Some fossil fungal form-taxa from the Maastrichtian and Palaeogene ages

Abstract

Some distinctive dispersed fungal spores of Maastrichtian to Oligocene age from the Amphitheatre Formation, Yukon Territory, Canada, the Iceberg Bay Formation at Strand Fiord and Kanguk Peninsula on Axel Heiberg Island, Northwest Territories, Canada, and the Deccan Intertrappean Beds of India are described. Except for Delitschia-type ascospores, all fungal spores are considered to be conidia. Five new fossil form-genera are described and have been named according to their affinities to modern taxa. Xylohyphites verrucosa, with affinity to Xylohypha, is described from permineralized fruits of Viracarpon. Ampulliferinites axelheibergi, Helicoonites goosii, Helicosporiates pirozynskii and Paleoslimacomyces canadensis are described for Ampulliferina-type, Helicoon-Helicodendron-type, Helicosporium-type and Slimacomyces-type conidia. New species in existing fossil genera including Piriurella alternariata, Diporicellaesporites icebergi, Brachysporisporites endophragmia and Pluricellaesporites excipularis are described for conidia with affinities to Alternaria, Diplococcium or Bispora, Endophragmia and Excipularia respectively. Conidia resembling those of the modern genus Uberispora are described but not named. Dicellaesporites delitschiapites sp. nov. is described for ascospores with affinity to Delitschia.

Full text

Mycal.
Res.
99
(5):
513-522
(1995)
Prinled
in
Greal
Brilain
Some fossil fungal form-taxa from the Maastrichtian and
Palaeogene agest
R.
M.
KALGUTKAR
1'"
AND
LYNNE
SIGLER
2
l
Instilule
of
Sedimenlary
and
Pelroleum
Geology,
Geological
Survey
of
Canada,
Calgary,
Alberla,
Canada
T2L
2A7
2
Universily
of
Alberla
Microfungus
Collection
and
Herbarium,
Devonian
Bolanic
Garden,
Edmonlon,
Alberla,
Canada
T6G
2El
Some
distinctive dispersed
fungal
spores
of
Maastrichtian to Oligocene
age
from
the
Amphitheatre
Formation,
Yukon
Territory,
Canada,
the
Iceberg
Bay
Formation
at
Strand
Fiord
and
Kanguk
Peninsula
on
Axel
Heiberg
Island,
Northwest Territories,
Canada,
and
the
Deccan
Intertrappean
Beds
of
India
are
described.
Except
for
Delilschia-type
ascospores,
all
fungal
spores
are
considered to
be
conidia.
Five
new
fossil
form-genera
are
described
and
have
been
named
according
to
their
affinities
to modem
taxa.
Xylohyphites
verrucosa,
with
affinity
to
Xylohypha,
is
described
from
permineralized
fruits
of
Viracarpon.
Ampulliferinites
axelheibergi,
Helicoonites
goosii,
Helicosporiates
pirozynskii
and
Paleoslimacomyces
canadensis
are
described
for
Ampulliferina-type,
Helicoon-Helicodendron-type,
Helicosporium-type
and
Slimacomyces-type
conidia.
New
species
in
existing
fossil
genera
including
Piriurella
alternariala,
Diporicellaesporiles
icebergi,
Brachysporisporites
endophragmia
and
Pluricellaesporites
excipularis
are
described
for
conidia
with
affinities
to
Alternaria,
Diplococcium
or
Bispora,
Endophragmia
and
Excipularia
respectively.
Conidia
resembling
those
of
the
modem genus
Uberispora
are
described
but not
named.
Dicellaesporiles
delitschiapiles
sp.
nov.
is
described
for
ascospores with
affinity
to
Delitschia.
513
In
recent years information
on
fossil fungi has increased
rapidly, providing
not
only significant data
on
the diversity of
fungi
in
the fossil record but also on the potential relationships
that exist among the major groups of fungi (Taylor &White,
1989). The study of fossil fungi
is
generally shunned
by
palaeobotanists and palynologists because of difficulties
encountered in identifying and interpreting dispersed material.
The geological record of fossil fungi
is
rich. Progressive
studies of their evolutionary history and diversity continue
to
demonstrate their importance. The application of fossil fungi
in
biostratigraphy has increased with discoveries of charac-
teristic and unique spores having limited stratigraphic ranges.
Remains of fossil fungi are also an important source of
information for interpreting past environments
if
their affinities
to extant species have been determined. As familiarity with
extant fungi
is
inevitably necessary
to
accomplish this purpose
successfully, collaborative work with mycologists can assist in
obtaining valuable results.
It
is
now
widely accepted that an emphasis
on
the
combined studies of fossil and living fungi
is
necessary
to
realize the
full
potential of fossil fungal palynomorphs as
predictors of past environments (Wolf, 1966a,
b,
1968, 1969;
Lange &Smith, 1971; Pirozynski, 1976; Ramanujam &Rao,
1978; Ramanujam &Srisailam, 1980; Jarzen &
Elsik,
1986;
Pirozynski
et
aI.,
1988; Kalgutkar &McIntyre, 1991).
Pirozynski (1976) stated
that'
evidence accumulates to support
•
Corresponding
author.
tGeology Survey of Canada contribution No. 24493.
the long-held view that the history of fungi
is
not
marked by
change and extinctions but
by
conservatism and continuity.
Thus fungi
in
fossil remains are likely
to
be sensitive
interpreters of past environments rather than index fossils'.
In
the Tertiary, many distinctive fungal spores show
affinities
to
modem
genera.
At
the species level, however, it
seems from accumulating work that fossil fungal forms rarely
correspond exactly to
modem
spores. This can be attributed
to
various factors including the rapid evolution in fungi
accompanied
by
ontogenetic and morphological changes, and
the total extinction of species
by
ecological, edaphic and biotic
changes which affected their substrata and consequently their
saprotrophic and parasitic modes of nutrition.
This paper describes some distinctive dispersed spores,
considered to be either ascospores or conidia of Palaeogene
and Maastrichtian age, encountered in specimens collected
from the Amphitheatre Formation, Yukon Territory, Canada,
the Iceberg Bay Formation at Strand Fiord and Kanguk
Peninsula, Northwest Territories, Canada and the Deccan
Intertrappean Beds of India
(Fig.
1).
Characteristics of spore
size and shape, septal nature and thickenings, wall sculpture,
absence or presence and number and nature of apertures
or
,pores', and other characters of diagnostic significance were
used to classify them in existing or new fossil form-genera and
to assess their possible affinities
to
modern taxa.
In
fossil
studies, an apparent opening or aperture in aspore
is
referred
to
as
a
'pore'
regardless of its functional attribute,
i.e.
as
germinalus or point of attachment. The form taxa of the
Sporae Dispersae (Sheffy &Dilcher, 1971;
Elsik,
1976) are

Fossil
fungi
514
Yukon Territory Arctic Islands
India
Arctic
Ocean
*Kangu~
••
"......._
Peninsula
•Strand
u:
L:J
,\
I
• I
•'{i •Amphitheatre Mountain
-1'&. I
I
~
•Whitehor e \
.~,
\
.
\"
Alaska
l40·W
Fig. 1. Map showing locations of samples collected from Amphitheatre Mountain (61 0
16'
N,
139°
21'
W),
Yukon Territory, Strand
Fiord (790
14'
N,
91°
21'
W)
and Kanguk Peninsula (79°
IS'
N,
92°
33'
W),
Northwest Territories and Mohgaon
Kalan
(220
30'
N,
79020'
E),
Chhindwara District, India.
grouped according to Saccardos spore group (Hawksworth,
Sutton &Ainsworth, 1983) for convenience.
MATERIALS
AND
METHODS
Samples were collected from the Iceberg
Bay
Formation on
Axel Heiberg Island at Strand Fiord (790
14'
N,
910
21'
W;
Early Eocene) and Kanguk Peninsula (790
15'
N, 92033' W;
late Palaeocene or early Eocene), from the Amphitheatre
Formation
in
southwestern Yukon Territory
(61
016'
N,
139021' W; late Eocene or early Oligocene) and from the
Deccan Intertrappean
Beds
of the Maastrichtian Mohgaon
Kalan locality, Chhindwara District of India (22 0
30'
N,
79° 20'
E).
The ages given are based on the associated
angiosperm pollen and fungal spores. The samples were
prepared by using standard palynological procedures, modified
by using aweak or diluted Schultze's solution to control the
rate of oxidation, atechnique found to improve the recovery
of fungal spores (Kalgutkar, 1993). The residues of unsieved
(UN),
+45 Iffil, +20
IJJT1
and -20
IJJT1
were mounted on
slides. Coordinates
of
all
figured specimens are from aCarl
Zeiss transmitted light microscope with camera attachment
located at the Institute of Sedimentary and Petroleum Geology
in
Calgary, Alberta.
All
slides bearing the figured specimens
including the holotypes will be permanently stored
in
the type
collection of the Geological Survey of Canada
in
Ottawa.
DESCRIPTIONS
OF
TAXA
Spore group: Amerosporae
Xylohyphites Kalgutkar &Sigler gen. nov.
Type
species:
XyIohyphites verrucosa
Conidia
in
chains formed
in
acropetal succession, simple,
ellipsoidal or cylindricaL pale brown, verrucose, aseptate,
occasionally I-septate; conidia tapering at each end but
sometimes with slightly protuberant hilum.
Derivation
of
name:
Affinity to
Xylohypha
(Fr.)
E.
W. Mason emend.
Deighton, 1960.

R.
M.
Kalgutkar and Lynne Sigler 515
Figs 2&3.
Xylohyphites
verrucosa.
Fig. 2, holotype. esc 106532,01-10,
112-1
x8'7, esc
loc.
01-10, Fig. 3, esc 106532, 01-10,
112'1 x8'7, esc
loc.
01-10. Note that in
all
legends the species name
is
followed
by
the esc type number, the slide number, stage
coordinates, the esc locality.
Bar
markers =10
IJm;
Figs
3,
6-14
and 16-22 are to the same scale (bar on
Fig.
14),
Figs
2,
4,
5and
15
are to the same scale (bar on
Fig.
IS).
Comments:
Although the cylindrical form of the conidia and
arrangement
in
chains initially suggested an arthroconidial
type of development, the presence of asingle bud-like
proliferation
(Fig.
3, arrow) suggested an affinity to a
dematiaceous blastoconidial fungus. Three modem genera
in
which blastic chains of dematiaceous conidia are formed from
more or less undifferentiated hyphae are
Xylohypha,
Bispora
Corda and
Septonema
Corda.
Xylohypha
is
separated from the
latter genera by having one-celled conidia (Deighton, 1960),
but species within the genus differ
in
this respect. Conidia of
those species which can be grown on agar tend to disassociate
with difficulty and there may be fragments of different length
and pattern of septation (Hughes, 1972; Padhye
et
al.,
1988;
Kwon-Chung, Wickes &Plaskowitz, 1989). Conidia of
Xylohyphites
differ from
all
modem species of
Xylohypha
in
being verrucose.
The occurrence of
Xylohyphites
verrucosa
as
asaprotroph
in
fruits of
Viracarpon
and its verrucose conidia
in
chains also
suggest apossible affinity with
Cladosporium.
This modem
genus
is
widely associated with plant materials of various
kinds including fruits. However, no evidence could be found
of aconidiophore among the tangled chains. Moreover, the
conidia lack the prominent hila and detachment scars of
typical
Cladosporium
conidia.
Xylohyphites
verrucosa
Kalgutkar &Sigler
sp.
nov.
(Figs
2,
3)
Holotype:
esc 106532, esc
loc.
DI-1O
Characteristics
as
in
generic diagnosis. Conidia
9-16
x
2'5-5'5
IJm.
Derivation
of
name:
Verrucose nature of conidia.
Occurrence:
In
permineralized fruits of
Viracarpon,
amonocotyledonous
fruit consisting of acephalium borne on apeduncle to which
compound drupes (phalanges) are attached (Kalgutkar
et
aI.,
1993);
collected from Deccan Intertrappean Beds of Mohgaon
Kalan
Locality, India; Age: Maastrichtian; Locality: 01-10.
Comments:
Conidia are similar to those of
Xylohypha
nigrescens
(Pers.)
E.
W.
Mason
in
size (mostly
10-13'7
X
3'7-5
Ilffi,
fide
Hughes, 1972) and shape, but differ
in
being verrucose.
Spore group: Didymosporae
or
Phragmosporae
Ampulliferinites Kalgutkar &Sigler gen. nov.
Type
species:
Ampulliferinites axelheibergi
Fungus filamentous; filaments
in
short or long, determinate
chains of conidia; filaments traversed by thin and thick-walled
septa which alternate along the chains; conidia
in
arthro-
conidial chains separated by dark, thick septa; didymosporous,
not or slightly constricted at median septum and with median
septum thinner, smooth, brown, cylindrical with truncate ends
except the terminal conidium which
is
rounded at its apex.
Derivation
of
name:
Affinity
Ampulliferina
B.
Sutton.
Comments:
Ampullijerinites
appears to be similar to modem
Ampullijerina
(Sutton, 1969) which has didymosporous,
catenate conidia that separate by fragmentation through the
thick, dark brown septa along its length of arthroconidial
chains.
Ampullijerinites
is
also characterized by the presence of
abasal
cell
with an attachment scar similar to one present

Fossil fungi
516
8
12
21
6
10
5
17
Figs
4-22.
Figs 4
&:
5.
Ampulliferinifes
axe/heibergi.
Fig. 4, hololype. esc 106520, 3473-19
(+
20), 96'9 x19'0, esc lac. C-189109.
Fig.
5,
esc 106521, 3473-19
(+
20),
100'6 x14'0, esc lac. C-189109. Fig.
6.
Dicellaesporifes
delifschiapifes,
holorype. esc 106524,
2614-126
(-
20), 98'0 x3'5, esc lac. C-112393. Figs 7
&:
8.
Diporicellaesporites
icebergi.
Fig.
7,
holotype. esc 106522, 2614-129
(-
20),
86'0 x15'4, esc lac. C-112397. Fig. 8, esc 106523,2614-126
(UN),
91'0 x5'9, esc lac. C-112393. Figs 9&
10.
Brachysporisporites
endophragmia.
Fig. 9, holorype. esc 106525, 2614-126 (UN), 92'1 x5'9, esc lac. C-112393. Fig.
10,
esc 106526,

R.
M. Kalgutkar and Lynne Sigler 517
in
Ampulliferina.
Bispora
also produces catenate two-celled
conidia, but the extension growth
is
blastic, the conidia are
more strongly constricted at the fragmentation septa, and the
conidia have very dark brown to black bands at each septum
(Ellis,
1971).
Ampulliferinites
axelheibergi
Kalgutkar &Sigler sp. nov.
(Figs
4,
5)
Holofype:
esc
106520,
esc
loc.
C-189109.
Chains comprising
5-8
conidia straight and unbranched and
slightly constricted at the thick separating septa. Filamentous
chains
79-240
!-1m
long. Didymosporous conidia occurring in
arthroconidial chains separated by thick, dark-brown septa
which alternate with the thin septa along the length of the
filament. Filaments short to long, determinate; subtended
by
abasal cell. Basal cell pale brown, aseptate, tapering towards
the base with
an
attachment scar left after the detachment of
the filament from the mycelium. Conidia two-celled, not
or
slightly constricted at septum, smooth, brown, truncate at
both thickened ends,
16-22
x
7-11
!-1m.
Detached conidia
were not found.
Occurrence:
Iceberg
Bay
Formation at
Kanguk
Peninsula,
Axel
Heiberg
island,
Northwest Territories; Age:
late
Palaeocene
or early
Eocene.
Locality:
C-112393.
Comments: Spores of
Dicellaesporites
delitschiapites
are generally
similar
to
the ascospores of the liVing loculoascomycetous and
coprophilous
De/itschia
in the presence of furrows which
appear similar to the distinctive germ slits. The ascospores of
modem
De/itschia
are generally surrounded
by
conspicuous
gelatinous sheaths. These probably were lost either during
fossilization or laboratory processing. Athin hyaline layer
around each spore of
Dicellaesporites
delitschiapites
is
discernible
under high magnification (not shown) although its gelatinous
nature may be questionable. Jarzen &
Elsik
(1986) have
described Delitschia-type spores with
good
illustrations from
river deposits, Luangwa Valley, Zambia.
Diporicellaesporites
Elsik,
1968
Type
species:
D. stacyi
Elsik
1968.
Diporicellaesporites
icebergi
Kalgutkar &Sigler sp. nov.
(Figs
7,
8)
Derivation
of
name:
After
its
type locality
on
Axel
Heiberg
Island.
Occurrence:
Amphitheatre
Formation,
Yukon
Territory; Age:
late
Eocene
or
early Oligocene.
Locality:
C-189109.
Comments:
Ampulliferinites
axe/heibergi
resembles Alternoseptites
e/ongatus
Rouse (1962) in having filaments indented at every
second septum, but in
A.
elongatus
the septa are not thickened
and filaments are
not
arthroconidial
in
nature.
Holofype:
esc
106522,
esc
lac.
C-I12397.
Conidia simple cylindrical, brown, smooth to minutely
verruculose, diporate, catenate,
1-3
septate; septa thicker than
conidium wall, about 2
!-1m
thick, dark brown, with narrow
septal pores
in
the centre. Septal thickening extending
laterally alittle along the spore wall
on
either side. Pores
or
areas of contact between conidia at
both
ends dark, simple,
slightly convex. Mature conidia
27-42
x
5-7
I-lffi. Two-celled
conidia
12-16
x
5-7
!-1m.
Dicellaesporites
Elsik 1968 emend. Sheffy &Dilcher, 1971
Derivafion
of
name:
After
Iceberg
Bay
Formation.
Comments:
Diporicellaesporites
icebergi
may be compared with
Diplococcium
Grove 1885
or
Bispora
in
which conidia are
predominantly two-celled and have abroad, dark-brown band
at the septum. The conidia of
both
Bispora
betulina
(Corda)
S.
Hughes and
D.
spicatum Grove are borne in chains through
pores and are rounded
at
each end or slightly convex. The
septa of
D.
spicatum are marked
by
conspicuous septal pores,
rather similar
to
those of
Diporicellaesporifies
icebergi
shown
in
Figs 7and
8.
There
is
no evidence of blastic conidial
development
in
the fossil specimen.
Derivafion
of
name:
Affinity
Delifschia
Auersw.
Type
species:
D. popovii
Elsik,
1968.
Holofype:
esc
106524,
esc
lac.
C-112393.
Occurrence:
Iceberg
Bay
Formation
at
Kanguk
Peninsula,
Axel
Heiberg
Island,
Northwest Territories; Age:
late
Palaeocene
or
early
Dicellaesporites
delitschiapites
Kalgutkar &Sigler sp. nov.
Eocene.
Locality:
C-I12393
and
C-I12397.
(Fig.
6)
Spores broadly
ellipSOid
with rounded ends, 2-celled,
aporate, brown, without aconstriction at the septum; septum
nearly median; spores with afurrow or germinal slit; germinal
slit lateral, slightly gaping and about two-thirds the length of
the spore; spore wall smooth, surrounded
by
an hyaline,
narrow layer. Spores
25-28
x
11-13
!-1m.
2614-126
(-20),
100'7 x6'8, esc
loc.
C-112393.
Fig. 11.
Pluricellaesporifes
excipularis,
hoJotype.
esc
106527,
2614-128
(UN),
85'7
x
9'0,
esc
loc.
C-112396.
Fig. 12.
Uberispora-type
A. esc
106530,
2614-126
(UN),
93'0 x
4'5,
esc
loc.
C-112393.
Fig. 13.
Uberispora-type
B.
esc
106531.
2614-134
(UN),
94'1
x
4'9,
esc
loc.
C-112402.
Figs 14 &15.
Piriurella
alternariafa.
Fig. 14, holotype.
esc 106518, 2614-126
(UN),
92'8 x
8'0,
esc
loc.
C-112393.
Fig. 15, esc
106519,
2614-126
(+
20),
92'9 x5'1, esc
loc.
C-112393.
Figs 16 &17.
Helicoonites
goosii.
Fig. 16, holotype.
GSC
97291.
2614-57
(+
20),
87'7 x13'5, esc
loc.
C-111847.
Fig. 17,
GSC
97292,
2614-57
(+
20),
79'0
x
15'7,
esc
loc.
C-111847.
Figs
18-20.
Helicosporiafes
pirozynskii.
Fig.
18,
holotype, esc
97294,
2614-57
(+
20),
76-0
x3'5, esc
loc.
C-111847.
Fig.
19,
esc
97295,
2614-57
(UN),
101'8 x6'9, esc
lac.
C-111847.
Fig.
20,
esc
97296,
2614-57
(+
20),
86'0 x12'4, esc
loco
C-111847.
Figs
21
&
22.
Paleoslimacomyces
canadensis.
Fig.
21,
holotype, esc
106528,
2614-126
(UN),
96'8 x
4'2,
esc
loc.
C-112393.
Fig.
22,
esc 106529, 2614-126
(UN),
94'8 x
14'4,
esc
lac.
C-112393.

Fossil fungi
Spore group: Phragmosporae
Brachysporisporites
R.
1.
Lange &
P.
H.
Sm.,
1971
Type
species:
B.
pyriformis
R.
T.
Lange
&
P.
H.
Sm.,
1971.
Brachysporisporites
endophragmia
Kalgutkar &Sigler
sp.
nov. (Figs
9,
10)
Holotype:
esc
106525,
esc
loco
C-112393.
Conidia simple, obovoid to clavate, brown, smooth. Conidia
2-4
septate, central cells darker than the subhyaline to hyaline
cell at the proximal end; cells with broad dark bands at the
septa; cell wall slightly notched and thickened at the septa;
basal cell with apore or an attachment scar at the end. Conidia
27-34
~m
long,
12-19
~m
thick in the broadest part and
5-6
~m
wide at the truncate base.
Derivation
of
name:
Affinity
Endophragmia
Duvernoy &
Maire,
1920
emend
Hughes,
1979.
Occurrence:
Iceberg
Bay
Formation
at
Kanguk
Peninsula,
Axel
Heiberg
Island,
Northwest Territories; Age:
late
Palaeocene
or
early
Eocene.
Locality:
C-112393.
Comments:
The conidia of
Brachysporisporites
endophragmia,
with their truncated ends and broad thickened septa, compare
favourably with conidia of the living
Endophragmia.
Hughes
(1979) revised
Endophragmia
after transferring several of its
species
to
other genera including anumber
to
Endophragmiella
B.
Sutton, (1973). The truncated proximal end and absence of
abasal protuberance separate the conidia of
Brachysporisporites
endophragmia
from those of
Endophragmiella.
Pluricellaesporites
Hammen,
1954
emend.
Elsik
&Janson.,
1974
Type
species:
P. typicus
Hammen,
1954.
Pluricellaesporites
excipularis Kalgutkar &Sigler sp. nov.
(Fig.
11)
Holotype:
esc
106527,
esc
lac.
C-112396.
Conidia solitary, multiseptate, dark brown except for terminal
and basal cells which remain distinctively hyaline
or
pale.
Conidia fusiform,
5-6
septate, smooth; septa
dark
about
4-5
~m
thick. Conidia truncate at point of attachment; basal
cell not porate; terminal cell with thin, entire and clear wall.
Conidia
46-48
x18'5
~m.
Derivation
of
name:
Named
for
apparent
affinity
to
Excipularia
Sacc.
Occurrence:
Iceberg
Bay
Formation
at
Kanguk
Peninsula,
Axel
Heiberg
Island,
Northwest Territories; Age:
late
Palaeocene
or
early
Eocene.
Locality:
C-112396.
Comments:
The conidia of
Pluricellaesporites
excipularis
appear
morphologically similar to those of the modem genus
Excipularia
in shape, septation and in the differentiated hyaline
basal and terminal cells.
Spore group: Staurosporae
or
Phragmosporae
Uberispora-type A
(Fig.
12)
Illustrated
specimen:
esc
106530,
esc
lac.
C-112393.
Conidia complex, solitary, light brown to subhyaline,
composed of three linear and two
or
more lateral cells; cells
518
equidistant, terminal cell semi-circular, basal cell triangular and
central cell rectangular; basal cell with
an
apparent apiculus.
Conidia thin-walled, subhyaline, characteristically with papil-
late lateral cells arising from the central cell at the septa.
Conidia multi-septate; septa dark and thick. Conidial surface
rough and scabrid; overall size about
30
x18
~m.
Uberispora-type B
(Fig.
13)
Illustrated
specimen:
esc 106531, esc
loc.
C-112402.
Conidia complex, solitary, terminal, brown; consist of three
linear and one or more lateral cells; cells smooth walled, upper
and basal cells rounded, middle cell compressed and seemingly
subrectangular. Lateral cells similar to those on the main axis,
not papillate or thin-walled. Basal cell with apore-like opening,
indicating the point of attachment of the conidium to a
conidiogenous
cell.
Conidia multi-septate; septa thick and
dark. Conidia 31 x
18
~m.
Occurrence:
Iceberg
Bay
Formation
at
Kanguk
Peninsula,
Axel
Heiberg
Island,
Northwest Territories; Age:
late
Palaeocene
or early
Eocene.
Locality:
C-112393.
Comments:
Both
Uberispora-type
Aand
Uberispora-type
B
spores are distinctive and peculiar in the fossil fungal
assemblage from this site.
Uberispora-type
Adiffers from
Uberispora-type
B
in
the nature of the cell wall, and in the
development of lateral cells.
In
the former, the surface
is
rough
and scabrid, and the spores have papillate, thin-walled lateral
cells whereas in
Uberispora-type
Bthe surface
is
smooth and
lateral cells are larger, thicker and non-papillate. They resemble
living
Uberispora
simplex
(Ichinoe) Piroz. &Hodges in their
lateral branching pattern and characteristic form. This affinity
was noted also by Drs
K.
Pirozynski (pers. comm. 1992) and
S.
J.
Hughes (pers. comm. 1992). Pirozynski &Hodges (1973)
established
Uberispora
with type species
U.
simplex
after
determining and redescribing the fungus
Arachnophora
simplex
Ichinoe originally described from Japan (Ichinoe, 1972).
Hughes (1979) also briefly described
Uberispora
as
having
Endophragmiella-like
proliferations after rhexolytic secession
of
the conidia.
Uberispora
simplex
is
characterized
by
acentral,
thick-walled, dark-brown angular
cell,
one apical and three
lateral hyaline, thin-walled conical cells and apale brown
conico-truncate basal
cell.
In
Uberispora-type
Aand
Uberispora-
type Bthe main axis consists of three cells and irregularly
budding lateral cells arise mainly from the central
cell.
Both
fossil types are larger than the conidia of
U.
simplex.
Spore group: Dictyosporae
Piriurella Cookson &Eisenack,
1979
Type
species:
Piriurella elongata
Cookson
&
Eisenack,
1979.
Piriurella
alternariata
Kalgutkar &Sigler sp. nov.
(Figs 14, 15)
Holofype:
esc
106518,
esc
loc.
C-112393.
Conidia arising singly
or
in clusters; multicellate, muriform,
solitary, ovoid to obclavate, rostrate, cicatrized
or
not, pale
brown to brown, smooth. Conidia with ashort conical beak
and
8-12
transverse and several longitudinal
or
oblique septa;
transverse septa more prominent and thicker than the

R.
M. Kalgutkar and Lynne Sigler
longitudinal or oblique septa; terminal conical beak about
9-11
~m,
broad with aconspicuous dark thickened tip probably
representing the point of origin (attachment scar) of the
successive spore in the conidial chain. Conidia when cicatrized,
with ascar at the proximal end at the point of attachment to
the conidiophore. Conidia
42-74
~
long,
18-27
~m
wide in
the broadest part.
Derivation
of
name:
Affinity to
Alternaria
Nees, 1817.
Occurrence:
Iceberg
Bay
Formation at Kanguk Peninsula, Axel
Heiberg Island, Northwest Territories; Age: late Palaeocene or early
Eocene.
Locality: C-1l2393.
Comments:
Cookson &Eisenack (1979) described
Piriurella
and
the type species
P.
elongata
from the Cretaceous of the
Eucla
basin, Australia
as
an algal form similar to modem
Lambertia.
Smith &Chaloner (1979) demonstrated the difference
in
structure between
Piriurella
and
Korshikoviella
schaefernai
(=
Lambertia
schaefernai)
and concluded that
Piriurella
is
aconidium
of
Alternaria.
Elsik
(1992) indicated its similarity to
Alternaria
and commented that it has priority in the event aform genus
is
created, assuming it
is
considered afossil and not amodem
contaminant.
As
PiriurelJa
is
considered to include fossil fungal spores
with
an
affinity to modem
Alternaria,
the Alternaria-type
conidia described here are described under
PiriurelJa.
However,
the senior author
is
not convinced that the type species,
Piriurella
elongata,
has the essential characters that distinguish
Alternaria
conidia,
i.e.
abroader basal region with transverse
and longitudinal septation and adistally tapering short or
cylindrical beak with or without adark thickened tip. The
antapical tapered portion described for
P.
elongata
appears like
aremnant or residual portion of aconidiophore or ahypha
subtending the spore. Re-examination of the holotype of
P.
elongata
is
necessary to resolve this question and
if
it should
prove to
lack
the essential characteristics of an
Alternaria
phragmospore, anew genus would be required for fossil
Alternaria-like
dispersed spores.
The taxonomy of modem
Alternaria
is
difficult due to the
variability in shape, size and septation of conidia even within
the species (Barron, 1983). Emphasis on sporulation patterns
in
species determination (Simmons &Roberts, 1993) makes
assessment of affinities between fossil forms and modem taxa
difficult. Solitary conidia, however, can possibly be identified
as
allied to
Alternaria
if
they are mature and have diagnostic
features such
as
the beaks similar to those found in modem
species. The muriform conidia with short conical beaks and
dark thickened tips suggest this affinity for
Piriurella
alternariata.
The conidia of
Piriurella
elongata
differ from those of
P.
alternariata
in
being elongate with arather long, tapered,
gradually differentiated, antapical end. Also, the spore
is
not
muriform and lacks athickened
beal<.
Trevedi &Verma (1970)
described
Alternaria
malayensis
from the Eocene of Malaya and
strongly suggested the affinity of its spores to the conidia of
Alternaria.
However, the description and illustrations
(p.
71,
pI.
1,
fig.
13; text-figs
14-16)
do
not
appear similar or even
closely related to the conidia of
Alternaria.
Spores of
Staphlosporonites
species described and illustrated by Takahashi
(1991,
pI.
1,
figs
10-11)
appear similar to the conidia of extant
519
Alternaria
with distally extended beaks. Ediger (1981)
described the new fossil genus
Transeptaesporites
to include
spores resembling the conidia of
Alternaria.
However, it
becomes asynonym of
Piriurella.
Spore group: Helicosporae
The characteristic and distinctively coiled, septate spores
found in the samples from Strand Fiord and Kanguk Peninsula
in
Axel Heiberg Island are easily recognized
as
conidia of
helicosporous fungi. The genera to which they are referred
belong to agroup of aero-aquatic Hyphomycetes, which have
peculiar conidia devised to trap
air.
This assists
in
their
dispersal when the substratum
is
submerged under water
(Webster, 1980). These fungi do
not
sporulate when
submerged but do so at the moist interface between air and
water. Helicosporous fungi have awide geographical
distribution and apparently form the most conspicuous fungal
biota on submerged leaves decaying under relatively anaerobic
conditions and on wood and bark lying on the ground
(Kalgutkar &Mcintyre, 1991).
In
an extensive account of the helicosporous Hypho-
mycetes, Goos (1987) commented on the occurrence of a
unique helicoid conidial structure which can be easily
recognized and used to circumscribe this group of fungi.
Extant helicosporous genera and species are classified on the
basis of the morphology and ontogeny of their conidiophores
and conidia and the nature and development of their mycelia.
However, when the fungus and its substratum are dispersed
during deposition
as
in the fossil sediments, the conidia are
usually the only part of the organism that can be recognized.
Three forms of helicosporous conidia are described which
resemble conidia of the modem genera of either
Helicoon
Morgan or
Helicodendron
PeyroneL
Helicosporium
Nees and
Slimacomyces
Minter. These affinities were suggested
by
Pirozynski (pers. comm. 1992) and Goos (pers. comm. 1993).
Although similar helirosporous fossil genera have been
described previously, including
Involutisporonites
R.
T.
Clarke
(1965) emend.
Elsik
(1968),
Colligerites
K.
P.
Jain
&
R.
K.
Kar
(1979), and
Helicominites
Barlinge &Paradkar (1979), the
morphological characteristics of the conidia from the present
samples are sufficiently distinct to require new genera.
Elsik
(1992) recorded conidia such
as
these, having acoiled axis
forming aplanispiral to conical helix, but has
not
yet validly
published anew genus.
Kalgutkar (1993) indicated how fossil fungal types, which
are morphologically similar to modem equivalents and have a
limited habitat preference, could be important
in
assessing the
palaeoenvironment
by
relating the known environment of the
extant taxa to similar fossil forms. In this context, the
importance or helicosporous fungi
in
palaeoenvironmental
interpretations has been demonstrated
by
Ramanujam &
Rao (1978), Rarnanujam &Srisailarn (1980), Gray (1985),
Sherwood-Pike (1988) and Kalgutkar &McIntyre (1991).
Helicoonites Kalgutkar &Sigler gen. nov.
Type
species:
Helicoonites goosii
Conidia simple, tightly coiled or twisted
in
three planes to
form an ovoid, ellipsoidal (doliiform) to cylindrical or beehive-

Fossil fungi
to barrel-shaped spiral; spirals made up of variable numbers of
ascending coils or gyres, with each successive gyre usually of
smaller diam.; filaments multiseptate, fuscous;
cells
rectangular.
Comments:
Described and illustrated previously
as
Helicoon-
Helicodendron-type
(Kalgutkar &McIntyre, 1991, their Figs
1-3). Although it
is
easy to identify ahelicosporous conidium
because of its distinctively twisted structure, the conidia of
some species of
Helicoon
and
Helicodendron
are
so
similar
in
appearance that it becomes impossible to separate them on the
basis of their morphology alone. The most significant character
separating the two extant genera
is
the mode of conidial
development.
In
Helicodendron,
the conidia are proliferating,
forming catenate chains, or tangled masses or clusters,
whereas
in
Helicoon
the conidia are produced
Singly.
In
both
genera, the conidia could be aerogenous or acropleurogenous
and doliiform with tightly coiled spirals.
In
fossil sediments,
however, during deposition such secondary conidial aggre-
gates are unlikely to be retained on the mycelial tissue or
conidiophores and are generally preserved
in
scattered
condition. Therefore, the tightly coiled conidia of species of
Helicodendron
are extremely difficult
to
distinguish from the
similar conidia of
Helicoon
(Kalgutkar &McIntyre, 1991).
Helicoonites
is
created to encompass
all
fossil helicosporous
conidia with adefinite ellipsoidal, ovoid or doliiform shape
and which are loosely to tightly coiled or twisted
in
three
planes.
Helicoonites
differs from
Involutisporonites,
Colligerites
and
Helicominites,
and from
Helicosporiates
gen. nov. and
Paleo-
slimacomyces
gen. nov. by its helically coiled filaments
in
ellipsoidal to beehive-shaped spirals. Except
Helicominites,
in
these genera the conidia are generally
flat
and coiled
in
one
plane.
In
Helicominites
the conidia are bent or twisted
in
different directions.
Goos
et
al.
(1986) mentioned the occurrence of anearly
650-yr-old specimen from an archaeological site near Beetley,
Norfolk, England, which appeared to be assignable to modem
Helicoon
richonis
(Boud.)
Linder (1929). The conidia of this
subfossil were slightly larger than those of the extant species,
but otherwise appeared similar
in
shape and colour.
Helicoonites
goosii
Kalgutkar &Sigler
sp.
nov.
(Figs
16,
17)
Holotype:
GSC
97291, GSC
loc.
111847.
Conidia
Simple,
smooth, tightly coiled
in
three planes to form
an ellipsoidal (doliiform) to cylindrical spore body; filaments
multiseptate, about 5
~m
wide, fuscous; cells rectangular, not
incurved at the septa, transverse pattern of minute, apparently
internal striae commonly present. Conidia 50-75 x37-45
~m.
Derivation
of
name:
In
honour of Dr
R.
D.
Goos.
Occurrence:
Iceberg
Bay
Formation at Strand Fiord, Axel Heiberg
Island, Northwest Territories; Age: early
Eocene.
Locality:
C-
111847.
Comments:
Helicoonites
goosii
appears more closely allied to
species of
Helicoon
(Goos
et
al.,
1986) such
as
H.
richonis
than
to species of
Helicodendron
(Goos
et
al.,
1985).
520
Helicosporiates Kalgutkar &Sigler gen. nov.
Type
species:
Helicosporiates pirozynskii
Conidia simple, pale brown to brown, smooth, helicoid;
spirals of loose to tightly coiled filaments; filaments slender,
multicellular. Conidia usually helically coiled
in
one plane or
somewhat cochleate.
Comments:
Previously described and illustrated
as
Helico-
sporium-type (Kalgutkar &McIntyre, 1991, their
figs
4-6). The
conidia of modem
Helicosporium
are very similar to those of
Helicomyces
Link.
The conidia of species of both genera have
conidial filaments that are septate or indistinctly septate and
broadly to narrowly cylindrical. However, the conidia of
Helicosporium
are
mostly brown
in
contrast to those of
Helicomyces
which are invariably hyaline.
Modem concepts of the genera also place emphasis on
development of conidia or conidiophores.
In
Helicosporium,
the conidia are borne on differentiated (macronematous)
conidiophores which are mononematous, tall, slender, brown,
septate,
Simple
or loosely branched and bear pale to dark,
coiled or somewhat cochleate, pleurogenous or acro-
pleurogenous conidia.
In
Helicomyces,
conidia are generally
hyaline, usually tightly coiled, aerogenous, and are borne
terminally on sessile denticles on repent mycelium, or on
short, simple, hyaline conidiophores which arise from the
mycelium
as
lateral branches.
Helicosporiates
differs from
Involutisporonites
in
having
conidial spirals made
up
of multicellular filaments consisting
of slender cells with sometimes indistinct septa. Spores of
Involutisporonites
usually have alimited number of spirals
consisting of thickened broad
cells
with distinct septa and a
porate or aporate terminal
cell.
In
Colligerites
multicellular
compactly coiled spores are differentiated into acentral region
consisting
of
generally smaller, rounded cells and an outer
region with larger, rectangular
cells.
This feature separates
Colligerites
from other fossil helicosporous genera including
Helicosporiates.
The multicellular and multiseptate spores of
Helicominites
are tortuous and coiled
in
very loose spirals with
narrow ends, hence
are
easily distinguished from the conidia
of
Helicosporiates.
Helicosporiates pirozynskii Kalgutkar &Sigler
sp.
nov.
(Figs
18-20)
Holotype:
GSC
97294.
GSC
loc.
111847.
Conidia
Simple,
pale brown to brown, smooth, helicoid.
Conidial filaments forming the spirals coiled
2-4
times
in
one
plane
(Figs
18 &
19)
or cochleate
(Fig.
20); filaments slender,
multicellular, smooth, about 5
~
wide; septa present, or
sometimes indistinct; cells cuboid to rectangular, not indented
at the septa. Conidia
30-40
~m
diam.
Derivation
of
name:
In
honour of Dr
Kris
A.
Pirozynski.
Occurrence:
Iceberg
Bay
Formation at Strand Fiord, Axel Heiberg
Island, Northwest Territories; Age: early
Eocene.
Locality:
C-
111847.
Comments:
Because of its fuscous and loosely to tightly coiled
conidia,
Helicosporiates
pirozynskii
appears more closely
affili-
ated to
Helicosporium
than to
He!icomyces.

R.
M. Kalgutkar and Lynne Sigler
Paleoslimacomyces Kalgutkar &Sigler gen. nov.
Type
species:
PaIeoslimacomyces canadensis
Conidia simple, solitary, helicoid, curved to hemi-circinate,
brown
to
fuscous, smooth. Conidia
2-3
septate; septa dark,
often thick; conidial filaments short and made up of
3-4
broadly curved cells;
cells,
except the apical
cell,
darkly-
pigmented; apical
cell
hyaline to pale brown.
Comments:
Conidia of
Paleoslimacomyces
show some mor-
phological similarity with the conidia of extant
Slimacomyces
monospora
(W.
B.
Kendr.) Minter which was originally de-
scribed
by
Kendrick (1958)
in
Helicoma
Corda.
In
arevision of
Helicoma,
Ellis
(1976) transferred it to
Troposporella
P.
Karst.
as
T.
monospora
(W.
B.
Kendr.) M.
B.
Ellis.
After re-examining
type material, Minter (1986) concluded that the fungus was
different from
Troposporella
in its dispersal mechanism and
in
having two types of cells
in
afilament for performing different
functions.
One
type, having thick-walled cells with thick
septa, had asurvival function, whereas the second type,
having cells with thinner walls with ordinary septa, germinated
quickly.
He,
therefore, placed this fungus in anew genus
Slimacomyces
with
S.
monospora
as
type. Goos (1987) concurred
that the biology and morphology of
S.
monospora
justify its
placement
in
adifferent genus.
Paleoslimacomyces
conidia are distinguished from spores of
all
fossil helicosporous genera
by
their distinctively short and
curved to partially circinate filaments.
Paleoslimacomyces canadensis Kalgutkar &Sigler
sp.
nov.
(Figs
21,
22)
Holotype:
esc
106528,
esc loc.
C-112393.
Conidia simple, solitary, generally curved, horse-shoe shaped,
smooth, brown to fuscous,
2-4
septate, usually with 3septa;
septa dark, often thick, slightly thicker than the
cell
walls and
with central perforations. Apical
cell
hyaline to pale brown,
other cells dark pigmented; terminal
cell
broadly cylindrical,
rounded; cells forming the coiled axis curved with their outer
periclinal
cell
walls much greater
in
length than the inner
periclinal walls. Spores
13-16
~m
wide; filaments
5-6
~m
thick.
Derivation
of
name:
From
its
occurrence
in
Canada.
Occurrence:
Iceberg
Bay
Formation
at
Kanguk
Peninsula,
Axel
Heiberg
Island,
Northwest
Territories;
Age:
late
Palaeocene
or
early
Eocene. Locality:
C-112393.
Comments:
Conidia of
Paleoslimacomyces
canadensis
show some
similarities with those of extant
Helicoma,
Helicomina
L.
S.
Olive and
Trochophora
R.
1.
Moore.
In
these genera, however,
the conidial filaments are strongly circinate, narrower than
those of
Paleoslimacomyces
canadensis,
and have several septa.
Although conidia of this new fossil species do not show the
presence of two types of thick-walled cells that characterize
the conidia of extant
Slimacomyces
monospora,
they appear
similar in their general morphological features.
The authors wish to thank Dr Art Sweet of the Geological
521
Survey of Canada for reviewing the manuscript and making
useful comments. Drs Vasu Nambudiri and William Tidwell
kindly let
us
use the prepared slides of the fungal material
from the Deccan Intertrappean beds of India. Drs William
C.
Elsik
R.
D.
Goos,
S.
J.
Hughes and Kris Pirozynski are thanked
for suggesting affinities to some modem taxa. Financial
support from the Natural Sciences and Engineering Research
Council, Canada, to
L.
Sigler
is
acknowledged.
REFERENCES
Barlinge,
S.
G.
&Paradkar,
S.
A.
(1979). Record of new fossil algal and fungal
fonns from the Deccan Intertrappean of Mohgaon-Kalan, M.P., India.
Botanique
10, 163-175.
Barron,
G.
L.
(1983).
The
Genera
of
Hyphomycetes
from
Soil.
Robert
E.
Krieger
Publishing Company: Malabar, Florida.
Clarke,
R.
T.
(1965). Fungal spores from Vennejo Formation coal beds (Upper
Cretaceous) of central Colorado.
Mountain
Geologist
2, 85-93.
Cookson,
I.
C.
&
Eisenack.
A.
(1979). Some algae from Cretaceous sediments
of Australia.
Neues
Jahrbuch
fur
Geologie
und
Paliiontologie
Monatshefte
2,
77-82.
Deighton,
F.
C.
(1960). African fungi.
I.
Mycological
Papers
78, 1-43.
Ediger,
V.
S.
(1981). Fossil fungal and algal bodies from Thrace
Basin,
Turkey.
Palaeontographica
179,
87-102.
Ellis,
M.
B.
(1971).
Dematiaceous
Hyphomycetes.
Commonwealth Mycologial
Institute:
Kew,
Surrey, England.
Ellis,
M.
B.
(1976).
More
Dematiaceous
Hyphomycetes.
Commonwealth
Mycological Institute:
Kew,
Surrey, England.
Elsik.
W.
C.
(1968). Palynology of aPaleocene Rockdale lignite, Milan
County, Texas.
I.
Morphology and taxonomy.
Pollen
et
Spores
10, 263-314.
Elsik.
W.
C.
(1976). Microscopic fungal remains and Cenozoic palyno-
stratigraphy.
Geoscience
and
Man 15, 115-120.
Elsik.
W.
c.
(1992). The morphology, taxonomy, classification and geologic
occurrence of fungal palynomorphs. Ashort course presented under the
auspices of the American Association of Stratigraphic Palynologists,
Inc.
26-28
February.
Goos,
R.
D.
(1987). Fungi with atwist: the helicosporous Hyphomycetes.
Mycologia
79, 1-22.
Goos,
R.
D
..
Abdullah,
S.
K.,
Fisher,
P.
J.
&Webster,
J.
(1985). The anamorph
genus
Helicodendron.
Transactions
of
the
British
Mycological
Society
84,
423-435.
Goos,
R.
D.,
Abdullah,
S.
K.,
Fisher,
P.
j.
&Webster,
J.
(1986). The anamorph
genus
Helicoon.
Transactions
of
the
British
Mycological
Society
87, 115-122.
Gray,
j.
(1985). Interpretation of co-occurring of megafossils and pollen: a
comparative study with Clarkia
as
an example.
In
Late
Cenozoic
History
of
the
Pacific
Northwest
(ed.
C.
J.
Smiley), pp. 185-244. American Association
for the Advancement of Science,
Pacific
Division: San Francisco, CA.
Hawksworth,
D.
L.,
Sutton,
B.
C.
&Ainsworth,
G.
C.
(1983). Ainsworth &
Bisby's
Dictionary
of
the
Fungi,
7th edn. Commonwealth Mycological
Institute:
Kew,
412 pp.
Hughes,
S.
j.
(1972). New Zealand fungi.
18.
Xylohypha
(Fr.)
Mason.
New
Zealand
Journal
of
Botany
10, 447-460.
Hughes,
S.
j.
(1979). Relocation of species of
Endophragmia
aud.
with notes
on relevant generic names.
New
Zealand
Journal
of
Botany
17, 139-188.
Ichinoe, M. (1972). japanese Hyphomycete notes.
V.
Transactions
of
the
Mycological
Society
of
Japan
13,
57---<'>5.
Jain,
K.
P.
&
Kar,
R.
K.
(1979). Palynology of Neogene sediments around
Quilon and Varkala, Kerala Coast, South India.
1.
Fungal remains.
The
Palaeobotanist
26, 105-118.
Jarzen,
D.
M. &
Elsik,
W.
C.
(1986). Fungal palynomorphs recovered from
recent river deposits, Luangwa Valley, Zambia.
Palynology
10,
35---<'>0.
Kalgutkar,
R.
M. (1993). Paleogene fungal palynomorphs from Bonnet Plume
Formation, Yukon Territory.
Contributions
to
Canadian
Paleontology,
Geological
Survey
of
Canada
Bulletin
444, 51-105.
Kalgutkar,
R.
M. &McIntyre,
D.
J.
(1991). HeJicosporous fungi and an early
Eocene pollen
flora,
Eureka Sound Group, Axel Heiberg Island, N.W.T.
Canadian
Journal
of
Earth
Sciences
28, 364-371.

Fossil
fungi
Kalgutkar,
R.
M,
Nambudiri,
E.
M
V.
&Tidwell,
W.
D. (1993).
Diplodiles
sweetii
sp. nov. from the Late Cretaceous (Maastrichtian) Deccan Inter-
trappean Beds of India.
Review
of
Palaeobotany
and
Palynology
77,
107-118.
Kendrick,
B.
W.
(1958).
Helicoma
monospora
sp.
nov. from pine litter.
Transactions
of
the
British
Mycological
Society
41,
446-448.
Kwon-Chung,
K.
J.,
Wickes,
B.
L.
8<
Plaskowitz,
J.
(1989). Taxonomic
clarification of
Cladosporium
trichoides
Emmons and its subsequent
synonyms.
Journal
of
Medical
and
Veterinary
Mycology
27,
413-426.
Lange,
R.
T.
&Smith,
P.
H.
(1971). The Maslin Bay
flora,
South Australia.
3.
Dispersed fungal spores.
Neues
Jahrbuch
fur
Geologie
und
Paliiontologie
Monatshefte
11,
663--{)81.
Linder, D.
H.
(1929). Amonograph of the helicosporous Fungi Imperfecti.
Annals of
the
Missouri
Botanical
Garden
16,
227-388.
Minter, D. (1986).
Slimacomyces
gen. nov.
Bulletin
of
the
British
Mycological
Society
20,
17-24.
Padhye, A
A,
McGinnis,
MR.,
Ajello,
L.
&Chandler,
F.
W.
(1988).
Xylohypha
emmonsii
sp. nov., anew agent of phaeohyphomycosis.
Journal
of
Clinical
Microbiology
26, 702-708.
Pirozynski,
K.
A(1976). Fungal spores
in
fossil record.
Biological
Memoirs
1,
104-120.
Pirozynski,
K.
A&Hodges,
C.
S.
Jr.
(1973). New Hyphomycetes from South
Carolina.
Canadian
Journal
of
Botany
51,
157-173.
Pirozynski,
K.
A,
Jarzen, D. M., Carter, A&Day,
R.
G.
(1988). Palynology
and mycology of organic clay balls accompanying mastodon bones -New
Brunswick, Canada.
Grana
27,
123-139.
Ramanujam,
C.
G.
K.
&
Rao,
K.
P.
(1978). Fungal spores from the Neogene
strata of Kerala
in
South India.
Proceedings
of
the
Fourth
International
Palynological
Conference,
Lucknow
(1976-1977) 1,
291-304.
Ramanujam,
C.
G.
K.
&Srisailam,
K.
(1980). Fossil fungal spores from the
Neogene beds around Cannanone
in
Kerala State.
Botanique
9, 119-138.
Rouse,
G.
E.
(1962). Plant microfossils from the Burrard Formation of western
British Columbia.
Micropaleontology
8, 187-218.
(Accepted
27 August
1994)
522
Sheffy, M
V.
&Dilcher, D.
L.
(1971). Morphology and taxonomy of fungal
spores.
Palaeontographica
133,
34-51.
Sherwood-Pike, M. A(1988). Freshwater fungi: fossil record and paleo-
ecological potential.
Palaeogeography,
Palaeoclimatology,
Palaeoecology
62,
271-285.
Simmons,
E.
G.
&Roberts,
R.
G.
(1993).
Alternaria
themes and variations (73).
Mycotaxon
48,
109-140.
Smith,
P.
H.
8<
Chaloner,
W.
G.
(1979).
Is
Piriurella
Cookson &Eisenack an alga
or afungus 7
Neues
Jahrbuch
fur
Geologie
und
Paliiontologie
Monatshefte
11,
701-704.
Sutton,
B.
C.
(1969). Forest microfungi.
I.
Ampul/iferina
persimplex
n.
gen.,
n.
sp. on leaves of labrador tea.
Canadian
Journal
of
Botany
47,
609--{)
16.
Sutton,
B.
C.
(1973). Hyphomycetes from Manitoba and Saskatchewan,
Canada.
Mycological
Papers
132,
1-143.
Takahashi,
K.
(1991). Fungal and algal palynomorphs from the Tokatan and
Kiritappu Formations of the Nemuro group, Eastern Hokkaido.
Japanese
Journal
of
Palynology
37,
151-168.
Taylor,
T.
N. &White,
J.
F.
Jr
(1989). Fossil fungi (Endogonaceae) from the
Triassic of Antarctica.
American
Journal
of
Botany
76,
389-396.
Trevedi,
B.
S.
&Verma,
C.
L.
(1970). Fungal remains from Tertiary coal bed
of Malaya.
Journal
of
Palynology
5,
68-73.
Webster,
J.
(1980).
Introduction
to
Fungi.
Cambridge University Press:
Cambridge,
U.K.
Wolf,
F.
A
(1966a).
Fungus spores
in
East African lake sediments.
The
Bulletin
of
the
Torrey
Botanical
Club
93, 104-113.
Wolf,
F.
A(1966
b).
Fungus spores
in
East African lake sediments.
II.
Journal
of
the
Elisha
Mitchell
Scientific
Society
82,
57--{)1.
Wolf,
F.
A(1968). Fungus spores in
Lake
Singletary sediments.
Journal
of
the
Elisha
Mitchell
Scientific
Society
84,
227-232.
Wolf,
F.
A(1969). Nonpetrified fungi
in
Late Pleistocene sediments from
eastern North Carolina.
Journal
of
the
Elisha
Mitchell
Scientific
Society
85,
41-44.