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American Association of Stratigraphic Palynologists
A Revision of Reduviasporonites Wilson 1962: Description, Illustration, Comparison and

Biological Affinities
Author(s): C. B. Foster, M. H. Stephenson, C. Marshall, G. A. Logan and P. F. Greenwood
Source: Palynology, Vol. 26 (2002), pp. 35-58
Published by: Taylor & Francis, Ltd. on behalf of the American Association of
Stratigraphic Palynologists
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A REVISION OF REDUVIASPORONITES WILSON 1962:
DESCRIPTION, ILLUSTRATION, COMPARISON
AND BIOLOGICAL AFFINITIES
C.B. FOSTER G.A. LOGAN

Geoscience Australia Geoscience Australia
P.O. Box 378 PO Box 378
Canberra ACT Canberra ACT
Australia 2061 Australia 2061
e-mail: clinton.foster@ga.gov.au e-mail: graham.logan@ga.gov.au.
M.H. STEPHENSON P.F. GREENWOOD

British Geological Survey Geoscience Australia
Keyworth, Nottingham, NG12 5GG PO Box 378
United Kingdom Canberra ACT
e-mail: m.stephenson@bgs.ac.uk. Australia 2061
e-mail: paul.greenwood@ga.gov.au.
C. MARSHALL
Australian Centre for Astrobiology

Department of Earth and Planetary Sciences
Macquarie University
New South Wales 2109
Australia
e-mail: cmarshal@els.mq.edu.au.
Abstract
Geochemical analyses of specimens of Reduviasporonites sug-chalastus is present in Capitanian to Griesbachian rocks spanning
gests that it is most likely of algal, rather than fungal origin. Asat
a least 10 million years, and outside the more narrow age span of
the Permian-Triassic boundary. The size of the constituent cells
probable alga, Reduviasporonites is unlikely to be integral to the
process of mass extinction occurring at or near the Permian- present in R. chalastus appears to be related to paleolatitude with
Triassic boundary, as suggested by Visscher and other workerslarge examples occurring in the paleotemperate Permian of China,
because it cannot have acted as a saprophytic metaboliser of dead
Russia, and Australia (Moura) and smaller specimens occurring in
vegetation resulting from that event. Moreover, it ranges outsidethe paleotropical and paleoequatorial Permian of northern Austra-
lia, Saudi Arabia, United Kingdom and Austria.
the postulated time of mass extinction by at least 10 million years.
Optical and electron microscopy of topotype material confirms
that Reduviasporonites Wilson 1962 is the senior synonym of
Chordecystia Foster 1979, and Tympanicysta Balme 1980. More-
INTRODUCTION
over the type species of the last two genera, assigned in 1999 to
Reduviasporonites by Elsik as R. chalastus (Foster) and R.
stoschianus (Balme), are conspecific. The type species, R. A 'spike' of supposed fungal spores occurs at the
catenulatus Wilson 1962, differs from R. chalastus in that its
mian-Triassic boundary in various locations, in close
constituent cells are significantly smaller, more rounded, and
ciation with a mass extinction event (Erwin 1993). This
have less well developed connecting areas (terminal rims) be-relationship has leadpalynologists (Eshet et al., 1995; Vis
tween cells. Brazilea helbyi forma gregata Foster 1979, recorded
et al., 1996; Elsik, 1999) to consider that the supposed f
only from the type material of R. chalastus, is likely to be a junior
may have been a saprophytic metaboliser of dead veget
synonym of that taxon.
formed during the extinction event. Elsik (1999) sugg
The stratigraphic occurrences of the species ofReduvia-sporonites
suggest that R. catenulatus is most common in the Wordianthat two of the taxa often associated with the 'fungal s
(Kazanian) of Oklahoma, though specimens of the size rangeChordecystia Foster 1979 from the Australian Rewan
associated with R. catenulatus are present very rarely in the Earlymation, and Tympanicysta Balme 1980 from the Ma
Triassic Mazzin Member of the Werfen Formation, Austria. R.
Shale, Greenland, are junior synonyms ofReduviasporo
Palynology, 26 (2002): 35-58

@ 2002 by AASP Foundation ISSN 0191-6122
36 PALYNOLOGY, VOLUME 26 - 2002
Wilson 1962, and may be of various

affirmed shapes depending on the
a fungal extent of its
origin for
is not clear if shrinkage.
topotype material was re-examin
this conclusion. In contrast, Krassilov et al.
gested that 'spores' assigned by Russian paly
Tympanicysta are more SYSTEMATIC
closely PALEONTOLOGY
allied to the zyg
algae, a view confirmed by Afonin et al. (20
Genus
For this study, we have Reduviasporonitestopotype
examined Wilson 1962 emend
Reduviasporonites catenulatus Wilson 1962 and
Reduviasporonites chalastus (Foster) Elsik 1999, includ- Reduviasporonites Wilson, 1962, p. 91-93; pl. 1, f
ing its junior synonym Tympanicysta stoschiana as well as
well-preserved specimens of these taxa from Greenland, Chordecystia Foster, 1979, p. 109-110; pl. 41, fig
Britain, China, Saudi Arabia, Russia, Australia, and Aus- text-fig. 22.
tria (see Plate Explanations for details). The aims of the Tympanicysta Balme, 1980, p. 22-24; pl. 1, figs.
project are to determine whether:
Type. R. catenulatus Wilson 1962, by original
1. the taxa are synonymous tion.
2. the taxa have fungal or algal origins Emended generic diagnosis. Microfossil, usually
ing chains, but also present as pairs of cells o
dispersed cells. Cell outline usually subcircu
MORPHOLOGICAL TERMINOLOGY subrectangular; more rarely flask-, spindle- or 'Y
Most cells contain an inner body of the same sha
The natural affinities of specimens described outer
in this
cell that may be adpressed to the inner surfa
paper are uncertain; they have been interpreted main
eithercell
as so that it is difficult to distinguish optic
being of fungal origin (Wood and Elsik, 1999; Elsik,
inner body may also occupy a smaller part of the c
1999; Visscher et al., 1996) or of possible algal origin shrinkage). Outer cell wall approxima
(?through
(Krassilov et al., 1999, Afonin et al., 2001). In view of this
1 Itm thick, though may be locally thicker; smoo
uncertainty a new terminology independent of mycologybody wall less than 1 [tm thick, smooth. Cells in
and phycology is adopted. New terms suggestedjoined are (see
by thickened parts of the outer cell wall; t
Text-Figure 1): be regular rim-like structures or simple irregular
ings. Cells may have up to three thickenings
branched
Cell: the main unit of chains; usually hollow and cylin-chains to form.
drical, but may be Y-shaped, ovoid or spherical. Generic description. Cell shape (see Text-Figure 2):
The two
Terminal rim: the, darkened, slightly protuberant known species ofReduviasporonites can be distin-
annu-
lar thickening usually present at the termini or guished partly by cell shape but this can also vary consid-
opposite
margins of the cell. erably within a single assemblage of one or other taxon. R.
catenulatus
Terminal plane: the usually circular or rectangular area is always subcircular or oval in outline, sug-
enclosed by the terminal rim, which is perpendicular inoriginal, uncompressed spherical or ovoid shape.
gesting an
orientation to the long axis of the cell. In R. chalastus, the most common outline by far is
Inner body: the thin walled body, internal to the subrectangular
cell, that and this is consistent with SEM evidence
suggesting
normally mimics the overall shape of the cell but whichthat most cells of these taxa were originally
cylindrical in shape with a length:width ratio of between
1:1 and 4:1. Also occurring in assemblages of R. chalastus
Length (distance between are flask shaped, ovoid and spherical cells. In the British
terminal rims)
and Saudi Arabian assemblages ofR. chalastus, rare spindle-
S"rninal
shaped specimens with constricted and protuberant termi-
Width rim nal rims occur (Plate 2, Fig. 17; Plate 5, Fig. 6). Very rare
are 'three way cells' which have three terminal rims; these
Terminal
face
commonly have a 'Y' shape (Plate 3, Fig. 4; Plate 4, Fig. 14;
Cell material Plate 5, Figs. 10, 14).
Cell Inner body
Outer cell wall: The outer cell wall of Reduviasporonites
Text-Figure 1. Terminology used in morphological isde-

between 0.5 and 1.5 Ctm thick. When well preserved the
scriptions. outer surface of the cell wall is smooth or very finely and
C.B. Foster, M.H. Stephenson, C. Marshall, G.A. Logan, P.F. Greenwood: A revision of Reduviasporonites Wilson 1962 37
regularly granulate; but when poorly preserved the surface the inner body are firmly attached to the inner surface of the
is irregularly punctate or intragranulate. The outer cell wall terminal rim. With further shrinkage these finally detach
seems particularly susceptible to pyritization and com- from the outer cell wall. Text-Figure 3 illustrates possible
monly displays a surface irregularly pitted with the rectan- stages of shrinkage.
gular marks of invasive pyrite growth. The color of the Cell material: Dark cell material has been observed
outer cell wall is often darker than other palynomorphs in within the inner body of most specimens of R. chalast
the same assemblage, and may contain more natural pig- though not as yet in R. catenulatus. In the former, this
ment than associated spores and pollen. present in most cells and appears to occur within the in
Many specimens show 'striations' in the outer wall of the body. The material was recorded by Balme (1980)
cell, usually orientated parallel to the long axis of the cell 'amorphous material'; the present study confirms it is,
(Plate 2, Figs. 2-3, 6), and in rare cases perpendicular to it. many cases, structureless, but in the best preserved spe
These may be due to shrinkage related to desiccation, and mens (Plate 1, Figs. 4-7; Plate 4, Fig. 1, 3) it appears dar
are most common in narrow cells or cells with constricted dense and granulate. In a number of specimens the materi
and protuberant terminal rims. is clearly enveloped by the delicate, shrunken inner bo
Inner body: An inner body is observed in most specimens (Plate 2, Fig. 11; Plate 3, Figs. 11-12;). Afonin et al. (200
of R. chalastus but not as yet in the many examples of R. who considered Tympanicysta to be of algal origin, int
catenulatus. In the former, the inner cell wall is usually less preted the dark material within the cells as 'preserved
than 0.5 [tm thick and its surface is smooth in the few chloroplasts'.
specimens where it is exposed. The inner body commonly Folding patterns (see Text-Figures 4 and 5): Thinner
has the same shape as the outer cell and may be adpressed walled specimens of R. chalastus have a characteris
to the inner surface of the main cell so that it is difficult to diagonal folding pattern (Text-Figure 4). Observation an
distinguish optically. Through shrinkage, it may occupy a modeling of the cell by the authors suggests that thi
smaller part of the cell cavity and may be twisted. Shrink- caused by twisting of the cell body about the long axis of
age first occurs in the middle parts; usually the termini of cell. Other folds, in either of the cell walls, commonl
--- -- --- --
40@
-- - - - -- - - - -
0.'. '3
5
Text-Figure 2. Cells inR. chalastus and R. ca

cell with narrow protuberant terminal rims,
terminal rims, (7) Spindle-shaped cell with p
delineate rectilinear shapes and

R. chalastus and thethese may
means by which cells join repres
is less clear
of residual attachment of the inner and outer cell walls.
in the former (Text-Figure 6).
Commonly short lunate folds, orientated perpendicular to Effects of processing on size of cells. In order to inves-
the long axis of the cell, occur close to the margins of ovoidtigate the effects of palynological processing techniques on
cells (Text-Figure 4). These are believed to result from the the size of cells, two aliquots of post-hydrofluoric acid-
attachment of the termini of the inner body to the innertreated (HF) residue from the DILM-1 well, Saudi Arabia,
surface of the outer body and as such may represent parts ofcontaining abundant R. chalastus, were subjected to differ-
a poorly developed terminal rim. ent oxidation and potassium hydroxide treatments. The
Within the cell wall of R. catenulatus, folding patterns results of these were compared with an untreated post HF
are rather random in chains of cells, or occasionally residue from the same shmple to determine whether size
subcircular and concentric in arrangement in the case of changes had occurred. The treatments were:
single cells (Text-Figure 5; Plate 6, Figs. 5, 7- 9, 33, 36).
Terminal rim: The terminal rim is an annular thickening 1. Aliquot 1, concentrated nitric acid for one minute then
or fold, occurring at the cell terminus, which appears to treatment with 5% potassium hydroxide solution;
allow the cells to connect into chains. The rim is usually 2. Aliquot 2, concentrated nitric acid for 30 minutes then
well-developed in R. chalastus but very rare in R. treatment with 10% potassium hydroxide solution.
catenulatus. The rim is firmly attached on its inner surface
to the termini of the inner body. Close to the base of the The treatments caused no obvious change in cell size or
terminal rim is an incipient weakness in the outer cell wall shape and so it was concluded that processing method
which allows it to be partially or wholly detached from the cannot be responsible for size variability from sample to
rest of the cell (Plate 3, Figs. 5, 14). The terminal rim is sample.
often discontinuous (Plate 1 Fig. 3; Plate 2, Figs. 13, 15).
Light microscopy indicates that the few specimens that Reduviasporonites catenulatus Wilson 1962
display 'end on' views of the terminal face have no Plate 6, figs. 1-36
aperture (Plate 1, Fig. 12; Plate 3, Fig. 7), but SEM images
(e.g. Plate 5, Fig. 13) suggest that other specimens have an Reduviasporonites catenulatus Wilson, 1962, p. 94-95; pl.
indistinct, raised boss-like feature in the centre of the 1, figs. 1-6; Morgan, 1967, pl. 1, fig. 1.
terminal rim.
Chain formation: Chains of cells were observed in all the Description. Microfossil, usually forming chains, but
assemblages studied, some reaching 15 cells in length. also present as pairs of cells or single dispersed cells. Cell
Occasionally the three terminal rims of 'Y'-shaped cells outline subcircular or subrectangular; probably sub-spheri-
allow several chains to develop from one cell. Chains in R. cal before compression. Outer cell wall approximately 1
catenulatus are more irregular in appearance than those of[tm thick, smooth; may be irregularly thickened at the
PLATE 1
Illustrated specimens are housed in the Commonwealth Palaeontological Collection (CPC) at Geoscience Australia, PO Box 37
ACT, Australia 2061, and are assigned a unique catalog number prefixed CPC. Specimens are first identified by slide code, th
Finder reference and finally by CPC number.
All specimens photographed using differential interference contrast (DIC); scale bar is 25[tm.
8-18 R. chalastus (Foster) Elsik 1999 from 37.62m, Bore

1-7 R. chalastus (Foster) Elsik 1999 from 262m, BHPP
Billawock No. 1, offshore Bonaparte Basin, Australia. hole N.S.30 (Moura), Bowen Basin, Queensland, Aus-
tralia.
1, Urgen 29.2, J46/2, CPC 36673.2, SGM2, 053, CPC

36674. 3, Urgen 29.2, K46/1, CPC 36675. 4, SGM4, 8, MFP 9801/2, P36, CPC 36678. 9, MFP 9801/2, N31,
T39/3, CPC 36676. 5, SGM5, M39/1, CPC 36677. 6, CPC 36679. 10, MFP 9801/1, U46/3, CPC 36680. 11,
SGM4, T39/3, CPC 36676 (detail of 4). 7, SGM5, M39/ MFP 9801/2, D51/2, CPC 36681. 12, MFP 9801/1,
1, CPC 36677 (detail of 5). P33/4, CPC 36682. 13, MFP 9801/2, S44, CPC 36683.
14, MFP 9801/2, 050, CPC 36684. 15, MFP 9801/1,
U42, CPC 36685. 16, MFP 9801/2, N46/1, CPC 36686.
17, MFP 9801/2, 048/1, CPC 36687. 18, MFP 9801/2,
S36, CPC 36688.
C.B. Foster, M.H. Stephenson, C. Marshall, G.A. Logan, P.F. Greenwood: A revision of Reduviasporonites Wilson 1962 Plate 1
- .. 2 ..... . . 3
-'
....1 0-.
,. $3I
margins; no inner body or cell material has been observed.

Cells in a chain, and particularly single unattached cells,
have regular concentric or subpolygonal fold patterns.
Method of cell-to-cell attachment not known, though some
chains appear to indicate that the convexity of part of a
given cell fits into a corresponding hemispherical concav-
ity in an adjacent cell. No terminal rims are present.
Dimensions. Mean length 14.0 [tm; mean width 14.7
[tm; standard deviation of lengths 2.4; standard deviation of
widths 1.9; maximum length 22 [tm; maximum width 22
[tm; minimum length 9 tm; minimum width 11 tm; mean
length to width ratio 0.96. Number of specimens measured
70.
Reduviasporonites chalastus (Foster) Elsik 1999

Plates 1-5
Chordecystia chalasta Foster, 1979, p. 109-110; pl 41,

figs. 3-9; text-fig. 22.
? Brazilea helbyi forma gregata Foster, 1979, p. 112;
32
pl. 41, figs. 1-2.
Tympanicysta stoschiana Balme, 1980, p. 22-24; pl. 1,
figs. 3-7; Afonin et al., 2001, p. 484-486, figs. 1; 2A-
C, E,F.
.2- Reduviasporonites stoschianus (Balme) Elsik, 1999, p. 40;
pl. 1, figs. 1-24.
20-6/69 4 ? Reduviasporonites stoschianus (Balme) Elsik, Wood and
Elsik, 1999, p.46-48; pl.1, figs 1-9; pl. 2, figs 1-7.
Text-Figure 3. Shrinkage of inner body in the cell cavity.

Description. Microfossil, usually forming chains but
(1) Inner body occupies most of the cell, (2) Inner body
shrunk but attached at both ends to the inner surface of also
the present as pairs of cells or single dispersed cells. Cell
outer cell wall, (3) Inner body in advanced state outline
of subrectangular, rarely oval, irregular or 'Y' -
shrinkage; envelopes cell material, (4) Inner bodyshaped,
ab- though most cells were probably originally cylin-
sent.
drical in shape before compression. Outer cell wall ap-
PLATE 2
All specimens photographed using DIC; scale bar is 25 jAm.
1-10 R. chalastus (Foster) Elsik 1999 from Outcrop 11,Locality

R. chalastus from outcrop at Conisborough, UK.
16-18
6.75 (see Balme 1980; p. 9, fig. 1), Kap Stosch, East
Greenland.
11, 5031, H47/2, CPC 36698. 16, 5031, B47/2, CPC

1, MFP 12114/3, 750-6, A52/3, CPC 36689. 2, MFP 36703. 17, 5031, F24/4, CPC 36704. 18, 5031, B27,
12114/3, 750-6, B9, CPC 36690.3, MFP 12114/3,750- CPC 36705.
6, J21/2, CPC 36691. 4, MFP 12114/3, 750-6, E24/1,
CPC 36692.5, MFP 12114/3,750-6, J36/1, CPC 36693. 12-15 R. chalastus from Hilton Beck, UK.
6, MFP 12114/3,750-6, B9, CPC 36690 (detail of 2). 7,
MFP 12114/3, 750-6, E6, CPC 36694. 8, MFP 12114/ 12, HIL 01, N26/2, CPC 36699. 13, HIL 01, D30/2,
3, 750-6, F23/1, CPC 36695. 9, MFP 12114/3, 750-6, CPC 36700. 14, HIL 01, G28/1, CPC 36701. 15, HI
E17, CPC 36696. 10, MFP 12114/3, 750-6, E35, CPC 01, B43/3, CPC 36702.
36697.
-? ~ I
.....::.::..
t
...
'mm"""
.... ....
.... .. . ..
-......13....
"16 -.17 -..

......9.....- .-. -..
i L.' ?- 1 1
......: . ...... . . . . : .. . . . . ..:

.. ............. ..........*
.ii.. i ...
.....:.. ..P...... : ...
........ : ..
...........
1 3?::;':"'"""""
6 "
.. ... ... .
1 2
Text-F
joined
ment
0.5 to
hyalin
or gr
often
has a
?2j...
agona
twist
Other
eate r
residual attachment of the inner and outer cell walls.
Commonly, short lunate folds, orientated perpendicular
to the long axis of the cell, occur close to the margins of
ovoid cells (see Text-Figure 5). These are believed to
result from the attachment of the termini of the inner body
to the inner surface of the outer body and as such may
Text-Figure 4. Fold patterns in R. chalastus. (1) Diagonal
represent parts of a poorly developed terminal rim.
folds in the outer wall of the cell. Interpreted as being
Dimensions. Mean length 79.4 [tm; mean width 38.6
caused by twisting of the terminal rims in opposing
[tm; standard deviation of lengths 43.2; standard deviation
directions, (2) Diagonal folds in the inner wall of the cell,
of widths
(3) Folds or dark areas interpreted as being caused by 23.1; maximum length 220 LAm; maximum width
127or
residual adpression of inner body to the outer cell wall [tm; minimum length 18 [im; minimum width 9 jtm
mean length to width ratio 2.2. Number of specimen
by folding in the inner body. Folds may, therefore,
delineate the position of the inner body when it is measured
not 230.
visible, (4) Folds close to the margins of ovoid cells,The
andlengths and widths of 300 specimens of R. catenu-
orientated perpendicular to the long axis of the cell, areand R. chalastus are shown on a scattergram (Text-
latus,
believed to result from the attachment of the termini of
Figure 7) that also delineates the size fields ofR. chalastus
the inner body to the inner surface of the outer body
andand
R. catenulatus.
as such may represent parts of a poorly developed termi-
nal rim.
proximately 0.5 rtm to 1.5 [tm thick, smooth, may be

slightly thickened in the regions of the terminal rims. The
- 1
inner body commonly has the same shape as the outer cell
and may be adpressed to the inner surface of the main cell 2
so that it is difficult to distinguish optically. The inner
body may also occupy a smaller part of the cell cavity
(?through shrinkage) and may be twisted. Detachment of
the two walls appears first to occur in the middle parts;
usually the termini of the inner body are firmly attached
Text-Figure 6. Chain formations in (1) R. catenulatus and
to the inner surface of the terminal rim, though detach- (2) R. chalastus.
Comparison. R. chalastus is distinguished from R. the outer cell walls are generally transparent, revealing the
catenulatus by the greater size of its cells (up to 10 times usually shrunken and twisted inner body, almost always
greater in most cases, Text-Figure 7), the generally rectan- attached to the inner surface of the terminal rim (Plate 4,
gular outline shape of its cells, the presence in its cells of Figs. 2, 4). A number of cells have poorly developed,
distinct inner bodies, often containing cell material, and by discontinuous terminal rims (Plate 4, Fig. 3).
the presence of terminal rims allowing regular articulation Moura, Bowen Basin, Australia: The two assemblages
of cells into chains. As noted by Elsik (1999) rare, small from the N.S.30 Moura Borehole, Queensland, Australia,
specimens of R. chalastus from the Mazzin Member are yield poorly preserved cells ranging in shape from very
similar in size to R. catenulatus (Text-Figure 7) but these regularly cylindrical to ovoid and spherical. Perhaps re-
have the terminal rims, cell material and distinct inner flecting this poor preservation, very few chains were noted.
bodies associated with R. chalastus. A number of cells display the diagonal folds thought by
Remarks. Our examination of topotype specimens of Foster (1979) to be characteristic of R. chalastus (e.g. Plate
the Greenland material originally described as Tympanicysta 1, Fig. 11). A few specimens (e.g. Plate 1, Fig. 12) display
stoschiana by Balme (1980) and topotype specimens of 'end on' views of the terminal face which indicate that, in
Australian material described by Foster (1979) as these specimens at least, no aperture is present.
Chordecystia chalasta, shows that they are synonymous. Perhaps due to poor preservation, none of the Moura
Careful examination of specimens ofBrazilea helbyi forma specimens display the inner body with clarity. In most
gregata Foster 1979 suggest that this is also likely to be cases, only folds in the inner body are visible due to the
either synonymous with R. chalastus, or a new species of increased opacity that this produces. In other specimens,
Reduviasporonites. Further work on better-preserved ma- folds in the outer wall of the cell caused by residual
terial is required to confirm this interpretation. attachment of the inner body are visible. Overall these fold
The relationship of 'fungal remains' assigned to patterns suggest that partially shrunken inner bodies are
Reduviasporonites stoschianus (Balme) Elsik 1999 by Wood present, usually attached to the interior surface of the
and Elsik (1999) to other species of Reduviasporonites is terminal rim.
being reviewed by us. A number of spherical to ovoid cells with folding pat-
Morphological variation in R. chalastus. Variation in terns suggesting cylindrical inner bodies (Plate 1, Figs. 10,
size, shape and preservation is evident within and between 16-17, 18), are closely similar to specimens figured and
assemblages ofR. chalastus. In this section these variations assigned by Foster (1979; pl. 41, figs. 1-2) to Brazilea
are described in detail. helbyi forma gregata Foster 1979.
Junggar, Northern Xinjiang, China: The five assemblages Kap Stosch, Greenland: The assemblages from Kap
from the Junggar section, China contain large numbers of Stosch, Greenland contain common elongate, cylindrical
cells in a variety of forms. Elongate, cylindrical cells are cells, but also ovoid and flask -shaped cells (Plate 2, Figs.
common, but ovoid, spherical, Y-shaped and flask-shaped 7-9). Although relatively well preserved, the outer cell
cells also occur (Plate 3, Figs. 1, 4, 8-10, 15). Overall the walls are generally relatively transparent, revealing the
cells are rather poorly preserved but this has rendered the usually shrunken and twisted inner body, almost always
outer cell wall relatively transparent so that the inner body attached to the inner surface of the terminal rim (Plate 2,
and cell material can be examined and photographed (Plate Figs. 3-4)
3, Figs. 2-3). A number of specimens illustrate the shrink- Billawock, offshore Bonaparte Basin, Northern Terri-
age of the inner body and its twisted appearance (Plate 3, tory (NT), Australia: One assemblage and six single grain
Figs. 2, 11-12). In a few specimens the twisted, shrunken mounts from BHP Petroleum Billawock No. 1 borehole,
inner body is shown to envelope the cell material (e.g. Plate NT contain well preserved cells ranging in shape from
3, Figs. 11-12). Other specimens illustrate the incipient very regularly cylindrical to ovoid (Plate 1, Figs. 1-3).
weakness in the outer cell wall which occurs at the base of Although the outer cell wall is relatively well preserved,
the terminal rim, so that partial (Plate 3, Fig. 14), and full and therefore less transparent than in specimens from
(Plate 3, Fig. 7) detachment of the terminal rim can occur. Moura, the inner bodies are nevertheless revealed in a few
In terms of range of shapes, preservational condition and specimens (Plate 1, Fig. 1) and granular cell material is
size, the Chinese assemblages are most similar to those of very distinct (Plate 1, Figs. 4-7). A number of specimens
Australia (Moura), Russia and Greenland. have poorly developed and discontinuous terminal rims
Russian Platform: The assemblage from the Russian (Plate 1, Fig. 2).
Platform, Vologodskaya Region contains common elon- Conisborough, UK: The assemblage from Conisborough
gate, cylindrical cells, but also ovoid and flask-shaped cells contains very well preserved, relatively small (Text-Figure
(Plate 4, Figs. 1, 3-4). Although relatively well preserved, 7) cells ranging in shape from cylindrical, to flask-shaped,
?3 . aUK (m n)
USA
140
... . . ........ . . 1
.7. ..
am *s atenim?ISOas
. .. . Ph
i7'7 L wr
Text-Figure 7. Scattergram of the lengths and widt
PLATE 3
All specimens photographed using DIC, except (5) and (6) which are SEM microphotographs; 5, scale bar = 20 1km,
all other scale bars are 25 1tm.
1-9, R. chalastus from outcrop approximately 170.5 m be-11, 640 3380 NLD36, P5/1, CPC 36707 (detail of Fig.
11-15 low the top of the Guodikeng Formation, north limb of 2). 12,640 3380 NLD36, P5/1, CPC 36707 (detail of 2).
the Dalongkou Anticline. 13, 640 3380 NLD36, C12, CPC 36716. 14, 640
3380NLD36, D17/4, CPC 36717. 15,640 3380 NLD36,
1, 640 3380 NLD36, D35, CPC 36706. 2, 640 3380 C20/4, CPC 36718.
NLD36, P5/1, CPC 36707. 3, 640 3380 NLD36, L35,
CPC 36708.4, 640 3380 NLD36, N21/4, CPC 36709. 10 R. chalastus (Foster) Elsik 1999 from outcrop approxi-
5, 640 3380 NLD36, mounted on Stub 7, CPC 36710. mately 192.5 m below the top of the Guodikeng Forma-
6, 640 3380 NLD36, mounted on Stub 13, CPC 36711. tion, north limb of the Dalongkou Anticline, Junggar,
7, 640 3380 NLD36, N40/2, CPC 36712 showing Northern Xinjiang, China.
detached terminal rim. 8, 640 3380 NLD36, C20/4,
CPC 36713. 9, 640 3380 NLD36, C20/4, CPC 36714. 10, 640 3381 NLD 37, E29/1, CPC 36715.
. .. . ..
. .
...
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14,?1?ii
.. ... l
. . . . . . . . . . . . . . . .
~~Mg.
. ....... . .lir B I
......... .. . . . . .... .. .
?" a........
. .. .. .. .. .. .... ..... ...... .. ..... ....
........ ............
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" " ........10.11.1 .. .... . .
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. . . . . ........ . .... .
10 12
spindle-shaped and ovoid.shape from cylindrical,

There are to flask-shaped,
more ovoid and Y-
flask- and
shaped cells than in other assemblages
shaped; the studied
cells are commonly joined in long chains of (P
Figs. 16-18). As with the Austrian
up to 20 cells (Plate 4). and Saudi
specimens, the outer cell and
The cell wall andinner
the inner body body have
wall are relatively thick r
thick walls and the inner bodies,
and usually smooth; thewhen
inner body andvisible,
outer cell are ar
completely adpressed toadpressed
theand inner cell wall.
difficult to distinguish A anu
optically, though
specimens display longitudinal 'striations'
number of specimens possibly
show small amounts of detachment
by shrinkage (e.g. Platebetween
2, theFig.
two wall18).
layers (Plate 4, Figs. 6, 9-10). Rarely
A few specimens (e.g. Plate 2, Fig.
specimens indicate 17)
high degrees display
of inner bot
body shrinkage
and outer layers and these
(Plate suggest that
4, Fig. 8). As in specimens fromthe inner
other locations, it isb
is smooth and clear that detachment
approximately 1 tm of the inner
thick,body occurswhich
last from the is
erably thicker than termini of seen
that the cell outer wall.
in Russian, Chi
Billawock specimens. The outer cell wall i
Conisborough specimens Palaeolatitude
is usually and size variations
smooth, in R. chalastus.
thoug
have a very regular, Text-Figure
very small 7 shows that the largest cells
granulate of R. chalastu
ornamen
just resolvable using oil
occurxl00 objective).
in assemblages Like
from Junggar Basin, China; t
K
body wall, the outer cellStosch,
wall Greenland,
in and the the Russian Platform. The pale
Conisboroug
mens is relatively thick.geographical positions of these localities are within th
Hilton Plant Beds, UK: paleotemperate
The assemblage latitudes (Text-Figure
from 8). Smaller
the ce
Plant Beds contains well occur in the paleotropicalrelatively
preserved, and paleoequatorial Permian
sma
Figure 7), mainly northern Australia,
ovoid cells. Many Saudi
ofArabia,
the UK and Austria.ha
cells This
trend may
poorly developed terminal rims.indicate a paleoenvironmental control on th
DILM-1, SaudiArabia: The
growth assemblage from
and development of the organism the R
that produced
I well, central Saudi
contains very well pr chalastus.
Arabia,
relatively small (Text-Figure 7), cells ranging in
from very regular cylindrical, to flask-shaped,
shaped and ovoid. The cell wall AGES OF THEthe
and MATERIAL
inner body
relatively thick and usually smooth; the inner b
outer cell are adpressed Australia-Bowen
and difficult and Bonaparte
to Basins
distingui
cally, though a few broken specimens (Plate 5, Fig
Examples
the relationship between the ofR. chalastus
inner body from the
andBowen Basin are f
outer
to be examined. type material from the Rewan Formation (Foster, 19
Tesero section, Austria: The four outcrop samples while those from the Bonaparte Basin, in northern Au
from the Permian-Triassic boundary beds at Tesero, lia (Text-Figure 8) are from the Blina Shale, intersect
Austria contain extremely rich assemblages of cells. a petroleum exploration borehole, BHPP Billawo
These are generally small (Text-Figure 7) and range in studied by Helby (1992).
PLATE 4
1-10 are photographed using DIC, 11-15 are scanning electron microphotographs; 1-10 scale bars are 25 tm; 11 scale ba
bar = 5 pm; 13 and 14 scale bars = 20 pm; 15 scale bar = 2 ptm.
1-4 R. chalastus (Foster) Elsik 1999 from 44.4m, Borehole No. 5, T1 6403434/2, N31/3, CPC 36723. 6, Ti 6403434/2,
453K, 3.5km, NE from Shartanovo Village, N33/4, CPC 36724. 7, T1 6403434/2, N6, CPC 36725.
Vologodskaya Region, Russia. 8, T1 6403434/2,016/3, CPC 36726.9, TI 6403434/2,
N8, CPC 36727. 10, T1 6403434/2, P15/3, CPC 36728.
1, 203.3Russia, T55/1, CPC 36719. 2, 203.3Russia, 11, T1 6403434/2, mounted on Stub 1, CPC 36729. 12,
D41/4, CPC 36720.3, 203.3Russia, L49/2, CPC 36721. Ti 6403434/2, mounted on Stub 1, CPC 36730. 13, Ti
4, 203.3Russia, F58/1, CPC 36722. 6403434/2, mounted on Stub 12, CPC 36731. 14, Ti
6403434/2, mounted on Stub 12, CPC 36732. 15, TI
5-15 R. chalastus from Lutite horizon within the Mazzin 6403434/2, mounted on Stub 1, CPC 36733, showing
Member at the Tesero section, Austria. break in the cell wall.
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..
: ....
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&mX ?- % ..'..-'
a -
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.%I%
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17-1 ..
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ii mm ... 11.. ..:ll.., ..: . ..:.. ..
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.:,CA?Xpml.-m:,-. -:. RL.I1 !%.:?-
(see Dolby
In both basins, R. chalastus occurs and Balme, 1
in assembla
signed to the Protohaploxypinus
younger than microcorpus
early Grie Z
emended by Foster (1982). Essential
The ageelements of th
of this zone i
versial, in that some authors regard it
the eponymous as eith
species,
Permian or earliest Triassic, or spanning
Triplexisporites the bou
playfordi
immediately
McLoughlin et al., 1997). Assemblagesunderlying
from this
Aratrisporites spp.. Based on this
of Foster Foster
(1982). Some et
w
facies
argued for a Late Permian age,variant oftheP.
but facies maymi co
first appearance of thesebut what
spores is most
from one importa
lycopod
is nevertheless noteworthy
and Helby
that the
(1973),
downward
Austra
pearance
Aratrisporites spp. and its of these
inclusion in thekey spe
P. mi
from
Zone in Helby et al. (1987, the
fig. uppermost
5) results from Cha
which on conodont eviden
error (Helby, pers. comm.).
Changhsingian
Independent age constraints (Wardla
for the P. microcor
are based on superposition. Faunas associate
palynological evidence th
older than early Changh
palynofloras from the underlying Dulhuntyisporap
Zone (APP 5; Price, 1997) range in age from
(Wordian) to Dzhulfian (see Foster and Archbold
In terms of the Kap
Russian Stosch,
Stages, Greenland
Jones and Che
suggest a correlation
of D. parvithola-bearing sed
Two samples,
the Newcastle Coal Measures, 750(675)-
Sydney Basin, eas
tralia, with the Late Tatarian Vjatian 6.75
from locality horizon (si
(see Bal
Russian Platform, based on Greenland,
East conchostracan
but faun
the
youngest age for the P. samples cameZone
microcorpus was not ide
relies o
logical correlation, and acceptance
belong of the
to the Protohaplo
Aratrisporites, which (1980)
occursconsidered
in marine to rocks
be ea
Griesbachian age in the palynological
Perth Basin ofcomparison
western
32
06
~7
16/0/144
Text-Figure 8. Palaeogeographic map showing positions of the samples studied. Map and sample positions based on Ziegler et al.
(1996). (1) Junggar Basin, Northern Xinjiang, China, (2) Russian Platform, (3) Kap Stosch, Greenland, (4) Conisborough and
Hilton Plant Beds, UK, (5) Tesero Section, Austria, (6) DILM-1, central Saudi Arabia, (7) Billawock, Bonaparte Basin, Australia,
(8) Moura, Bowen Basin, Australia.
Arctic islands, and on the occurrence of Otoceras woodwardi Jiucaiyuan Formation, are wholly Triassic in age. Li et al.
boreale Spath and Glyptophiceras triviale Spath. Balme (1986) reported Alisporites spp., Pteruchipollenites spp.,
(1980) also reported R. chalastus from the succeeding Klausipollenites spp., Protohaploxypinus limpidus, P.
Griesbachian Taeniaesporites Association. Utting and ovaticorpus, Lueckisporites virkkiae, Limatulasporites
Piasecki (1995) reported (but did not illustrate) R. chalastus fossulatus, and Lundbladispora in the lower and middle
(as T. stoschiana) from the Karstryggen Formation (corre- parts of the formation but noted an increase in the abun-
lated with the Capitanian by Henderson & Mei, 2000), and dance of spores, accompanied by the appearance of
the younger Schuchert Dal Formation which is dated Taeniaesporites spp. and Equisetosporites spp., in the
Dzhulfian by ammonoids (Nassichuk, 1995). These occur- upper part of the formation. Li et al. (1986) considered
rences indicate that R. chalastus has a relatively long strati- that these changes heralded the onset of the Triassic.
graphic range in the region. Significantly, Li et al. (1986) did not record Aratrisporites
in the Guodikeng Formation but did report it from the
overlying Jiucaiyuan Formation along with abundant
Tesero section, Austria spores including Lundbladispora nejburgii and L.
willmotti. Li et al. (1986) considered that conchostracans
The three samples TI 6403434, T2 6403435 and T3 in the top 30m of the Guodikeng Formation also indicated
6403436, are from a lutite horizon within the Mazzin a Triassic age.
Member at the Tesero Section, Austrian Alps. The Mazzin Ouyang et al. (1999) and Ouyang and Norris (1999)
Member is separated from the latest Permian Bellerophon suggested that the top 40-50 m of the Guodikeng Forma-
Formation by the Tesero Member and the basal Triassic tion is Early Triassic in age, on the basis of the occurrence
conodont marker species H. parvus occurs 130cm above of an assemblage containing Lundbladispora spp.,
the base of the Tesero Member in the Bulla Section (Cassinis Aratrisporites spp. and Lunatisporites spp.. Ouyang and
et al., 2000), and has been reported from the lower Mazzin Norris (1999) considered that the appearance of
Member (see Broglio Loriga and Cassinis, 1992) confirm- Aratrisporites has special significance because, based on
ing a Triassic age for the Mazzin Member. Based on other the palynological work of Ouyang and Utting (1990), its
included fossils, the Mazzin Member has been assigned a first appearance is 2.7m above the base of the Triassic at
Griesbachian age by Assereto et al. (1973), Broglio Loriga Meishan. Ouyang et al. (1999) noted species such as
et al. (1983) and No6 (1987). Lueckisporites virkkiae and Scutasporites unicus (as S.
Recent evidence (Looy et al., in press, see ODP Excur- xinjiangensis) in the lower and middle parts of the Guodikeng
sion Field Guide, Italia 2001; p. 95) suggests that 'fungal Formation, which they therefore consider to correlate with
remains' (?R. chalastus) appear to occur over relatively the Tatarian of the Russian Platform.
short interval extending approximately 12m below, and Samples collected for the present study from the lower
approximately 15m above, the Permian-Triassic bound- part of the Guodikeng Formation, confirm the presence of
ary. L. virkkiae and Scutasporites spp.. Conchostracans from
the same samples, assigned by Prof. Shen Yan-bin (pers.
comm.) to the following species: Beijianglimnadia sp.,
Junggar, Northern Xinjiang, China Bipemphigus cf. gennisi Novozhilov 1965 and Polygrapta
chatangensis Novozhilov 1946, also suggest a Tatarian
Five samples, 640 3380, 640 3381,640 3417, 640 3390 age. On going studies of palynofloras from the upper part
and 640 3400 were taken from the lower part of the of the lower Guodikeng Formation, by an international
Guodikeng Formation (Lower Canfanggou Group) on group coordinated by one of us (CBF), are attempting to
both limbs of the Dalongkou anticline. Although the define the age more precisely.
Permian of the Junggar Basin is continental, and therefore Apart from paleontological evidence, paleomagnetic
difficult to relate to the marine basal Triassic stratotype evidence (Jin and Shang, 2000) which places the Illawara
section at Meishan, a number of lines of evidence suggest Reversal within the Lucaogou Formation (of the Upper
that the Guodikeng Formation spans the Permian-Triassic Jijicao Group; see Ouyang et al., 1999) further suggests that
boundary. the Upper Jijicao and Lower Canfanggou Groups are
Li etal. (1986) described the palynomorphs, megaspores, Capitanian or younger in age.
megaflora, bivalves, ostracods, conchostracans and ver- Ouyang and Utting (1990) reported R. chalastus (as T.
tebrates of the Guodikeng Formation. They considered stoschiana) from the top 40 m of the Changhsing Forma-
that all but the top 30 m of the Guodikeng Formation is tion at Meishan, but not from the Griesbachian Chinglung
latest Permian in age while the top 30 m, and the overlying Formation. Ouyang and Norris (1999) noted the presence
ofR. chalastus in the Flowerpot Formation has been dated

Griesbachian on reptile fossil re-
Jiucaiyuan
the Junggar section. mains as Kazanian
The present in age (Chudinov,
study1965). exten
downward, as R. chalastus occurs abundantly
part of the Guodikeng Formation.
Hilton Plant Beds, UK
Russian Platform Illustrated specimens from the Hilton Plant Bed are from
a palynological strew slide given to one of us (CBF) by
This sample, collected by Dr. O. P. Yaroshenko, origi- Prof. H. Visscher in 1978. Although not illustrated here, Dr.
nates from the lower Vetlugian Series at a depth of 44.4 m G. Warrington has also recorded R. chalastus from the
in Borehole No. 453K, 3.5 km NE from Shartanovo Vil- Hilton Plant Bed at Hilton Beck, 0.5 miles west of Hilton
lage, Vologodskaya Region. The lower part of the Vetlugian Village, Cumbria. Clarke (1965) implied a Late Permian
on the Russian Platform is correlated with the Otoceras and'Zechstein' age for the Hilton Plant Beds on the basis of
comparison with palynofloras described by Klaus (1963),
Ophiceras zones on the basis of vertebrate, palynological,
ostracod and conchostracan evidence (Lozovskiy, 1992) Leschik (1956), Grebe and Schweitzer (1962) and
and is therefore Griesbachian in age. Scharschmidt (1963). Stoneley (1958) considered the
macroflora of the bed to be of Late Permian 'Thuringian'
age. On the basis of recent correlation using conodonts
DILM-1 well, Saudi Arabia Henderson and Mei (2000) have suggested a Wuchiapingian
age for the Zechstein.
This sample originates from the informally named 'basal
Khuff clastics' at a depth of 7827.9 feet in borehole DILM-
1, central Saudi Arabia. Direct dating of this unit is by Conisborough, UK
palynology and palaeobotany only; though some upper
limit dates are available from the overlying Khuff Forma- A single sample (73862 ) was collected by Dr. B. E.
tion. El-Khayal and Wagner (1985) suggested a Zechstein- Balme from a 'grey claystone of the Lower part of the
equivalent age for the basal Khuff clastics macroflora at the Permian succession above the Carboniferous' at
Ash Shuqqah section. Stephenson and Filatoff (2000) con- Conisborough, Yorkshire, UK. Balme (pers. comm.) co
sidered the basal Khuff clastics to be Tatarian or younger in lated the assemblage with those of the Zechstein, w
age (possibly Changhsingian). The lower part of the Khuff implies a possible Wuchiapingian age for the assemb
(?basal Khuff clastics) in the Hawtah Field is dated as Late (Henderson and Mei, 2000).
Tatarian (van der Eem in Al-Jallal (1994)).
Summary of age information

Greer County, Oklahoma, USA
The ages suggested for the localities studied and for o
Two samples were collected by Dr. B. E. Balme from the published occurrences of R. chalastus and R. catenu
Flowerpot Formation of Greer County, Oklahoma. The are summarised in Text-Figure 9. The ranges indicate
PLATE 5
1-12 are photographed using DIC, 13-15 are scanning electron microphotographs; 1-12 scale bars are 25 [tm; 13 scal
15 scale bars = 20 tm.
1-15 R. chalastus (Foster) Elsik 1999 from core sample from 8, DILM-1 7827.9, F43/4, CPC 36741. 9, DILM-1
depth 7827.9 feet, Borehole DILM-1, central Saudi 7827.9, K35/1, CPC 36742. 10, DILM-1 7827.9, 042/
Arabia. 2, CPC 36743. 11, DILM-17827.9, 045/2, CPC 36744.
12, DILM-1 7827.9, 045/2, CPC 36745. 13, DILM-1
1, DILM-1 7827.9, D31/1, CPC 36734. 2, DILM-1 7827.9, mounted on Stub 6, CPC 36746 showing detail
7827.9, N30, CPC 36735. 3, DILM-1 7827.9, K49/2, of terminal rim. 14, DILM-1 7827.9, mounted on Stub
CPC 36736.4, DILM-1 7827.9, Q30/2, CPC 36737. 5, 6, CPC 36747. 15, DILM-1 7827.9, mounted on Stub 6,
DILM-1 7827.9, 045, CPC 36738.6, DILM-1 7827.9, CPC 36748.
W33, CPC 36739.7, DILM-1 7827.9, C33, CPC 36740.
.... ....... . ...
?":
? : ." . . . . .. ...... ... .? ... ... ' .. ... ... '? ?i I . iCi-
.....................
?: '} i i "
? '..... .. . . . . . .....
Aw:
? ?: : -.?:. IN,.
r..y
? "?? '?
': . .'
.....
ii::::i....
!. ii~ ' . ... .,.., ..........

." .......
. .... . . .. h.Z.
although R. catenulatus Our studies

has of potential foodstratigraph
a restricted sources (for the sup
that of R. chalastus is saprophyte), of both
relatively woody spanning
long, tissue and pollen from
at
Myr in
the Permian underlying Late Permian Groedner Sandstone,as
(Capitanian-Chansinghian)
tinuing into the Early Triassic. Significantly
813C isotopic composition of -22%o, which isits
signifr
heavier. Thus,
not restricted to the narrow it is unlikely
sections that R. chalastus
spanning Pe
Triassic boundary. saprophytic organism deriving its energy from w
tissue and pollen.
Because our specimens ofR. chalastus are from the
GEOCHEMICAL ANALYSIS Triassic Werfen Formation it might be argued that
should be depleted isotopically in b13C, as a res
secular change. However, older, Late Permian a
The carbon isotopic composition (813C) of R. chalastus,
and of its apparent food sources, suggests itblages
is notwith
a R. chalastus from the Junggar Basin
saprophyte. Moreover the wall composition ofR. chalastus,
study) are also depleted in 813C, with values of-30 to
The molecular composition of the wall material
determined by gas chromatography mass spectrometry
chalastus was investigated by micro-FTIR and pyr
(GCMS ) and micro-Fourier Transform Infrared Spectros-
copy (micro-FTIR), is consistent with that of an GCMS.
alga. The fossils were first cleaned by either mild
The palynological assemblage from the Werfenmal treatment or the use of organic solvents (aceto
Forma-
tion (TI 6403434) is almost entirely made up of R.
dichloromethane ) to remove extraneous surface ad
chalastus, but was further concentrated by using aorganic
+25 itmmatter. The FTIR analysis was performed w
magna-IR
sieve fraction, and the purity then confirmed using a low 760 equipped with a Nicolet microscope a
power objective. The 813C isotopic composition of from
signal R. the material was characterised by prom
hydroxyl,
chalastus is -32%0. If R. chalastus was a saprophyte living aliphatic and carbonyl groups. Aliphati
on decaying vegetation, its isotopic compositionbands
wouldpresent included symmetric and asymmetric
be slightly heavier than its food source (Koch et tions
al., 1994).
at 2930 cm-1 (vas CH2 + CH3), 2850 cm-1 (vs CH
This study Other records
CO 0AC
0)
-i 0) 0)
o00 0
0 0)
0 C
-_ "_ '- 0) _ 2 --
01-
v0
C\I t m
0)0 0) CD
c
Location (all records for
? . 0 00al
chalastus except for # for0 R.0D %I- 0c

2. ma_ _C00)c 73
catenulatus).OD,
- asEa " C-O
. -
O
M, as -
=3 C C 0 06 0 -0
-2 ED
>
CO 40m 0E->
m a) C
CO -tm
L- Cm CD I
0 CO
C 0 L- I
V) C W
- 0
CO a :3 I a 0 m
O 2 2' OC C
T0 O%-radto
ICS scal Tethysscale Trditiona
standard
Griesbach.c
Changh. Dorhamia
ICS scale Tethys Mchiaing.
scale Tslta~ndar zhulfan Taaria
Capitanian Midian
Griesbach. **
Wordian Murgabian Kazania
Changhs. Dorshamian
cnWordian * * * *
Murgabian Kazanian
Roadia Kuberand. mian
* * * * *
Roadian Kubergand. Ufimian) C
Wuchiaping. Dzhulfian Tatarian* **
Capitanian Midian*
Text-Figure 9. Table of ages suggest

catenulatus. The correlation of the IC
case of other published studies, ages
Reduviasporonites stoschianus (Balm
and deformational vibrations at 1450 cm-1 (aCH2) and 1380 usually the main aromatic products detected from algaenan,
cm 1 (aCH2) (Text-Figure 10). but typically in low concentration relative to aliphatic
Pyrolysis was performed on -0.3 mg of fossils at 600'C/ hydrocarbons. Their high relative abundance may be due to
10s using a CDS (Chemical Data Systems, Inc.) pyroprobe the antiquity (approximately 250 Ma) of these fossils or
interfaced to a HP 6890/5973 GCMS. The products were alternatively may reflect non-indigenous organic material
detected by monitoring selected ions diagnostic of com- that was not removed by the pre-treatment procedures.
mon compound classes. The main aliphatic pyrolysates The presence of algaenan biomacromolecules is indica-
produced were a homologous series of n-alkenes/alkanes tive of an algal origin. Furthermore, the results show little
from Cg to - C21 (Text-Figure 1 la). The n-alkenes are specific evidence for fungal biopolymers such as cellulose,
products of the open pyrolysis process and arise from the mannans and/or glucans.
same saturated long chain macromolecular precursors as The biogeochemical data, presented here, strongly sup-
the n-alkanes. These straight chain compounds most likely ports an algal origin for R. chalastus.
account for the aliphatic C-H bands measured by FTIR.
They are similar, although of relatively shorter average
chain length, to the n-alkene/alkane series typically de- CONCLUSIONS
tected from the highly aliphatic biomacromolecule
'algaenan' of many species of Chlorophyceae (Tegelaar et 1. Geochemical evidence suggests that R. chalastus
al., 1989; see also Al-Arouri et al., 1999 for discussion of algal and not fungal origin.
specific examples) and are attributed to a macromolecular
structure based on polymethylenic chains, linked primarily 2. R. chalastus is unlikely to be integral to the proc
via ether bonds. mass extinction occurring at or near the Perm
Several distributions of aromatic pyrolysates were also Triassic boundary as suggested by Eshet et al. (1
detected in high abundance (Text-Figure 1 1b). These in- and Visscher et al. (1996) because:
cluded alkylbenzenes, alkylnaphthalenes and alkylphenols,
the latter group are also reflected by the large -OH band * it ranges outside the postulated time of the m
measured by FT-IR (3400 cm-1'). Alkylphenols are not par- extinction,
ticularly source specific, but have been detected from vari- * as a probable alga rather than a fungus, it cannot
ous types of algae as well as higher order plants, lignin and have acted as a saprophytic metaboliser of dead
even melanoidin like compounds (see Al-Arouri et al., 1999 vegetation resulting from the extinction event.
for more comprehensive discussion). The absence of
methoxyphenols and isoprenoids suggests there is minimal 3. Chordecystia Foster 1979 and Tympanicysta Balme
lignin and melanoidin input, respectively. Akylbenzenes are 1980 are junior synonyms of Reduviasporonites
Wilson 1962.
4. Reduviasporonites stoschianus (Balme) Elsik 1999 is a

junior synonym of Reduviasporonites chalastus (Fos-
ter) Elsik 1999. Reduviasporonites catenulatus Wilson
1962 differs from R. chalastus in being smaller, in
having more rounded constituent cells and in having
less well developed terminal rims.
>C=O
5. Brazilea helbyi forma gregata Foster 1979, recorded
Svs CH 2+CH,=3 c only from the type material of R. chalastus, may also be
OH I CH2+CH3 a junior synonym of R. chalastus.
vas CH2+CH 3
si-o 6. The size of the constituent cells present in R. chalastus
appears to be related to paleolatitude, with large ex-
4000 3500 3000 2500 2000 1500 1000
amples occurring in the paleotemperate Permian of
Wavenumbers (cm-')
China, Russia, Sverdrup and Moura, Australia and
smaller cells occurring in the paleotropical and
Text-Figure 10. FT-IR spectra of R. chalastus. Major ali-
paleoequatorialand
phatic signals are observed at 2800-3000 Permian of northern Australia, Saudi
1300-
1500 cm-. Arabia, UK and Austria.
(a) m/z 57
C E7
C9 ] C3 1 CI7
(b) TIC
B C P1 C31 D F2
9 litI N NDFIh -" -- il Im ii - -.Im .ILL - -- -
Retention Time
Text-Figure 11. (a) reconstructed m/z 57 and (b) TIC from the 6000C pyroprobe GC-MS analysis of R. chalastus. Cn = n-
B = benzene (B 1 = methylbenzene); P = phenol; N = naphthalene; DBF = dibenzofuran.
PLATE 6
All specimens are photographed using DIC; scale bar for all figures is 25 [tm.
1-36 R. catenulatus Wilson 1962 from outcrop samples from WM 2504-1, T36, CPC 36766. 19, Flowerpot WM
Greer County, Oklahoma, USA. 2504-1, J27/3, CPC 36767. 20, Flowerpot WM 2504-1
N34/4, CPC 36768. 21, Flowerpot WM 2504-1, M29
1, Flowerpot WM 2504-1, G41, CPC 36749. 2, Flow- 2, CPC 36769.22, Flowerpot WM 2504-1, M24/1, CP
erpot WM 2504-1, H40/4, CPC 36750. 3, Flowerpot 36770. 23, Flowerpot WM 2504-1, E23, CPC 36771
WM 2504-1, G44, CPC 36751. 4, Flowerpot WM 24, Flowerpot WM 2504-1, H10/2, CPC 36772. 25
2504-1, T12/2, CPC 36752. 5, Flowerpot WM 2504-1, Flowerpot WM 2504-1, E36, CPC 36773. 26, Flower
S33, CPC 36753.6, Flowerpot WM 2504-1, T7/1, CPC pot WM 2504-1, J11/2, CPC 36774. 27, Flowerpo
36754. 7, Flowerpot WM 2504-1, T22/1, CPC 36755. 66398, Y41/4, CPC 36775. 28, Flowerpot WM 2504-
8, Flowerpot WM 2504-1, S15, CPC 36756. 9, Flower- G36, CPC 36776. 29, Flowerpot WM 2504-1, G16
pot WM 2504-1, S35, CPC 36757. 10, Flowerpot WM CPC 36777. 30, Flowerpot WM 2504-1, G39/2, CP
2504-1, T20, CPC 36758. 11, Flowerpot WM 2504-1, 36778. 31, Flowerpot WM 2504-1, D43, CPC 36779
E19, CPC 36759. 12, Flowerpot WM 2504-1, E19/4, 32, Flowerpot 66398, Y41/4 (detail of Fig. 27), CPC
CPC 36760. 13, Flowerpot WM 2504-1, E15, CPC 36775. 33, Flowerpot WM 2504-1, L24/4, CPC 3678
36761. 14, Flowerpot WM 2504-1, E19, CPC 36762. 34, Flowerpot WM 2504-1, N23/4, CPC 36781. 35
15, Flowerpot WM 2504-1, E18, CPC 36763. 16, Flowerpot WM 2504-1, L27/4, CPC 36782. 36, Flow
Flowerpot WM 2504-1, N25, CPC 36764. 17, Flower- erpot WM 2504-1, C34, CPC 36783.
pot WM 2504-1, M39/1, CPC 36765. 18, Flowerpot
I:
, ..
Ill. -
it .. -
.It--.-!I .A%-
. ...
. .- -
,::%i. - . -.1.
.
Ill--
I -.. .
- .. . .
... ... .- ... .
'? -
? I-
: -
` - -
'.;e%--.%1.-'
.-'. : I I , ,--t-.%(V-,.k
% ,
.??-;'--.`---??:- .'t, -'. , .%, "':
- . . -."??.-'.'-'
. . -:? ,1 1,?! !;' -..'...-
, -, " , - .. - -`---,.r
- - ....
1 -,..t U
.A 7 -.:
.-.,.a 7-
...-..
.U .. ... : .: I .......
Itip, - :- ::::.. litili-.-AM's '--.-,LW'-,ac.%-
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7. The stratigraphical ASSERETO,

occurrences of
R., BOSELLINI, A., FANTINI the
SESTINI, N., andspec
Reduviasporonites SWEET, W.C.
suggest that R. catenulatus is
common in the 1973 The
Wordian Permian-Triassic boundaryof
(Kazanian) in the southern
Oklahom Alps
(Italy). In: Logan, A. and Hills, L.V. (eds.) The
though specimens of the size range associated with
Permian and Triassic systems and their mutual bound-
catenulatus are present very rarely in the Austr
ary, Canadian Society of Petroleum Geologists (Mem-
Mazzin Member. R. chalastus occurs in Late Permian
oir 2), Calgary, Alberta: 176-199.
and Early Triassic rocks but may be present outside BALME, B.E.
narrow sequences spanning the Permian-Triassic 1970 Palynology of Permian and Triassic Strata in the Salt
boundary. Range and Surghar Range, West Pakistan. In:
Kummel, B., and Teichert, C. (eds.) Stratigraphic
boundary problems: Permian and Triassic of West
ACKNOWLEDGMENTS Pakistan. University Press of Kansas, Department of
Geology Special Publication 4: 306-453.
BALME, B.E.
This paper relies heavily on access to rock samples and
1980 Palynology of Permian-Triassic boundary beds at
palynological strew slides containing R. chalastus from
Kap Stosch, east Greenland. Meddelelser om Gron-
many different parts of the globe. For this generous exchange
land, udgivne af kommissionnen for Kidenskabelige
of material we are grateful to: B.E Balme, and R. HelbyUndersogelser i Gronland, 200: 1-37.
BALME, B.E., and HELBY, R.J.
(Australia), Prof. H. Wopfner (Germany), O.P. Yaroshenko
1973 Floral modifications at the Permian-Triassic bound-
(Russia), Prof. H. Visscher (Netherlands), G. Warrington
(UK), Prof Shen Yan-bin (China) and the late Prof. L.R. ary in Australia. In: Logan, A. and Hills, L. V. (eds.)
The Permian and Triassic systems and their mutual
Wilson (USA). Russian material was collected by CBF
boundary, Canadian Society of Petroleum Geolo-
under the DIST-funded Australia-Russia exchange scheme:
gists (Memoir 2), Calgary, Alberta: 433-444.
Chinese samples were collected by CBF as part of an ARC
BROGLIO LORIGA, C., and CASSINIS, G.
funded research project Grant No. A39906543 awarded to Permo-Triassic boundary in the southern Alps
1992 The
Prof. Ian Metcalfe. For geochemical analyses ofR. chalastus
(Italy) and in Periadriatic regions. In: Sweet, W. C.
and discussion we are grateful to Prof. R.E Summons (MIT) Zunyi, Y, Dickens, J. M., Hongfu, Y. (eds.) Permo-
and C. Boreham (Geoscience Australia). Foster, Logan, and
Triassic events in the eastern Tethys, Cambridge
Greenwood publish with the permission of the CEO of University Press: 78-97.
Geoscience Australia. M.H Stephenson publishesBROGLIO LORIGA, C., GOCZAN, F., HAAS, J. LENNER, K.,
with the
NERI, C., ORAVECZ S., POSENATO, R., SZABO, I., and
permission of the Executive Director, British Geological
TOTH MAKK, A.
Survey. The Saudi Arabian Oil Company (Saudi Aramco)
1990 The Lower Triassic sequences of the Dolomites
kindly gave its permission to use a sample from the DILM-
(Italy) and the Transdanubian Mid-Mountains (Hun-
1 well for this study. We are grateful to our reviewers whose
gary) and their correlation. Memorie di Scienze
suggestions improved this paper. Geologiche, Universita di Padova, 17: 41-103.
BROGLIO LORIGA, C., MASETTI, D., and NERI, C.
1983 La Formazione di Werfen (Scitico) delle Dolomiti
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A Revision of Reduviasporonites Wilson 1962: Description, Illustration, Comparison and

Taylor & Francis, Ltd., American Association of Stratigraphic Palynologists are

C.B. FOSTER G.A. LOGAN

M.H. STEPHENSON P.F. GREENWOOD

Australian Centre for Astrobiology

Palynology, 26 (2002): 35-58

Wilson 1962, and may be of various

Text-Figure 1. Terminology used in morphological isde-

Text-Figure 2. Cells inR. chalastus and R. ca

delineate rectilinear shapes and

8-18 R. chalastus (Foster) Elsik 1999 from 37.62m, Bore

1, Urgen 29.2, J46/2, CPC 36673.2, SGM2, 053, CPC

margins; no inner body or cell material has been observed.

Reduviasporonites chalastus (Foster) Elsik 1999

Chordecystia chalasta Foster, 1979, p. 109-110; pl 41,

Text-Figure 3. Shrinkage of inner body in the cell cavity.

All specimens photographed using DIC; scale bar is 25 jAm.

1-10 R. chalastus (Foster) Elsik 1999 from Outcrop 11,Locality

11, 5031, H47/2, CPC 36698. 16, 5031, B47/2, CPC

"16 -.17 -..

......: . ...... . . . . : .. . . . . ..:

proximately 0.5 rtm to 1.5 [tm thick, smooth, may be

Text-Figure 7. Scattergram of the lengths and widt

" " ........10.11.1 .. .... . .

spindle-shaped and ovoid.shape from cylindrical,

.% .-? ..i?".I..,.:.(,...,......,.,..,...,...,...:..I.. - ..I,.,...,.....,...,.?.-".'i.'-.'.'..'-"-.'.-."'A?.'.'.."-.',

.. - . ... ... . . . ........

:::: ........... m-w.

? ---... ... ...... .. .

..... .. . .?:,.?,.?,,,. ., """"""""""''

-?:-.... .1 11,11 A -... .. .0

.:,CA?Xpml.-m:,-. -:. RL.I1 !%.:?-

ofR. chalastus in the Flowerpot Formation has been dated

Summary of age information

.... ....... . ...

!. ii~ ' . ... .,.., ..........

although R. catenulatus Our studies

This study Other records

chalastus except for # for0 R.0D %I- 0c

Text-Figure 9. Table of ages suggest

4. Reduviasporonites stoschianus (Balme) Elsik 1999 is a

9 litI N NDFIh -" -- il Im ii - -.Im .ILL - -- -

-- -. - ... -, ? . "?:... ...'. .... . 11 7g&,, H

-.'. :11 .. ., ::. . .:?t .

M-m 1??; w L:'.- ........ ..::L.. .1.

. . . . . . I:. : : %.. . - . . ?v? %"' .1" ,

... I ... . . "' - ... .. - .. - ... " - m..! ............ .. m ......

IW F . ?? ??` .... . ."'I"',

I::. 1% - -.11-1 -1-11 RM '--'--"---'---'--...--.-..-.-..--...--'.'- --'?:,.,.,....;....,.".......,.,:.,...."..,.- .1 .... .. .:.. . %.

.:m. . . -,." .", 1-. ,...

I is;-,i I.' . .. ? --:-? . ?:!- . ...

I .. ............ ...t . ..........

..;' % ... , . .. 1- -. 1 . ... .. ... %- !.,.. . .. . :

. : . %. ?. -. -. , ., ?."- ,.l 1-:t -,. -l`% .;I .: It

7. The stratigraphical ASSERETO,

EL-KHAYAL, A.A., and WAGNER, R.H. JIN, Y., and SHANG, Q.

ing the Permian-Triassic graphic

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