriales, Sordariales, Xylariales, Dothideales, Pleosporales, and M e l a n o m matales) have been reported f r o m freshwater habitats. O r d e r s w i t h 10 o r m o r e representatives f r o m freshwater are the H a l o s p h a e r i a l e s , H e l o t i a l e s , P l e o s p o r a l e s a n d
S o r d a r i a l e s . These orders represent f o u r d i s t i n c t l y different e v o l u t i o n a r y lines: the
m a r i n e p y r e n o m y c e t o u s H y m e n o a s c o m y c e t e s , the d i s c o m y c e t o u s H y m e n o a s c o m y cetes, the L o c u l o a s c o m y c e t e s , a n d the p y r e n o m y c e t o u s H y m e n o a s c o m y c e t e s ,
respectively.
T h e r a t i o o f species o f P l e c t o m y c e t e s : D i s c o m y c e t e s : P y r e n o m y c e t e s : L o c u l o a s c o mycetes is 6:94:71:71. A n interesting difference between the F W A a n d m a r i n e A s c o mycetes is the absence o f D i s c o m y c e t e s i n m a r i n e habitats. O f 255 ascomycete t a x a
k n o w n f r o m m a r i n e habitats ( K o h l m e y e r a n d V o l k m a n n - K o h l m e y e r 1991), o n l y one
belongs i n the D i s c o m y c e t e s . T h e F W A differ f r o m the m a r i n e A s c o m y c e t e s not o n l y
i n the relative p r o p o r t i o n s o f t a x a i n m a j o r ascomycete g r o u p s but also i n species
c o m p o s i t i o n . O n l y eight species have been r e p o r t e d f r o m b o t h freshwater a n d m a rine ( i n c l u d i n g b r a c k i s h ) habitats: Aniptodera
chesapeakensis,
Halosarpheia
retorquens, Halosarpheia
viscosa, Luttrellia
estuarina, Nais inornata, Phaeosphaeria
halima, Savoryella lignicola, a n d Zopfiella latipes. It appears that some aspect o f the
m a r i n e e n v i r o n m e n t has been an extremely effective barrier to the c o l o n i z a t i o n o f
m a r i n e habitats b y freshwater A s c o m y c e t e s a n d vice versa. T h i s barrier(s) has effectively isolated the t w o mycotas a n d it appears that they have e v o l v e d differently (See
E v o l u t i o n Section).
Geographical distribution
T h e r e are so few c o m p r e h e n s i v e studies o f the F W A o f different g e o g r a p h i c a l areas
that little c a n be said about this t o p i c . A t this t i m e , d i s t r i b u t i o n patterns are largely
c o l l e c t o r - l i n k e d . Nais inornata, Pseudohalonectria
lignicola a n d Wettsteinina
niesslii have each been reported f r o m at least three w i d e l y separated g e o g r a p h i c a l regions
( T a b l e 1) suggesting that they are w i d e l y d i s t r i b u t e d . C l e a r l y m u c h m o r e w o r k is
needed i n this area.
Ecology
F W A c a n be d i v i d e d i n t o t w o b r o a d e c o l o g i c a l groups based o n their n u t r i t i o n : (1)
parasites a n d endophytes o f aquatic m a c r o p h y t e s a n d algae, a n d (2) saprophytes o n
dead plant m a t e r i a l . A variety o f aquatic macrophytes, i n c l u d i n g Juncus,
Phragmites,
Scirpus, a n d Typha, serve as substrata f o r F W A . T a x a i n the L o c u l o a s c o m y c e t e s
were first n o t e d i n o r i g i n a l descriptions a n d t a x o n o m i c treatises ( M i i l l e r 1950, H o l m
1957, M u n k 1957, L e u c h t m a n n 1984), but little was reported a b o u t their e c o l o g y .
T h e first systematic effort to collect F W A o n aquatic m a c r o p h y t e s was b y I n g o l d
(1951, 1954, 1955) a n d I n g o l d a n d C h a p m a n (1952). These p i o n e e r i n g studies were
f o l l o w e d b y those o f W e b s t e r a n d L u c a s (1961), D u d k a (1963), P u g h a n d M u l d e r
(1971), A p i n i s et a l . (1972a, b), T a l i g o o l a et a l . (1972) a n d M a g n e s a n d H a f e l l n e r
20
(1991). A l t h o u g h I n g o l d (1955) predicted a r i c h p y r e n o m y c e t o u s F W A m y c o t a o n
aquatic m a c r o p h y t e s , this g r o u p has been largely i g n o r e d . L i t t l e is k n o w n a b o u t any
aspects o f the b i o l o g y a n d ecology o f this g r o u p , especially the nature a n d
specificities o f their interactions w i t h their hosts. M a g n e s a n d H a f e l l n e r (1991) recently collected A s c o m y c e t e s o n emergent plants s u r r o u n d i n g a l p i n e lakes to determ i n e s u b s t r a t u m r e l a t i o n s h i p s . A m o n g 52 t a x a , they f o u n d one large g r o u p o f species that appeared to be t o t a l l y u n s p e c i a l i z e d a n d that o c c u r r e d o n a v a r i e t y o f p l a n t
hosts. T h e y f o u n d 17 species that appeared to be substrate specific but they d i d n o t
consider these species to be p a r a s i t i c . Seven species o c c u r r e d o n t w o d i s t a n t l y related
plants, Carex a n d Equisetum.
S o m e F W A o c c u r o n l i v i n g s u b m e r g e d stems o f m a c rophytes ( I n g o l d a n d C h a p m a n 1952, I n g o l d 1955, W e b s t e r a n d L u c a s 1961, D u d k a
1963, D e n n i s 1968, A p i n i s et a l . 1972b, N a n n f e l d t 1985). W h e t h e r these fungi are
parasitic o r e n d o p h y t i c o n their hosts has not been d e t e r m i n e d . A n u m b e r o f species
o f L o c u l o a s c o m y c e t e s have been reported f r o m dead m a c r o p h y t e s ( D u d k a 1963,
A p i n i s et a l . 1972a a n d b , but whether they were actively g r o w i n g o n their hosts before they died o r c o l o n i z e d t h e m s a p r o p h y t i c a l l y after death o f the hosts is n o t
k n o w n . I n several studies, the succession o f fungi o n emergent m a c r o p h y t e s has
been f o l l o w e d ( P u g h a n d M u l d e r 1971, A p i n i s et a l . 1972a a n d b , T a l i g o o l a et a l .
1972). F o r Typha latifolia, d u r i n g the early stages o f leaf emergence, the m y c o t a was
d o m i n a t e d b y species o f yeasts a n d dematiaceous h y p h o m y c e t e s . A f t e r the leaves
d i e d , h o w e v e r , species o f L o c u l o a s c o m y c e t e s (Phaeosphaeria,
Paraphaeosphaeria,
Leptosphaeria
a n d Nodulosphaeria)
became p r o m i n e n t ( P u g h a n d M u l d e r 1971).
A s c o m y c e t e s were present i n early (Massarina a n d Wettsteinina)
a n d late (Ophiobolus, Lasiosphaeria
a n d Passeriniella)
stages o f succession o n s u b m e r g e d c u l m s o f
Phragmites australis ( A p i n i s et a l . 1972b). Massarina arundinaceae
was t h o u g h t to
assist i n the w e a k e n i n g o f c u l m s o f P. australis w h i c h results i n eventual breakage
at o r b e l o w the water surface ( A p i n i s et a l . 1972b). H o w effective F W A are i n dec o m p o s i n g m a c r o p h y t e s a n d whether they serve as a n u t r i t i o n a l resource to d e t r i t i v o r o u s invertebrates needs to be d e t e r m i n e d . C r i t i c a l studies o f the life cycles o f
F W A o n aquatic m a c r o p h y t e s are needed to elucidate their e c o l o g i c a l r o l e .
T h e m a j o r i t y o f F W A have been reported f r o m dead plant substrata. W h e t h e r this
reflects a true p r e p o n d e r a n c e o f saprophytes over parasites a n d endophytes o r the
s u b s t r a t u m bias o f collectors is not k n o w n . C e r t a i n l y there have been m o r e studies
o f aquatic fungi o n dead p l a n t substrata, p a r t i c u l a r l y a l l o c h t h o n o u s debris, t h a n o n
l i v i n g , a u t o c h t h o n o u s substrata. T h u s the p r o b a b i l i t y o f detection m a y have
favored s a p r o p h y t i c f o r m s . M a n y o f the s a p r o p h y t i c F W A i n f l o w i n g water have
been r e p o r t e d f r o m w o o d y debris c o m p a r e d to o n l y a few f r o m leafy debris ( T a b l e
1). T h e d i f f e r e n t i a l occurrence o f F W A o n these substrata has been discussed prev i o u s l y (Shearer 1992) a n d m a y be due to factors such as s u b s t r a t u m l o n g e v i t y
and/or nutrition.
W o o d as a s u b s t r a t u m for aquatic fungi has been studied relatively intensely ( D u d k a
1963, Shearer 1972, W i l l o u g h b y a n d A r c h e r 1973, L a m o r e a n d G o o s 1978, M i n o u r a
a n d M u r o i 1978, Shearer a n d v o n B o d m a n 1983, Shearer a n d C r a n e 1986, R e v a y
a n d G o n c z o l 1990, Shearer a n d Webster 1991) a n d n u m e r o u s F W A have been rep o r t e d f r o m these studies. F W A have also been r e p o r t e d f r o m water-saturated w o o d
21
i n c o o l i n g towers ( E a t o n a n d Jones 1970, 1971a a n d b , E a t o n 1972, E a t o n 1976).
I n a d d i t i o n , m a y new t a x a o f F W A have been described f r o m w o o d i n freshwater
(Webster 1959, Webster a n d L u c a s 1961, Webster 1965, T u b a k i 1966, Jones and E a t o n
1969, E a t o n a n d Jones 1970, I q b a l 1972, A d u l l a h a n d W e b s t e r 1983, M i n o u r a a n d
M u r o i 1978, Shearer 1978, Shearer a n d C r a n e 1978a, Shearer a n d C r a n e 1980a a n d
b , A b d u l l a h et a l . 1981, F i s h e r a n d P e t r i n i 1983, F i s h e r a n d W e b s t e r 1983, M o n o d
a n d F i s h e r 1983, Shearer 1984, Shearer 1989a a n d b , W e b s t e r et a l . 1991, C r a n e et
a l . 1992, H y d e , i n press).
T h e r o l e o f F W A i n e n z y m a t i c a l l y d e g r a d i n g w o o d is l a r g e l y u n k n o w n . C u l t u r a l
studies o f seven l i g n i c o l o u s F W A a n d a n a n a m o r p h o f a F W A ( Z a r e - M a i v a n a n d
Shearer 1988a) revealed that they were a l l e n z y m a t i c a l l y able to degrade a w i d e range
o f w o o d c o m p o n e n t s i n c l u d i n g starch, x y l o s e a n d cellulose. W i t h the e x c e p t i o n o f
t w o Nectria species a n d the a n a m o r p h o f TV. lugdunensis, Heliscus lugdunensis Sacc.
a n d T h e r r y , a l l species f o r m e d soft-rot cavities i n b a l s a , ash a n d c o t t o n w o o d . I n another study o f the same species ( Z a r e - M a i v a n a n d Shearer 1988b), a l l were f o u n d
to p r o d u c e c o u p l e d cellulases, but species d i f f e r e d i n their a b i l i t y to decay w o o d a n d
f o r m soft-rot cavities. T h e Nectria species, i n c l u d i n g the a n a m o r p h , H.
lugdunensis,
a n d Nais inornata caused little weight loss i n ash a n d c o t t o n w o o d b l o c k s c o m p a r e d
to f o u r other species. Nectria species are often the first F W A to appear o n n e w l y
submerged w o o d a n d disappear at a relatively early stage o f decay ( W i l l o u g h b y a n d
A r c h e r 1973, Shearer a n d v o n B o d m a n 1983). Since the Nectria species tested thus
far are u n a b l e to degrade w o o d t h r o u g h the f o r m a t i o n o f soft-rot cavities, it is l i k e l y
that they use components o f w o o d other t h a n secondary walls such as proteins, starch
a n d soluble sugars. These substances o c c u r i n relatively s m a l l quantities c o m p a r e d
to the cellulose, hemicelluloses a n d lignins o f secondary walls a n d they disappear
relatively early i n the d e c o m p o s i t i o n process b y l e a c h i n g a n d m i c r o b i a l u t i l i z a t i o n .
T h i s m a y explain, i n part, the early disappearance o f Nectria species f r o m w o o d ( W i l l o u g h b y a n d A r c h e r 1973, Shearer a n d v o n B o d m a n 1983). T h e early disappearance
o f these species also m a y be b r o u g h t a b o u t b y the antagonistic activities o f m o r e
c o m p e t i t i v e species. I n a study o f h y p h a l interactions a m o n g l i g n i c o l o u s f u n g i , the
same Nectria species discussed above were a m o n g the species least resistant to the
antagonistic activities o f other fungi (Shearer a n d Z a r e - M a i v a n 1988).
B y virtue o f their a b i l i t y to soften w o o d by the f o r m a t i o n o f soft-rot cavities, l i g n i c o l o u s F W A m a y p l a y an i m p o r t a n t role i n increasing the p a l a t a b i l i t y o f w o o d to
stream invertebrates. D u d l e y a n d A n d e r s o n (1982) indicate that softening is necessar y before submerged w o o d c a n be used b y invertebrate scrapers a n d gougers. T h e y
also suggest that w o o d borers are p a r t i a l l y dependent o n m i c r o b e s f o r n u t r i t i o n . P e reira et a l . (1982) have f o u n d f u n g a l h y p h a e a n d spores associated w i t h w o o d i n the
gut contents o f w o o d borers, gougers a n d grazers. B i r c h w o o d sticks p r e c o l o n i z e d
w i t h the F W A , Pseudohalonectria
lignicola, a n d then p l a c e d i n a stream s h o w e d evidence o f w o o d g o u g i n g after eight weeks o f s u b m e r s i o n (Shearer u n p u b l i s h e d ) a n d
Asellus sp. was successfully m a i n t a i n e d i n the l a b o r a t o r y o n P. lignicola g r o w i n g
o n b i r c h w o o d (Shearer and M a t t i n g l y unpublished). T h e importance o f leaf-degrading
fungi to stream invertebrates has been w e l l - d o c u m e n t e d ( S u b e r k r o p p 1992); whether
l i g n i c o l o u s F W A p l a y a s i m i l a r role awaits further i n v e s t i g a t i o n .
22
R e c e n t l y , it was f o u n d that l i g n i c o l o u s aquatic f u n g i were generally m o r e antagonistic a n d resistant to a n t a g o n i s m t h a n their f o l i i c o l o u s counterparts (Shearer a n d Z a r e M a i v a n 1988). These d a t a s u p p o r t e d the hypothesis that f u n g i o n l o n g - l a s t i n g substrata s u c h as w o o d have m o r e c o m p e t i t i v e strategies t h a n those o n less l o n g - l a s t i n g
substrata s u c h as leaves ( P u g h 1980). I n c l u d e d i n this study were 10 l i g n i c o l o u s a n d
2 foliicolous F W A . B o t h the foliicolous a n d Nectriaceous lignicolous F W A were p o o r
c o m p e t i t i o r s c o m p a r e d to the other F W A . A m o n g the m o s t c o m p e t i t i v e l i g n i c o l o u s
F W A were f o u r species o f Pseudohalonectria.
Species i n this genus were subsequently
f o u n d to be antagonistic at a distance against a w i d e range o f f u n g a l t a x a suggesting
that they p r o d u c e a diffusible i n h i b i t o r ( A s t h a n a a n d Shearer 1990). C u l t u r a l antago n i s m is not always correlated w i t h antagonism i n the field (Webber a n d H e d g e r 1986).
H o w e v e r field experiments w i t h P. lignicola a n d P. adversaria d e m o n s t r a t e d that
these species were able to defend c o l o n i z e d substrata against i n v a s i o n b y other species w h i l e a w e a k l y antagonistic species, N. haematococca,
was n o t (Shearer a n d B a r t o l a t a u n p u b l i s h e d ) . W i l l o u g h b y a n d A r c h e r (1973) suggested earlier that c o m p e t i t i o n m i g h t o c c u r between l i g n i c o l o u s F W A because r a r e l y m o r e t h a n one o r t w o species c o l o n i z e d a single t w i g . I n a d d i t i o n , o f the five F W A r e c o r d e d f r o m 41 twigs
i n their study, t w o species, Ceratostomella
sp. a n d Mollisia
sp. never o c c u r r e d t o gether. F i s h e r a n d A n s o n (1983) f o u n d that Massarina aquatica, a l i g n i c o l o u s F W A ,
produces a n t i f u n g a l substances i n c u l t u r e o n n a t u r a l substrata. I n recent field experiments w i t h the a n a m o r p h o f M. aquatica, Tumularia aquatica (Ingold) D e s c a l s
a n d M a r v . , this species excluded competitors f r o m c o l o n i z e d w o o d (Shearer a n d W e b ster u n p u b l i s h e d ) . These d a t a suggest that F W A , t h r o u g h their a n t a g o n i s t i c a c t i v i ties, m a y p l a y a n i m p o r t a n t r o l e i n o r g a n i z i n g freshwater l i g n i c o l o u s f u n g a l c o m m u nities.
L i t t l e is k n o w n a b o u t the response o f F W A to e n v i r o n m e n t a l parameters s u c h as
temperature, seasonality, water c h e m i s t r y , f l o w rates, a n d degree o f submergence.
F a c t o r s i n f l u e n c i n g ascospore f o r m a t i o n , l i b e r a t i o n , d i s p e r s a l , attachment a n d germ i n a t i o n are also largely u n k n o w n . Intraspecific a n d interspecific interactions o f F W A
are also largely u n s t u d i e d .
Evolution
It is l i k e l y that m o s t F W A have evolved f r o m terrestrial ancestors v i a a variety o f
e v o l u t i o n a r y p a t h w a y s . O n e p a t h w a y is as pathogens, endophytes a n d saprophytes
o f w e t l a n d a n d a q u a t i c p l a n t s . A s these plants i n v a d e d freshwater habitats, they n o
d o u b t b r o u g h t their associated m i c r o o r g a n i s m s w i t h t h e m . F u n g a l species able to
survive a n d adapt to aquatic habitats m a y have been ancestral to present-day species
that o c c u r o n freshwater m a c r o p h y t e s . I n c l u d e d i n this g r o u p are p r i m a r i l y t a x a i n
the D i s c o m y c e t e s a n d L o c u l o a s c o m y c e t e s . A second route to freshwater m a y have
been o n r i p a r i a n vegetation such as tree a n d shrub litter. T h i s debris carries w i t h
it a considerable c o m p l e x o f fungi capable o f a d a p t i n g to freshwater. I n c l u d e d i n
this g r o u p are species o f D i s c o m y c e t e s , H y p o c r e a l e s , S o r d a r i a l e s , H a l o s p h a e r i a l e s
a n d L o c u l o a s c o m y c e t e s . A t h i r d route to freshwater m a y have been t h r o u g h the r u n o f f o f r a i n w a t e r a n d sediments. I n c l u d e d i n this g r o u p m a y be species o f E u r o t i a l e s ,
Sordariales and Hypocreales.
23
K o h l m e y e r a n d K o h l m e y e r (1979) suggested that extant marine fungi c o u l d be d i v i d e d
i n t o t w o groups based o n their o r i g i n s . P r i m a r y m a r i n e fungi ( H a l o s p h a e r i a l e s , S p a thulosporales) e v o l v e d d i r e c t l y f r o m a m a r i n e ancestor, p r o b a b l y one c o m m o n to
b o t h m a r i n e fungi a n d red algae. S e c o n d a r y m a r i n e fungi o r i g i n a t e d f r o m terrestrial
ancestors a n d are m o s t l y s a p r o p h y t i c . K o h l m e y e r a n d K o h l m e y e r (1979) agree w i t h
Saville (1968) that parasites are m o r e ancient t h a n saprophytes a n d they consider that
the most ancient types o f m a r i n e A s c o m y c e t e s are S p a t h u l o s p o r a l e s a n d some parasitic Sphaeriales o n m a r i n e R h o d o p h y t a .
Some F W A t a x a i n c l u d e d i n the Halosphaeriales and presumably considered by K o h l meyer a n d K o h l m e y e r (1979) to be o f p r i m a r y o r i g i n i . e . , f r o m a m a r i n e ancestor,
(e.g. Aniptodera,
Halosarpheia,
Nais) o c c u r c o m m o n l y i n freshwater. T h e occurrence o f these t a x a i n freshwater c o u l d m e a n that (1) some m a r i n e A s c o m y c e t e s m i grated f r o m seawater to freshwater, o r (2) some o f the t a x a n o w i n c l u d e d i n the
H a l o s p h a e r i a l e s are not o f p r i m a r y o r i g i n , but e v o l v e d f r o m terrestrial o r freshwater
species. A c c o r d i n g to the first i d e a , these species c o u l d have m i g r a t e d to freshwater
f r o m seawater b y i n t e r t i d a l m i x i n g o r b y t r a v e l l i n g o n the bodies o f m i g r a t i n g waterf o w l . A p l a u s i b l e argument, h o w e v e r , c o u l d be made for their terrestrial o r i g i n a n d
subsequent m i g r a t i o n to freshwater a n d then to seawater. A constant d o w n s t r e a m
m o v e m e n t o f ascomycete substrata occurs a n d it is not unreasonable to expect that
some terrestrial A s c o m y c e t e s a d a p t e d to freshwater a n d then seawater v i a this route.
A s c o m a t a i n the above genera are m a d e o f t h i n - w a l l e d p s e u d o p a r e n c h y m a a n d are
b e a k e d , features w h i c h are characteristic o f several families i n the S o r d a r i a l e s . T h e y
also share the l i g n i c o l o u s habitat w i t h members o f the Lasiosphaeriaceae i n the S o r dariales. A d a p t a t i o n s to freshwater m a y have i n c l u d e d disappearance o f the ascus
a p i c a l apparatus, release o f ascospores t h r o u g h the deliquescence o f asci, m o d i f i c a t i o n o f physes to f o r m chains o f enlarged cells (catenophyses), a n d presence o f viscous ascospore appendages. I n the genus Aniptodera,
for e x a m p l e , v a r i o u s degrees
o f m o d i f i c a t i o n o f the ascus apex c a n be seen f r o m A. chesapeakensis
w i t h a distinct
a p i c a l pore a n d t h i c k e n i n g o f the w a l l o f the ascus apex to A. rosea w i t h no pore
o r t h i c k e n i n g . I n A. chesaspeakensis,
ascospores m a y be f o r c i b l y discharged or m a y
accumulate outside the beak after d i s s o l u t i o n o f the asci. E n v i r o n m e n t a l c o n d i t i o n s
influence the m e t h o d o f ascospore l i b e r a t i o n . I n A. rosea, asci deliquesce early to
release ascospores. C o m p e l l i n g evidence to support either hypothesis about the evo-
Fig. 1. Asci and ascospores of Aquadiscula aquatica from leaves of Acer rubrum submerged in LaRue
Swamp, Union County, Illinois, x 680. Fig. 2. Ascospore of Caryospora callicarpa from wood submerged
in Jordan Creek, Illinois. Note thin layer of gelatinous material surrounding the ascospore. x 870. F i g .
3. Ascospores of an unidentified species of Massarina from wood submerged in Jordan Creek, Illinois.
The thick gelatinous sheath surrounding the ascospore is stained with India ink. x 1,190. F i g . 4. A s c o spore of an unidentified ascomycete from a decorticated oak twig submerged in the River Teign, Devon,
England. A thin outer layer appears to separate from the ascospore wall to form a balloon-like structure,
x 1,700. F i g . 5. Ascospore of Phaeosphaeria typharum from Typha latifolia, St. A n t h o n y Head, C o r n wall, England. The broad gelatinous sheath surrounding the ascospore is stained with India ink. x 1,133.
F i g . 6. Ascospore of Aniptodera chesapeakensis with unfurling appendage from wood submerged in Jordan Creek, Illinois, x 704. F i g . 7. Filiform ascospore of Ophioceras sp. 652-1 from wood submerged in
Allee Creek, Barro Colorado, Panama. x 8 0 0 .
24
25
l u t i o n o f these t a x a does n o t n o w exist. M o r e intensive c o l l e c t i n g o f F W A m a y reveal the existence o f other t a x a related to the H a l o s p h a e r i a l e s , but because o f possible convergent e v o l u t i o n , t r a d i t i o n a l m o r p h o l o g y - d e r i v e d systems o f classification
m a y not be indicative o f the true e v o l u t i o n a r y relationships. B i o c h e m i c a l a n d / o r m o lecular techniques m a y p r o v e to be m o r e i n f o r m a t i v e a n d useful.
F W A have adapted m o r p h o l o g i c a l l y to a q u a t i c habitats i n a v a r i e t y o f w a y s . O n e
type o f m o d i f i c a t i o n involves the presence o n ascospores o f v i s c o u s , sticky a p p e n d ages w h i c h m a y enable ascospores to stick o n t o substrata a n d r e m a i n attached i n
the face o f water m o v e m e n t . Species i n Aniptodera,
Aquadiscula,
Ceriospora,
Ceriosporopsis,
Halosarpheia
a n d Nais a l l have ascospores e q u i p p e d w i t h viscous
appendages. T h e appendages o f species o f Halosarpheia
a n d Aniptodera
( F i g . 6).
u n w i n d to f o r m extremely l o n g threads w h i c h m a y help entangle the ascospore w i t h
substrata (Shearer a n d C r a n e , 1980). M a r i n e A s c o m y c e t e s have evolved a fascinating
array o f ascospore appendages i n v o l v i n g the fragmentation a n d / o r extension o f outer
ascospore walls (Jones a n d M o s s 1978). N o n e o f the appendages r e p o r t e d thus far
for F W A result f r o m these m e c h a n i s m s , rather they are s i m p l e , a p i c a l a n d m u c i l a g i n o u s . O n e e x c e p t i o n m a y be a n u n d e s c r i b e d F W A w i t h ascospores whose outer w a l l
separates f r o m the inner to f o r m a b a l l o o n - l i k e structure a r o u n d the ascospore ( F i g .
4). T h e ascospores o f several l o c u l o a s c o m y c e t e t a x a are e q u i p p e d w i t h gelatinous
sheaths (Caryospora, Leptosphaeria,
Massarina,
Phaeosphaeria,
Pleospora,
Trematosphaeria) (Figs. 2-3,5) o r a p i c a l gelatinous appendages (Ascalgilis,
Lophiostoma,
Rebentischia,
Wettsteininia).
Since most o f the f o r e g o i n g genera c o n t a i n terrestrial
species w i t h ascospore sheaths or appendages, aquatic representatives m a y have been
pre-adapted. W h e n i n t r o d u c e d into water, terrestrial ancestors w i t h ascospore sheaths
or appendages m a y have been better able t h a n species l a c k i n g these features to attach to substrata i n water a n d thus were m o r e l i k e l y to succeed.
M a n y F W A have l o n g f i l i f o r m ascospores ( F i g . 7) w h i c h assume a s i g m o i d shape
i n water ( T a b l e 3). T h e s i g m o i d spore f o r m as a n a d a p t a t i o n to a q u a t i c habitats has
been discussed frequently (Webster a n d D a v e y 1984, W e b s t e r 1987). T h e s i g m o i d
shape is t h o u g h t to increase the area o f o r t h o g o n a l p r o j e c t i o n a n d the l o n g , f i l a mentous f o r m is thought to enhance entanglement w i t h substrata. S i g m o i d ascospores
m a y represent convergent e v o l u t i o n i n that they are f o u n d i n t a x a i n the D i s c o m y cetes {Apostemidium,
Loramyces,
Niptera, Obtectodiscus,
Vibrissea) a n d P y r e n o mycetes (Ophioceras, Gaeumannomyces,
Plagiosphaeria,
Pseudohalonectria)
and L o culoascomycetes (Ophiobolus). A g a i n , since the terrestrial taxa i n some o f these genera
have f i l i f o r m ascospores, it is difficult to resolve whether p r e - a d a p t a t i o n o r convergent e v o l u t i o n o r b o t h have o c c u r r e d .
A s c o m a l a n d ascus structure a n d f u n c t i o n d o not seem to be m o d i f i e d f o r aquatic
habitats. T h e r e is little difference i n these structures between a q u a t i c a n d terrestrial
counterparts. A s m e n t i o n e d p r e v i o u s l y deliquescent asci are present i n some genera
(Aniptodera,
Halosarpheia,
Nais). It is t h o u g h t that this is a n a d a p t a t i o n to water
as ascospores c a n be washed f r o m a s c o m a t a a n d f o r c i b l e discharge is not necessary.
I n some genera that are p r o b a b l y i m m i g r a n t s , such as Chaetomium
and
Zopfiella,
asci are t y p i c a l l y deliquescent. A n interesting s i t u a t i o n exists i n
Pseudohalonectria
(Shearer 1989a) a n d Ophioceras, ( C o n w a y a n d K i m b r o u g h 1978)in w h i c h the asci,
26
Table III. Spore adaptations of freshwater Ascomycetes.
A.
Adaptation of teleomorph and anamorph.
F i l i f o r m ascospore/branched conidium - Vibrissea
flavovirens
F i l i f o r m ascospore with gelatinous sheath/sigmoid conidium - Loramyces
Sheathed ascospore/knobbed
conidium - Massarina
Sheathed ascospore/sigmoid conidium - Massarina
sp.
Sheathed ascospore/tetraradiate conidium - Massarina
B.
juncicola
aquatica
sp.
Adaptation of teleomorph.
Sigmoid or f i l i f o r m ascospores - Apostemidium ( a l l species), Gaeumannomyces
graminis, Gorgoniceps boltonii,
Ophioceras ( a l l species), Pseudohalonectria ( a l l
species), Niptera excelsior, N. melanophaea, N. pulla, Coleosperma lacustre,
Loramyces macrospora, Obtectodiscus aquaticum, Vibrissea ( a l l species),
Cudoniella clavus, Plagiosphaeria
nilotica.
Gelatinous appendage on ascospore - Aniptodera chesapeakensis, A.
lignatilis,
Aquadiscula aquatica, Ascalgilis
bipolaris,
Ceriophora sp., Ceriospora caudaesuis, Ceriosporopsis sp., Halosarpheia lotica, H. parva, H. retorquens, H.
viscosa, Lophiostoma arundinis, L. appendiculata, Rebentischia unicaudata, Nais
inornata, Phaeosphaeria
herpotrichoides.
Gelatinous sheath on ascospore - Loramyces macrospora, Leptosphaeria acuta,
Massarina amphibia, M. aquatica, M. australiensis,
M. tetraploa,
Nectriella
lacustris,
Nimbomollisia melatephroides, N. eriophori, Phaeosphaeria alpina, Ph.
caricinella,
Ph. culmorum, Ph. eustoma, Ph. typharum, Pleospora gaudefroyi, P.
palustris,
P. scirpicola,
P. submersa, Pyrenophora typhaecola, Wettsteinina
niesslii.
C.
Adaptation of anamorph.
Aeroaquatic conidium - Hyaloscypha
lignicola,
H. zalewski
Branched conidium - Hymenoscyphus africanus, H. splendens, H. tetradadius,
varicosporoides,
Miladina lechithina, Nectria lugdunensis
Sigmoid conidium - Hymenoscyphus inberbis,
sp., Orbilia sp., Pezoloma sp.
Mollisia
uda, Mollisia
Helicosporous conidium - Hymenoscyphus paradoxum, Lambertella
gigantea, Tubeufia paludosa.
H.
sp., Nectria
tubulosum,
Mollisia
w h i c h separate f r o m the ascogenous h y p h a e a n d assume a s i g m o i d a l shape, are discharged f r o m the a s c o m a t a . I n some cases, f i l i f o r m ascospores are released inside
the a s c o m a t a a n d ooze out t h r o u g h the b e a k , a n d i n s o m e cases the ascospores are
f o r c i b l y shot f r o m the a s c o m a t a . I n submerged culture, b o t h asci a n d ascospores
are released s i m u l t a n e o u s l y . T h e retention o f ascospores together i n asci p r e s u m a b l y
w o u l d be advantageous i f m a t i n g types were necessary. A l l o f the species w i t h this
type o f discharge pattern, however, are h o m o t h a l l i c . T h e larger size o f the ascus c o m p a r e d to a single ascospore m a y i m p r o v e the chances o f i m p a c t i o n o n a substrate.
T h e m a r i n e ascomycete, Lignincola
laevis H o h n k , also liberates asci f r o m ascogenous
h y p h a e p r i o r to ascospore discharge.
N u m e r o u s F W A m a y have adapted to a q u a t i c habitats v i a their a n a m o r p h s . A n a m o r p h s w i t h b r a n c h e d , tetraradiate, s i g m o i d , h e l i c o s p o r o u s , a n d m u l t i c e l l u l a r airt r a p p i n g c o n i d i a have been reported for F W A (Table 3). O n l y five species have b o t h
ascospores a n d c o n i d i a m o r p h o l o g i c a l l y specialized for the a q u a t i c h a b i t a t . T h e sigm o i d c o n i d i a l f o r m appears to have e v o l v e d i n several different e v o l u t i o n a r y lines;
it occurs i n the D i s c o m y c e t e s , P y r e n o m y c e t e s a n d L o c u l o a s c o m y c e t e s . C o m p a r a b l e
a d a p t a t i o n b y a n a m o r p h s has not o c c u r r e d i n m a r i n e A s c o m y c e t e s . O n l y four s i m i lar t e l e o m o r p h - a n a m o r p h connections have been m a d e for m a r i n e A s c o m y c e t e s , t w o
w i t h tetraradiate, one w i t h s i g m o i d , a n d one w i t h helical c o n i d i a ( K o h l m e y e r a n d
V o l k m a n n - K o h l m e y e r 1991). T h i s is not surprising given that seawater a p p a r e n t l y
selects against c o n i d i a l forms (Shearer 1972).
27
Conclusions
Studies o f F W A are f e w , b u t t h e y i n d i c a t e t h a t F W A c o m p r i s e a diverse g r o u p c o n sisting o f i n d w e l l e r s a n d i m m i g r a n t s . A l t h o u g h there is s o m e o v e r l a p , F W A d i f f e r
t a x o n o m i c a l l y f r o m the m a r i n e A s c o m y c e t e s . F W A v a r y i n t h e i r degree o f m o r p h o l o g i c a l a d a p t a t i o n t o the a q u a t i c h a b i t a t a n d h a v e a d a p t e d m o r p h o l o g i c a l l y p r i m a r i ly b y a s c o s p o r e m o d i f i c a t i o n s a n d / o r c o n i d i a l s p e c i a l i z a t i o n . T h e r e are t w o m a j o r
e c o l o g i c a l g r o u p s o f F W A : parasites o r e n d o p h y t e s o f a q u a t i c a n d w e t l a n d m a c r o p h y t e s , a n d s a p r o p h y t e s o f d e a d p l a n t m a t e r i a l . A d d i t i o n a l s t u d y is needed r e g a r d i n g the systematics, g e o g r a p h i c a l d i s t r i b u t i o n , b i o l o g i c a l a c t i v i t i e s , life histories a n d
responses to e n v i r o n m e n t a l factors o f this c h a l l e n g i n g g r o u p .
Acknowledgements
Appreciation is expressed to D r . A l a n Parker for assistance with literature retrieval and to D r . J . Webster
and D r . J . L . Crane for reviewing the manuscript.
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33
Usnea aurantiacoatra ( J a c q . )
Bory
T w o c o l l e c t i o n s o n v o l c a n i c r o c k , m o s t l y a b o v e 100 m a l t i t u d e (96-09, 111-13). A n
A n t a r c t i c species.
Verrucaria famelica D a r b . *
R a t h e r u n c o m m o n o n v o l c a n i c r o c k , also o n concrete (79-02). T h i s species is k n o w n
o n l y f r o m the A n t a r c t i c r e g i o n . T h e m a t e r i a l f r o m D e c e p t i o n I s l a n d agrees w e l l w i t h
t y p e m a t e r i a l ( S o u t h S h e t l a n d I s l a n d s , N e l s o n I s l a n d , S k o t t s b e r g 11.1.1902, B M ) .
Verrucaria maura W a h l e n b . ex A c h . * *
O n e c o l l e c t i o n o n v o l c a n i c r o c k i n the salt-spray z o n e (117-01). A c o s m o p o l i t a n
species.
Xanthoria candelaria ( L . ) T h . F r .
C o m m o n , m o s t l y o n w o o d , but also o n i r o n , asphalt paper a n d v o l c a n i c r o c k (29-04).
A
c o s m o p o l i t a n species.
Xanthoria elegans ( L i n k ) T h . F r .
R a t h e r u n c o m m o n , o n v o l c a n i c r o c k o n l y (96-03). A c o s m o p o l i t a n species.
Acknowledgements: W e would like to thank D r . D . Galloway and D r . R . Huxley for their permission
to investigate partly unprocessed historic collections and D r . P . Diederich for the identification o f the
Arthonia, the Trimmatothelopsis and the parasites. D r . S. Stenroos is thanked for the identification o f
the Cladonia. The investigations were supported by the Netherlands Marine Research Foundation (S.O.Z).,
which is subsidized by the Netherlands Organization for Scientific Research ( N . W . O ) .
5.
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