International Journal of Coal Geology, 3 (1984) 205--255 205

Elsevier Science Publishers B.V., Amsterdam -- Printed in The Netherlands

CHANGING PATTERNS OF PENNSYLVANIAN COAL-SWAMP

VEGETATION AND IMPLICATIONS OF CLIMATIC CONTROL
ON COAL OCCURRENCE

TOM.L. PHILLIPS and R U S S E L A. P E P P E R S

Department of Botany, University of Illinois, Urbana, IL 61801 (U.S.A.)
Coal Section, Illinois State Geological Survey, Champaign, IL 61820 (U.S.A.)
(Received March 28, 1983; revised and accepted September 13, 1983)

ABSTRACT

Phillips, T.L. and Peppers, R.A., 1984. Changing patterns of Pennsylvanian coal-swamp
vegetation and implications of climatic control on coal occurrence. Int. J. Coal Geol.,
3: 205--255.

Improved regional and interregional stratigraphic correlations of Pennsylvanian strata

permit comparisons of vegetational changes in Euramerican coal swamps. The coal-swamp
vegetation is known directly from in situ coal-ball peat deposits from more than 65
coals in the United States and Europe. Interpretations of coal-swamp floras on the
basis of coal-ball peat studies are extended to broader regional and stratigraphic patterns
by use of coal palynology. Objectives of the quantitative analyses of the vegetation in
relation to coal are to determine the botanical constituents at the peat stage and their
environmental implications for plant growth and peat accumulation. Morphological
and paleoecological analyses provide a basis for deducing freshwater regimes of coal
swamps.
Changes in composition of Pennsylvanian coal-swamp vegetation are quite similar
from one paralic coal region to another and show synchrony that is attributable to
climate. Paleobotany and paleogeography of the Euramerican province indicate a moist
tropical paleoclimate. Rainfall, runoff and evapotranspiration were the variable climatic
controls in the distribution of coal-swamp vegetation, peat accumulation and coal re-
sources. In relative terms of climatic wetness the Pennsylvanian Period is divisible into
five intervals, which include two relatively drier intervals that developed during the
Lower--Middle and Middle--Upper Pennsylvanian transitions. The climate during Early
Pennsylvanian time was moderately wet and the median in moisture availability. Early
Middle Pennsylvanian was drier, probably seasonally dry-wet; late Middle Pennsylvanian
was the wettest in the Midcontinent; early Late Pennsylvanian was the driest; and late
Late Pennsylvanian was probably the wettest in the Dunkard Basin. The five climatic
intervals represent a general means of dividing coal resources within each region into
groups with similar botanical constituents and environments of peat accumulation.
Regional differences in basinal geology and climate were significant variables, but the
synchronous control of paleoclimate was of primary importance.

INTRODUCTION

The paleogeography during the Carboniferous and the numerous studies

of floras including coal-swamp vegetation have established that the Eurameri-

0166-5162/84/$03.00 © 1984 Elsevier Science Publishers B.V.

206

can coal belt had a wet tropical climate (Wegener, 1929; Chaloner and
Creber, 1973; Schopf, 1973, 1975, 1979; Frakes, 1979; Habicht, 1979;
Scotese et al., 1979; Ziegler et al., 1979, 1981; Smith et al., 1981; Morel
and Irving, 1981; and Zharkov, 1981). More in situ and anatomically well
preserved vascular plant deposits occur in coal seams of Carboniferous
age than in deposits of any other geological period. Coat-ball peats have
been found in more than 65 coal seams with those in Europe occurring
largely in the Westphalian A-C and those in the United States limited to
the Middle and Upper Pennsylvanian (Phillips, 1980). Quantitative studies
of the botanical constituents of coal and subsequent paleoenvironmental
implications became feasible on a broad scale a b o u t a decade ago and are
strongly complemented by a much larger data base from coal palynology.
The anatomically preserved plants from coal-ball peats provide the best
paleobotanical evidence that coal-swamp plants of the tropical belt were
the same from the Ukraine to Oklahoma. Interregional similarities also
have been noted in the compression floras and in coal-spore floras. These
have allowed correlations between western Europe and the proposed Penn-
sylvanian stratotype section in the Appalachians (Gillespie and Pfefferkorn,
1979) and between western Europe and the Illinois Basin Coal Field (Peppers,
1979b), which is our primary reference area for interregional comparisons
of coal-swamp vegetation. It is important to note that these two independent
correlations are consistent with each other. Nevertheless, there are few
detailed stratigraphic correlations of coals between these two Pennsylvanian
coal regions in the United States (Wanless, 1939; Kosanke, 1973).
In this overview of Euramerican coal-swamp vegetation and its relation
to climate and coal we summarize pertinent revisions in stratigraphic cor-
relations of the United States and Europe. We present approximate cor-
relations of coal seams in the Morrowan of the Illinois Basin Coal Field
and the lower Middle Pennsylvanian in northeastern Tennessee. This time of
change in coal-swamp vegetation is relevant to our thesis of synchronous
primary climatic control in the United States and Europe. Although the
evidence for a tropical paleoclimate in the Euramerican coal belt is over-
whelming, we provide a brief review of temperate to cooler climatic regimes
and coal-swamp vegetation of Angaran and Gondwanan provinces. Similarly,
while the climatic trends in the Euramerican coal belt may be more evident
in coal swamps, it is important to clarify how coal-swamp floras differed
from those of other wetlands, the lowlands and uplands and how, in turn,
these floras responded to inferred climatic changes.
Our previous summary of coal-swamp vegetational patterns during the
Pennsylvanian in the Illinois Basin Coal Field (Phillips et al., 1974) was
concerned primarily with the recognition and documentation of major
changes rather than their causes. Two principal times of major change
approximately divide the Pennsylvanian into Lower, Middle and Upper
Series. The most drastic paleobotanical change occurred during the Des-
moinesian--Missourian (Middle--Upper Pennsylvanian) transition. Dominant
 207

lycopod trees, characteristic of most Euramerican coal swamps, became

extinct and were supplanted predominantly by tree ferns. The early interval
of more subtle vegetational changes, beginning at the Lower--Middle Penn-
sylvanian transition, consisted of reduction in total abundance and fluc-
tuation in dominance of lycopod trees; there were subsequent introductions,
marked evolution, and increases in abundance of tree ferns, seed plants
and small herbaceous plants. W e considered only the drastic Middle--Upper
Pennsylvanian transition from lycopod-dominated to tree fern-dominated
coal swamps as probably Euramerican in scope. W e attributed this transition
to a broad, abrupt climatic change across the coal belt. Much of the new
data presented herein pertains to the early Middle Pennsylvanian which
also represents a time of synchronous but less severe change across m u c h of
the Euramerican tropics.
Our emphasis on primary paleoclimatic control of the coal-swamp vege-
tation specifically relates to the trends in freshwater regimes (net rainfall,
runoff and evapotranspiration). If our interpretations are correct, there
were two major intervals of drier climate and three of typically wet Penn-
sylvanian climate, all of different magnitudes. The times of change from
wet to less wet correspond approximately to established epoch boundaries
of the Lower--Middle and Middle--Upper Pennsylvanian and to the West-
phalian A--B and Westphalian--Stephanian boundaries in western Europe.
If paleotectonics were controlling or important contributory factors in the
changing wetness trends of the paleoclimate, then the onset of drier con-
ditions is a clue to the timing of some major tectonic events during the
Pennsylvanian.
The feasibility of comparing coal-swamp vegetation and environmental
implications from one region to another and ultimately interpreting paleo-
climatic trends depends on a number of factors that we develop in sub-
sequent parts of our paper. The relatively small number of genera and
species of coal-swamp trees that contributed the bulk of peat had very long
stratigraphic ranges for the most part. Thus, environmental implications
suggested by a dominance of certain kinds of trees are applicable over much
of Pennsylvanian time. The Euramerican coal belt remained in the paleo-
tropics throughout the Pennsylvani~n, thus the key climatic variable was
net available freshwater.
The diversity of growth and reproduction among woody seed plants,
heterosporous arborescent lycopods and homosporous tree ferns provided
different kinds of responses to changes in relative wetness. In turn, the
generally dominant heterosporous lycopods represent a spectrum of very
wet to relatively drier or seasonally wet-dry adaptations for sexual repro-
duction in wetlands. So far as we know, none of the lycopods had the
capability of vegetative propagation. Interpretations of strategies for growth
and reproduction should be consistent with data derived from paleoecological
analyses of c o m m u n i t y assemblages in peat profiles, with the composition
of peat in terms of preservation and organ components, and, in general,
with estimates of identified bituminous coal resources on a regional basis.
208

Because coal-swamp forests began and ended in aquatic environments

with a minimal accumulation of clastic sediment, the successive habitats
of forests also became environments of deposition and the edaphic sub-
strates. Progressive changes in freshwater regimes during the temporal
span of a coal swamp produced a cycle of in situ growth, peat deposition,
and substrate penetration. As a result, the peat provides the most direct
means of comparing the growth and reproductive implications of successive
forest communities with the depositional environments.
An example of the kind of deductive logic developed for interrelating
these sources of evidence to paleoctimate is the following: the kinds of
trees that were adapted to the wettest coal-swamp conditions (standing
water and net rising water table) should have been dominant in the wettest
stratigraphic intervals; should result in greater net peat accumulation because
the wettest regime is most conducive to submergence and peat preservation;
should result in peat composition of high shoot-to-root ratios and less
fusain for the same reason; and should produce the bulk of the peat in the
intervals of maximum coal resources.
Coal-swamp communities from which a coal seam is derived are repetitious
in succession with usually one or sometimes two kinds of communities
much more frequent than others; such communities dominate the seam
vegetation as a whole. In turn, the similarity of major kinds of coal-swamp
communities from seam to seam indicates that there should be predictable
regional patterns of qualities of coal resources during most of the Penn-
sylvanian. Although the five proposed intervals of climatic wetness repre-
sents a general means of grouping coal resources with similar peat origins
within each region, there are exceptions.

METHODS AND SOURCES OF DATA

This summary of studies on the ecology of Pennsylvanian coal-swamps

of the Euramerican coal belt draws heavily on the results of our research
team based at the University of Illinois and the Illinois State Geological
Survey where the paleobotanical preparations and specimens are housed.
The techniques for study and quantitative analyses of coal-ball peats are
those given by Phillips et al. (1976, 1977) and by Phillips and DiMichele
{1981). The stratigraphic patterns of coal-swamp vegetation were reported
by Kosanke (1947), Phillips et al. (1974), Peppers (1979a), Phillips and
Peppers (1979), and Phillips {1980, 1981). The morphological, systematic,
and paleoecological relations of lycopods in coal swamps were established
by DiMichele (1979a,b, 1980, 1981), DiMichele et al., (1979), DiMichele
and Phillips {1979), Leisman and Phillips (1979), and Phillips (1979).
Studies of coal-ball peats from specific coals and sites include those by:
Phillips (1976) -- lower Westphalian A of England; Phillips and Chesnut
(1980) -- lower Middle Pennsylvanian of Eastern Kentucky; ( R a y m o n d ,
1980) -- lower Desmoinesian of Iowa; Eggert and Phillips {1979, 1982) --
 209

Staunton Formation of Indiana; Phillips et al. (1977), DiMichele and Phillips

(1980), and Phillips and DiMichele (1981) -- Herrin (No. 6) Coal Member
of the Illinois Basin Coal Field; and Galtier and Phillips (1979) --Missourian
of the Illinois Basin Coal Field and Stephanian A of France.
The palynology of coals in the Illinois Basin Coal Field has been studied
by Schopf (1938), Kosanke (1947, 1950, 1964), Guennel (1952, 1958),
Winslow (1959), Peppers (1964, 1970, 1979a,b), and Peppers and Pfefferkorn
(1970). Spore data from Tennessee coals are based on one channel sample
from each coal analyzed; all samples are from west of the Pine Mountain
Thrust Fault. They were collected by Cropp (1963) who reported on the
spore genera from some of the coals. The identified Pennsylvanian bitu-
minous coal resources of the United States were compiled and plotted
stratigraphically by region by Phillips et al. (1980).

STRATIGRAPHIC CORRELATIONS

In order to compare the coal-swamp vegetation patterns of the Illinois

Basin Coal Field with other coal regions, reasonably accurate stratigraphic
correlations of Upper Carboniferous strata are essential. We briefly indicate
some of the recent changes and persistent problems because they have a
significant bearing on the interregional recognition of similar patterns of
change. A summary correlation chart is shown in Fig. 1.

PENNSYLVANIAN SYSTEM

LOWER MIDDLE UPPER SERIES

MORROWAN ~OK~"~ OESMOINESIAN MISSOURIAN ~VIRGILIAN MID-CONTINENT
SERIES
CASEYV,LLE ASSOTT CARSONOALE MO ESTO ONO I .ATTOON ILLINOIS
FORMATIONS
POTTSVILLE ALLEGHENY CONEMAUGH I GAHELA
MONON- APPALACHIAN
FORMATIONS
UPPER C A R B O N I F E R O U S SYSTEM
WESTPHALIAN STEPHANIAN EUROPEAN
A , B , (; I V SERIES
UPPER U.S.S.R.
MIDDLE CARBONIFEROUS
CARBONIFEROUS SYSTEMS
U.S.S.R.
BASH K I R IAN MOSCOVIAN KASIMOVlAN iGZE LIAN DIVISIONS
ISGS1982

Fig. 1. Correlation of time-stratigraphy of the Pennsylvanian System in the United

States with that of the Carboniferous of Europe and the U.S.S.R.

United States

Several major correlations of the Pennsylvanian have appeared since

those of Wanless (1939), Moore et al. (1944) and Read and Mamay (1964).
A generalized correlation is presented in "Paleotectonic Investigations
of the Pennsylvanian in the United States" (McKee and Crosby, 1975)
and numerous regional correlations appear in the U.S. Geological Survey
210

Professional Papers, 1110A-L and l l l 0 M - D D (1979). In a compilation of

Pennsylvanian coal resources of the Midcontinent and Appalachian coal
regions, Phillips et al. (1980) assembled coal correlations based on earlier
studies and on information from geologists working in various regions.
The American Association of Petroleum Geologists is currently sponsoring
the preparation of a series of charts showing "Correlation of Stratigraphic
Units of North America" (COSUNA).

Morrowan--A tokan--Desmoinesian boundaries

Defining boundaries in the Midcontinent series, particularly in relation
to the Atokan Series in the Western Interior Coal Region, is necessary for
correlations to other regions. The Lower--Middle Pennsylvanian boundary
was designated in the Midcontinent as between the Morrowan and Atokan
(Bradley, 1956). The Morrowan Series actually correlates with the Lower
and lower Middle Pennsylvanian in the Appalachian Coal Region (Rice,
1977). Cobb et ah (1981) adopted a similar position for the correlation.
A somewhat higher position for the top of the Morrowan was adopted
for the correlation of Lowland Basins and Arches (Shaver et al.,in press) as
part of the C O S U N A series. The Lower--Middle Pennsylvanian boundary
correlates with about the top of the Caseyville Formation in the Illinois
Basin Coal Field. The Morrowan has been considered equivalent to about
the lower half of the CaseyviUe Formation in the IllinoisBasin Coal Field
(Kosanke et al., 1960) or the entire Caseyville (Hopkins and Simon, 1975)
and to the lower half of the Pottsville Series (Moore et al., 1944; M c K e e
and Crosby, 1975). The Morrowan--Atokan boundary was placed consider-
ably higher in Indiana, at the base of the Brazil Formation (equivalent
to the upper part of the Abbott Formation in Illinois)based on ostracod
studies by Shaver and Smith (1974).
As originally defined (Keyes, 1893), the Desmoinesian Series includes
the lower part of the Pennsylvanian of Iowa and thus the entire Cherokee
Group. Ravn (1981, Ravn et al.,in press) found that spore assemblages in
coals in the lower part of the Cherokee indicate that the coals are correlative
with those in the upper part of the Abbott Formation of Illinoiswhich was
classified as Atokan (Shaver and Smith, 1974; Hopkins and Simon, 1975;
M c K e e and Crosby, 1975). Thus, the upper part of the Abbott Formation,
which is correlative with the Brazil Formation (Peppers and Popp, 1979),
could either be Atokan or Desrnoinesian in age.
Ravn (1979, 1981, Ravn et al., in press) and geologists with the Iowa
Geological Survey, in preparing their portion of the C O S U N A correlation,
have proposed that the Desmoinesian be redefined to extend d o w n to but
not include the highest occurrence of the spore Dictyotriletes bireticulatus,
just below the coal underlying the Seville Limestone Member. Peppers
(unpublished data) has observed this species in an Iowa coal correlated with
the Rock Island (No. 1) Coal M e m b e r of Illinoisthat underlies the Seville
Limestone. However, the highest occurrence of D. bireticulatus in the
 211

Illinois Basin Coal Field is in the Herman Coal Member just above the
Seville Limestone of Illinois and in "Coal II" (name n o w discontinued)
of Indiana just above the Perth Limestone Member (correlative with the
Seville). The lowest Fusulinella-bearing stratum in Iowa (the Seville Lime-
stone), remains in the Desmoinesian, b u t in Indiana Fusulinella begins
lower in the limestone between the Lower Block Coal Member and the
Upper Block Coal Member in the Brazil Formation. Shaver et al. (in prepa-
ration) place the t o p of the Atokan slightly higher, at the base of the Seville
Limestone.
In this paper we are adopting the lower Desmoinesian boundary proposed
in Iowa -- just below the R o c k Island (No. 1) Coal -- which also conforms
to the position of Hopkins and Simon (1975).

Eastern K e n t u c k y
A correlation of Pennsylvanian coal beds, coal zones and other strati-
graphic units in the six coal-reserve districts of eastern Kentucky was pre-
sented by Rice and Smith {1980). Correlations between districts were
based mainly on the recognition of key marine beds, coal seams, and sand-
stone deposits. The Lower and Middle Pennsylvanian Series were undifferen-
tiated. The Upper Cumberland District, the only district that lies east of the
Pine Mountain Thrust Fault, contains the Upper Path Fork coal bed, an
important source of lower Middle Pennsylvanian coal-ball peats. Correlation
of stratigraphic units on either side of the fault is particularly difficult.

Illinois Basin Coal Field and northeastern Tennessee

Palynological correlations between coals of the Illinois Basin Coal Field
and those of northeastern Tennessee and eastern Kentucky regions of the
Appalachians are necessary for stratigraphic alignment and for quantitative
comparisons of the spore floras for synchrony of major changes beginning
near the Lower--Middle Pennsylvanian boundary. The northeastern Ten-
nessee analyses are completed and presented in Fig. 2. The boundary equiva-
lent to the Westphalian A-B of Europe was placed near the base of the
A b b o t t Formation in Illinois and between the H o o p e r coal bed and the
overlying Poplar Creek coal bed in the Crooked Fork Group of Tennessee.
This boundary is based on the beginning of the range of Endosporites
globiformis (Fig. 3) which first appeared in coals in Illinois, Great Britain
and the Ukraine (Teteryuk, 1976) immediately after Schulzospora rara
disappeared. In Tennessee S. tara last occurs between the Morgan Springs
coal bed and the H o o p e r coal bed above. Since only one sample of the
H o o p e r coal bed was studied, the beginning of the range of E. globiformis
would be more reliable than the absence of S. tara in a single sample. An
alternate interpretation is to rely on the range of S. rara and place the
Westphalian A-B boundary between the Morgan Springs and H o o p e r coal
beds or in the lower part of the Breathitt Formation. Gillespie and Pfeffer-
korn (1979) placed the Westphalian A-B boundary in the upper part of
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214

the New River Formation in the proposed Pennsylvanian stratotype in

West Virginia.

Palynological correlations. The histograms (Fig. 2) representing the relative

abundance of spores from coals in the Illinois Basin Coal Field were derived
from analyses of more than 800 coal samples; those in Tennessee are from
only one sample per coal. The tentative correlation of the coals is based on
a comparison of stratigraphic ranges of selected species (Fig. 3) as well as
on major changes in relative abundance of dominant species of Lycospora.
More coals are present in strata equivalent to the upper Westphalian C of
northeastern Tennessee than in the Illinois Basin Coal Field. The two strati-
graphic columns of Fig. 2 correspond to their respective spore profiles.
Direct correlation of one specific coal to another is n o t intended.
Some taxa appeared slightly earlier in the Illinois Basin Coal Field than
in Tennessee in the lower part of the Middle Pennsylvanian (Fig. 3) while
the reverse is observed in the middle part of the Middle Pennsylvanian.
A number of taxa that were useful in correlating Pennsylvanian strata in
the Illinois Basin Coal Field with Upper Carboniferous strata in Europe
were not observed in the Tennessee coals; for example, the beginning of
the range and base of the epibole of one of these taxa, Torispora, are in-
dicators of the Westphalian C in Europe. Only the two upper coals on
Cross Mountain contain Radiizonates difformis, thus, its stratigraphic
range in northeastern Tennesee is not known.
The similarities of quantitative changes of the three major species of
Lycospora in the two regions (Fig. 2) and the range data provide the most
acceptable general correlation. If the changing species dominance did not
take place simultaneously, it was nearly synchronous. Coals with spore
assemblages dominated by Lycospora micropapillata are known in the part
of the Illinois Basin Coal Field that extends into Kentucky b u t are not
present in Illinois. The western Kentucky coals with abundant L. micro-
papillata were sampled from several diamond-drilled cores in Union and
Crittenden Counties and, in turn, correlated eastward (96 km) to L. micro-
papillata-dominated coals in Butler County, Kentucky {unpubl. data).

Vegetational patterns. The combination of ranges and abundance of spore

taxa indicate evolutionary diversity and paleoecological trends in the coal-
swamp vegetation. The correlation between the Illinois Basin Coal Field
and northeastern Tennessee permits the first detailed quantitative com-
parisons of the vegetational patterns of the two regions. The miospore
floras in the coals of both regions were generally dominated by Lycospora.
Principal patterns of quantitative changes in the three Lycospora species
are similar in the two regions (Fig. 2). The main stratigraphic trends are
a progressive decrease in abundance of L. pellucida, a dominant miospore
of the Lower Pennsylvanian, and a concurrent increase in abundance of
L. granulata, a dominant miospore of the upper Middle Pennsylvanian.
 215

Both species of Lycospora are attributed to Lepidophloios (from Lepi-

dostrobus oldhamius kinds Qf cones); Lepidophloios harcourtii and L.
hallii were the respective parent trees. The shift from one species to another
morphologically involves a change from a thin barked trunk to a thick
barked one and from a flanged microspore to one with little flange develop-
ment. The inferred changes of Lepidophloios populations may be evo-
lutionary ones rather than a replacement of one species by another. After
the major increase in abundance of Lycospora granulata in the middle
Morrowan, there was an interval of abundant Lycospora micropapillata.
At present the plant sources of L. micropapillata include two different
lycopods, Paralycopodites brevifolius and Lepidophloios haUii. Lycospora
micropapiUata is the characteristic microspore of Paralycopodites. The
interval of abundance of such microspores coincides with the dominance
of Paralycopodites in the coal-ball peats. Some L. micropapillata spores
have origins in the determinate cone tips of Lepidophloios where smaller
abortive microspores are found with different ornamentation (Felix, 1954).
These differ from the typical L. granulata spores of the cone. A number
of important taxa gradually increased in abundance during the relative
shift in dominance among the three species of Lycospora, but the major
change came during and after the L. micropapillata expansion. This included
the appearance and gradual expansion of marattiaceous tree ferns and
different rates of increase in the herbaceous lycopods, calamites and cordaites
between the two regions.
The subtle changes of the early Middle Pennsylvanian coal-swamp vege-
tation suggest synchronous but more gradual and fluctuating change than
the Middle--Late Pennsylvanian transition. Regional differences in spore
abundances and beginning ranges are evident. The introductions and evo-
lution of plants during the early Middle Pennsylvanian established the
m a i n floristic components of subsequent coal swamps and represent a
significant time of plant evolution within the coal swamps. The vegetational
patterns do not reflect a prevalent kind of climatic regime for the whole
interval.

Middle--Upper Pennsylvanian boundary

The Middle--Upper Pennsylvanian boundary was defined by the U.S.
Geological Survey (Bradley, 1956) as being correlative with the Allegheny--
Conemaugh boundary in the Appalachians and the Desmoinesian--Missourian
boundary in the Midcontinent. These boundaries in the Appalachian and
Midcontinent regions have been equated by some authors such as McKee
(1975), Arkle et al. (1979), and Rice et al. (1979a). In some cases the
boundary in the Appalachians is expressed as the Breathitt--Conemaugh
(Rice et al., 1979a,b; Cobb et al., 1981) or the Lee and Breathitt--Conemaugh
(McDowell et al., 1981). Numerous treatments indicate that the Allegheny--
Conemaugh boundary is lower than the Desmoinesian--Missourian boundary
(Wanless, 1939; Moore et al., 1944; Kosanke et al., 1960; Shaver et al.,
1970; Rice et al., 1979b).
216

In the stratotype study area (Arndt, 1979) the Middle--Upper Pennsyl-

vanian boundary is relocated higher between the Charleston Sandstone (with
the "Mahoning" Sandstone as the uppermost unit) and the Conemaugh
Formation. However, the "Mahoning" Sandstone is shown as being a little
younger than the Mahoning Formation of the type Conemaugh. The intent
seems to be to raise the Middle--Upper Pennsylvanian boundary for the
Appalachians and this would make it closer to that of the Desmoinesian--
Missourian.
The major changes in the coal-swamp vegetation t o o k place simultane-
ously, or nearly so, in the Midcontinent and Appalachian regions. Such
patterns of change noted in the coal-spore floras form the basis for com-
paring boundaries of the Middle--Upper Pennsylvanian in the two regions.
Palynological investigations (Schemel, 1957) of the Upper Freeport coal
bed (top of the Allegheny Formation) indicated that Lycospora was the
dominant spore and that Lycospora was reduced to 2% or less in the Brush
Creek coal bed near the base of the Conemaugh Formation. Thymospora
pseudothiessenii became rare in the Brush Creek Coal and disappeared at
this time of Lycospora diminution in the Western Interior Coal Region
and Illinois Basin Coal Field. The abundance of Endosporites and Florinites
increased markedly in the Brush Creek coal bed and in coals in the Missourian
Series in the Illinois Basin Coal Field.
The Mahoning coal bed is between the Upper Freeport and Brush Creek
coal beds and in the Conemaugh Formation. Samples of the Mahoning
coal bed from Ohio and Pennsylvania were examined for spore content
and, as in the Upper Freeport coal, it is dominated by Lycospora, and
Thymospora pseudothiessenii is an important constituent. Thus, the Ma-
honing coal bed contains a typical upper Desmoinesian spore assemblage,
and the Allegheny--Conemaugh boundary is slightly older than the Des-
moinesian--Missourian boundary.

Western Europe

Cantabrian of Spain
The most widely used classification of Carboniferous rocks in western
Europe came o u t of the 1935 International Congress of Stratigraphy and
Geology; subsequent emendations were made by the Subcommission on
Carboniferous Stratigraphy. At the Seventh Carboniferous Congress in
Krefeld (1971) the Cantabrian Stage was included as a stage between the
Westphalian and Stephanian Series. This addition was based on Wagner's
{1966) description of a flora containing Westphalian D as well as Stephanian
elements in northern Spain. A Cantabrian Stage has not been d o c u m e n t e d
in North America.

Miospore zonation
Major palynological correlations among Carboniferous coal basins in
western Europe have been made (Liabeuf et al., 1967; Butterworth, 1969;
 217

Coquel et al., 1970, 1976; Loboziak, 1971, 1972, 1974; Van Wije and
Bless, 1974; Coquel, 1976; Loboziak et al., 1976; Butterworth and Smith,
1976; Bless et al., 1977). On the basis of these studies, a miospore assem-
blage zonal scheme for western Europe has been proposed (Clayton et al.,
1977).

U.S.S.R.

Bashkirian--Moscovian boundary of Donets Basin

Spore zonal correlations have been made between western Europe and the
Donets Basin (Owens et al., 1978), but no section showing the Middle
Carboniferous stratigraphic nomenclature of the Donets Basin was included.
Wagner and Higgins (1979} pointed out the difficulty in correlating the
position of the Bashkirian--Moscovian boundary in the Donets Basin with
the European stages; they placed the boundary, based largely on plant
fossils, goniatites, and microfaunas, in the middle Westphalian A. The
Bashkirian--Moscovian boundary coincides with the Westphalian B--C
boundary, according to Soviet stratigraphers working in the Donets Basin
(Wagner et al., 1979} and by Bouroz {in Bouroz et al., 1978). Phillips (1981}
took an intermediate position placing the boundary in the upper West-
phalian B because Lyginopteris is absent from coal balls from at least as low
as coal ks in the C:s suite. Lyginopteris is present in coal-ball peats as high
as the Katharina Seam {Westphalian A--B boundary); it extends into the
lower Westphalian B (Patteisky, 1957) in compression floras of western
Europe and to the Lower--Middle Pennsylvanian boundary in the Appa-
lachians (Gillespie and Pfefferkorn, 1979). A comparison of miospore
ranges in the Donets Basin (Inosova et al., 1975; Teteryuk, 1976), in western
Europe {Clayton et al., 1977) and the Illinois Basin Coal Field (Peppers,
1979b) indicates that the Bashkirian-Moscovian boundary is probably
just above the middle of the Westphalian B.

North American--Western Europe correlation

Compression floras
Among the early correlations between North America and Europe based
on plant megafossils were those of Jongmans and Gothan (1934), Bertrand
(1935) and Bode (1958). The last two authors did not recognize the pres-
ence of Stephanian rocks in the Illinois Basin Coal Field. A more recent
correlation based on plant compressions is that from the proposed Penn-
sylvanian stratotype study in the Appalachians by Gillespie and Pfefferkorn
{1979). The Westphalian A--B boundary is recognized as being just below
the New River--Kanawha boundary (Lower--Middle Pennsylvanian bounda-
ry) and a Westphalian C--D boundary was indicated just below the Kanawha
--Charleston Sandstone boundary. The Westphalian--Stephanian boundary
was placed in the lower half of the Conemaugh.
218

Spore floras
Generalized palynological correlations between western Europe and
North America have been made (Potoni~ and Kremp, 1954, 1956; Hacque-
bard et al., 1960; Bharadwaj, 1960; Butterworth, 1964, 1969; Helby, 1966;
Alpern and Liabeuf, 1969; Butterworth and Smith, 1976), primarily on the
bases of ranges of spore genera. Recently the Pennsylvanian System in the
Illinois Basin Coal Field was divided into miospore assemblage zones using
biostratigraphic ranges and abundance of certain taxa, and these zones
were used in making correlations with the Upper Carboniferous of western
Europe (Peppers, 1979b). Major changes from this study include shortening
the Westphalian B and C in Illinois and lowering the Westphalian C--D
boundary from near the top to near the base of the Spoon Formation.

PALEOFLORISTIC PROVINCES OF THE LATE PALEOZOIC AND CLIMATIC CON-

TROL

The first world-wide differentiation of floral provinces occun'ed during

the Carboniferous and Permian (Fig. 4) (Chaloner and Lacey, 1973; Chaloner
and Meyen, 1973) as paleocontinents progressively approached the pole
to pole Pangean configuration that was fully established in the Triassic.

UPPER
CARBONIFEROUS PERMIAN

cOROAITES

I
,,, EURAMERICAN
t-I ITREE FERNS
~¢ LYCOPODStPTERIDOSPERMS

GLOSSOPTERIS

GONDWANAN
Fig. 4. Diagrammatic representation of the differentiation of major floristic provinces
in the late Paleozoic, with the important kinds of coal-swamp trees indicated in each
province (modified from Chaloner and Lacey, 1973, and Chaloner and Meyen, 1973).
 219

The comparative vegetational patterns of the dominant kinds of forests

in the Gondwanan and Angaran provinces furnish part of the evidence
that the Euramerican coal belt was paleolatitudinally, or at least meteoro-
logically, tropical.
The dominance of gymnosperms (woody seed plants) progressively
developed in all three floristic provinces during the late Paleozoic. Domi-
nance of Euramerican wetlands by arborescent lower vascular plants during
the Pennsylvanian was a unique tropical p h e n o m e n o n that included both
the zenith and demise of aquatically adapted, heterosporous lycopod trees.
Lycopod forests characteristized most coal swamps and many other wet-
lands during the Early and Middle Pennsylvanian except in physiologically
drier swamps that became the domain of gymnosperms and ferns. The most
drastic changes were the extinctions of the dominant lycopod genera and
further expansion of the tree ferns and seed ferns. The principal cause for
this change is considered to be a markedly drier climatic regime that de-
veloped across the tropical belt as paleocontinents collided and orographic
barriers developed east of each major coal region except for Spain. Paleo-
geographic mapping of the land masses during the Carboniferous and Permian
indicates that the paleolatitudinal disposition of Laurussia (North America,
Greenland and Europe) did not change significantly during the Pennsyl-
vanian (Habicht, 1979; Ziegler et al., 1981).
The onset of Euramerican coal-swamp expansion apparently coincided
with glaciation in the Gondwanan province (Mississippian--Pennsylvanian
transition) (Bouroz et al., 1978), development of the Glossopteris flora
began during the Late Pennsylvanian. During the Early Pennsylvanian there
was a significant loss of tree forms of lower vascular plants (especially
lycopods) and a concurrent expansion of pteridosperms and then cordaitean
gymnosperms in the Angaran province. This change is indicative of rapid
cooling accompanying continental drift into higher latitudes (Meyen, 1977).
While approximate paleolatitudes of the three floristic provinces have
been indicated by geomagnetic data (Smith et al., 1981) and the paleo-
climatic zones can be generally corroborated with "climate sensitive" sedi-
ments (Frakes, 1979; Habicht, 1979; Zharkov, 1981; Ziegler et al., 1981),
the kinds of forests of each of the floristic provinces provide overwhelming
documentation of the bracketing of a tropical Euramerican coal belt by the
temperate Angaran and Gondwanan provinces. Woody trees of the temperate
zones were gymnosperms that uniquely combined reproductive and vege-
tative dormancy which is indicated in their woods by seasonal growth rings
(Kr~usel, 1964; Lepekhina, 1972; Meyen, 1977; Gould and Delevoryas,
1977). These trees probably were deciduous.
In the tropical zone, trees were woody, semi-woody (lycopods) and
herbaceous (tree ferns) and diversely representative of all the major kinds
of vascular plants. The lower vascular plants that were semi-woody to
woody were primarily heterosporous and adapted for reproduction in moist
to semi-aquatic habitats (Phillips, 1979); with neither seasonal dormancy
220

of vegetative growth nor of reproduction they were restricted to a warm

moist climate and consequently could not and did not exhibit seasonal
growth rings. The absence of growth rings in the gymnosperms of the tropi-
cal lowlands and uplands indicates that even the wet-dry seasonality ex-
pected in some tropical areas was apparently not severe enough to evoke
such a pattern of growth (Chaloner and Creber, 1973); however, one can
hardly regard the upland vegetation as well explored and documented
in this regard. Thus, the comparison of the dominant tree forms in coal
swamps of the Euramerican coal belt with those of the Permian coal swamps
of the Angaran and Gondwanan provinces is a contrast between diverse,
evergreen, tropical, lower vascular plants that were not major wood pro-
ducers and the deciduous, temperate, woody gymnosperms - - A n g a r a n
cordaites and Gondwanan glossopterids.
The paleoclimatic interpretations derived from a more detailed analysis
of the Euramerican coal-swamp vegetation are based on the premise that the
coal belt was tropical throughout Pennsylvanian time. Consequently, the
principal climatic variable and control on the tropical coal-swamp vegetation
was the freshwater regime (net rainfall, r u n o f f and evapotranspiration).
We do not subscribe to the idea that most or even many of the coal-swamp
environments of economically important coal seams were brackish or man-
grove-like, despite their frequent burial by marine sediment. Some coal
swamps were mangrove-like, but the balance between slight to moderate
brackishness and freshwater flushing is still relevant to climatic control
as detected by vegetational composition. The potential effects on climate
from changes in sea level resulting in periodic marine invasion or extensive
withdrawal are also improtant.
Emphasis on the freshwater control of coal-swamp vegetation and peat
accumulation does not resolve the many differences among basinal charac-
teristics such as subsidence rates, proximity to sediment sources and seas
or topography. Stratigraphic analyses of swamp vegetation must be under-
taken on a region by region basis in order to assess the impact of regional
differences that also involve gradients of wetness.

TROPICAL EURAMERICAN FORESTS

Forest ecosystems of the Euramerican coal belt were of more diverse

tree composition than those of temperate regions. Dominance patterns
along the wet to dry gradients provide perspective for interpreting vege-
tational patterns of the coal swamps and the underlying paleoc!imatic con-
rols. The generalized environments are divided along relative moisture
gradients (wetland, lowland and upland) with coal swamps as a particular
kind of wetland. Lycopods, tree ferns and calamites (lower vascular plants)
dominated most of the wetland forests; pteridosperms, cordaites and coni-
fers (gymnosperms) formed the principal forests of the lowlands and uplands
(Fig. 5).
 221

GYMNOSPERMS LOWER VASCULAR PLANTS

........~.~ ..........................................................................:>..... ~::::..:::. ~

<:i:>~ ....... I~ :i!!i!i!;i!S :~i!i!i:!.!i:!¢!:!.!-!:i:!!:i!:i:i~:'.:s:!
S v:.:::.:: .........:i:
~ :'~!1 ............ :~. : ........................:~:~:~:::~ ~.::". ......... ...::.:: . ~

iiiiiiiiiiiiiiiiiiii 00 Iiiii:7, Ei ;::ic::i'::~c/~//

~,.>
!!!!!::::.... ,..:: ~::..~ ,.. ~ :.:: ~ -.... Z
............~ . . : . . ~ ;.,- . . . . . .. . . . . . . . . . . . . .
;::'O ':::!::"~2' R S :.:D.-: ....~>. . . : ...... ...,. Z
C,.; .+:::: (~. -- ~ --:--~ "

iiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiii ii:i
UPLANDS LOWLANDS |OTHER C O A LSWAMPS
~¥ETLANDS

Fig. 5. Diagrammatic representation of the distribution of dominant kinds of vascular

plants in tropical Euramerican habitat-environments.

Forests in these varied e n v i r o n m e n t s e x h i b i t e d d i f f e r e n t a d a p t a t i o n s

and responses t o t h e same p a l e o c l i m a t i c changes. Speciation evidently
o c c u r r e d m o r e f r e q u e n t l y in d r y l a n d s t h a n in wetlands and resulted in the
periodic or r e p e a t e d i n t r o d u c t i o n o f tree species into wetlands. Conifers
a p p a r e n t l y evolved in the uplands during the Pennsylvanian, b u t the o t h e r
five m a j o r kinds o f trees had already evolved b y Pennsylvanian time and
o c c u r r e d in b o t h lowlands and wetlands, including coal swamps. In t u r n ,
all t h e i m p o r t a n t tree genera of Pennsylvanian coal swamps were p r e s e n t
in t h e earliest Westphalian coal swamps k n o w n f r o m p e a t deposits a l t h o u g h
some genera were relatively rare.

Stratigraphic ranges of major coal-swamp genera

T h e principal tree genera of E u r a m e r i c a n coal swamps were always rela-

tively small in n u m b e r {Fig. 6). Polysporia is the o n l y h e r b a c e o u s p l a n t
s h o w n in Fig. 6. E x c e p t for t h e l y c o p o d s o n l y o n e or t w o tree genera f r o m
each o f the m a j o r p l a n t groups c o n t r i b u t e d the bulk o f t h e p e a t and spores
o f t h o s e groups. All the genera had long stratigraphic ranges, as did m a n y
o f their species. Lepidodendron, Paralycopodites and Lepidophloios ac-
c o u n t e d f o r m o s t o f the biomass o f coal-ball peats and spore floras o f the
Early and Middle Pennsylvanian age and b e c a m e e x t i n c t at the Middle--
L a t e P e n n s y l v a n i a n transition.
222

SYSTEM PENNSYLVANIAN
u.s.A. LOWERI M I D D L E UPPER
MIDCONTINENT MORROWAN A~OKA~ DESMOINESIAN NISSOLIRIAN~IRGILIAN
w E STP H A L I A N STE P H A N I A N
WESTERN EUROPE
AI a I cI D
~_E , , = ' / o o , o H L O , O S . . . . . . . . . . . . . . .
L EPmOCA.POA, MA J OR

LEPIDODENDRON LYCOPOD
,~CHLA~'YOOCARPO~V o~ . . . . -

PARALYCOPODIrES . . . . . . . . . . . . . . . . EXTINCTIONS
LEPIDOSTROBUS
S/G/LLARIA
MAZOCARPON

POLYSPORM

PS,4RONIUS

CORDA I TES ::::::::::::::::::::::::::::::::::::::::::::5::.9.:::.::::.:A::.:::.:!:':::.-.::::..::.:::::::::1

CALAM/TES

MEDULLOSA ~ / / / / / / / / / / / / / / / / / / / / ~

L YGINOP TERIS

I MAJOR INTRODUCTIONS AND EXPANSIONS

Fig. 6. Stratigraphic ranges of major genera of trees in Euramerican coal swamps, based
on both coal-ball peats and coal-spore floras.

Tree ferns of the genera, Megaphyton or Caulopteris, in the compression

record, and Psaronius, when anatomically preserved, were rare in Lower
Pennsylvanian strata. While tree ferns span the entire Pennsylvanian, the
evolutionary development of these herbaceous trees occurred largely out-
side coal swamps and they progressively expanded in lowlands or wetland
habitats prior to becoming abundant in coal swamps.
The generic name, Cordaites, actually includes two kinds of cordaitean
assemblages. The stem genera are known as Mesoxylon and Pennsylvanioxylon
and they respectively produced Mitrospermum and Cardiocarpus ovules.
Mesoxylon occurs earlier in the coal swamps than Pennsylvanioxylon and
both occur in some Middle and Upper Pennsylvanian coal-ball peats. Cor-
daites were the only gymnosperms that were d o m i n a n t in any coal swamps
of the Pennsylvanian and, as discussed later, they indicate physiologically
drier conditions resulting from several different causes.
Calamites includes three stem genera with Arthropitys the only moder-
ately abundant tree form in coal swamps. Calamites, the only trees that
exhibited vegetative propagation, were much more abundant in clastic-
rich wetlands than in coal swamps.
 223

MeduUosa was the largest pteridosperm tree in the coal swamps; Lyginop-
teris was a robust liana or small tree. Both genera were well represented in
Pennsylvanian floras from wetlands to lowlands. Lyginopteris was c o m m o n
in coal swamps during the Early Pennsylvanian prior to extinction. Medullosa
was relatively rare in Early Pennsylvanian coal swamps.

Generalized habitats

Wetlands. Distinctions need to be made between coal-swamp environments

with their in-situ plant assemblages and low mineral content, and those
represented by transported assemblages in the clastic deposits between
coal seams. The assumption that compression floras in coal measures are
indicative of the coal-swamp floras is a fallacy propagated by the general
abundance of lower vascular plants in wetlands and the widespread occur-
rences of all five major kinds of vascular plants from coal swamps to low-
lands. As the actual in situ coal-swamp forest assemblages of the Penn-
sylvanian become better known, it becomes evident how different the
vegetation was from the array of floras deposited between coal seams. The
coal-swamp communities were far less diverse and there are probably more
taxonomic differences between the specific kinds o f abundant trees found
inside and outside of the coal-swamp environments than there are simi-
larities; some cosmopolitan species are well-known exceptions. The trees
were usually smaller than the record-size specimens found in clastic deposits.
These differences are consistent with the low nutrient, acidic, and oxygen
deficient semi-aquatic environments of growth and peat accumulation
of coal swamps.

Coal swamps. L y c o p o d s dominated the Early and Middle Pennsylvanian

coal swamps, and cordaites were abundant only during the early Middle
Pennsylvanian; cordaites also grew in the uplands. Tree ferns dominated
most of the Late Pennsylvanian coal swamps and were abundant in the low-
lands at the same time.

Other wetlands. Clastic-rich swamps and other freshwater wetlands were

also dominated by lycopods during the Early and Middle Pennsylvanian.
Calamites (sphenopsids) were much more abundant in other wetlands
than in coal swamps b u t were sometimes dominant. The diversity of lyco-
pods was greater in clastic deposits than in coal swamps. During the Late
Pennsylvanian, tree ferns, pteridosperms, calamites and some surviving
lycopods characterized the wetlands in such a way as to diminish distinctions
between lowland floras on the one hand and coal-swamp floras on the other.

Lowlands
Pteridosperrns formed the principal forests of the Pennsylvanian lowlands;
tree ferns became subdominant by the Late Pennsylvanian. All the major
224

kinds of plants were represented in lowland floras which resulted in the most
diverse assemblages known for the Pennsylvanian.

Uplands
The drier habitats and those more remote from environments of deposi-
tion have yielded the fewest floras; consequently, the vegetational patterns
and actual areal extent of upland forests are more conjecture than fact.
It is generally assumed that cordaites were more abundant in the uplands
than elsewhere and that conifers evolved in such remote areas. Cordaites
and Megalopteris pteridosperms occur in deposits of Early and early Middle
Pennsylvanian age that are widely scattered among major coal regions of
the United States. Conifers were rare on the whole. They first occurred
during the early Middle Pennsylvanian (Westphalian B) of Europe (Scott,
1974) but became more c o m m o n during the Late Pennsylvanian (Stephanian)
of both the United States and Europe (Florin, 1945).

Floristic migration

Consistent vegetational patterns that may be interpreted as migration

of florisfic elements from uplands to lowlands and wetlands occurred during
the two major times of change to drier climate in the Pennsylvanian coal
belt.

Early--Middle Pennsy lvanian

The environmentally significant cordaites-Megalopteris flora probably
migrated from upland to lowland habitats with well-drained soils, hence the
widespread occurrences of floras regarded as upland in character. Megalop-
teris floras were most abundant where Pennsylvanian strata occur above
pre-Pennsylvanian karst surfaces in which calcareous or terra rossa-type
softs had formed. This unusual distribution feature occurs in all the localities
west of the Appalachians (Cross, 1977). It has been suggested that the
widespread occurrence of Megalopteris floras in the Midcontinent and
western part of the Appalachian Plateau would be consistent with the near
synchrony of the initiation of Pennsylvanian sedimentation in these regions
(Cross, 1977). The known stratigraphic range of the Megalopteris floras
has been expanded downward to the Early Pennsylvanian (New River or
Westphalian A time) by Cross {1977) and by Leafy (1981). The most wide-
spread Megalopteris zone, Zone 7 of Read and Mamay (1964), is in the
lower Middle Pennsylvanian {lower Kanawha or lower Westphalian B).
In the coal swamps the major changes in composition began near the
Westphalian A--B boundary with fluctuations in the kinds of dominant
lycopods and expansion of cordaites. There were subsequently a diminution
in lycopods and progressive introductions and expansions of seed plants,
tree ferns and herbs from lowlands or other wetlands. The expansion of
cordaites in coal swamps concurrently with their migration into lowland
 225

areas is considered particularly important. From a floristic point of view,

the diversity of coal swamps was increased both by introductions and by
evolution within the swamps. The ~kinds of trees and herbs that appeared
and/or expanded during the early Middle Pennsylvanian became principal
survivors of the more severe change at the Middle--Late Pennsylvanian
transition. Lyginopteris, a cosmopolitan genus of coal swamps and other
wetland and lowland habitats, became extinct.

Middle--Late Pennsylvanian
During the early Late Pennsylvanian, upland floras again migrated into
lowland areas of deposition, especially in the drier regions. Conifers, along
with cordaites and some Callipteris sensu lato, were locally abundant and
widespread in such environments for the first time (Elias, 1936; Moore et
al., 1936; Cridland and Morris, 1963; Cridland et al., 1963). Florin (1945)
has summarized the occurrences of conifers in the Euramerican province.
The principal, dominant, l y c o p o d tree genera, Lepidodendron, Lepi-
dophloios and Paralycopodites, and their closely related forms in the other
wetlands, became extinct at the Middle--Late Pennsylvanian transition.
Psaronius tree ferns and Medullosa seed ferns became the dominant and
subdominant trees in the Late Pennsylvanian swamp forests; concurrently
they had the reverse order of importance in the lowlands. Cordaites were
also abundant in the lowlands.

CHANGING PATTERNS OF COAL-SWAMP VEGETATION

The stratigraphic patterns of coal-swamp vegetation are presented in

two ways: (1) rise-decline patterns (in outline form) based on spore flora-
peat comparison, and (2) a generalized pattern that will be refined.

R ise-decline patterns

The rise-decline of plant groups outlined in Fig. 7 compares spore-flora

distribution data from the Illinois Basin Coal Field with an appropriate
stratigraphic source of coal-ball peats from anywhere in Euramerica. This
allows us to outline intervals in which the same kinds of coal-swamp trees
were synchronously most abundant in the Illinois Basin Coal Field spore
floras and much of Euramerica. At the onset of the Westphalian A coal
swamps, spore floras and the coal-ball peats from the Union Seam of Lanca-
shire demonstrate that lycopods were at their maximum dominance. No
coal-ball deposits are known a~ the Middle--Late Pennsylvanian boundary
when the abrupt demise of the dominant lycopods occurred. Coal-ball
peats indicate strong dominance of lycopods below the b o u n d a r y and
strong dominance of tree ferns above the boundary; spore floras provide
the detailed changes.
226

PENNSYLVANIAN UPPER
CARBONIFEROUS
M IOSPORES (COAL) MI DCONTINENT WESTERN EUROPE PEAT (COAL BALLS)
U.S.A.

VIRGIL IAN

STEPHANIAN

MISSOURIAN
~TREE F E R N S ~ 8 0 %
MAXIMUM

DECLINE
~::'~.!':.: LYCOPODS":,?~::] 82%

DECLINE
~'/.~ CORDA ITE S///~ WESTPHALIAN D ,~/.-~.COROAtTES7/'/~ I%
DESMOINESIAN

MAXIMUM 6%
,~//~. CORDA ITE S///~ 7 3%
ATOKAN WESTPHALIAN C

EXPANSION
~ T R E E FERNS~
WESTPHALIAN B

EXPANSION MORROWAN
~///zCORDAITES//v~ ~/_//~CORDAITEST//~ I0%

WESTPHALIAN A
MAXIMUM
[,:,:~:~,LYCOPODS~!~ ~?:':~' LY C0 PODS~ 96% i":

Fig. 7. Diagrammatic comparison of the stratigraphic rise-decline patterns of lycopods,

cordaites and tree ferns in Euramerican coal swamps, based on miospores from the coals
of the Illinois Basin Coal Field and peat data from pertinent Euramerican coal-ball
occurrences.

Psaronius tree ferns began contributing abundant spores to coal deposits

during the early Middle Pennsylvanian, but there was apparently a time lag
between their proximity to coal swamps and their actual abundance in
coal-ball peats. All the Missourian coal-ball peats were greatly dominated
by tree ferns, but some small, scattered, deposits of coal balls in the Virgilian,
outside the Illinois Basin Coal Field, indicate that pteridosperms, calamites
or cordaites (France) may also have been very abundant, if not dominant,
in some places. On the whole, tree ferns were dominant during the Late
Pennsylvanian.
The cordaitean interval was produced by the combination of t w o different
kinds of environmental phenomena. The expansion of cordaites took place
near the Lower--Middle Pennsylvanian boundary as a Euramerican-wide
pattern. The coal-ball source nearest this change is the Katharina seam of
 227

the Ruhr in which cordaites were subdominant trees. The termination of

this interval overlapped the onset of major marine transgressions across
the Western Interior Coal Region into the Illinois Basin Coal Field in the
early Desmoinesian (late Westphalian C). The maximum of cordaitean de-
velopment in the Westphalian C and early D, as shown by spore floras and
coal-ball peats, reflects only the pattern seen in the Midcontinent area.
Cordaites dominated some of the Iowa and Kansas coal swamps during the
early Desmoinesian and were subdominant in the Illinois Basin Coal Field
at the same time. On the basis of spore and Peat data, cordaites declined
in abundance simultaneously in the Western and Eastern Coal Regions of
the Interibr Province. Cordaites continued to occur (about 1% biomass)
in all subsequent coal swamps known from coal-ball peats.

Generalized patterns

The basic pattern of coal-swamp vegetation is summarized in Fig. 8.

Lycopods dominated the coal swamps during the Early and Middle Penn-
sylvanian except for the brief regional dominance of cordaites in the Kansas-
Iowa area. There were successive changes in the dominance of lycopod
genera during the lycopod interval. Lepidodendron dominated the West-
phalian A coal-swamps, Lepidophloios was moderately abundant, and
Sigillaria was not uncommon. At the onset of vegetational change during the
Early--Middle Pennsylvanian (approximately Westphalian A--B boundary)
two kinds of patterns appeared in the swamps across the early Middle
Pennsylvanian (Westphalian B and C). On the basis of coal-ball data coal
swamps were dominated either by Paralycopodites or by Lepidophloios,
and Mesoxylon (Cordaites) was usually subdominant. Concurrently there
was an increase in diversity represented, in part, by the expansion of MeduUosa
seed ferns, then by Psaronius tree ferns and by the appearance of the herb,
Polysporia. The onset of this vegetational change is not as abrupt as the
second one at the Middle--Late Pennsylvanian transition but there was a
subsequent progressive diminution in the abundance of lycopods commen-
surate with the expansion of other groups.
Beginning with the middle Middle Pennsylvanian (late Westphalian C)
Lepidodendron and then Lepidophloios with Lepidodendron became domi-
nant in the swamps. During the early Desmoinesian these genera were
abundant along with cordaites and dominated the coal floras of the Western
Interior Coal Region by middle Desmoinesian (middle Westphalian D) time.
Concurrently with the end of the Euramerican cordaitean interval (West-
phalian B and early C) and the onset of regional marine encroachment in
the Interior Coal Province, Psaronius tree ferns and to a lesser extent Medul-
losa seed ferns expanded in abundance to subdominant levels. Subsequently,
they increased in importance toward the Late Pennsylvanian.
The extinction of the three dominant lycopod swamp:genera at the
Middle--Late Pennsylvanian transition resulted in Psaronius-Medullosa swamp
228

PENNSYLVANIAN UPPER 1
MtDCONTINENT 3ARBONIFEROUS[ Basic Pattern
U. S . A . NESTERNEUROPE
I

VIRGILIAN I
STEPHANIAN
POLYSPORIA MARSHLANDS
Mi SSOURIAN

WESTPHALIAN D
DESMOINESIAN .O.si%O i MOST
TENS,VE
E
~ 1 MIDCONTINENT CORDAITES

ATOKAN WESTPHALIAN C co.o.,T,,

WESTPHALIAN B ~-~.,,.INTRODUCTION OF TREE FERNS

MORROWAN

WESTPHALIAN A ~ ,,,,~-EXTINCTtON OF LYGINOPTER/$

9o ~ 50 ~o no o
PERCENT VOLUME OF PEAT

Fig. 8. Generalized patterns of coal-swamp vegetation in the Euramerican coal belt

showing the important kinds of plants that contributed the bulk of the peat biomass
and spore floras to the coals from the lower Westphalian A to the Upper Stephanian
(modified from Phillips, 1981). Further explanation of this figure is given in the text.

forests th at were significantly different in structure from the earlier l y c o p o d

swamp forests. L y c o p o d , cordaitean and calamitean trees had tall habits
with main trunks capable of branching; t he y bore linear leaves. T he lycopods
were supported mostly by bark with relatively little wood; the cordaites
and calamites were w o o d y but minor contributors to the peat by com-
parison. The Psaronius-Medullosa trees were smaller in stature with a co-
lumnar (unbranched) t r unk and large fronds.
The frond-bearing smaller trees were ideal lower canopy trees and, when
dense, could p r o b a bl y form an almost closed canopy at a height of 5 to 10
meters. The tree ferns were largely s u p p o r t e d by r o o t mantles and scleren-
c h y m a (no wo o d or bark), and the seed ferns were supported as much by
their sclerenchyma as by their structurally weak vascular tissue. Survivors
f r o m the Middle--Late Pennsylvanian transition included Sigillaria (lyco-
pods) and calamitean trees.
Intercalated within the coal sequences of the Upper Pennsylvanian are
numerous, very thin and usually b o n y coals that do n o t represent fern
forests. On the basis of spore floras, these deposits were f o r m e d by ephemeral
 229

marshlands, some of which were dominated by Polysporia. None of the coal-

ball deposits from minable coals in the Upper Pennsylvanian have yielded
peats indicative of a predominantly marshland character.
The basic pattern depicted (Fig. 8) is consistent with the collective data
from other regions, and strongly supports the idea of synchronous, similar
kinds of changes across the Euramerican coal belt. Having encountered
one regional divergence from the basic pattern (dominant and subdominant
cordaites in the Interior Coal Province), we are seeking other differences
that may occur regionally. The stratigraphic occurrence of this regional
pattern and implied differences in moisture availability between the Appa-
lachians and the Illinois Basin Coal Field suggest that other vegetational
differences are likely, particularly in the kind of dominant lycopod of the
late Middle Pennsylvanian in the Appalachians. The inference from coal
resources is that the Appalachians were not as wet as the Midcontinent
during most of the late Middle Pennsylvanian. Because of insufficient data,
the question cannot be resolved. At present the stratigraphic patterns ex-
amined from peat studies and palynology are from the lower Westphalian A
up to the lower Virgilian. The limits are due to the absence of earliest and
latest Pennsylvanian strata in the Illinois Basin Coal Field, and the fact
that quantitative analyses of the youngest coal-ball deposits are from the
Illinois Basin Coal Field.

PALEOCLIMATIC INTERPRETATIONS

The paleoclimatic interpretations derived from the stratigraphic patterns

of Pennsylvanian coal-swamp floras represent an approximation of relative
wetness. Revisions and refinements will no d o u b t be necessary, and more
detailed data on intervals and regions are still needed to establish these
patterns across the Euramerican belt and to determine regional divergences.
Nevertheless, it is useful at this stage to summarize the general wetness
trends in the United States and then provide examples of how these trends
have been determined from the paleobotanical and coal data.
The climatic trends of the tropical Euramerican belt are viewed as a
series of alternating wetter to drier pulses with each drier interval becoming
more severe than the previous (Fig. 9). The first dry interval was in the early
Middle Pennsylvanian, the second in the early Late Pennsylvanian, and
the onset of the Permian probably as the beginning of the third. The major
changes in the coal swamps occurred during the first and second drier
intervals; coal-swamp development essentially came to a close with the
Permian. The wetter intervals constitute the norm for the Pennsylvanian;
generally the Pennsylvanian coal belt had quite a moist climate. The term
dry or drier interval is relative and not meant to convey arid or semi-arid.
The climatic regime of the Early Pennsylvanian (specifically the Westphalian
A) is regarded as the median reference point for wetter and drier. While it
seems likely that rainfall was seasonal in the Early and early Middle Penn-
230
PE N N S Y L V A NIAN
LOWERJ MIDDLE
W E S T P H A L IA N
I UPPER
STEPHANIAN
A Icl D
F-

I..-
t--
w

co
t~

Fig. 9. Diagrammatic representation of a generalized wet-dry curve for Euramerican

coal swamps during the Pennsylvanian, beginning in the early Westphalian A equivalent.
The first drier interval occurred in the early Middle Pennsylvanian (Westphalian B and
early C) with the second drier interval in the early Late Pennsylvanian (early Stephanian).
The five proposed stratigraphic-climatic intervals are based on environmental implications
of coal-swamp studies.

sylvanian, during the remaining Pennsylvanian, seasonality can not be

demonstrated from our data. Zangerl and Richardson (1963) have suggested
seasonality in their studies of black-shale from the middle Desmoinesian
and Parrish {1982} in modeling the Westphalian climate presented a mon-
soonal circulation.
One of the means of developing a relative scale of wetness to convey
the different magnitudes of each of the five stratigraphic-climatic intervals
was by the use of the identified bituminous coal resources of the eastern half
of the United States, compiled for each major coal region by Phillips et al.
(1980). The regional data for the central Appalachians, plotted on a semi-
log scale, form the outline of the basic curves that were modified to accom-
modate the Lower Pennsylvanian resources from the southern Appalachians
and the greater resources of the Midcontinent from the Desmoinesian.
This leads to a vertical exaggeration of the wettest and driest intervals,
which is desirable for purposes of extracting relative ecological amplitudes
(Fig. 10) of the dominant lycopods on the basis of their stratigraphic abun-
dances and relative ranges on the driest to wettest axis.
The slopes of the wet-dry curve are comparatively significant (Fig. 9).
The onset of the first drier interval was moderately rapid geologically, and
the return to wetter conditions was quite gradual, continuing during much
 231

of the Middle Pennsylvanian. Consequently, the vegetational change at

the end of the first dry interval was n o t as distinct as at the beginning.
In contrast, the Middle--Late Pennsylvanian change in climate was precipi-
tous from the wettest to the driest in a short geologic time. The return of
wet conditions in Monongahela time in the Dunkard Basin also appears to
have been abrupt.
The Appalachians are generally considered the wettest region in the coal
belt of the United States, with the Illinois Basin Coal Field and the Western
Interior Coal Region progressively less wet westward. The regional differ-
ences are n o t entirely consistent through time; there were probably gradients
within the Western Interior Region that were almost as great as those be-
tween the Western Region and the Appalachians during Desmoinesian
time. The wetter intervals are actually the most subjective in magnitude,
because they are partly based on American coal deposits, and cannot be
attributed generally to Europe.
The drier intervals are more diagnostic for Euramerica as a whole because
the vegetation simultaneously changed in the same way. It is the marked
deviation from a typically wet climate that divides the Pennsylvanian into
five stratigraphic-climatic intervals. The moderate wetness during the Early
Pennsylvanian was more than adequate to maintain coal swamps; numerous
basinal characteristics determined whether coal swamps developed under
such climatic conditions. There can be biases in our perception of how
increasingly wet conditions became, on the basis of coal resources alone.
However, the relationships between identified bituminous coal resources
and the dominance patterns of each of the lycopod trees indicate that the
coal resources provide a reasonable approximation of wetness during the
Early and Middle Pennsylvanian. We do n o t have comparable botanical
indicators to check the relative wetness implied by coal resources in the
tree-fern dominated coals in the Upper Pennsylvanian. The extinction of
Lepidophloios, Lepidodendron, Paralycopodites and their relatives in other
wetlands indicates the severity of the drier kind of climate t h a t ensued at
the Middle--Late Pennsylvanian transition.

ENVIRONMENTAL INDICATORS OF COAL SWAMPS

The environmental implications of coal-ball peat deposits, extended with

coal palynology, can be only deduced. Consequently, it is necessary to
explore numerous lines of circumstantial evidence and the resulting con-
sistency or inconsistency of a given interpretation. Some of the evidence
is descriptive morphology of growth and reproduction; some is quantitative.
Biological interpretations of d o m i n a n t trees provide the keystone for estab-
lishing a line of deductions on a given parameter such as relative wetness.
Controls of the distribution of tropical semi-aquatic forest vegetation
are well suited to interpretations of freshwater trends indicative of paleo-
climate. In the Pennsylvanian coal belt the a b u n d a n t kinds of trees were
232

few in number, stratigraphically long ranging and indicative of basically

different reproductive and growth strategies. The gymnosperms were pri-
marily adapted to terrestrial and physiologically drier environments; their
abundance in coal swamps reflects stressful parameters that they tolerated
and that impeded the dominance of highly specialized semi-aquatic, hetero-
sporous lower vascular plants. Some of the lower vascular plants, particu-
larly the dominant lycopods, were primarily adapted for growth and sexual
reproduction in freshwater swamps. The changing abundance patterns
of the lycopod genera provide the most sensitive guides to the freshwater
regime of plant growth and peat accumulation so long as they dominated.
The least specialized group reproductively were the homosporous tree ferns.
With a totally herbaceous habit, a massive r o o t mantle system for support
and aeration, and diversity of species, Psaronius exhibited the broadest
ecological amplitude of any of the Pennsylvanian swamp trees. Psaronius,
like the gymnosperms, was not confined to wetlands.

Growth and reproduction

The dominance of semi-woody lycopod trees during the Early and Middle
Pennsylvanian and herbaceous tree ferns during the Late Pennsylvanian
reflects different combinations of growth and reproductive strategies that
have similar attributes in coal swamps but different survival values. Both
kinds of vegetative growth were rapid in comparison to w o o d y trees, and the
nutrient requirements were probably less limiting. The root systems were
fundamentally different in that spreading, central, stigmarian systems
with lacunose rootlets occurred in the lycopods, whereas massive buttressing,
aerenchymatous, adventitious r o o t mantles characterized Psaronius tree
ferns. The latter system allowed responses to changing freshwater regimes.
The lycopods, with a living bark and photosynthetic leaf cushions, had the
capability of withstanding physiological water stress by dropping many of
their leaves; some species also had deciduous branches. The slower growing
cordaitean trees probably also periodically shed leaves in response to water
stress and exhibited the capability for axillary branching and adventitious
bud developments on the trunks. In some of the Desmoinesian swamps
aerenchyma developed even in the basal stems of cordaites.
Lycopods, tree ferns and cordaites as dominant or subdominant plants
during the Pennsylvanian, contributed proportionally very large amounts
of root material to the peat. In contrast, the seed ferns and calamitean
trees consistently contributed disproportionally smaller amounts of root
material and larger amounts of fusain. These reflect differences in both
growth habits and in habitats; probably neither group was dominant in
coal swamps on a whole seam basis during the Pennsylvanian. Among the
lycopods, the nearest analog to them in distributional patterns (and probably
ecological amplitude) is Sigillaria.
The lycopods of coat swamps exhibited a broad spectrum of hetero-
 233

sporous reproductive morphology which has been functionally interpreted

by Phillips (1979) and by DiMichele et al. (1979). Lepidophloios and Lepi-
dodendron trees produced boat-like megasporangium-megasporophyll struc-
tures that were aquatic analogs of seeds. The inference that these trees
were the most aquatically adapted and indicative of the wetter swamp
regimes is tested with subsequent data sets. At the other extreme was Sigil-
laria, adapted in the development and dispersal of its megasporophylls and
ultimately megaspores to repeated reproduction in drier conditions. Between
the wetter and the drier adaptations were Paralycopodites (Phillips, 1979;
DiMichele, 1980) and the robust herb Polysporia (DiMichele et al., 1979),
which were morphologically less specialized in their reproduction. These
genera exhibit overlapping ecological amplitudes on a wet to dry gradient,
and the environmental sequence indicated by additional analyses is Lepi-
dophloios, Lepidodendron, Paralycopodites, Polysporia and Sigillaria (Fig.
10). The relevance of these genera to Lycospora spores of the coal floras
is critical because the three dominant kinds of trees of the Early and Middle
Pennsylvanian were Lycospora producers with some Lepidodendron species
producing Cappasporites. Polysporia produced Endosporites, and Sigillaria
dispersed Crassispora.

EXTINCTION

WETTEST DRIEST

Fig. 10. Diagrammatic representation of wet-dry relationships among lycopod genera

showing their overlapping ecological amplitudes. Extinction of Lepidophloios, Lepi-
dodendron and Paralycopodites during the Middle-Late Pennsylvanian transition indicates
reduction in the freshwater regime to at least the position of the arrow on the relative
wetness scale.

Ecological amplitudes of lycopods

In order to convey graphically the relatively wet to dry habitat relation-

ships among the lycopod genera that are used for environmental inter-
pretations, curves indicating their ecological amplitudes are shown in Fig.
234

10. These are tentative and might change some with additional analyses.
Lepidophloios and Lepidodendron show the greatest overlap on the wettest
end of the scale. Polysporia and Sigillaria largely overlap on the driest end
and Paralycopodites occupies a smaller part of the spectrum in the middle.
The overlaps reflect a degree of co-occurrence in assemblages. Other taxa
can be added to the graph as analyses continue.
During the wet Early and late Middle Pennsylvanian, the dominant plants
were Lepidodendron and Lepidophloios. During the drier early Middle
Pennsylvanian, Paralycopodites was dominant in some swamps and Poly-
sporia appeared. At the Middle--Late Pennsylvanian transition, the shift
(Fig. 10) to severely drier conditions resulted in the extinction of Lepi-
dophloios, Lepidodendron and Paralycopodites. Polysporia and Sigillaria
survived. The extinction point was determined by the environmental limits
of the plants to reproduce. Had the dominant lycopods survived the drier
climate of Conemaugh and equivalent time, they probably would have
flourished again in Monongahela time. Instead, tree ferns with less special-
ized reproduction and vegetative morphology, highly adaptive for coal
swamps, expanded and dominated these Late Pennsylvanian habitats.

Community structure and analyses

Peat profiles analyzed according to the techniques of Phillips et al. (1977)

and Phillips and DiMichele (1981) provide a means of examining successive
assemblages or communities sequentially within coal seams, comparing
their relationships to each other and to geological variables, and assessing
the hypothesized environmental conditions by means of community analysis
techniques such as polar ordination and dominance diversity curves. These
results are then compared to regional and stratigraphic changes of dominance
of the genera.
Peat profiles consisting of histograms of identifiable biomass of lycopod
shoot debris, zone by zone for the Herrin (No. 6) Coal Member of Illinois,
illustrate some of the kinds of patterns seen in Early and Middle Pennsyl-
vanian coal swamps (Fig. 11). The histograms are normalized for 100 percent
lycopod aerial biomass to emphasize lycopod tree sequences by genera.
Composition of the peat sequence, based on total biomass, is 49 percent
Lepidophloios hallii, 12.5 percent Lepidodendron, and 2.5 percent Sigillaria.
The remaining portion of the floras is made up of a b o u t an equal amount
of tree ferns and seed ferns. This and other plots by Phillips et al. (1977)
and by Phillips and DiMichele {1981) indicate repetitive dominance usually
by one species (in this case Lepidophloios hallii) and consequently its
dominance on a whole seam basis. Also, a directional biotic succession is
lacking. The relative percentage of genera in such profiles varies from site
to site in the Herrin Coal, so that Lepidodendron became almost as abundant
as Lepidophloios in places or Lepidophloios was even more strongly domi-
nant in other places. The greatest abundance of Lepidophloios in the profiles
 235

ZONES
IOO

90

80

70

6O

so

~ 40

30

20

I0

io ~o ~o To 9o Ho 13o Eso 17o ~9o 210

Lineor Dislonce From Bottom of Seom, (cm)
ooo=N,o Por tings
Cool
(~ Maximum Fusoin Zone = 27.5%

Fig. 11. Relative percent of identified lycopod stem genera in Sahara vertical section 4,
based on coal-ball peat profiles from the Herrin (No. 6) Coal Member in southern Illinois.
Lepidophloios is the dominant lycopod tree in most of the coal-ball zones and dominates
the seam vegetation on the whole. Selaginella is the only herbaceous lycopod show~
in the peat profile (modified from Phillips and DilVlichele, 1981). For further explanation
see text.

studied occurs at sites near (within 9 km) the Walshville channel, t he major
stream system through t he swamp (DiMichele and Phillips, 1980).
P r o n o u n c e d changes in the kinds and abundances of l y c o p o d - d o m i n a t e d
forests in the Herrin profiles ~sually occurred at mineral-rich or clastic
bands which indicate abiotic disruptions of freshwater regimes - drops in
water level, flood, fire, and changes in nut r i ent supply. Diminutions in the
a m o u n t s of l y c o p o d peat and changes in do m i nance occur f r o m below t o
above such mineral bands. The change of d o m i n a n c e is usually from Lepi-
dophloios to a n o t h e r l y c o p o d or f r om one Lepidodendron species to another.
Concurrently, tree ferns became more abundant; seed ferns were usually
m o s t a b u n d a n t in p r o x i m i t y to the mineral bands, immediately above and
below.
The m o s t unusual z one in the profile shown (Fig. 11) is the Sigillaria-Para-
lycopodites assemblage (actually two successive communities, with the
Paralycopodites c o m m u n i t y located below t hat of Sigillaria). This pair of
assemblages is sandwiched between t w o clastic bands located a b o u t where
236

the so-called "blue band" occurs in the Herrin. The zone contains the maxi-
mum fusain content for the profile. In an adjacent profile (Phillips and
DiMichele, 1981) at the same mine, there were three coal-ball zones between
the pair of clastic bands in which an abundance of Lepidophloios-Lepi-
dodendron is at the bottom, Paralycopodites is abundant in the middle,
and the fusain-rich Sigillaria zone is at the top. Phillips and DiMichele
(1981) interpreted this sequence as a progressively wet to drier "succession"
consistent with the reproductive biology of the plants (Phillips, 1979).
Small amounts of Polysporia also occur with Paralycopodites or Sigillaria
in the profiles. While the fusain-rich zone is consistent with inferred drier
or alternating wet-dry environment for Sigillaria, the two plant groups
that consistently contributed disproportionately more fusain in the Herrin
and other coals were the seed ferns and sphenopsids.
Polar ordination of Herrin coal-swamp assemblages by Phillips and Di-
Michele (1981) consistently shows a spread of lycopod-dominated com-
munities in which Lepidophloios represented one extreme and Sigillaria
the other; Lepidodendron species and Paralycopodites were in between the
two extremes as shown in Fig. 10. Assemblages with abundant Psaronius
tree ferns overlap most of the other c o m m u n i t y types (i.e., Psaronius ex-
hibits a broader ecological amplitude than any of the other genera).
Dominance-diversity curves (Phillips and DiMichele, 1981) for the lyco-
pod communities plotted in the polar ordination also showed Lepidophloios
and Sigillaria as extremes. Lepidophloios-dominated assemblages were plot-
ted as an almost vertical line with little area under the curve, which indicated
strong dominance, little resource sharing, and low diversity. These properties
are consistent with the interpretation that Lepidophloios forests stood in
water, and lower canopies and ground cover were absent. The plots for the
Sigillaria assemblage produced a broad curve with a large area below, indica-
tive of weak dominance, great resource sharing, and high diversity of plant
assemblages. These factors describe a highly structured forest with a lower
canopy of seed ferns and tree ferns as well as an extensive ground cover.
Lepidodendron-dominated assemblages, especially if Medullosa seed ferns
were included, usually also had a well structured or layered forest com-
munity but were rarely as diverse as the Sigillaria assemblages. The Sigillaria
c o m m u n i t y is the rarest kind in the Herrin Coal, and the Lepidophloios is
the most common.
The dominance of Lepidophloios communities in the Herrin Coal on a
whole seam basis is interpreted as indicative of the predominance of standing
water and net rising water table in the peat swamp. Drier or wet<lry fluctu-
ations (Sigillaria zone and perhaps some of the mineral-rich bands indicative
of peat degradation) were relatively infrequent. The relationship of Lepi-
dophloios, Lepidodendron, and Sigillaria to a wetter or drier climate is
supported by patterns of miospore distribution of the genera during the
trend toward a drier climate in the early Middle Pennsylvanian (Peppers,
1979a).
 237

Regional abundance of L ycospora

Maps depicting the relative abundance of major miospore taxa in the

Springfield and Herrin Coals in the Illinois Basin Coal Field are in prepara-
tion, and one of these is used to show the regional abundance of Lycospora
in the Herrin Coal (Fig. 12). The above two coals contain the largest identi-
fied bituminous coal resources in the United States. Major paleochannel
systems contemporaneous with peat deposition influenced the abundance
patterns of Lycospora. A b o u t 90 percent of the Lycospora was produced
by Lepidophloios hallii.
The Walshville channel in the Hevrin Coal had a pronounced influence
on the distribution of Lycospora. Lycospora is most abundant and accounts
for more than 85 percent of the miospore floras in some areas near the
channel except in the northeastern portion of the basin where the area of
largest abundance is displaced to the south and east. That area was the
location of a shallow lake or flood basin (Treworgy and Jacobson, 1979).
The thickest coal deposits are generally close to the Walshville channel
(Smith and Bengal, 1975a; Smith and Stall, 1975). The abundance of Lyco-
spora granulata as well as the Lepidophloios in peat profiles, previously
mentioned, indicate that Lepidophloios was the major contributor to the
deposits. The environmental inference is that the wettest part of the swamp
was along the flanks of much of the channel and the shallow lake area. A
similar pattern occurs near the Galatia channel sandstone in the Springfield
Coal in the southeastern part of the Illinois Basin Coal Field. Lycospora
is more abundant and the Springfield Coal is thicker (Wanless et al., 1970;
Smith and Bengal, 1975b) adjacent to the Galatia channel (unpublished
map).

Organ composition of peat deposits

A reasonable estimate of the shoot-root ratio of biomass of the living

lycopod trees is 70 to 80% shoot system (trunk, branches, leaves) and 20
to 30% r o o t system. Since the shallow root systems are buried in situ by
natural growth they are less subject to exposure or loss by degradation;
the aerial portions of the trees are more subject to decomposition except
when burial or submergence is fairly rapid. In general, if other factors re-
main the same, the higher the water table, the less the degradational loss
of fallen trees. In fluctuations of wet-dry seasonality the loss would be
substantial. Knowing the morphology and dominance of the swamp trees
during the Early and Middle Pennsylvanian, we can use the shoot-to-root
ratios of peat deposits as a general index to relative trends in wetness. These
trends should be consistent with environmental inferences from the domi-
nant trees.
R o o t material is proportionally more abundant in the peat than in the
original biomass of intact lycopod trees during the Early and Middle Penn-
238

5 . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . i
I
/
\ t

I
I

/
J
J

/
t

i'

ky~

~Present limit of Herr

Walshville channel
- - 6 0 - - P e r c e n t Lycospora ', / ~"o~ ~
• Sample site '~,,~':~' ~'e°~ocky
~-:'~=--~ Lo ke

Fig. 12. Relative a b u n d a n c e o f L y c o s p o r a in the Herrin ( N o . 6) Coat in the Illinois

Basin Coal Field based on analyses o f a b o u t 7 0 channel samples. For further e x p l a n a t i o n
see text.

sylvanian. Roots account for about half (or slightly more) of the lower
Westphalian A peats from western Europe. As previously mentioned, the
Early Pennsylvanian is regarded as moderately wet, certainly with a surplus
of freshwater for swamp maintenance. The relatively high amount of aerial
 239

plant degradation may have been due to wet-dry seasonality, which is

suggested by other lines of evidence (Broadhurst et al., 1980) in the Lanca-
shire coal measures. R o o t peats comprise up to 75% of the biomass during
the Westphalian B and C (early Middle Pennsylvanian) in Europe and the
Appalachians b u t are quite variable. The extremely high r o o t composi-
tion and mixture of Lepidophloios and cordaites are consistent with wet-
dry seasonality. In the upper Middle Pennsylvanian (Westphalian D) of the
midcontinent, roots usually account for less than one half of the peat com-
position and as little as 27% at some sites in the Herrin Coal (Phillips et al.,
1977). The interpretation of these trends is that the climate was relatively
wet during Early Pennsylvanian, drier during the early Middle, and wettest
during the late Middle Pennsylvanian.

Seasonal wet-dry inferences

The likelihood of wet-dry or dry-wet seasonality during the Early and
early Middle Pennsylvanian, respectively, seems quite consistent with our
vegetational and peat-composition data, b u t the lines of reasoning are not
firm enough to independently d o c u m e n t this assumption. In the lower
Westphalian of Europe even the r o o t peats were exposed to degradation and
served as a substrate for small rhizomatous ferns. The mixture of Sigillaria,
Lepidodendron and Lepidophloios, which we can not examine from in
situ sequences, would be consistent with wet-dry seasonality resulting in
abundance of such a variety of lycopods. On the whole, Lepidodendron
dominated.
While not very abundant in the lower Westphalian A, gymnospermous
cordaites became more abundant, and often subdominant, in the West-
phalian B and C (lower Middle Pennsylvanian). They exhibited t w o peculiar
features in their r o o t systems. The roots lacked aerenchyma (indicative of
aquatic adaptation) and exhibit eccentric growth rings, that were n o t annual
growth rings in the sense of a wet-dry or warm-cold seasonality; the stems
do n o t exhibit such interruptions of growth. In general, the cordaites became
abundant in coal swamps only where t h e conditions were physiologically
drier or brackish, where excessive evaporation occurred or perhaps where
seasonally wet-dry climate (with emphasis on the drier season) existed.
The last explanation would be the most consistent with the high root-peat
composition of the lower Middle Pennsylvanian. The cordaites without
r o o t aerenchyma (Cridland, 1964) may have flourished in the dry season
b u t were barely tolerant as shown by interruptions of root growth during
the flood season. The nearest modern analog to the eccentric growth rings
in the cordaitean roots may be incipient cypress r o o t knees (Kramer et al.,
1952) that develop only in prolonged seasonal flooding and not in brief or
permanent flood-level regimes (Whitford, 1956). The dominant lycopod
that grew with these cordaites was Lepidophloios, a standing water indicator.
In some swamps of the same interval, Paralycopodites was dominant, and
240

a minor development of the cordaites and higher shoot to r o o t ratios oc-

curred.

Stratigraphic patterns o f identified coal resources

A compilation of identified bituminous coal resources for each of the

major coal regions of the Pennsylvanian of the United States was undert aken
by Phillips et al. (1980). The objectives were to compare the stratigraphic
relationships between the inferred paleoclimatic interpretations of patterns
of l yco p o d dominance and the quantitative changes in coal deposits. We
recognize that the caveats necessary to make such approxi m at e comparisons
are numerous. The basic hypothesis to be considered is: if there are pre-
dictable qualitative relationships between peat composition and environ-
mental implications, then there should be quantitative relationships. Both
the peat and coal are wetness indicators; each provides a measure of climatic
trends stratigraphically in each region where coal swamps developed.
The division of the coal resource chart in Fig. 13 shows the southern
Appalachian coal fields separated because of limited stratigraphic corre-
lations. The resources pl ot t ed are for regional seams (or groups in Alabama)
with more than l 0 s short tons of identified bituminous coal. The anthracite
deposits of Pennsylvania were omitted because of correlation problems,
and areas with relatively small coal resources were not included, except
those within the major coal regions.
The general stratigraphic trends coincide with the patterns of l y c o p o d
dominance in the Lower and Middle Pennsylvanian: an abundance peak in
the Lower Pennsylvanian, a decline in the lower Middle Pennsylvanian, and
a marked rise in the upper Middle Pennsylvanian. The resources of the lower
Middle Pennsylvanian (and the paleobotanical data) do n o t convey a simple
trend. While there is a decline in resources in the lowermost part of the
Middle Pennsylvanian, there is a subsequent rise. During the middle Middle
Pennsylvanian there is a slight decline in coal resources of the Appalachians;
at the same time there is a rise in abundance of coal resources in the Mid-
c o n t i n e n t coalfields. In the Appalachian coal region there are more major
coal seams in the lower Middle Pennsylvanian than in any other interval.
While the onset of the early Middle Pennsylvanian is regarded as a drier
interval, there is a subsequent increase in resources, which indicates pro-
gressively wetter conditions or perhaps a diminution in seasonality. We are
somewhat uncertain how to interpret the shifts evident from the study of
coal resources and available paleobotanical data. The stratigraphic patterns
may be the consequence of gradually increasing wetness of the climate
during Westphalian C time and then a westward shift of rainfall b e y o n d the
Appalachians to the Midcontinent area. As previously mentioned, the late
Middle Pennsylvanian is regarded as probably the only time in which the
Eastern Interior Basin was wetter on the whole than the Appalachians.
The marked diminution of coal resources at the Middle--Upper Pennsyl-
 241

INTERIOR COAL PROVINCE A P P A L A C H IAN

~- Western Region ,Eostern Region COAL REGION
U) I0 IO 10 9 I 0 It 1010 10 9 I0 I0 ,10 9
I I I I
I .%
e~

.................~ii!iiiiii~i~i~!~iii~i~i~i~iiii~i~!!~iii~i~i!i~iii~i;~;~ii1
......................

, iiiiiiiiiiiiiiiiiiiiiiii!!iiii
, ......................................

= ~ ~ .........i•i!!i!!i•i:•i!•,!!!i•i!i!,!!•i:•i•!ii••••!i i

............ ::::::::::::::::::::::::::::::::::: ~i

WARRIOR, PLATEAU, £ND ................ ~::iiiii~!~!!::i?iii!ii:::::ii:;!:iiii:ii:iii[iliiiii)

i:i:,
CAHABA COAL FIELDS . . . . . . . . . i;i,i;iiiiiiii!i!;iiii:iiii:i!iii!.::::::::::::::::::::::
........~iiiili!iiiii!ii!iiii!i!i!iiiiiiiiiii:i!
.
.... , ................ iiiiiiii

o
--J . .

Fig. 13. Stratigraphic patterns of identified bituminous coal resources for the major
Pennsylvanian coal regions of the United States. Regional resources are plotted on a log
scale for seams or coal groups (Alabama) with >108 short tons (compiled by Phillips et
al., 1980). Lycopod dominance of coal-swamp vegetation diminished somewhat with a
concomitant shift in relative importances of genera near the Lower--Middle Pennsyl-
vanian boundary; this coincides with the diminution of coal resources. Subsequently,
Lepidodendron and then Lepidophloios became the dominant lycopods in the Interior
Coal Province as coal resources increased to a maximum level. All the dominant lycopod
tree genera became extinct during the Middle--Upper Pennsylvanian transition where
coal resources abruptly diminish.

vanian transition is consistent with the climatic interpretation derived from

l y c o p o d patterns. This was probably exacerbated by extensive marine
transgressions that further reduced the refuge areas for freshwater adapted
reproduction of lycopods. The stratigraphic diminution of coal resources,
first in the Western Coal Region, then in the Illinois Basin Coal Field, and
finally in the Appalachians is consistent with reduced moisture from the
east. On the basis of coal resources alone, the climate during deposition
of the upper part of the Conemaugh (Casselman Formation) and equivalents
may have been more severely dry than the lower; there are no significant
coal resources presently reported for this stratigraphic interval, although
coal basins in Maryland are currently being assessed.
The inferred onset of very wet conditions, beginning with the Pittsburg
coal bed in the Dunkard Basin, is abrupt; this return to the typical Pennsyl-
242

vanian coal-swamp climate is no d o u b t a complicating factor in delineating

a Pennsylvanian--Permian boundary in that area (see Barlow, 1975). While
this major series of coal deposits is mostly limited to the eastern extreme
of the coal belt in United States, consistent with an eastern source of mois-
ture, there is evidence of an initial broader development of coal swamps
on across the Midcontinent area.
There are several important coal resources in the Virgilian of Kansas and
Iowa, and, despite the lack of comparable Virgilian outcrops in the Illinois
Basin Coal Field, there were probably a few significant coal deposits prior
to erosional loss. Virgilian strata younger than the Pennsylvanian strata
described elsewhere in the Illinois Basin Coal Field have been discovered
in a graben in Union County in western Kentucky. A core drilled for the
Kentucky Geological Survey encountered two relatively thick coal seams;
one approximately 4.3 m thick, and the other 1.5 m thick, within a 19-m
interval. Coals immediately above and below this interval are thin, which
is typical for the Upper Pennsylvanian coals in the Illinois Basin Coal Field.
Preliminary palynological examination of the thickest coal indicates an age
approximately equivalent to the lower part of the Monongahela Formation
of the Appalachian Coal Region.

Stratigraphic patterns of petrographic composition

Regional stratigraphic analyses of the petrographic composition of 30

of the major coal seams in West Virginia by Grady {1979) included coals
from each of the proposed stratigraphic-climatic intervals. This is the first
extensive, regional, stratigraphic analysis of petrographic composition
available for comparison with paleoclimatic interpretations. Grady (1979)
also presented paleo-environmental interpretations that are quite consistent
with the general patterns presented herein. We have summarized some of
his general observations, quoted a few of his summaries, and refer the reader
to this highly significant contribution.
Grady divided the coals and environmental interpretations into three
groups which he termed lower, transitional or middle, and upper. The lower
group is Lower Pennsylvanian from the Pocahontas No. 3 coal bed to the
Sewell. The transitional or middle group is lower Middle Pennsylvanian
from the Gilbert to the Cedar Grove coal bed and the upper is both upper
Middle and Upper Pennsylvanian from the Coalburg coal bed to the Waynes-
burg coal bed. The Coalburg was included in both transitional and upper
groups on different characteristics.
Grady (1979, p. 272), comparing each of the three different swamp
environments stated, "Following each stage of inorganic sediment deposi-
tion, nearly the same conditions of organic accumulation were repeated, al-
lowing another seam to form with only slight compositional differences from
the seam below." He noted (p. 266) that, "Mineral matter content is signifi-
 243

cantly higher in coals in the upper half of the Pennsylvanian system; the
Coalburg seam and above. Inertinite, and probably exinite, is higher in the
lower Pennsylvanian [Lower and lower Middle] coals; the Coalburg seam
and below. Vitrinite content is generally lowest in the coals of the middle
Pennsylvanian Kanawha Formation.
In the Lower Pennsylvanian coals the increased inertinite and exinite
are not concentrated in layers in the seams and Grady's interpretation that
the water table dropped frequently, as in short droughts, is consistent with
the wet-dry seasonality inferred by Broadhurst et al. (1980) for the Lancashire
coal fields. The lower Middle Pennsylvanian coals (Gilbert through Coalburg)
have a wide range in inertinite and slightly more mineral matter, which
could be due to flooding (Grady, 1979) or to exposure and degradation of
peat according to Cecil et al. (1979). The lower vitrifiite content of the
lower Middle Pennsylvanian coals would be consistent with higher root and
degraded plant content noted in the coal-ball peats. In the coals of the
upper half of the Pennsylvanian (upper Middle and Upper Pennsylvanian),
as treated by Grady, there are somewhat greater variations in maceral com-
position than in the lower or transitional group even with the exclusion of
the Coalburg. It is suggested that the Stockton-Lewis coal bed to Upper
Freeport coal bed form a natural grouping of coals. The principal variants
among the Upper Pennsylvanian coals are the uppermost major coals in
each group, the Elklick coal bed in the Conemaugh Formation and the
Waynesburg coal bed in the Monongahela Formation.
In comparing proposed stratigraphic-climatic intervals to divisions of
the West Virginia coals, based on petrographic composition, the same
groupings are recognized by Grady (1979) for the Lower and lower Middle
Pennsylvanian; an upper Middle grouping could also be designated. The
Upper Pennsylvanian coals are different but do not exhibit a coherent
pattern.

E U R A M E R I C A N CLIMATE AND COAL

Warm equable climates for the Euramerican coal belt were suggested by
paleobotanists interested in the origin of coal (Potoni~, 1909; White and
Thiessen, 1913; Zalessky, 1914; White, 1931; Jongmans, 1954; Krausel,
1964; Schopf, 1973), but few accepted or explicitly advocated a tropical
climate as did Zalessky (1914) or related it to continental drift (Meyen,
1969, 1970, 1973; Schopf, 1973, 1975). Paleogeographers and paleoclima-
tologists have advanced tropical interpretations for the Pennsylvanian coal
belt to the point that differences largely concern disposition of the paleo-
equator across North America (Fig. 14) and Europe, the position of Gond-
wana (Ziegler et al., 1981; Smith et al., 1981; Morel and Irving, 1981) and
the lithospheric plate collisions between Laurussia and Gondwana (Ziegler
et al., 1979).
 244

EXPLANATION - Suggested Paleoequators

Highlands A DE~I~IOINESIA N
1"--7 Lowlands (Dott and Batten, 1976)
Coastal plains B WESTPHALIAN C-D
Cool basins (presentextent (Bombach and others, 1980)
EEE~] Shallow marine C DESMOINESIAN- VIRGILIAN
Evaporites (Heckel, 1977, 1980)

Fig. 14. Paleogeographic m a p of coal regions in the United States and adjacent areas for
the Middle Pennsylvanian (modified by John Shepard from Scotese et al., 1979, ac-
cording to Averitt, 1975, Dott and Batten, 1976, and Van Houten and Brown, 1977).

Orographic barriers in the paleotropics

Proposed paleoequators for North America generally arc diagonally

across the United States through or near the Canadian Maritime Province
to Europe. From Mississippian to Permian time proposed paleoequators
in map sequences are generally similar. The divergences occur within the
range shown in Figure 14. The paleoequator of Bambach et al. {1980) and
Ziegler et al. (1981) is more typical of those published.
 245

The "collision model" proposed by Ziegler et al. (1979) and the expand-
ed paleoclimate model from Robinso n (1973) suggest plausible paleotectonic
events for changing the wetness of the tropical coal belt but not for all
paleotropical areas. According to Ziegler et al. {1979), Gondwana rotated
clockwise and collided with Laurussia during the Pennsylvanian, resulting
in the formation of the Ouachita, Appalachian, Mauritanide and Hercynian
foldbelts. The rise of highlands east of each of the major coal regions re-
sulted in orographic barriers to the moist equatorial easterlies.
If the "collision model" is applied to the paleoclimatic patterns of change
inferred from the coal-swamp vegetation, there would be several major
tectonic pulses or episodes of orogeny with temporary "rainshadows".
Wetter intervening intervals imply reduction of barriers to moist easterlies
by erosion. This general kind of a model, coupled with an increasingly
emergent paleocontinent and diminished moisture sources, poses a plausible
alternative to extraterrestrial control (Meyen, 1973). The pulselike drier
intervals were progressively more severe and the basic explanations for the
markedly drier Permian tropics are also applicable to Early-Middle and
Middle--Late Pennsylvanian transitions on a different scale.
Where orographic barriers were lacking or paleotectonic histories were
different, there should be quantitative exceptions to the paleoclimatic and
vegetational changes seen in the tropics. The gradual Changes in the com-
pression floras of the Cantabrian of Spain (Wagner and Prins, 1979) may
reflect the less severe coastal position without highlands to the east. China
exhibits a different kind of change at the Westphalian--Stephanian boundary
(Li and Yao, 1982) from a Euramerican compression flora to a Cathaysian
flora -- which sets it apart as a separate floral province. However, the coal-
swamp vegetation of the Permian of southwestern China contains the same
kinds of trees as does the Upper Pennsylvanian of Euramerica (Tian, 1979,
and pers. commun., 1982). One could regard the Cathaysian Province as
the Permian extension of tropical coal-swamp vegetation.

Regional gradients in wetness

According to the paleogeographic map of Bambach et al. (1980) for the

Middle Pennsylvanian (Fig. 14), the major coal regions of the United States
were located just south of the paleoequator which crossed the northern
part of the Forest City Basin through Iowa, Missouri and Kansas. This places
highlands east of the coal swamps as a potential orographic barrier. This is
consistent with the east--west gradient with less moisture generally evident
from the Appalachians to the Western Interior Coal Region and beyond.
Although the Western Interior and Appalachian coal regions have elongated
configurations with SW--NE trends, the regional gradients of wetness within
each were of different magnitudes. The expanded paleoclimatic model of
Robinson (1973) is interpreted by Ziegler et al. (1979) as predicting a broad
rainy zone along the eastern side of the coal-swamp region and a markedly
246

narrow zone within the interior. This model is consistent with apparent
vegetational gradients seen from the northeastern Oklahoma shelf to the
northern part of the Forest City Basin. The dominance of cordaites in some
coal swamps of Kansas and especially Iowa during the early Desmoinesian
may reflect a combination of some brackishness, moderate rainfall, and
severe evaporation rates. During the Permian, evaporite beds developed in
these states (Zharkov, 1981).
During the Desmoinesian, the wettest interval in the Interior Coal Prov-
ince resulted in the Illinois Basin Coal Field having the largest coal swamps
in the Euramerican Pennsylvanian. These coal swamps are n o w represented
by the Springfield {No. 5) and Herrin (No. 6) Coal Members and their
equivalents. Probably even larger coal-swamps, as represented by the Crowe-
burg-Colchester-Lower Kittanning coals existed when the coal regions were
connected (Wanless, 1939). The occurrences of the largest regional coal
swamps in the Illinois Basin Coal Field relate to basinal characteristics as
well as to freshwater availability. Rainfall in the Interior Coal Province as
well as in the Appalachian coal region increased during the Desmoinesian.
The ancient Michigan River System, draining areas in the northern Appa-
lachians and eastern Canadian Shield (Pryor and Sable, 1974) substantially
augmented the freshwater supply across the Illinois Basin Coal Field. This
sustained water supply could account for the very strong dominance of
Lepidophloios vegetation, high shoot to r o o t ratios, and low fusain con-
tent in the coal-ball peats.
The drastic change in climate at the Middle--Late Pennsylvanian transition
should have resulted in at least slight differences in coal-swamp floras from
region to region if the moisture sources were to the east and were reduced
by orographic barriers. The eastward trend in moisture reduction is evident
in the diminution of identified coal resources, first in the Western Interior
Coat Region, then in the Illinois Basin Coal Field, and finally in the Appa-
lachians. The marked reduction of key spore taxa, such as Lycospora and
Thymospora pseudothiesseni, occur concurrently at the D e s m o i n e s i a n -
Missourian boundary in the Interior Province, and they became rare by
earliest Conemaugh time (Brush Creek coal) in the Appalachians.

ACKNOWLEDGMENTS

For generous help in field work, stratigraphic problems and access to

information and materials we thank Matthew J. Avcin and R o b e r t L. Ravn
formerly of the Iowa Geological Survey; Elso S. Barghoorn, Harvard Uni-
versity; Manfred Barthel, Humboldt-Universit~t zu Berlin, D.D.R.; Robert
W. Baxter, University of Kansas, Lawrence; C. Blaine Cecil, U.S. Geological
Survey, Reston; Donald Chesnut, Kentucky Geological Survey; Alan Donald-
son, West Virginia University; Philip J. DeMaris, Debbie Gaines and John
Nelson, Illinois State Geological Survey; Muriel Falron-Demoret, University
of Liege, Belgium; Donald L. Eggert, Indiana Geological Survey; Samuel A.
 247

Friedman, Oklahoma Geological Survey; William H. Gillespie,U.S. Geologi-

cal Survey, Charleston; Jean Galtier and John Holmes, University of Mont-
pellier,France; Robert M. Kosanke, U.S. Geological Survey, Denver; Robert
E. McLaughlin, University of Tennessee, Knoxville; Hermann W. Pfefferkorn,
University of Pennsylvania, Philadelphia; Anne Raymond, University of
Chicago; John Shepard, Shell Oil Company, N e w Orleans; Cedric Shute,
British Museum of Natural History, London; Natasha S. Snigirevskaya,
Komarov Botanical Institute,Leningrad; W.D. Ian Rolfe, Hunterian Museum,
Glasgow.
We wish to specially acknowledge the participation and contributions to
these studies by William A. DiMichele, University of Washington at Seattle.
This cooperative research project was supported in part by N S F Grant
E A R 78-12954 entitled, "The Quantitative Analysis Of Pennsylvanian
Coal-Swamp Vegetation in Relation to Coal" to the Paleobotany Laboratory,
University of Illinois,and by the IllinoisState Geological Survey.

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