6r-
A PRELIMINARY REVIEW OF THE BIOSTRATIGRAPHY OF THE UPPERMOST
PERMIAN, TRIASSIC AND LOWERMOST JURASSIC OF GONDWANALAND
by
H. M. Anderson and J. M. Anderson.
CONTENTS
A. INTRODUCTION
B. RADIOMETRIC AGES, STANDARD STAGES AND AMMONOID ZONES OF THE
UPPER PERMIAN, TRIASSIC AND LOWER JURASSIC OF THE WORLD.
C. CHARTS 1-22
(a) Tetrapod Vertebrate Palaeontology (charts 1-5)
(b)
(c)
(d)
Chart 1 The vertebrate fauna of the Cistecephalus, Lystrosaurus and
Cynognathus zones of the Beaufort 'Group', Karroo Basin, South Africa,
with notes on the invertebrate fauna, microflora, macroflora and geology of
these zones.
Chart 2 The vertebrate fauna of the Molteno, Red Beds and Cave Sandstone
'Formations' of the Stormberg 'Group', Karroo Basin, S. Africa, with notes
on the invertebrate fauna, microflora, macroflora and geology of theEe
formations.
Chart 3 The vertebrate fauna of the Triassic of S. America, India and the
Ntawere and Manda 'Formations' of southern Africa.
Chart 4 The uppermost Permian, Triassic and lowermost Jurassic vertebrate
faunas of the Northern Hemisphere countries (excluding India) and of the
remainder of the Gondwana faunas not covered in charts 1, 2 and 3.
Chart 5 Uppermost Permian, Triassic and lowermost Jurassic tetrapod
vertebrate evolution chart of the world.
Microflora (charts 6-8)
Chart 6 Microfloral range chart for the uppermost Permian and lower half of
the Triassic of the Sydney Basin, eastern Australia. l
Chart 7 Microfloral range chart of the uppermost Permian, Triassic and ower
Jurassic of the Bowen, Clarence-Moreton & Surat Basins of Queensland,
eastern Australia.
Chart 8 Microfloral range chart of the Upper Permian and the lower half of the
Triassic of the Salt Range and Trans-Indus Ranges of W. Pakistan.
Macro/lora (charts 9-12)
Chart 9 Diagramatic key to the genus Dicroidium.
Chart 10 Macrofloral range chart of the uppermost Permian and Triassic of the
Sydney Basin and the Clarence-Moreton Basin of eastern Australia.
Chart 11 Macrofloral lists of the Triassic of the Karroo Basin, South Africa (i.e.
of the Upper CynognathUS zone and of the Molteno).
Chart 12 Macrofloral lists of all described and listed floras of the Triassic of
southern South America.
Geology and General Palaeontology (charts 13-20) .
Chart 13 Western Australia (Perth, Canning and Carnarvon Basins).
Chart 14 South Australia, Tasmania and Victoria.
Chart 15 Bowen, Galilee and Cooper Basins, eastern Australia.
Chart 16 Sydney Basin, eastern Australia.
Chart 17 Clarence-Moreton Basin, eastern Australia.
Chart 18 Southern South America.
Chart 19 India.
Chart 20 Antarctica, Southern Africa (excluding the Karroo Basin) and
Madagascar.
(e) General (charts 21-22)
Chart 21 Correlation chart of the uppermost Permian, Triassic and lowermost
Jurassic 'formations' of Gondwanaland.
Chart 22 Map of the Gondwana continents showing the known distribution of
the Triassic (including uppermost Permian and lowermost Jurassic) outcrop
ard subcrop. The known Permian basins are also given.
2
D. GENERAL DISCUSSION AND CONCLUSIONS.
1. The present state of classification and nomenclature of the animal and plant life
of the Gondwana Triassic.
2. The present state of accuracy of interbasinal correlations within Gondwanaland
and of the tie up with the Standard Stages and ammonoid zones of the Northern
Hemisphere.
3. Discussion regarding the diversity and relative abundances of the tetrapod
vertebrates and the vascular plants in Gondwanaland during the Triassic.
(a) Diversity of fauna and flora in the world today (given here as a basis for
comparison).
(b) Introductory notes on the diversity of the tetrapod vertebrates and the
vascular plants of the Gondwana Triassic.
(c) The tetrapod fauna of the Lystrosaurus zone of South Africa and its
supposed equivalents; the Fremouw Fm. of Antarctica and the Panchet Fm
of India.
(d) Brief consideration of the presently known diversity of the tetrapod fauna
of the Gondwana Triassic as a whole.
(e) The macroflora and micro flora of the Molteno 'Formation' of South Africa
and its supposed equivalents elsewhere in Gondwanaland.
(f) Brief consideration of the macroflora and microflora of the Gondwana
Triassic as a whole.
4. Discussion concerning the possibility of arriving at a reasonable estimate of the
evolution of the tetrapod vertebrates and the vascular plants in Gondwanaland
during the Triassic.
5. The relationship between the evolutionary patterns of the tetrapod vertebrates
and the vascular plants of the Gondwana Triassic.
E. ACKNOWLEDGEMENTS.
F. BIBLIOGRAPHY.
G. NOTES ADDED AT PROOF STAGE.
A. INTRODUCTION
This review pager was born out of a desire to
visualize in some Mtail the picture of the evolving
Gondwana Triassic landscape and the story of life
and ecology upon it; a desire to provide a general
setting in which to view the details of the work in
which we are actively involved at the present-the
macro flora (H.M.A.) and microflora a.M.A.) of
the Molteno 'Formation' of the Karroo Basin of
Sou th Africa.
The need to superimpose all the available
information regarding lithology, tectonics, animal
and plant life of the Triassic of Gondwanaland is
recognised and attempted.
It is also recognised that any such attempted
review will on publication already be out of date
and inaccurate. Firstly, it is virtually impossible to
gather together all the mass of relevant literature.
Secondly, a large amount of information lies
dormant either for indefinite periods in mining and
oil company files, in unpublished theses and simply
in the unpublished ideas and accumulated
knowledge of individual persons; or for shorter
periods in the hands of editors and publishers.
We hope, however, that in presenting this
review it will catalyze its own obsolescence.
We have chosen to present the review in the
form of charts. This we have done for two chief
reasons.
1. Far more information can be presented in far
less space.
2. The information can be presented more clearly
and succintly and is easily referred to.
Vagueness, blanketing the unknown or unsure
facts, is avoided.
The format which we would theoretically have
liked to follow in this review is presented below
and is followed by a very brief resume of what we
have actually managed to do.
Theoretically desired format.
I. Reconstruction of Gondwanaland (on a global
scale with its relationship to the Northern landmass
shown) including the known Triassic outcrop and
subcrop in detail, the areas of Phanerozoic
deposition and orogenesis, the stable Precambrian
basement areas, the possible lines of Palaeolatitude
and possible climatic belts.
Although only the Triassic (or what it is
supposed to encompass at the time) should be
plotted on the distribution map, it would be
desirable to include a study of the uppermost
Permian and lowermost Jurassic beds in the
following sections, for obvious reasons. Firstly, it is
not at all clear just where the lower and upper
Triassic boundary fits in the various Gondwana
'basins'. Secondly, it is desirable to know just what
happens at these boundaries, whether any
particularly dramatic events occur or whether in
fact there is simply a normal continuum of change.
II. Each Triassic Basin (or remnant basin)
discussed and/or illustrated in turn under the
following headings:
(a) Breakdown into groups, formations and
members.
(b) Isopach map; cross sections.
(c) Post-depositional folding, faulting, intrusions,
erosion and possible original extent of
deposits.
(d) Account of the underlying and overlying
strata.
(e) Range charts (as opposed to evolution charts,
as they refer only to a restricted area and
restricted time intervals) for the different
animal and plant groups where a sufficient
number of productive 'formations' renders this
meaningful.
(f) Each 'formation' elaborated on fully under the
following headings:
(1) Isopach map, facies map, cross sections.
(2) Detailed lithological and structural
account.
(3) Detailed palaeontological account of-
Vertebrates; .
Invertebrates;
Macroflora;
Microflora;
Trace fossils
giving full classified lists of all species (or
types) present, their relative abundance,
peculiarities of distribution, preservation
and lithological matrix.
(4) Possible conditions of deposition
(physiographic and climatic).
III. Evolution charts (to the same scale as the
range charts- see lIe-and correlated as accurately as
possible with the Standard Northern Hemisphere
Stages and ammonoid zones) for the vertebrates,
invertebrates, macrotlora and microtlora.
IV. Correlation chart (to the same scale as the
evolution charts and again tied in as accurately as
possible with the Standard Northern Hemisphere
Stages and ammonoid zones) incorporating all
known 'basins' and 'formations'; with the predomi
nan t palaeontological and lithological character
istics of each 'formation' included in code form.
V. Appendix giving all available absolute dates
and palaeomagnetic data (with critical assessments
of their derivation, validity and usefulness) used in
compiling correlation charts and reconstructions of
Gondwanaland.
VI. An account of the Standard Stages and
Ammonoid Zones established for the Triassic of
the N. Hemisphere, and on which all worldwide
correlations of Triassic strata should be based.
Brief resume of format followed in this review.
Due to the lack of a great deal of necessary
information, some in the published literature but
unavailable to us; some lying dormant in literature
unavailable to the public at large; some investigated
but not yet put to paper; and a good proportion
not yet investigated at all; the format followed in
this review falls far short of that just proposed, but
which should, with concerted effort be possible in
the not too distant future. Further difficulties are
the very obvious lack of standardization at present
of the use of the geological terms-group,
formation and member; and of the nomenclature
applied to the families, genera and species of the
various animal and plant groups.
3
For the sake of clarity we have indicated
below the coverage of the present review in the
same order as that of the proposed format, but it
must be noted that it is not in the order in which
this review is actually presented (see contents).
I. A composite map of the Gondwana continents
(chart 22) but not a detailed reconstruction of
Gondwanaland, has been compiled. (New Zealand,
New Guinea and the northern halves of South
America and Africa are not included in this
composite map, nor are they considered in the
review). On it we have plotted the known Triassic
(including in some cases the uppermost Permian
and lowermost Jurassic) outcrop and subcrop, and
named all the particular 'basins', boreholes, and
palaeontological localities mentioned in the charts.
The underlying 'Permian' basins (where possible)
are also included. Since we have not attempted a
reconstruction of Gondwanaland, we have also not
given the possible lines of latitude and the climatic
belts of the Triassic.
(We refer the reader here to Robinson (1971)
who provides the most comprehensive attempt,
known to us, to reconstruct the Triassic world with
its lines of latitude.)
II. We have. attempted in varying degrees to
discuss each Gondwana Triassic 'basin' as indicated -
in the proposed format, but fall far short of the
ideal, especially on the geological side. (Charts
1-20, excluding 5 and 9.)
III. It is at present only possible to attempt an
evolu tion chart for the tetrapod vertebrates (chart
5). This we have done considering the whole
world.
IV. A correlation chart (chart 21) to the same
scale as the evolution and range charts and
matched as accurately as possible with the
Standard Stages and ammonoid zones of the N.
Hemisphere, has been compiled. This chart covers
the majority of the 'Triassic' basins indicated in the
distribution map and incorporates the 'formations'
of the Triassic as well as those possibly or probably
belonging to the uppermost Permian and lower
most Jurassic. The predominant palaeontological
characteristics for each 'formation' are included in
code form.
V. The known absolute dates relevant to the
Upper Permian, Triassic and Lower Jurassic of the
world are discussed and their present usefulness in
the compilation of a Triassic timescale and in
correlations is assessed.
Since we have not attempted a reconstruction
of Gondwanaland, it has not been of use to review
the relevant available palaeomagnetic data.
VI. A discussion on the Standard Stages and
ammonoid zones of the Upper Permian, Triassic
and Lower Jurassic is presented.
4
B. RADIOMETRIC AGES, STANDARD STAGES
AND BIOSTRATIGRAPHICAL UNITS (ZONES)
ESTABLISHED FOR THE UPPER PERMIAN,
TRIASSIC AND LOWER JURASSIC OF THE
WORLD
REFERENCES:
Radiometric ages
Upper Permian - Smith 1964}
Triassic - Tozer 1964
Lower Jurassic - Howarth
1964
in "Geological Society
Phanerozoic time-scale
1964".
Standard stages and biostratigraphical units
Upper Permian- Ruzhentsev & Sarycheva 1965
(Translation by Brown 1968).
Triassic - Tozer 1967; Silberling & Tozer 1968.
Lower Jurassic - Arkell 1956.
Correlation of old zones of Spath (1934) and new
zones of Tozer (1967) for the Lower Triassic- Balme
(1968).
RADIOMETRIC AGES ESTABLISHED FOR THE
UPPER PERMIAN, TRIASSIC AND LOWER
JURASSIC.
A. Upper Permian. Smith (in "Geological Society
Phanerozoic time-scale 1964")
1. The Permz"an-Trz"assz"c boundary
"Most of the lavailable isotopic dates which
can be related to the age of the Permian-Triassic
transition are from the New England intrusive
complex of eastern Australia. The rocks of this
complex yield isotopic ages ranging from 225 to
253 m.yrs. ... and are progressively younger
northwards. They cut sparsely fossiliferous beds
thought to range in age up to Artinskian or early
Kungurian, but are generally believed ... to have
been associated in time with the Hunter-Bowen
oroge~y. An early phase of this orogeny took place
at the end of Upper Marine (low-mid to mid
Kungurian) deposition, and the isotopic age of the
earliest of the New England granites is consistent
with intrusion at this time. Later phases occurred
in mid-Newcastle (= upper stage of Upper Coal
Measures) time, but the main compression
undoubtedly followed deposition of the highest
Newcastle beds (? mid to late Tatarian), as
evidenced by the widespread non-sequence and
local unconformity between these beds and the
overlying continental beds of the Lower Narrabeen
Series (low-mid to mid-Bunter). Since the Lower
Narrabeen Series was subjected to only minor
epeirogenic movements and does not appear to
have been affected by plutonic activity, it appears
likely that even the youngest of .dated rocks-The
Stanthorpe Granite and Carpenter's Gully
molybdenite with isotopic ages of
225 m.yrs.-are pre-early-Bunter. The most likely
time of intrusion of these bodies is thought to be
during or immediately following the main phase of
the Hunter-Bowen orogeny which ... took place at
the end of the Permian Period. This is therefore
tentatively placed at 225 m.yrs."
2. The Kazanz"an- Tatarz"an boundary
"No available age-determinations bear directly
on the age of this boundary, which can thus be
estimated only by interpolation. In the type-area,
the Kazanian stage is represented by a complex
series of marine and semi-continental deposits in
which three or four major cycles of sedimentation
are separated by non-sequences or unconformi ties.
By contrast, the Tatarian stage of the type-area is
represented by only a few hundred feet of
continental beds and may have been relatively
brief. An age of 230 m. yrs. is tentatively suggested
for this boundary."
3. The Kungurz"an-Kazanz"an boundary (i.e. the
base of the Kazanian stage)
"The age of this boundary can only indirectly
be inferred from a single isotopic age determina
tion-that of 241 m. yrs. on sylvine from the
Verkhnekamensk Formation, Solikamsk, U.S.S.R.
The sylvine is about high-mid-Kungurian, and
provides the only isotopic age directly referable to
the adopted standard sequence. Analysis of both
K-Ar and Ca-K ratios of a sample of allegedly
unrecrystallized sylvine gave concordant
results, and because of this the authors consider
the isotopic age to be accurate. Kulp (1961,
p.lll0) however, believes that the calculated age
should probably be regarded as minimal because of
the readiness of sylvine to recrystallize, and in view
of this an age of about 240 m .yrs. is suggested for
the Kungurian-Kazanian boundary."
B. Triassic. Tozer (in "Geological Society
Phanerozoic time-scale 1964") See Table I
"Little information is available that can be
used with confidence to construct a Triassic
time-scale. The base of the system is variously
dated in the range 220 to 238 m.yrs., the age of
the boundary with the Jurassic is also uncertain,
but may be placed at about 190 to 200 m. yrs. Apart
from a Russian determination giving a minimum
age of 200 m. yrs. for the Middle Triassic, the only
radiometric data available are for the Upper
Triassic. "
Only 3 dates within the Triassic are available.
Ii is clear that the Triassic and its stages are in
fact very poorly dated. The ages used in our
various charts are those presented, (as the best
present estimates) in the summary of the ages
presently available in the "Geological Society
Phanerozoic time-scale 1964".
C. Lower Jurassic. Howarth (in "Geological
Society Phanerozoic time-scale 1964") See Table II
For the Lower Jurassic only four reasonably
reliable radiometric dates (all based on biotite;
K-Ar) are available.
Because of the anomaly of the 193 m.yrs. date
for the Palisade sill, (see Triassic ages) which occurs
within the Upper Triassic, being younger than the
Hotailuh botholith (194 m.yrs.) which is sup
posedly close to the Triassic-Jurassic boundary, a
best estimate of 190-195 m.yrs. is given for the
Triassic-J urassic boundary.
TABLE I. RADIOMETRIC AGES FOR THE TRIASSIC.
Palisade sill
(Intrudes Stockton Fm.
of Newark Group)
Chinle Fm.
( ores)
(Petrified Forest M.
Pitchblende ore)
Mt. Yatyrgvart
(? rock type)
New jersey
- U.S.A.
Colorado
Plateau -
U.S.A.
Russia
Middle to Upper Norian
(see chart 5)
± Mid-Norian
(see chart 5)
?
TABLE II RADIOMETRIC AGES FOR THE LOWER JURASSIC.
Takteetna Alaska probably near the Bajocian-
intrusion Toarcian boundary.-Howarth 1964
Topley British { of onm"'n ",ot'''''phi,
intrusion Columbia - position. Both these intrusions
Canada. may fit anywhere within the L.
jurassic, though the top of the
Billiton Indonesia L. jurassic is favoured for the
granite Topley intrusion.-Howarth 1964
Hotailuh British fairly accurately placed as
batholith Columbia - being close to the Triassic-
Canada Jurassic boundary.- Howarth 1964
TABLE III. STANDARD STAGES AND AMMONOID ZONES OF THE UPPER PERMIAN.
Stages & ages presented in
"Geological Society
Phanerozoic time-scale 1964"
225 m.yrs (top)
Tatarian
(continental beds)
230 m.yrs. (base)
Kazanian
(endemic marine fauna
not useful for
correia tions)
240 m.yrs. (base).
presently accepted Substages
standard stages.
(As presented in Ruzhentsev & Sarycheva 1965)
Dzhulfian
Capitanian
Guadalupian
Wardian
193 m.yrs.
Many dates
ranging from
5
20 m.yrs.-218 m.yrs.
200 m.yrs.
170 m.yrs.
179 m.yrs.
181 m.yrs.
194 m.yrs.
Ammonite
zones.
Cyclolobus
Timorites
Waagenoceras
D. Conclusions regarding the radiometric ages
available
STANDARD STAGES AND BIOSTRATI
GRAPHICAL UNITS (ZONES) FOR THE UPPER
PERMIAN, TRIASSIC AND LOWER JURASSIC.
I t is clear that very little in the way of
absolute radiometric ages are available for the
Upper Permian, Triassic and Lower Jurassic, and
certainly none that are accurately tied in with the
standard marine sections.
On our various range charts we have included
the Triassic as occupying 32~ m.yrs. (from
225-192~ m.yrs.), based on the best evidence
available. Although this gives us a convenient
working basis for the time being, not too much
reliance must be placed on it.
A. Upper Permian. Ruzhentsev & Sarycheva (1965).
(Translation by Brown, 1968)
The Tatarian and Kazanian of the Russian
Platform (W. of Urals) and Southern Urals used in
the "Geological Society Phanerozoic time-scale
1964" as the standard . stages for the Upper
Permian are considered unsuitable by Ruzhentsev
& Sarycheva 1965. This is because the Tatarian is
of continental origin, and the Kazanian has an
endemic marine fauna, and neither are practical for
in tercon tinen tal correlations.
6
The type sections of the Dzhulfian stage occur
in Nakhichevan, on the left bank of the Araks
River, S. of the Caucasus Mountains, between The
Black and Caspian Seas (i.e. Trans-Caucasia).
Although the ammonoid assemblages of this stage
are most diversified and best represented in
Trans-Caucasia, corresponding assemblages are also
found in The Salt Range (W. Pakistan), Himalayas,
New Caledonia (Pacific island E. of Australia),
Greenland and Madagascar. The zone fossil
Cyclolobus is widespread.
The Guadalupian type section occurs in W.
Texas in the Guadalupe and Glass Mountains.
These sections abound in fossils, including
ammonoids. In Mexico the fauna is even more
abundan t and well preserved. In The Glass
Mountains the stage is sub-divided into the two
sub-stages, Capitanian and Ward ian, with the zone
fossils being Tim o rites and Waagenoceras
respectively.
The resolving power of the Upper Permian
(with a duration on present estimates of 15 m.yrs.)
ammonoid zones is only of the order of 5 m.yrs. in
comparison with that of ± 1 m.yrs. for the Triassic
and Jurassic.
B. Triassic. Tozer (1967); Silberling and Tozer
(1968)
"In Alpine Europe, the classical region for the
study of the Marirye Triassic, only a dozen or so
distinct ammonoid faunas are represented, and all
bu t one of these are in the Middle and Upper
Triassic. Moreover, few of the zones characterized
by these faunas are in contiguous succession, and
some of them, because of the stratigraphic and
structural complexity of this region, are not in
directly determinable sequence. Attempts to fill
out the zonal succession of the Triassic, as known
from the Alpine region, began as long ago as 1895,
when Mojsisovics, Waagen and Diener added to it
ammonoid zones from the older parts of the
Triassic in the Salt Range and Himalayas of the
Indian region.
"A recent 'standard' ammonoid zonation for
the Triassic is that included in the correlation chart
of Triassic formations in North America (Reeside
and others, 1957) and in the Treatise on Inverte
brate Paleontology (Ark ell and others, 1957, p.
1124); it incorporates a similar mixture of Alpine,
Salt Range and Himalayan zones and adds to these
still other Lower and Middle Triassic zones from
Asia Minor, Timor, and N. America. This com
posite scheme, comprising 30 Triassic zones,
suggests a number of successive distinct ammonoid
faunas expectable in the Triassic and indicates the
usage made of the various stage and 'series names,
but it does not constitute a standard of comparison
that is objectively applicable to anyone region.
Rather, it is more in the nature of a listing of
ammonoid faunas from various places in the world
in what was believed to be their order of age.
"The marine Triassic rocks of western and
arctic North America are well endowed with
ammonoid faunas, which at one place or another
represent nearly every part of Triassic time.
Provincialism among North American faunas is in
some cases great enough to warrant separate zonal
designation of roughly correlative strata in dif
ferent parts of the region, but in general the lateral
continuity of faunas is good enough so that the
many local sections preserving parts of the faunal
sequence can be pieced together in an overlapping
fashion into a sequential framework ... . At
present, about 35 distinct biostratigraphic units of
demonstrably different age can be recognised on
the basis of ammonoid faunas in the Triassic of
North America, and some of these units combine
still more refined local faunal sequences. Hence,
using current estimates of the duration in years of
the Triassic System, the average resolving power of
the zonal units is 1.0 to 1.5 m.yrs., though the
actual time span of the individual units is of course
probably widely variable." (Silberling & Tozer
1968).
It is added that changes and additions to the
scheme are expected and that it is presented
mainly as a temporary standard to which reference
can be made.
"The faunal succession of the Triassic in
north-eastern British Columbia and in the Arctic
Islands of Canada complement one another and
together form by far the most complete sequence
of Triassic ammonoid faunas known from anyone
region. Because of its importance, Tozer (1967)
has prepared a fully documented summary of
present knowledge regarding this succession and
has established a standard sequence of Canadian
Triassic ammonoid zones." (Silberling & Tozer
1968).
It is this standard sequence presented by Tozer
(1967) for the Triassic of Western and Arctic
Canada which we have incorporated in our various
range charts.
N.B. Although Silberling & Tozer (1968)
suggest that the average resolving power of the
Triassic ammonoid zones is between 1.0 and
1.5 m.yrs. (see above) it would appear that from
the best estimate at present of the duration of
Triassic (i.e. 32Y2 m. yrs.) this resolving power is in
fact very close to 1 m. yrs.
C. Lower Jurassic-Arkell (1956)
The standard stages and biostratigraphical
units (zones) for the Jurassic, based on
ammonoids, are derived from N.W. Europe
(England, France and to a lesser extent Germany).
The zones for the upper half of the Jurassic System
are based on the English Sequence, and the lower
half predominantly on the French Sequence. The
scheme involving 58 ammonoid zones (Lower
Jurassic-17; Middle Jurassic-21; Upper
Jurassic-20) first devised in 1850 has remained
essentially the same since that date. The ammonoid
zones of the Jurassic, which on current estimates
occupied 56Y2 m.yrs. are thus on average resolved
to a little under 1 million years each, which is very
much the same as for the Triassic.
D. GENERAL DISCUSSION AND
CONCLUSIONS
We do not intend to give here a summary of
the information contained in each of the 22 charts.
Their direct perusal by the reader will be far more
worth while. Instead we intend simply to expand
on some interesting points of discussion that come
to light.
(N.B. Wherever we mention Triassic, the
uppermost Permian and lowermost Jurassic must
be considered under discussion as well.)
If one is eventually to construct a reasonably
detailed and accurate evolving picture of the
landscape and life of the Gondwana Triassic, there
are two related prerequisites which need to be
lifted to a level of some degree of sophistication-
1. The classification and nomenclature of the
animals and plants of the Gondwana Triassic.
2. The interbasinal correlation of the Geological
strata (groups, formations and members) and
the correlation of these strata with those of
the Northern Hemisphere.
1. The present state of classification and nomen
clature of the animal and plant life of the
Gondwana Triassic.
Fish and invertebrates are found very rarely in
the Triassic of Gondwanaland. They have been
cursorily' treated in the charts, and are not
discussed here. Non-vascular plants are included in
the macro floral and microfloral charts, but are
otherwise ignored.
(a) Tetrapod Vertebrates (refer to charts 1-5)
The creation of a very large number of
artificial species and genera in the past has greatly
hindered progress in the understanding of the
diversity, natural relationships, evolution and
ecology of the tetrapods of the Triassic of
Gondwanaland. However, taxonomic reviews in the
last decade or so have rectified the position to a
point where it has been possible to attempt to
compile an evolutionary and relative abundance
chart involving all described genera of tetrapods for
the Triassic of the whole world. (See chart 5.) Very
much more collecting and revision work is neces
sary, however, before any such chart can be
regarded as truly reliable. At present it merely
presents a good initial impression of the major
events that occurred.
(b) Macroflora (refer to charts 9-12, 19,20)
The state of nomenclature of the fossil leaves
found in Gondwana Triassic beds is very confusing.
Generic names for a fair number of commonly
found types have become fairly standard, but very
poorly circumscribed specific designations are still
far too freely used. It is therefore not possible at
present to attempt to compile a general evolu
tionary and relative abundance chart for the
Gondwana Triassic plants. The presentation of lists
of all standing genera and specIes (within the best
classification available) for each particular plant
bearing 'formation' is all that can be presently
given. In the case of the Sydney and Clarence-
7
Moreton basins of Australia (see chart 10) a
sufficient number of productive 'formations'
occur, to present the assemblage lists in the form
of a range chart. No relative abundances could be
included.
(c) Microflora (spores and pollen grains of vascu
lar plan ts) (Refer to charts 6-8)
Here the situation is in some respects better
and some respects worse than in the case of the
macro flora. In Australia where most of the work to
date has been done a reasonable degree of
uniformity of useage of generic and specific names
has been arrived at and it is possible to draw up
range charts (including general indications of
relative abundances) for the Sydney Basin and for
the Queensland basins (see charts 6 and 7). A range
chart has also been drawn up for the Salt Range
(W. Pakistan) (see chart 8). A fairly high degree of
conformance of results exists between these 3
charts. The few Triassic spore/pollen assemblages
that have been described from other Gondwana
areas are very similar to those of Australia and the
Salt Range. The generic and specific names that
have been applied to the forms described, however,
are very different. As a result direct comparisons of
the microfloras via assemblage lists are largely
impossible to assess.
2. The present state of accuracy of interbasinal
correlations within Gondwanaland and of the tie
up with the Standard Stages and ammonoid zones
of the Northern Hemisphere.
(Refer to chart 21 which is based on a
consideration of all palaeontological and other
information presently available.)
The continued accumulation of new in
formation will almost certai~y necessitate several
significant and many minor changes in the correla
tion chart we have attempted to compile. The
results of work in the following areas will be
particularly valuable in this regard.
(i) Comprehensive collecting and taxonomic
revision of the animal and plant groups.
(ii) Rigorous mapping and definition of geological
'groups' and 'formations'.
(iii) Recognition of 'formation' contacts
(transitional, non-depositional or erosional).
(iv) The meaning and significance of the magni
tude of the faunal and floral breaks encoun
tered.
(v) Rates of deposition enabling more accurate
plotting of the various 'formations' within the
available time scale.
(vi) Radiometric age dating of inter or intra
'formational' igneous events.
In view of the lack of marine strata, other
than in the Lower Triassic of W. Australia,
Madagascar and the Salt Range, direct correlations
with the N. Hemisphere Standard Stages and
ammonoid zones are very difficult.
Roundabout correlation via tetrapod verte
brates and microfloras is the present basis of the tie
8
In of the Gondwana and N. Hemisphere
'formations'. With improved knowledge of world
Triassic tetrapod faunas and floras these correla
tions will become very much more reliable.
3. Discussion regarding the diversity and relative
abundances of the tetrapod vertebrates and the
v.ascular plants in Gondwanaland during the Trias
SIC.
(a) Diversity of fauna and flora in the world today
(given here as a basis for comparison)
Invertebrates (Barnes 1963). ± 1 000 000 species.
(i.e. ± 95% of total animal species).
Vertebrates (Carrington 1962). ± 40 000 species.
Fish >20 000 species
Amphibians ± 1 500 species
(frogs, toads, salamanders, Apoda)
Reptiles ± 6 000 species
(snakes, lizards, turtles, crocodiles,
Tuatara)
Birds ± 8 500 species
Mammals ± 5 000 species
Flora (Vascular plants only)
(Foster & Gifford 1959; de Wit 1966;
Phillips 1951).
families genera species
Psilopsida ~ 2 4
Lycophyta 4 ± 900
Sphenophyta 1 1 ± 30
Filicophyta 11-21 ± 300 ± 10 000
Cycodophyta 1 9 ± 80
Ginkgophyta 1 1 1
Coniferophyta 7 53 ± 500
Gnetales 3 3 ± 80
Angiosperms 250-350 ± 10000 ± 200 000
It would be very valuable to present a fairly
detailed account of the distribution and natural
habitat of the above mentioned plants and animals.
An estimation could then be arrived at, of what
number of these might reasonably be expected to
be fossilised. This quite obviously cannot be done
here. We merely present the above figures to give
an impression of the very great diversity of plants
and animals today. In the plant kingdom, the
Angiosperms, comprising the bulk of today's flora,
were not yet in existence during the Triassic (or
not in the depositional areas at any rate). On the
other hand, the seed ferns, predominant in the
Triassic depositional areas, are now extinct.
(b) Introductory notes on the· diversity of the
tetrapod vertebrates and the vascular plants of
the Gondwana Triassic
(See figures provided above)
In comparison with the continental fauna and
flora of today, that of the Gondwana Triassic
appears to have been extremely limited in diversifi
cation. Before such a conclusion is justifiable,
however, we must consider what proportion of the
total plant and animal life might have been
preserved.
In the first instance, we are dealing only, or
very predominantly so, with the life of the
depositional areas, i.e. the lowland plains and the
adjacent deltaic areas and in fewer instances, inland
plains. Only very rarely are sediments with
included plant and animal remains preserved in the
hilly terrain of the upland areas. In considering the
tetrapod vertebrates of the Triassic of the whole
world only one such hill fauna is known, the ±
mid-Norian fissure fillings of the Bristol Channel
area, England (see chart 4). These fissures occurred
in the hilly Carboniferous Limestone country of
the time.
Present knowledge is such that it is not
possible to assess what proportion of the animal
and plant species occupying the Gondwana
depositional basins of any particular period during
the Triassic would actually have been preserved.
Nor is it yet possible to determine how accurately
the relative abundances of the fossils collected
reflect the true proportions of the various forms
living at the time.
These assessments will only be possible when
far more detailed work has been accomplished (for
each fossiliferous 'formation' within each
Gondwana Triassic basin) regarding the following:
1. The subtleties of the depositional environ
ment.
2. The details of lithology and sedimentological
structures associated with the plant and animal
remains.
3. The particular characteristics regarding the
association of species, their relative abundance
and their attitude, completeness and state of
preservation at each fossiliferous locality.
4. Precisely where in the depositional areas of
today fossiliferous accumulation of similar
character occur.
In order to give an indication of the diversity
and relative abundance of the presently known
species of Gondwana Triassic tetrapod vertebrates
and vascular plants, we will consider in some detail
what are probably the best known tetrapod bearing
and plant bearing 'formations' and theIr supposed
equivalents, respectively.
(c) The tetrapod fauna of the Lystrosaurus zone
of South Africa (see chart 1) and its supposed
equivalents; the Fremouw Fm. of Antarctica
(see chart 4) and the Panchet Fm. of India (see
chart 3)
THE LYSTROSAURUS ZONE OF S.
AFRICA (see chart 1). The information contained
below is based on Kitching (1968) and amplified
by recent discussions with him. (Kitching pers.
comm. 1971). See acknowledgements.
9
TABLE IV. THE TETRAPOD VERTEBRATES OF THE LYSTROSAURUS ZONE OF SOUTH AFRICA.
Reptiles
Lystrosaurus
Pro colophon
Myosaurus
Owenetta
Prolacerta
Scaloposaurids
Galesaurids
Pro terosuc hus
Moschorhinus
Amphibians
Lydekkerina
Uranocentrodon
Fish- None yet found.
No. of
species
± 6 *
1
1
1
a few
a few
1
1
1
No. of
specimens
abundant
common
8
1
5
±8
+ 40
±9
± 12
common
several
Size and
general appearance
young-medium grown Amphibious
hippopotamus herbivore
large lizard terrestrial
herbivore (or
insectivore )
mouse Terrestrial
herbivore
terrestrial
lizard- small carnivores &
leguaan insectivores
large leguaan- amphibious
small crocodile carnivore
squat leopard terrestrial
carnivore
skull length aquatic
± 5-10 cm insectivore
skull length aquatic
± 50 cm carnivore
(? piscivore)
* This figure of ± 6 species was based on Kitching's personal opinion (per. comm. 1971) and Cluver (in press) 1971.
(See chart 1) We have since receivedCluver'spaper in which he has now decided on retaining 9 species of Lystrosaurus.
Refer to chart 22 for distribution of Lystro
saurus zone. The outcrop of the zone ('formation')
is quite clearly the most extensive of any
Gondwana Triassic 'formation'. It occupies most of
the outcrop area indicated outside the broken line
on the map, i.e. below the Molteno 'Fm.'. (The
Cynognathus zone is of restricted distribution
occurring only in the southern half of the Triassic
exposure between the Lystrosaurus zone and the
Molteno 'Fm.'). The existing remnant of the
Lystrosaurus zone measures about 600 km x
300 km. The depositional area of the beds of this
zone probably originally extended over the entire
Karroo Basin indicated on the map, i.e. a vast plain
roughly equivalent in area to the largest lowland
river basins of today, e.g. the Amazon, Congo or
Mississippi.
The -Lystrosaurus zone fauna described here is
based on some 1-2 000 specimens (it must be
noted that of the more common forms-Lystro
saurus, Procolophon and Lydekkerina-only the
better material has been retained) collected from
89 localities scattered throughout the outcrop area.
The first descriptions of elements of this fauna are
over 100 years old, but the most systematic
collections have been made by ] ames Kitching over
the last 15 years or so. "Even so, most areas
around the outcrop have not been thoroughly
combed as yet." (Kitching pers. comm. 1971).
Ly strosaurus
By far the most abundant form. Found at
nearly all the localities around the outcrop. Usually
found complete in the northern half of the outcrop
where several animals are often found in close
proximity to one another, and as skulls and
scattered postcranial bones in the southern half.
Procolophon
Occurs at many localities widespread in the
southern areas of the outcrop sometimes as
complete well preserved specimens in mudstones
and siltstones and sometimes as fragmentary
material in mud-pellet congloll}erates at or near the
base of sandstone lenses. OccUrs more rarely in the
northern area from only a few localities and here
always as fragmentary material in mud-pellet
conglomerates at the base of sandstone lenses.
Where good material has been collected in the
south, whole colonies of animals appear to have
been overwhelmed by mud. As many as 6
specimens may be found in a single horizon within
an area of 2 sq. yards. In the mud-pellet
conglomerates, in north and south, the frag
mentary material is abundant and many animals
are represented at any particular locality.
Myosaurus
All eight known specimens have been collected
from an area 12 ft. x 12 ft. at a single
locality-Harrismith Dongas-at the north end of
the outcrop. The skulls were well preserved, but
most of the skeletal material was missing.
Owenetta sp.
The single specimen, from the Lystrosaurus
zone found to date is an almost complete animal
and derives from a locality near Bethulie in the
. mid-western part of the outcrop.
10
Prolacerta
The five specimens, mostly complete animals,
have been collected from widely scattered localities
around the outcrop.
Scalopsaurids
The specimens have mostly been found from
widely scattered localities along the southern
portion of the outcrop. A single specimen has been
found in the Orange Free State in the north
western part of the outcrop. Some of the
specimens are complete skulls and skeletons and
so~e only skulls.
Galesaurids
Galesaurus-one specimen from near Middel
burg in the south-west and one from near Bulwer
in the north-east. Skulls only.
G I 0 chinodontoides-Two specimens from
Harrismith in the north and one from near Bethulie
in the mid-west. Skulls only.
Micrictodon-one specimen from Harrismith in
the north.
Nanictosaurus-one specimen from near
Graaff-Reinet in the south-west. Skull only.
Notictosaurus-Six specimens have been found
to date, all from the south-west portion of the
outcrop. One specimen was found near Graaff
Reinet and the oth1r five from two localities near
Burghersdorp. Four of these (complete specimens
of an adult and three immature animals) derive
from a single block from one of the two localities.
Nythosaurus-One specimen from near Smith
field in the mid-west. Skull only.
Platycraniellus-One specimen from near
Harrismith in the north and one from near Bethulie
in the mid-west.
Thrinaxodon-± 30 specimens, nearly all
complete, have been found to date. About half the
specimens have come from a single locality at
Harrismith in the north and the other half from the
farm Newcastle (on Oliviershoek Pass) about 25
miles to the south of Harrismith. Two specimens
have been found from near Bethulie in the
mid-west. At the two main localities Thrinaxodon
is found very closely associated with Lystrosaurus
and Lydekkerina. In assessing the distribution
patterns of the Galesaurids it must be noted that
taxonomic revision of the group is urgently needed
and that some of the genera at least will certainly
become synonymous with Thrinaxodon.
Proterosuchus
The ± 9 specimens have been found at
localities scattered widely throughout the outcrop.
Portions of the postcrania occurred with some of
the specimens.
Moschorhinus
The ± 12 specimens have been found at
localities scattered widely throughout the outcrop.
Most specimens consist of isolated complete skulls.
In only one case was the complete animal found.
Lydekkerina
The numerous specimens have been collected
from only two localities i.e. Harrismith (in the
north) and from the farm Newcastle (on
Oliviershoek Pass), about 25 miles to the south of
Harrismith. These are the same two localities at
which Thrinaxodon is commonly found. All
specimens are complete and often found very
closely associated with Lystrosaurus and
Thrinaxodon.
Uranocentrodon
All the specimens (which are complete and
well preserved) have been found at a single locality
near Senekal in the north-west. All occurred in a
single horizon with specimens of Lystrosaurus
occurring in horizons immediately above and
below.
THE FREMOUW 'FORMATION' OF ANTARC
TICA (see charts 4 and 22)
The information contained below is drawn
directly from the results of James Kitching's recent
collecting trip (1970/71 season). (Colbert et al
1971.) Earlier collections consist of very frag
mentary material and add nothing to the overall
picture.
Lystrosaurus-The most prominent form but not
overwhelmingly so as in South Africa. Skulls and
scattered skeletal elements are found separately.
Pro colophon-Several specimens, all complete with
skulls and skeletons.
Small anomodonts-Three specimens, 1 complete
and two with skulls and parts of the skeleton.
Thrinaxodon-Several specimens all complete.
Prolacerta-like forms-± Four specimens, all com
plete.
Lydekkerina-Much fragmentary material. Only
one skull (more or less complete) has been found.
The faunal assemblage known to date is clearly
very similar indeed to that of the Lystrosaurus
zone. The only elements not common with the
Lystrosaurus zone forms are the small anomo
donts. Preparation of all the specimens is still
necessary before detailed comparisons can be
made.
The specimens have been collected from a
number of localities from an outcrop area far
smaller than that of the South African outcrop (see
chart 22).
THE PANCHET 'FORMATION' OF INDIA (see
charts 3 and 22)
The fauna is very poorly known as yet. (For
reference see chart 3.)
Ly strosaurus - Predominant (± 24 skulls and skeletons)
Procolophon I
Pro terosuehus
Braehyops I
Indo brae h y ops
Very little material to date
Other than in the case of Brachyops and
Indobrachyops, which are amphibians unrelated to
the two forms known from South Africa, the fauna
could well have been derived from any single
locality in the Lystrosaurus zone.
CONCLUDING REMARKS ON THE FAUNA OF
THE LYSTROSAURUS ZONE AND ITS EQUI
VALENTS
(1) The faunal assemb~age~ fro~ the
Fremouw and Panchet 'formatIOns are stIll very
poorly known but are clearly closely related in
overall characteristics and age to that of the
Lystrosaurus zone.
(2) Although the Lystrosaurus zone faunal
assemblage is the best known of any tetrapod
bearing Triassic 'formation' (inclu.ding. up~erm~st
Permian and lowermost JurassIc) It IS stlll,
according to Kitching (pers. comm. 1971), far
from comprehensively studied. Most areas of the
outcrop have only received sparse attention. From
the very small number of specimens known of the
more rare elements and the very restricted
presently known distribution. ~f certain forms
(particularly Uranocentrodon) It IS clear that there
remain significant gaps in our knowledge of the
fauna.
(3) Very little detailed geological study ~f
the Lystrosaurus zone has been undertaken and. It
is not yet possible therefore to assess the subtletIes
of the depositional environment. .
1 I
at these localities were actually equally common
but that since their natural habitat is not aquatic or
amphibious as with Lystrosaurus and Lydekkerina
they are less commonly the victims of mud floods.
It is obvious that once detailed notes are made
regarding the sediments and structures · directly
associated with the animal remains that it will be
possible to draw conclusions of the above nature
with more confidence. With a great number of
localities studied in this way the reconstruction of
a good overall picture of the depositional environ
ment should be possible.
(5) In a reconstruction of Gondwanaland the
three 'formations' discussed above lie at the apices
of a triangle whose sides are very roughly 2-3 000
miles long. (The distances obviously vary according
to the different suggested possible Gondwanaland
reconstructions.) If we assume the reconstruction
of the Upper Permian world, with lines of latitude
included, as given in Robinson (1971) (the most
comprehensive attempt known to us to date) to be
roughly correct we arrive at the following latitudes
for the three 'formations'-
Lystrosaurus zone (± {)0-:-65°S)
Fremouw 'Fm'. (± 800 S)
Panchet 'Fm'. (± 65°S)
We might thus envisage a considerable con
tinental area spanning some 20° of latitude (the
latitude values might be incorrect) whose depo
sitional basins appear to have supported a fauna
very limited in diversity and dominated throughout
by Lystrosaurus.
(6) The presently known tetrapod launa ot
the Lystrosaurus zone numbers ± 20 species, that
of the Lystrosaurus zone plus its equivalents, the
Panchet and Fremouw 'Formations', ± 23 species.
With more comprehensive collecting the total tally
might possibly reach twice I this figure. What
proportIOn of the total fmina inhabiting the
depositional basins of Gondwanaland during
Lystrosaurus zone times does this represent? How
many species left no trace of their existence? We
cannot at present attempt to answer these
questions.
(4) The attitudes and state of preservatIOn of
the various faunal elements of the Lystrosaurus
zone, and their relationships one to another as seen
in the field offer certain clues to the environment
and overall characteristics of the fauna, e.g. the
very close association of complete specimens of
Lystrosaurus and Lydekkerina at both localities at
which Lydekkerina occurs suggests that these
forms were overwhelmed by silt and mud perhaps
whilst going about their daily business in the (d) Brief consideration of the presently known
flooded flood plain of a meandering river system. diversity of the tetrapod fauna of th e Cond-
It is possible that the elements found more rarely wana Triassic as a whole (see charts 1- 5)
? lowermost Jurassic
r U. Norian-
Rhaetian
~ L- M. Nod,n
Triassic l Ani,i,n
Scythian
? uppermost Permian
TABLE V. THE TETRAPOD FAUNAS OF THE BEST KNOWN PRODUCTIVE
GONDWANA "TRIASSIC FORMATIONS"
Total
Species
Cave Sandstones 10
Red Beds 15
Ischigualasto 15
Fm
Manda Beds 15
Cynognathus 30
zone
Lystrosaurus 20
zone
Amphibian
Species
1
1
8
2
Reptile
Species
10
13
14
14
22
18
Mammal
Species
2
12
In the above table we have selected a
succession of Gondwana Triassic tetrapod-bearing
'formations' giving rough counts of the number of
species of tetrapods presently known from each.
The numbers are based on what are considered to
be valid natural species. The 'formations' chosen
are those bearing the 'richest' and best known
faunal assemblages of all tetrapod-bearing
Gondwana Triassic 'formations'. In the case of the
4 South African 'formations' the presently known
assemblages are the result of a considerable
(although by no means comprehensive) amount of
collecting over a number of decades. The Manda
and Ischigualasto 'Fm.' collections are more
limited.
(e) The macroflora and microflora of the Molteno
'Formation' of South Africa and its supposed
equivalents elsewhere in Gondwanaland.
THE MACROFLORA (VASCULAR PLANTS) OF
THE MOLTENO 'FORMATION' OF S. AFRICA
Refer to chart 11 for a fairly detailed account
of this flora and its peculiarities of distribution
according to the 21 localities (from which more or
less comprehensive collections have been made)
scattered around the outcrop (see chart 22).
After detailed consideration of the ± 3 350
selected specimens! of leaves (and sphenophyte
stems) collected from these 21 localities we
estimate the presence of ± 87 reasonably clearly
definable 'species'. As can be seen in the case of
Dicroidium (see diagrammatic key on chart 9) the
delineation of 'species' is by no means always easy.
However, in most of the other genera the task is
simpler. (One of us (H.M.A.) is at present preparing
a comprehensively illustrated account of the
Molteno Flora, which should be published in
1972.)
With further detailed collecting from the
presently known localities and many more,
resulting as it should in the discovery of the very
rare elements of the flora, our estimate is that the
eventual number of 'species' will not exceed 125.
THE MACROFLORA (VASCULAR PLANTS) OF
THE IPSWICH 'GROUP' OF E. AUSTRALIA
Refer to chart 10 for an account of the flora.
The total number of leaf species (including
sphenophyte stems) presently valid in the literature
on the Ipswich 'Group' (including the Esk flora) is
approximately 80. The Ipswich flora forming the
bulk of this total, was last revised in 1947 by Jones
and de Jersey. The revision was, for that time, very
comprehensive. The Esk flora was last dealt with
by Walkom in 1928. With further detailed
collecting and revision we presume that the total
flora will roughly match that of the Molteno in
diversity of 'species'.
At least half, and probably a lot more, (it is
not possible to judge accurately from the literature
available) of the Ipswich flora is common with that
of the Molteno.
THE MACROFLORA (VASCULAR PLANTS) OF
THE 'FORMATIONS' IN ARGENTINA AND
BRAZIL PRESUMED TO BE EQUIVALENTS OF
THE MOLTENO (see chart 12)
With the limited knowledge presently avail
able, it is of course very difficult to assess (as is
suggested on the correlation chart-chart 21)
whether all the plant-bearing 'formations' of
Argentina and Brazil are in fact of the same age.
Nor is it really possible at present, to assess how
closely in age they match the Molteno. There is,
however, no good evidence to suggest that the
above mentioned South American and South
African 'formations' are not equivalent in age and
for the present this is the best assumption.
Considering the apparent widespread similarities of
conditions in the depositional basins of Gondwana
land at anyone time (based on many independent
lines of evidence, e.g. the discussion just held on
the Lystrosaurus zone and its equivalents) this
conclusion appears justified.
The total number of species of leaves (and
sphenopayte stems) presently standing in the
literature- for all the Argentina and Brazil
'formations' is approximately 140 (see chart 12). It
is assumed once again that; with revision and
further comprehensive collecting, the number of
'species' will probably roughly match that of the
Molteno. It is extremely difficult to judge just how
many species are in common with the Molteno
and/ or Ipswich floras since a great many have been
inadequately described and illustrated to date.
However, it appears that well over half of the
Molteno 'species' occur in S. America.
CONCLUDING REMARKS ON THE MACRO
FLORA OF THE MOLTENO AND ITS SUP
POSED EQUIVALENTS
(1) Excellent floras, undescribed as yet but
obviously closely comparable to those of the
Molteno, Ipswich and S. American 'formations',
occur in many other Gondwana basins.
(2) Comprehensive collections of the flora
from all plant-bearing Molteno age 'formations',
and their description in the light of extensive
taxonomic revision is necessary.
(3) We can at present only make a very rough
estimate of the total number of vascular plant
'species' that were preserved in all Gondwana
depositional basins during Molteno times. Con
sidering what we presently know it is difficult to
imagine that the number will exceed 200.
(4) What proportion of the species of vascu
lar plants actually growing in the depositional
basins during Molteno times does the preserved
flora represent? Again, as in the case of the
Lystrosaurus zone and its equivalents, very limi ted
geological detail is known concerning the
'formations' in question. No attempts have been
made to give detailed accounts of the lithology and
sedimentary structures or the peculiarities of leaf
associations and relative abundances in the various
horizons or portions of an horizon within the
section at any particular fossil locality. In other
words no attempts have been made to assess the
details of the original growing and depositional
conditions at and nearby the plant localities.
Until this is done and until the overall
depositional regime of each 'formation' is known,
we can hardly hope to assess wi th confidence the
proportion of the total plant species, of the basins,
that might have been preserved. Nor can we hope
to be really aware of the true relative abundances
of the species involved.
THE MICROFLORA (VASCULAR PLANTS) OF THE
MOLTENO AND ITS SUPPOSED EQUIVALENTS
(1) The Molteno 'Formation' figures are
based on the detailed study of numerous samples
from many Molteno localities made by one of us
UMA.) over the past few years. The micro flora,
which has received no attention in the available
literature to date, will be published along with the
macroflora in 1972.
(2) The Potrerillos 'Formation' figures are
based on our estimations of the number of 'forms'
present, based on the photographic plates in Jain
(1968). Jain described 54 genera and 95 species
from about 20 samples from a single exposure
(covering about 160 m of strata). Considering only
consistently recognisible, distinct basic 'forms' (i.e.
not including as distinct the various abnormalities,
or the different stages of dehiscence, splitting,
opening and folding) we consider that only ± 34
'forms' are present.
(3) The Ipswich 'Group' figures are based on
de Jersey (1962 and 1970b) (see chart 7).
(4) Microfloras of supposed equivalent age to
the three 'formations' just mentioned have been
very briefly described, or merely mentioned, in the
13
literature from many other Gondwana basins (see
chart 21). No details for the purposes of this
discussion can be gleaned from these references,
however.
(5) In considering microfloras we can only
legitimately talk of distinct morphological 'forms'
rather than 'species'. This holds particularly for the
non-striate disaccate pollen grains presumed to
derive mainly from Dicroidium; and the Cyca
dopites-type pollen grains presumed (by compari
son with modern day forms) to be the
predominant pollen-type of the Cycads and
Ginkgos. The morphological possibilities are
limited to such an extent in these two pollen
groups that, in the case of the Molteno micro flora
only 7 and 4 distinct 'forms' can be recognised
respectively. In comparison, in the Molteno macro
flora, 25 reasonably distinct Dicroidium 'species'
and 19 Cycad + Ginkgo 'species' can be recognised
(see charts 9 and 11). In total only some 64 spore
and pollen 'forms' can be recognised as compared
to approximately 87 leaf 'species'.
(6) Although microfloral assemblages repre
sent predominantly the plants growing in the
immediate or close vicinity, contamination of the
flora can be expected to be effected by the influx
by wind or water of spore and pollen types derived
from plants growing outside the depositional basin.
(7) In both the Potrerillos and Ipswich
microfloras about 74 of the 'forms' are clearly
closely comparable with 'forms' from the Molteno.
(8) Considering the number of recognisable
'forms' presently known and taking into account
the great similarities between microfloras in the
different areas of Gondwanaland, the total micro
flora of the Molteno and its equivalents might be
. ~,
TABLE VI. THE MICROFLORA (VASCULAR PLANTS) OF THE MOLTENO AND ITS SUPPOSED EQUIVALENTS
Microfloral groups (with possible and/or Approximate No. of 'forms' !
probable parent plant groups)
Trilete & monolete cavate spores of
probable lycophyte origin
Other trilete and monolete spores
(Sphenophyta, Filicophyta and
additional Lycophyta).
Striate disaccate pollen grains
(possibly Glossopteris but perhaps of
other gymnospermous origin).
Non-striate disaccate pollen grains
(mainly Dicroidium but also probably
including other gymnospermous plants).
Cycadopites and other miscellaneous
gymnospermous pollen grains.
Alete forms (possibly including both
spores and pollen grains).
Total
Molteno
'Fm';
S. Africa
2
35
2
7
5
13
64
Potrerillos Ipswich
'Fm'; Cacheuta 'Grp';
Basin; S. E. Australia
America
3 4
13 25
3
7 (Several)
4 1
4 5
34 35 +?
14
expected eventually to approach 125 distinct
'forms'.
(f) Br£ef cons£dera#on of the macroflora and
m£croflora of the Gondwana Tr£ass£c as a
whole.
(1) The Molteno and supposed equivalent
'formations' are by far the most productive as
regards macrofloral (leaf) remains. Only in E.
Australia (see chart 21) does there exist a
reasonable scatter of plant-bearing 'formations'
through the rest of the Triassic. It appears that it is
from this region that most of the story of
Gondwana Triassic macrofloras will have to be
told.
Other than from the Ipswich 'Group' very little
systematic description of E. Australian floras has,
however, so far been undertaken. Townrow (pers.
comm. 1969) has over the past ten years or so
made extensive collections of many of the floras
under question, but has not, as yet, published the
results of most of his labours.
(2) As with the macroflora we have to turn
to E. Australia for the most complete succession of
good spore and pollen-bearing Triassic 'forma
tions'. The bulk of our present knowledge of
Gondwana Triassic microfloras is based on work
done in Australia (see charts 6 and 7).
)
4. Discussion concerning the possibility of arriving
at a reasonable estimate of the evolution of the
tetrapod vertebrates and the vascular plants in
Gondwanaland during the Triassic.
(1) We can at best only hope to discuss that
part of the evolutionary story which took place in
the lowland areas. How much disjointing of the
story will have been effected by the arrival of new
elements from the upland areas, unknown
previously in the lowland areas, and not evolved
from any of the forms occurring there, will only
become evident with much more precise work.
(2) A study of the correlation chart (chart
21) will give some indication of what periods of
Triassic time are represented by strata bearing
tetrapod vertebrates, macrofloras and microfloras.
It must- be clearly noted, however, that the
correlation chart is based on present evidence, and
we feel that it will look significantly different on
revision in years to come.
(3) As a consequence of the faunas and floras
being so constant from one depositional area to
another it is theoretically possible to compile the
evolutionary story by making use of all the
available information from all the 'formations' no
matter in which basin or country they occur.
(4) In this way tetrapod-bearing 'Formations'
can be pieced together to occupy most of the
Triassic succession; Microflora-bearing 'formations'
occur through most of the Triassic excluding the
Norian; Macroflora-bearing 'formations' offer
much the same overall coverage as do the
microfloras, but in a more restricted sense.
5. The relationship between the evolutionary
patterns of the tetrapod vertebrates and the
vascular plants of the Gondwana Triassic (see
charts 5 and 6-8).
That a very definite relationship between the
evolutionary patterns of the plants and tetrapods
exists, is clearly shown by comparing the tetrapod
evolution chart (chart 5) and the micro floral range
charts (charts 6-8). Extensive, major floral and
faunal changes occur roughly simultaneously at 3
levels within the period covered by this review; (a)
somewhere near the top of the Permian (but below
the Lystrosaurus zone and its equivalents); (b)
somewhere within the upper half of the Scythian;
and (c) somewhere near the base of Rhaetian.
Precisely how rapidly these changes occurred and
whether the plant changes preceded the vertebrate
changes or vice versa, or whether they occurred
virtually concurrently cannot be assessed at
present.
E. ACKNOWLEDGEMENTS
We would like to thank Arthur Cruickshank,
James Kitching and Chris Gow, members of the
Bernard Price Institute for Palaeontological
Research, without whose willing help the com
pilation of the vertebrate lists for southern Africa
would not have been possible.
Arthur Cruickshank helped considerably in the
recommendation of literature and with general
guidance and criticism throughout.
James Kitching deserves a special debt of
gratitude. He is at present working on the
enormous task of compiling full comprehensive
lists of all reptiles found from each known fossil
locality from the Beaufort Beds (upper half of the
Permian and lower half of the Triassic) of South
Africa. This study involves some 10 000 recognis
able specimens (predominantly housed in S.
Africa) from 5-600 localities. He will be
presenting, therefore, an extremely detailed
account of the reptile life of the several faunal
zones of the Beaufort beds. He has been perfectly
willing to impart to us any information we wished
to have. We would also like to thank him for
letting us incorporate the results of his very recent
(1970/71 season) and highly successful collecting
trip to Antarctica (see reptile list on chart 4 and
localities on chart 22) even though our review will
possibly appear before his own published account.
Chris Gow helped with the compilation of the
vertebrate lists and qualifying notes for the Red
Beds and Cave Sandstones of South Africa-see
chart 2. Much of the information contained in this
chart is unpublished and is based on his own
personal knowledge.
We would also like to acknowledge with
thanks the following persons-Hugh Allsop, Basil
Balme, Peter Barrett, Joseph Bonaparte, Andre
Keyser, Frederico Lange, Christian Mendrez, Edgar
Riek, John Townrow, Alick Walker, Hans Welkae,
with whom fruitful discussions have been held,
from which certain information has been drawn.
Marcia Orelowitz compiled the bibliography as
part of the requirements for her Diploma in
Librarianship (Witwatersrand University). For her
work and ever cheerful presence we thank her most
deeply.
Janet Cronje helped enormously in drawing up
the charts. A great deal of the art work including
the map (chart 22), the tetrapod evolution chart
(chart 5) and the microfloral range charts (charts
6-8) is hers. Her artistic soul has helped us to
endure.
Anna Benecke, busy on her Ph.D. thesis,
stopped to help us with corrections and tidying up.
The manuscript for the written part of this
review and the bibliography were typed by Eileen
Barrett, Pat Sharland and Sue McCarthy.
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