INSIDE: THE LATE PALEOZOIC ECOLOGICAL-EVOLUTIONARY  LABORATORY, A LAND-PLANT FOSSIL RECORD 
 PERSPECTIVE  
PLUS:  PRESIDENT’S COMMENTS, SEPM RESEARCH CONFERENCE 
SUMMARY, SEPM AND STEPPE, SEPM AT 2015 AAPG “ICE”
December 2014     |     3
The Sedimentary Record
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CONTENTS
4 The late Paleozoic ecological-evolutionary 
  laboratory, a land-plant fossil record perspective  
11 President’s Comments 
12 Summary:  SEPM Research Conference on  
  Autogenic Dynamics of Sedimentary Systems
13 SEPM a Global Network of Support
 SEPM and STEPPE
14 Upcoming 2015 Conferences
15 SEPM at the 2015 AAPG International Conference  
  and Exhibition (ICE)
The Sedimentary Record (ISSN 1543-8740) is published quarterly by the 
Society for Sedimentary Geology with offices at 4111 S. Darlington, Suite 100, 
Tulsa , OK 74135-6373, USA.
Copyright 2014, Society for Sedimentary Geology. All rights reserved. Opinions 
presented in this publication do not reflect official positions of the Society.
Published under Creative Commons (CC BY-NC 4.0)
The Sedimentary Record is provided as part of membership dues to the 
Society for Sedimentary Geology.
Cover image: Examples of precociously appearing Methusela taxa. 
(1) Dichophyllum moorei, Garnett, Kansas, early Late Pennsylvanian. 
Baxter and Hartman, 1954. (2) Dicroidium jordanensis, Dead Sea 
Region, Jordan, late Permian. Kerp et al., 2006. (3) Dioonitocarpidium 
sp., King County, Texas, late early Permian. DiMichele et al., 2001. 
(4) Manifera talaris, late early Permian, King County, Texas. Looy and 
Stevenson, 2014. Scale bars, I cm.
4     |     December 2014
The Sedimentary Record
The late Paleozoic 
ecological-evolutionary 
laboratory, a land-plant fossil 
record perspective  
Cindy V. Looy1, Hans Kerp2, Ivo A.P. Duijnstee1,3 and William A. DiMichele4
1 Department of Integrative Biology and Museum of Paleontology, University of California, Berkeley, CA 94720, looy@berkeley.edu, duijnstee@berkeley.edu 
2 Forschungsstelle für Paläobotanik, Institute for Geology and Palaeontology, University of Münster, Germany, kerp@uni-muenster.de
3 Department of Earth Sciences, Utrecht University, The Netherlands
4 Department of Paleobiology, NMNH Smithsonian Institution, Washington, DC 20560, dimichel@si.edu
INTRODUCTION 
In this essay we examine the fossil record of land plants, focusing 
on the late Paleozoic. We explore the nature of this record in 
terms of what is preserved, where, why and with what biases. 
And as a consequence, how it can be used to answer questions 
posed at various spatial and temporal scales, what cautions we 
must consider when interpreting it, and what surprises it may 
hold. Generally speaking, the record of terrestrial plants is rich 
and reveals clear directional trends in phenotypic complexity, 
biodiversity, and ecosystem organization. It also has reasonably 
well understood taphonomic biases. It must be used with 
considerable caution, however, when researching time and 
location of evolutionary innovations and the development of 
ecological structure and interactions.
THE LATE PALEOZOIC LABORATORY
 Earth experienced a 70-million-year period of intermittent 
glaciation (Montañez and Poulsen 2013) from the middle 
Mississippian to early Permian. This interval is characterized by 
105-year glacio-eustatic cycles (Heckel 2008), superimposed on 
longer, 106-year scale intervals of global warming and cooling 
(Birgenheier et al. 2009). These are further superimposed, in 
the equatorial regions, on a long-term, 107-year scale trend of 
warming and increasing aridity (Montañez et al. 2007, Tabor 
and Poulsen 2008). Consequently, the world of the time had 
many similarities to that of today, captured in the fossil and 
geological records. The Earth’s continental landmasses, however, 
were aggregated into the supercontinent of Pangea, which 
differed greatly from today’s high elevation world of dispersed 
continents (Figure 1).
 The Pennsylvanian and Permian are known for vast coal 
deposits, which formed in extensive peat swamps. In the tropics, 
these wetlands were populated by old, evolutionarily conservative 
plant lineages, the subjects of dioramas in natural history museums 
and illustrations in nearly every paleo-textbook. At the same time, 
however, large areas of the tropics harbored more evolutionarily 
derived plants adapted to seasonal drought (e.g., DiMichele 
2014). There were also distinct north- and south-temperate 
floras segregated into wetland and drought tolerant assemblages, 
but subject to strong seasonal temperature contrasts (Rees et al. 
2002). Such areas also tended to be populated by more derived 
evolutionary lineages.
 Our understanding of the origin and spread of major late 
Paleozoic plant groups is based on these patterns of ecosystem-
scale and biogeographic-scale patterns. Some of the groups 
originating in the late Paleozoic are still important today, such as 
conifers. Others, now extinct or diminished, dominated many 
pre-angiosperm, Mesozoic ecosystems. These include cycadalean, 
peltaspermalean, and corystospermalean seed-plants and ferns 
such as the osmundaleans and primitive filicaleans (Lidgard and 
Crane 1990). Until recently, many of these groups were thought 
to have had Mesozoic or latest Paleozoic origins. Over the past few 
decades, however, some have been found in Paleozoic deposits, often 
as isolated occurrences, suggesting that significant evolutionary 
innovation took place in parts of the terrestrial landscape poorly 
represented in the fossil record. This is not a matter for despair, 
however. Such patterns may mean we cannot easily or confidently 
“stack up” the record for a direct, temporal reading. Nonetheless, 
through linkage of sedimentological and ecological factors to 
patterns of spatial and temporal plant distribution, we can still infer 
a lot about the locus and nature of the evolutionary process.
December 2014     |     5
The Sedimentary Record
THE OVERPRINT OF 
TAPHONOMY
Rule #1: Plants are crystalized climate 
 “Ja, man kann die Pflanzendecke das 
 kristallisierte, sichtbar gewordene Klima 
 nennen, in dem sich so manche Züge 
 deutlicher zeigen als in den Angaben 
 unserer Instrumente.” 
 Wladimir Köppen (1936, p.6)
This may be translated: “Yes, one may call 
vegetation materialized, visible climate, 
in which quite a few climate traits are more 
readily discernible than in the readings of 
our instruments”, or, the part in bold above, 
somewhat more graphically as “plants 
are crystalized visible climate” (Claussen 
1998). There are few more compelling 
rules for understanding the fossil record 
of land plants. And it is safe to assume 
that terrestrial plants have conformed to 
this axiom since their earliest appearances, 
which should strongly condition our 
interpretations of their spatial and temporal 
distributions and evolutionary patterns.  
 In the Pennsylvanian-Permian, perhaps 
the best examples of this are the striking 
differences in taxonomic composition 
among equatorial Euramerican, equatorial 
Cathaysian, south-temperate Gondwanan 
and north-temperate Angaran assemblages 
(Figure 1, Wnuk 1996). At a spatially 
more refined level, several compositionally 
distinct biomes have been recognized in the 
Euramerican floral realm, each associated 
with physical indicators of greater seasonal 
dryness (Falcon-Lang and Bashforth 
2004, Tabor et al. 2013). Within the best 
known of these biomes, the wetlands, 
environmental preferences have been 
determined for particular taxa or lineages 
(e.g., DiMichele and Phillips 1996a) that 
can be traced back to the earliest radiations 
of terrestrial plants (Bateman et al. 1998).  
The other fundamental controls
 There are other important taphonomic 
factors that strongly influence interpretation 
of the land-plant macrofossil record 
(Gastaldo and Demko 2011). Taphonomic 
rule #2 is that short-term preservation of 
plant remains is most likely to occur under 
a background of perhumid to wet sub-
humid conditions (terminology of Cecil 
2003), though dry sub-humid and even 
arid climates may harbor some habitats 
where preservation is possible.
 Taphonomic rule #3 is that plant 
macrofossils rarely can be recycled by 
reworking. Impressions or fragile coalified 
compressions are easily destroyed, 
exceptions being wood or wood-like 
resistant tissues. Thus, the plant macrofossil 
record preserves fine levels of temporal 
resolution and high stratigraphic integrity. 
In practice, however, a collection of 
plant fossils is usually analytically time 
averaged by sampling (Behrensmeyer et 
al. 2000). This happens mostly because 
of the difficulty of tracing a “T0” time 
horizon (Johnson 2007) laterally for any 
distance unless it is tied to an “event” of 
determinable short-term duration, say 
an ash fall (Wing et al. 1993; Opluštil et 
al. 2014). Parautochthonous and some 
allochthonous assemblages generally 
represent either members of the same 
community or plants that lived in close 
proximity to the depositional environment, 
in time and space. 
 Rule #4: Plant organic matter will be 
destroyed rapidly by the combined actions 
of physicochemical (e.g., mechanical 
breakage, fire, slow oxidation) and biotic 
agents (e.g., microorganismal decay, 
roots), particularly if on or above the 
soil surface, or in the soil vadose zone 
of water table fluctuation (Gastaldo and 
Demko 2011). Consequently, most of 
the plant macrofossil record represents 
Figure 1: The Late Paleozoic supercontinent, Pangea. Four major floral zones are indicated, 
tropical Euramerica and Cathaysia, and temperate Angara and Gondwana. 
Paleogeography after Scotese (1997)
Figure 2: Plant evolutionary innovation and environment, Late Paleozoic. A. General pattern based on an early, incomplete knowledge of the 
fossil record. Major evolutionary innovations appear in stressful, extrabasinal habitats and track climate changes, moving into basinal habitats, 
where preservational potential is highest, during the progressive global warming and drying of the late Paleozoic. B. Emerging pattern with 
an increased sampling of the fossil record. Seemingly precocious floras change our general view depicted in A. Precocious appearances reflect 
climate oscillations and accompanying tracking by plants, bringing new forms initially temporarily into the window of preservation during drier 
episodes. Or – origination of clades forming new plant biomes outside the window of preservation, PO – precocious occurrence of fossil floras, 
CO – common occurrence of fossil floras.
6     |     December 2014
The Sedimentary Record
wetland assemblages or vegetation fringing 
standing water bodies, growing within 
channels or on wet floodplains (Scheihing 
and Pfefferkorn 1984), where chances of 
preservation are highest.  
 Rule #5: Most deep-time, fossil-plant 
accumulations will be confined to what 
were, at the time, actively subsiding 
basins. Even if preserved for the short-
term, organic deposits must be protected 
from decay on intermediate time scales 
of thousands to tens of thousands of 
years to permit sufficient subsidence and 
burial below the level of active erosion. 
Intermediate-term preservation is most 
likely where ocean transgressions or inland 
water bodies could flood the site of burial. 
This must be followed or accompanied 
by tectonic creation of accommodation 
space, permitting deeper burial and 
protection from erosion on million-year 
time scales. Thus, except in unusual tectonic 
circumstances (e.g., Opluštil 2005), the 
late Paleozoic terrestrial record contains 
primarily lowland deposits, leaving much 
room for speculation about what was going 
on evolutionarily and ecologically elsewhere. 
 That fossil floras occur throughout 
most of the Phanerozoic is empirical 
documentation that there is potential for 
the preservation of plant remains when 
the conditions are right. Conditions for 
intermediate and long-term burial of 
epicontinental sediments (e.g., Davies 
and Gibling 2013) are favorable in 
Pennsylvanian-Permian basins, leaving 
a reasonably good record. Within these 
deposits, organic remains of plants from 
wetlands and localized high-moisture 
habitats are best represented, including 
swamp, peri-lacustrine, lagoonal fringe, 
coastal mudflat, floodplain and stream 
corridor habitats. 
 From a climatic perspective, the best 
record of lowland vegetation comes 
from times of perhumid to wet sub-
humid climate, which most favor the 
first step of fossilization: short-term 
preservation. Due to unfavorable 
conditions for short-term preservation, 
the plant record from dry sub-humid to 
arid conditions is very limited. There are 
also few records of true “upland” floras, 
those from continental interiors or other 
places where erosion was the dominant 
sedimentological force on intermediate 
and long-term, million-year time scales.
Precocious occurrences: Methuselah taxa
 Of greater interest than Lazarus taxa, 
from an evolutionary perspective, are 
precocious taxonomic occurrences, 
millions to tens of millions of years 
preceding otherwise well-established 
ranges. Unexpectedly “old” occurrences 
like these lead us to suggest the term 
“Methuselah” taxa for those with a much 
older origin than assumed possible, given 
the bulk of earlier existing observations. 
Upon re-evaluation of all data, the epithet 
‘precocious’ really only exists in the eye 
of the myopic beholder, and turns out to 
mean nothing more than “inconceivably 
old’’, just like Methuselah in Hebrew 
Scripture. In the plant fossil record, these 
Methuselah genera and species typically 
occur in seasonally dry environments, 
often in deposits sandwiched among those 
with typical wetland floras. They also are 
composed of or contain many derived 
elements of evolutionary lineages, implying 
a linkage between environmentally 
“peripheral” habitats and major innovation 
in plant evolution (DiMichele and Aronson 
1992).
 Among the most noteworthy Methuselah 
occurrences is the callipterid peltasperm 
Dichophyllum (Cover, 1), from the early 
Late Pennsylvanian of Kansas (Cridland 
and Morris 1963). This occurrence, in a 
seasonally dry, channel complex (Feldman 
et al. 2005), falls within the midst of 
the Midcontinent USA coal measures 
and is conifer-dominated; an assemblage 
quite unlike that of shales associated with 
surrounding coal beds. This occurrence 
caused considerable debate about the age of 
the deposit, leading some biostratigraphers 
to argue for Permian age (e.g., Bode 1975). 
Since this time, other Late Pennsylvanian 
callipterid occurrences have been 
documented, but these are rare and none 
are as old as this. 
 Several other noteworthy examples of 
Methuselah occurrences include: (1) Four 
species of the corystosperm Dicroidium 
from the late Permian of Jordan (Cover, 
2), then equatorial Pangea, in a floodplain 
deposit formed under seasonally dry 
climate (Kerp et al. 2006). This genus is 
a characteristic element of late Early to 
Late Triassic high-latitude Gondwanan 
floras. (2) Dioonitocarpidium, a cycad-like 
reproductive structure typical of the Late 
Triassic and Early Jurassic of central Europe 
(Cover, 3). It occurs in a late early Permian 
deposit from Texas, in association with 
a peculiar assemblage, deposited under 
seasonally dry climate (DiMichele et al. 
2001). (3) Voltzian conifers, a derived 
group (Cover, 4), also occur in seasonally-
dry habitats of the Texas late early Permian 
(Looy 2007, Looy and Stevenson 2014). 
Their earliest prior occurrence was late 
Permian of central Europe. (4) The 
seed-bearing structure of highly derived, 
typically Mesozoic Peltaspermales has 
been reported from isolated occurrences 
in latest Pennsylvanian equatorial regions 
of Europe and North Africa (Kerp et al. 
2001), and the early Permian of China (Liu 
and Yao 2000) and the Urals (Naugolnykh 
and Kerp 1996, Kerp 1996). The species, 
Peltaspermum retensorium, was found at 
several localities in the same Angaran 
horizon, a chance basinal occurrence of a 
rarely found “upland” plant associated with 
a flora indicating seasonal moisture stress. 
(5) Another peltasperm, Germaropteris 
martinsii, from dryland settings of late 
Permian age (Lopingian) of Central 
and Southern Europe (Kustatscher et 
al. 2014), was recently reported from 
early Permian seasonally dry deposits in 
southern France (Galtier and Broutin 
2008) and from allochthonous offshore 
settings in Texas, presumably derived 
from coastal, mangrove-like habitats 
(Erik Kvale, personal communication, 
2014 – specimens examined by Kerp and 
DiMichele). Other precociously appearing 
conifers include (6) the “Mesozoic” genus 
Podozamites from seasonally dry early 
Permian deposits of Texas (DiMichele et 
December 2014     |     7
The Sedimentary Record
WHAT’S HAPPENING 
“OUT THERE” AND HOW 
DO WE KNOW?
 Due to the climatic and taphonomic 
factors discussed above, much of 
the natural experimentation that 
characterized Paleozoic plant evolution 
seems to have occurred outside of areas 
or time windows with the best chances 
for preservation. These include basins 
during the times they experienced 
climates unfavorable for short-term 
preservation and extrabasinal regions, 
lowland and true upland (Pfefferkorn 
1980). How can we tell if major 
evolutionary breakthroughs occurred 
in such places? Fortunately, plants 
faithfully reflect climate. Because climate 
is generally insensitive to tectonic 
regime, particularly subsidence, basins 
are sometimes subject to drier climate 
at the same time they experience 
conditions conducive to intermediate-
term preservation. When that happens, 
plants from habitats that rarely become 
fossilized will appear as isolated, 
seemingly anomalous occurrences. 
Stratigraphic anomalies
 There has long been attention, particularly 
among marine invertebrate biostratigraphers, 
to occurrences of taxa outside of previously 
known temporal ranges. Given such names 
as “Lazarus” taxa (Jablonski 1986) for those 
appearing well beyond inferred range termini, 
equally important are cameo appearances well 
before known ranges. In either case, these 
appearances strongly imply significant biases 
in the record or in the patterns of organismic 
distribution on the landscape. Such evidence 
is particularly powerful where the occurrences 
straddle extinction boundaries, indicating 
unsuspected earlier existence and/or survival 
in unseen areas. Regarding plant evolution, 
precocious occurrences may indicate 
evolutionary innovation at times and in 
places outside of our detection abilities, and 
can carry significant implications regarding 
climate and habitat. 
8     |     December 2014
The Sedimentary Record
al., 2001) and Late Pennsylvanian of New 
Mexico (Mamay and Mapes 1992). (7) 
Walchian conifers, rare but known from 
the Late Pennsylvanian equatorial regions 
(e.g., Kerp 1996, Hernandez-Castillo et al. 
2001), have been reported from Middle 
Pennsylvanian age localities, two in the 
Illinois Basin, a sinkhole in limestone at 
the basin margin (Plotnick et al. 2009) 
and a channel fill within a seasonally 
dry landscape (Falcon-Lang et al. 2009), 
and two from allochthonous deposits 
in New Mexico (Lucas et al. 2013). (8) 
A number of genera reported from the 
early Permian seasonally dry habitats of 
southwestern Euramerica, most notably 
Comia, Supaia and Compsopteris, are both 
significantly more abundant and have 
much broader distributions in the late 
Permian of Angaraland and Cathaysia 
(Mamay et al. 2009, Halle 1927). (9) The 
enigmatic gigantopterids, abundant in 
the late Permian of China occur in early 
Permian seasonally dry environments of 
southwestern Euramerica (DiMichele et al. 
2005), the Arabian Peninsula (Berthelin et 
al. 2003), Sumatra (Booi et al. 2009) and 
Venezuela (Ricardi-Branco 2008).
PRECOCIOUS OCCURRENCES 
AND PLANT EVOLUTION 
 Three patterns stand out when 
considering the significance of precocious, 
Methuselah occurrences. (1) These taxa 
nearly always appear in deposits formed 
under seasonally dry background climates, 
even if the fossils themselves are from wet 
substrate sites, consistent with constraints 
on short-term preservation. (2) The taxa 
are almost always among the more derived 
members of their respective evolutionary 
lineages at some taxonomic level. (3) The 
earliest host deposits tend to be “one-
offs” – single deposits or thin stratigraphic 
horizons – found in basinal lowlands or in 
allochthonous, offshore deposits, reflecting 
taphonomic controls. 
 This pattern may be contrasted with 
Paleozoic wetland communities dominated 
by evolutionarily less-derived lycopsids, 
pteridosperms, marattialean tree ferns, 
cordaitaleans and sphenopsids. These floras 
show long-term compositional conservatism 
and intra-assemblage species turnover 
strongly constrained by evolutionary-lineage 
ecological centroids (DiMichele and Phillips 
1996b), a pattern reflective of “phylogenetic 
niche conservatism” (e.g., Prinzing 
2001; Wiens 2004). Such conservatism 
led Knoll (1985) to refer to swampy 
lowlands throughout geological history as 
“museums”. They are characterized by long-
term persistence of ecological organization 
and evolutionary innovation and of 
taxonomic composition and ecomorphic 
characteristics. When disrupted by major 
environmental disasters, they are recolonized 
from “outside” species pools, restructured 
and, subsequently, again demonstrate 
conservatism for millions of years. 
 When considered together we draw 
two conclusions from these patterns 
(summarized in Figure 2). First, evolution 
of major body-plan innovations (meaning 
ancestor-descendant divergence reflected 
in higher, traditional-Linnean ranks) 
occurred more commonly in environments 
that were environmentally challenging to 
established plant lineages and unfavorable 
for organic preservation on the short-
term and intermediate-term time scales. 
Such environments, likely, were of 
initially low diversity and encompassed 
new and different resources that were 
available for use. Increasing drought and 
temperature stress, in particular, may have 
simultaneously limited range expansion of 
existing plants and created opportunities 
for innovation. Initially permissive, survival 
likelihood of variant forms was enhanced 
due to relaxed natural selection. Second, 
we first see the results of such innovation 
when environmental change in the 
lowlands, caused by increased seasonality 
of rainfall and perhaps temperature, 
permit these lineages to move into and 
occupy basinal areas temporarily. Based 
on the low number and typically singular 
appearance of Methuselah taxa we infer 
that conditions permitting their basinward 
biogeographic shifts most often occurred at 
times when intermediate-term preservation 
was unlikely. This makes them rare to start 
with, and the deposits difficult to find, 
even if present, thus causing initial myopia 
in the eye of the paleobotanical beholder 
(i.e. the pattern seen within the window of 
preservation in Figure 2 A). In the longer-
term, evolutionarily derived lineages became 
dominant in basinal lowlands. They did so 
not by displacing the incumbent, ancestral 
forms, but by replacing them as long-lasting 
environmental change opened basins to 
long-term colonization (DiMichele and 
Bateman, 1996). Consequently, whenever 
fossiliferous sites are found outside of 
preservation-friendly regions or in settings 
of generally drier climates, seemingly 
precocious occurrences will result (Figure 2 
B). Plants appearing well before previously 
known stratigraphic ranges should be 
expected rather than considered anomalous. 
 We interpret these patterns to suggest that 
the window for innovation in ecologically 
permissive environments is brief and the 
survival of new forms declines as resource 
pools are occupied (e.g. in the extreme, 
Valentine 1980; DiMichele and Bateman, 
1996). Intrabiome and intra-species-
pool turnover tend to be dominated by 
niche-conservatism and within-clade, 
near ancestor-descendant replacements, 
reflected by paired intra-generic extinction 
and origination. The result is minor 
compositional fluctuation at the level of 
the dominant lineages through time during 
which assemblages became hide-bound 
and niche construction (Odling-Smee et al. 
2013) was a rare phenomenon. The existing 
hegemony was broken-up by periodic, 
extrinsically induced disruptions (i.e., 
Vermeij 1993).
 We also note an inversion between 
the generalized evolutionary patterns in 
December 2014     |     9
The Sedimentary Record
marine invertebrates and land plants. The 
onshore-offshore pattern of evolutionary 
innovation and radiation in marine 
invertebrates (Jablonski et al. 1983) actually 
may contribute to high amounts of Lazarus 
taxa. There, heterotroph innovations occur 
in shallow marine environments within 
the window of preservation followed by 
radiation outside this window into the 
deep. Lazarus taxa wander back into the 
preservational window after ecological 
crises. Exactly the opposite happens with 
autotrophs in the terrestrial realm. Major 
innovations happen outside the window of 
preservation, with subsequent migration, 
and sometimes radiation into the window 
following environmental change. So one can 
expect this process to produce the opposite 
of Lazarus taxa, the apparent precociously 
appearing Methuselah taxa.
ACKNOWLEDGEMENTS 
 We thank Rudolf Serbet, Division of 
Paleobotany, Natural History Museum and 
Biodiversity Research Institute, University 
of Kansas (KUPB), for providing the 
image of Dichophyllum moorei. Reviews by 
Richard Bateman and Cortland Eble are 
greatly appreciated.
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Accepted November 2014