Impact of an icehouse climate interval on
tropical vegetation and plant evolution
Hermann W. Pfefferkorn
1
, Robert A. Gastaldo
2
, William A. DiMichele
3
1Department of Earth and Environmental Science, University of Pennsylvania, Philadelphia, PA 19104-6316, USA
email: hpfeffer@sas.upenn.edu
2Department of Geology, Colby College, 5820 Mayflower Hill, Waterville, ME 04901-8858, USA
email: robert.gastaldo@colby.edu
3Department of Paleobiology, MRC-121, National Museum of Natural History,
Smithsonian Institution, Washington, DC 20560, USA
email: dimichel@si.edu
ABSTRACT: Complex plant ecosystems first experienced the effects of major glaciation during the Late Paleozoic Ice Age. The general
response of Carboniferous tropical vegetation to these climatic fluctuations, especially the transitions from greenhouse to icehouse condi-
tions (ice age sensu lato) and a return to warm times, now can be characterized based on large paleobotanical data sets originally collected
to solve stratigraphic and paleoecologic questions. These data come primarily from North America and central Europe, which at the time
were part of a single continental mass situated in the tropics. At the onset of icehouse conditions, innovation (speciation leading to novel
forms and ecologies) occurred in environments subjected to perhumid (everwet) climates, while floras in better drained, drier, and more
seasonal conditions remained dominated by holdovers/survivors from older biomes. This pattern is termed the ‘Havlena effect’. During the
height of the ice age, glacial-interglacial cycles produced large sea-level fluctuations and concomitant climatic changes, such that signifi-
cant areas of continents in the tropics were alternately covered by shallow seas or densely vegetated terrestrial coastal plains. In spite of the
repeated destruction of wet lowland habitats during each marine transgression and their further fragmentation that accompanied a climate
change from humid to sub-humid, seasonally dry conditions, most of the species and the basic configuration of the plant communities in the
wetland biome remained stable. This resilience demonstrates that glacial-interglacial cycles by themselves are not responsible for either
extirpation or extinction of these biomes. At the transition from icehouse-to-greenhouse conditions, dry-biome forms, which had been
evolving outside of the taphonomic preservational window, became dominant across basinal landscapes while wet landscapes retained
their ‘conservative’species composition. This pattern is termed the ‘Elias effect’. Thus, environmental threshold-crossing marked both the
beginning and end of this cold interval, with the loci of response in different environmental settings. In contrast, the minor systematic
changes that occurred during glacial-interglacial cycles did not influence the composition or structure of tropical lowland vegetation sub-
stantially.
INTRODUCTION
The latest Mississippian–Pennsylvanian–Permian (Late Paleo-
zoic Ice Age or LPIA of Montañez and Poulson 2013) and the
Neogene-Quaternary are the only times in Earth history when
complex vegetation characterized by large, arborescent plants
covered the land during icehouse conditions (= ice age sensu
lato). In addition, the Late Paleozoic is the only interval in
Earth history during which the responses of vegetation to shifts
and transitions between these various climate states (green-
house to icehouse and icehouse to greenhouse) can be studied
(Gastaldo, DiMichele and Pfefferkorn 1996), leaving the LPIA
as the time interval where its effect on terrestrial plant cover
provides a model system. Currently, the planet has not experi-
enced an icehouse-to-greenhouse transition, but it can be expected
sometime in the future. Thus, paleobotanical investigations into
the biotic responses to these abiotic transitions have important
implications for understanding the current icehouse climate and
the resultant changes in vegetation and agriculture should hu-
man activity trigger the onset of greenhouse conditions.
Late Paleozoic paleobotanical data are most complete for the
paleotropics (Gastaldo, DiMichele and Pfefferkorn 1996) for
several reasons. Widespread Carboniferous tropical peats
formed the coal deposits that fueled the industrial revolution and
are still utilized to produce significant amounts of electricity to-
day. Macrofloral and palynological remains are well preserved in
the peat deposits themselves and in the associated siliciclastic and
carbonate sediments that envelope these organic accumulations
(e.g., Scott and Rex 1985; Gastaldo, Pfefferkorn and DiMichele
1995). Numerous paleobotanical publications based on data from
these widespread and intensively mined beds and associated
strata form the basis of the current contribution (text-fig. 1).
Three kinds of vegetational transitions between climate states are
considered in this paper. Although glacial deposits are known
from the latest Devonian (e.g., Isaacson et al. 2008; Brezinski et
al. 2009) and earliest parts of the Mississippian, we focus our at-
tention on the vegetational response during the LPIA. The LPIA
began in the earlier Visean (Middle Mississippian) and experi-
enced a second pulse of glaciation in the Serpukhovian (Late
Mississippian), when ice caps formed around the South Pole in
Gondwana (Ziegler 1989; Crowley and Baum 1992; Grossman et
al. 2008; Gulbranson et al. 2010). The first example (text-fig. 2,
A) highlights the transition from greenhouse-to-icehouse condi-
tions at the initiation of the LPIA. Subsequently, glacial-intergla-
cial cycles at the height of that ice age (text-fig. 2, B and C) are
Stratigraphy, vol. 14, nos. 1–4, text-figures 1–4, pages 365–376, 2017 DOI: 10.29041/strat.14.1-4.365-376 365
considered based on two examples. Finally the ice-
house-to-greenhouse transition in the latest Pennsylvanian is
evaluated (text-fig. 2, D).
Vegetational changes accompanying these climate-state transi-
tions commonly are recorded by the succession of tropical low-
land paleofloras in the very thick sections of Euramerica.
Milankovitch-type glacial-interglacial cycles are well docu-
mented in many Carboniferous basins (Wanless and Shepard
1936; Heckel 1995, 2008; Gastaldo, Purkyová and Šimnek
2009; Eros et al. 2012; Cecil, DiMichele and Elrick 2014; van
den Belt, van Hoof and Pagnier 2015), and associated
vegetational changes have been studied intensively. Visean–
Serpukhovian records are found in the Upper Silesian and
Lublin-Volhynian basins that straddle the borders between the
Czech Republic and Poland and Poland and Ukraine, respec-
tively (Purkyová 1970; Kmiecik 1995; Kotasowa and Migier
1995; Gastaldo et al. 2009; Gastaldo, Purkyová and Šimnek
2009). Temporal changes in paleofloras in these basins can be
compared with similar patterns found in correlative basins
across the Late Paleozoic paleo-equatorial region. These in-
clude Morrowan– Wolfcampian (Bashkirian–Asselian) succes-
sions in the Appalachian (Gillespie and Pfefferkorn 1979;
Wagner and Lyons 1997; Blake et al. 2002; Gastaldo et al.
2004) and Illinois basins (Phillips et al. 1974; DiMichele,
Pfefferkorn and Phillips 1996; DiMichele, Phillips and Nelson
2002) in the USA and later Langsettian–Sakmarian (Bash-
kirian–Sakmarian) successions in a number of European basins
(e.g., Kerp and Fichter 1985; Cleal and Thomas 2005; Opluštil
and Cleal 2007; Wagner and Álvarez-Vázquez 2010; Cleal et al.
2012; Opluštil et al. 2016). In addition, late in the Middle Penn-
sylvanian (~Moscovian–Kasimovian; Westphalian–Stephanian;
text-fig. 2), a period of major warming occurred accompanied
by ice melting and sea-level rise (Fielding, Frank and Isbell
2008; Heckel 2008; Rygel et al. 2008). This interval was
marked by a major floral turnover and has been investigated in-
tensively, particularly in North America (Phillips and Peppers
1984; DiMichele and Phillips 1996a; Kosanke and Cecil 1996;
Peppers 1996, 1997; Falcon-Lang et al. 2011), and can be com-
pared with patterns in Europe. Hence, the extensive literature on
Carboniferous paleobotany and biostratigraphy and its linkage
to independently documented environmental changes provide
the database on which the patterns discussed herein are built and
framed in a paleoclimatic and evolutionary perspective.
CHANGES DURING THE ONSET OF ICEHOUSE CON-
DITIONS: HAVLENA EFFECT
The onset of icehouse conditions in the earliest Serpukhovian
after the late Visean warm interval (Iannuzzi and Pfefferkorn
2002; Pfefferkorn, Alleman and Iannuzzi 2014) was marked in
the paleotropics by a transition from semiarid-to-subhumid,
wet/dry climatic cyclicity to a predominantly everwet, hu-
mid-to-perhumid climate (following the terminology of Cecil
2003). Although terrestrial facies from this stratigraphic inter-
val are present in many parts of the world, the thickest sections
366
Hermann W. Pfefferkorn et al.: Impact of an icehouse climate interval on tropical vegetation and plant evolution
TEXT-FIGURE 1
Partial palinspastic map of the tropical area during the Pennsylvanian (Carboniferous, 310 Ma), modified after Crowley and Baum 1992, showing areas
where data were generated, used, or cited: 1-Texas, 2-Kansas, 3-Illinois, 4-West Virginia, and 5-Alabama in the United States, 6-Upper Silesian Coal Ba-
sin in the Czech Republic and Poland, and 7-Lublin-Volhynian Basin in Poland and Ukraine. Paleo-equator (E–E’) is shown against the present-day con-
tinental outline for ease of orientation.
are preserved in the USCB (Upper Silesian Coal Basin), part of
a fore-deep basin that straddles the boundary between Poland
and the Czech Republic (Gastaldo et al. 2009; Gastaldo,
Purkyová and Šimnek 2009). The LCB (Lublin Coal Basin)
in Poland and Ukraine also preserves a virtually complete, even
though condensed, succession of intercalated marine and terres-
trial beds at this transition.
Purkyová (1959a, b) and Havlena (1961) identified a pattern
in plant-macrofossil distribution in stratigraphic sections of the
Czech part of the USCB that appeared contradictory at the time.
The roof shale floras preserved directly above a coal seam,
which in Havlena’s terminology in German is flöznah or
‘stratigraphically near the seam,’ represented the wettest envi-
ronments and were comprised of species similar to those occur-
ring in younger, overlying strata (e.g., Westphalian). However,
fossil floras recovered from the siliciclastic-dominated parts of
the successions higher above coal seams (flözfern or ‘strati-
graphically distant from the seam’) consisted of species known
from older parts of the stratigraphic section (e.g., Visean).
These latter floras represent somewhat drier environments as
shown by their autoecological features and their sedimentologic
context (Gastaldo 1996). Hence, the more mesic ‘flözfern’ flo-
ras were populated by survivors from earlier times, giving them
a late Visean aspect; in contrast, the ‘flöznah’ assemblages were
of a younger aspect. Thus, collections made in ice-house strata
but deposited during more seasonally dry time intervals within
a single Milankovitch cycle consist of floras that appear to be
‘old’, due to close evolutionary linkages with forms that were
dominant during the prior greenhouse interval (text-fig. 3, A). It
is clear that the everwet and seasonally dry biomes and their re-
spective floras occupied climatically or ecologically disparate
environments within the same broad, geographic region
(Gastaldo 1996). The phylogenetic relationships of taxa in these
two separate floras indicate that the wet environments were the
loci of evolutionary innovation, when earlier it had been con-
centrated in seasonally dry settings. Here, we name this pattern
the Havlena effect (text-fig. 2, A) after Vaclav Havlena
(1928–1984).
At the onset of the icehouse climate, the plants of the seasonally
dry biome survived in abundance in environments character-
ized by periodic drought stress, which remained part of the
tropical lowlands. Later, as icehouse conditions became more
fully developed, the lowlands experienced long-term oscilla-
tions between humid/perhumid climates and seasonally drier
climates, reflecting oscillations in polar ice volume (Montañez
and Poulson 2013). These oscillations were tracked by their re-
spective biomes; however, there were significant extinctions of
the older elements of the seasonally dry flora during the first
few glacial-interglacial oscillations of the early Serpukhovian
(Gastaldo et al. 2009).
At the same time, wetland assemblages persisted with the same
systematic composition and diversity through multiple gla-
cial-interglacial cycles, clustered into several separate, succes-
sive intervals of ~1.8-myr duration. Species turnover during
these times was 10% between successive glacial-interglacial
cycles and involved the less common taxa (Gastaldo et al.
2009); replacement species were derived from within the
coastal lowlands. Species turnover exceeded 30% during two
stratigraphic intervals and appears to have happened ‘rapidly’
in each case. However, Gastaldo, Purkyová and Šimnek
(2009) demonstrated that species extirpation and extinction
367
Stratigraphy, vol. 14, nos. 1–4, 2017
TEXT-FIGURE 2
Time intervals (A–D) under consideration in relation to intervals of cold
climate (= icehouse conditions or ice age sensu lato) and warm climate (=
greenhouse conditions). Strong but different effects on tropical lowland
floras are seen in intervals A and D, while vegetation during the gla-
cial-interglacial cycles at the height of the ice age (intervals B and C)
shows only minimal effects. Boundaries of icehouse and greenhouse times
and nature of glaciation after Isbell et al. (2003a, b) and Fielding, Frank
and Isbell (2008). Ages from the International Chronostratigraphic Chart
(IGC 2016.04).
during these intervals of elevated turnover were coincident with
widespread tropical warming, occurring during Southern Hemi-
sphere deglaciation, and reflected in the deposition of thick ma-
rine zones (201 m maximum thickness). Each marine zone
consists of multiple, condensed maximum flooding surfaces
(e.g., Naneta and Enna marine zones), identified using
gamma-ray logs and representing upwards of 500 kyr per ma-
rine zone (Gastaldo, Purkyová and Šimnek 2009).
As the ice age intensified towards maximum glacial coverage at
the Mississippian-Pennsylvanian boundary, plants of the
Early–Middle Mississippian seasonally dry biome continued to
survive in tropical latitudes. Their biogeographic range likely
was pushed to the western margin of Pangea during glacials and
returned to more central regions during interglacials, when
equatorial Mean Annual Temperature stabilized and seasonal
rainfall conditions returned. Much of the biome underwent ex-
tinction, although many taxa survived into the Pennsylvanian
and perhaps even the Permian. Some of the taxa that seemingly
disappeared from seasonally dry paleofloras apparently sur-
vived in extrabasinal refugia. These species reappear as Lazarus
taxa in the latest Carboniferous and earliest Permian, as cli-
mates dried in the western tropics.
The increasing dominance of the wet facies in tropical basins
during glacial phases of Milankovitch-controlled cycles agrees
with the prediction that climate belts contract towards the equa-
tor at the onset of an ice age, due to restriction and a much better
defined ITCZ (inter-tropical convergence zone; Pfefferkorn
1995; Cecil et al. 2003; Peyser and Poulsen 2008). As the
everwet tropical environments spread rapidly to dominance
during these phases of glacial-interglacial cycles, evolutionary
innovation occurred in this newly created, unoccupied ecologi-
cal space. Competition was minimal for the taxa that could live
in this new habitat and new morphological innovations sur-
vived to dominate the landscape. This Havlena effect conforms
to what Erwin (1992) termed a ‘novelty radiation’, one that oc-
curs under reduced competition for resources and permits major
body plan innovation.
PERSISTENCE OF VEGETATION DURING THE
HEIGHT OF THE ICEHOUSE INTERVAL
The Black Warrior Basin, Alabama, is a foreland basin with a
very thick section of Morrowan age strata (Bashkirian;
Langsettian, lowermost Westphalian; text-fig. 2, B). Through
thirteen glacial-interglacial depositional cycles representing
~1.3 myr (Pashin and Gastaldo 2009), megafloras and
palynological assemblages of the wetland portion of each cycle
do not exhibit any systematic differences and do not allow the
succession to be subdivided biostratigraphically (Gillespie and
Rheams 1985; Eble and Gillespie 1989). This is in spite of the
complexity of spatial distribution shown by the floras in any
given wetland landscape (Gastaldo et al. 2004). The
vegetational persistence at this temporal scale is similar to that
reported by Gastaldo et al. (2009) for Serpukhovian floras in
the Silesian Basin.
The Carbondale Formation of the Illinois Basin is of late
Desmoinesian age (late Moscovian; Asturian, latest West-
phalian; text-fig. 2, C). It also exhibits a pattern of vegetational
persistence in the face of repetitive changes in climate and sea
level. A minimum of 10 intervals of sea-level rise/fall and ac-
companying climatic changes are recorded by recurring pack-
ages of marine and terrestrial rocks (cyclothems), attributed to
glacial eustacy (Wanless and Shepard 1936; Heckel 1986; Ar-
cher and Kvale 1993). These Pennsylvanian terrestrial wetland
deposits are rich in plant fossils. They are preserved as coal
balls (concretions formed during peat stage; Phillips, Avcin and
Berggren 1976) or as dispersed pollen and spores in coals, as
well as adpressions in roof shales of coal beds and in floodplain
deposits preserved below coal beds, laterally equivalent to
paleosols (Gastaldo, Pfefferkorn and DiMichele 1995). Thus,
patterns of change are well documented at several levels of de-
tail, and the overall pattern is one of persistence of vegetational
structure (DiMichele, Phillips and Nelson 2002; Pfefferkorn et
al. 2008). This is particularly true of patterns in taxonomic dom-
inance, patterns of species co-occurrence, and in the centering
of species abundance in particular habitats (DiMichele,
Pfefferkorn and Phillips 1996; DiMichele and Phillips 1996a, b;
DiMichele, Phillips and Nelson 2002). Wetland species turn-
over during this time was <10% between successive cycles and
generally involved the less common taxa; the dominant tree
forms recur from one cycle to the next.
There is a strong correlation between climate and sedimentary
depositional patterns, which can be used to infer several drivers
of change in this system. At a coarse scale, indicators of wet
(coals and bauxites) vs dry (evaporites, vertisols, and caliches)
conditions have been used to configure paleogeographic maps
and infer prevailing climates (Cecil 1992). At a finer scale of
resolution, climate can be inferred from the relationship be-
tween rainfall and sediment-transport potential, with low sedi-
ment transport predominating during the wettest (coals) and
driest (chemical rocks) periods and maximum transport occur-
ring during times of seasonality (clastic rocks; Cecil 1992; Cecil
and Dulong 2003) or at the inflection points between climate
states (Gastaldo and Demko 2011). In addition, increasingly
finer resolution of climate dynamics is emerging from studies of
paleosols (Driese and Ober 2005; Catena and Hembree 2012;
Rosenau et al. 2013 a, b) and the association of floral evidence
with specific depositional environments in a sequence-strati-
graphic context (Bashforth et al. 2014, 2016; Archer et al. 2016;
DiMichele et al. 2016). Such patterns are used to infer changes
in climate associated with particular vegetation types.
It is clear from such lithologic-climatic linkages that tropical
vegetation was affected by both periodic marine transgressions
over large parts of the continent and by associated climatic
changes involving rainfall patterns and seasonality, along with
possible temperature regimes. The wetlands, in particular, were
severely fragmented and restricted in extent by these climate
and sea-level oscillations (text-fig. 4). Nonetheless, wetland
biomes reassembled regularly during the wettest phase of each
glacial-interglacial cycle during the height of the LPIA.
CHANGES DURING WANING OF ICEHOUSE
CONDITIONS: ELIAS EFFECT
A pronounced change from, on-average, a wetter to a drier cli-
mate occurred at the Desmoinesian-Missourian boundary
(~Moscovian-Kasimovian, text-fig. 2, D; Phillips et al. 1974;
Mapes and Gastaldo 1986; Cecil 1992; DiMichele and Phillips
1996a). This interval signals a period of extensive deglaciation
and global warming (Crowley and Baum 1992; Isbell et al.
2003a), marking the end of an icehouse interval. During this
change in climate state, entire taxonomic groups of land plants
disappeared from the Euramerican tropics, and wetland ecosys-
tems were extensively reorganized (DiMichele and Phillips
1996a; Pfefferkorn et al. 2008). Concomitantly, many of these
368
Hermann W. Pfefferkorn et al.: Impact of an icehouse climate interval on tropical vegetation and plant evolution
369
Stratigraphy, vol. 14, nos. 1–4, 2017
TEXT-FIGURE 3
Patterns of apparent ages (indicated by oblique arrows) recognized in tropical wetland floras (white leaves) and somewhat better drained and seasonably
drier, tropical mesic habitats (gray leaves) in local stratigraphic sections of the Carboniferous at three different times: A (greenhouse to icehouse transi-
tion), C (during glacial intervals of icehouse conditions), and D (icehouse to greenhouse transition; see also text-fig. 2, A, C, D); dates in Ma from Interna-
tional Chronostratigraphic Chart, IGC 2016.04. Apparent biostratigraphic discrepancies of tropical Carboniferous vegetation at the beginning and end of
an icehouse interval demonstrate several key features. Innovation (i.e., origination of new taxa) occurred in wetlands at the initiation of the icehouse condi-
tions (Havlena effect at A, early Namurian = Serpukhovian). During the height of the icehouse (C), floras preserved in the different tropical lowland set-
tings do not show any distinct differences. At the beginning of the next greenhouse interval (D), innovation occurs in the more seasonably drier settings
with the appearance of floras with a more modern (i.e., Permian) composition (Elias effect). The situation in Acan be compared to the tropics at the begin-
ning of our present icehouse interval in the Tertiary (beginning of Neogene); the situation in C is comparable to the Pleistocene and the Recent, whereas the
situation at D models the consequences of the possible, and often predicted, transition from icehouse to greenhouse conditions in the future.
major groups survived in peat-forming wetlands of the
Cathaysian tropical microcontinents where high rainfall pre-
vailed (Tian et al. 1996; Hilton and Cleal 2007; Wang et al.
2012). Following the transition to greenhouse conditions, gla-
cial episodes returned, albeit with smaller ice volumes in
near-field records across the Southern Hemisphere continents
(Crowley and Baum 1992; Isbell et al. 2003b). These episodes
are reflected in higher sea levels and lower variance in
cyclothemic deposits of the North American Midcontinent
(Heckel 2008; Rygel et al. 2008). Paleobotanical patterns in the
western tropics exhibit a return to a wetland flora, albeit of dif-
ferent taxonomic composition, along with the occasional pres-
ervation of a seasonally dry biome in the lowland basins (Elias
1936; Moore, Elias and Newell 1936; Mapes and Gastaldo
1986; Broutin et al. 1990; DiMichele and Aronson 1992; Tabor
et al. 2013b; Looy and Hotton 2014; DiMichele et al. 2016).
These latter plants possibly extended their biogeographic range
from areas in western Pangea, where permanent populations re-
sided (Falcon-Lang, Kurzawe and Lucas 2016).
Prior to the end of the icehouse climate, vegetation in
peat-forming mires of Europe and North America was domi-
nated by six or more species of lycopsid trees, with subordinate
tree fern and pteridosperm (gymnosperm) species. The change
at the Desmoinesian-Missourian boundary (Phillips et al. 1974)
is marked by a regional extirpation/extinction in Euramerica
that claimed nearly two-thirds of the existing mire species, in-
cluding 87% of the tree forms and 33% of the ground cover
(DiMichele and Phillips 1996a). For a short stratigraphic inter-
val after the extinction, palynological analyses (Kosanke and
Cecil 1996; Peppers 1996, 1997) indicate that dominance pat-
terns varied from coal-to-coal in the local succession, with
changing dominant groups that included small lycopsids
(Polysporia), previously rare tree lycopsids (Sigillaria),
sphenopsids, and tree ferns (DiMichele and Phillips 1996b). We
refer to this time as a ‘species lottery’. After several cycles of
peat deposition and sea-level fluctuation, a variety of opportu-
nistic tree-fern species became the dominant mire elements
(Willard and Phillips 1993; Willard et al. 2007). Similar pat-
terns occurred in flood-basin vegetation as identified from
roof-shale floras (Pfefferkorn and Thomson 1982). This consti-
tutes a reorganization of the ecological landscape within the
broader context of the lowland-wetland biome. The dominant
species of Late Pennsylvanian wetlands, although different
from those found stratigraphically below, were drawn from the
same evolutionary lineages that occurred in these habitats dur-
ing the Middle Pennsylvanian.
Prior to this Desmoinesian-Missourian wetland vegetational re-
organization, assemblages had begun to appear in the lowlands
that were quite distinct in their species composition. Dominated
by gymnosperms, particularly cordaitaleans, the Middle Penn-
sylvanian assemblages included taxa that were not to become
dominant floristic elements until the Late Pennsylvanian and
Permian. These included such things as conifers (Falcon-Lang
et al. 2009; Plotnick et al. 2009; van Hoof et al. 2013) and
Taeniopteris (Brongn.) (Bashforth et al. 2016), and possible
Sphenopteris germanica Weiss or a similar form (DiMichele et
al. 2016). They occurred in areas quite distant from contempo-
raneous high-elevation loci, indicating the presence of these
plants in the basinal lowlands during dry parts of glacial-inter-
glacial cycles. Conifer-rich floras are found more commonly
and more fully developed in the Late Pennsylvanian lowlands,
invading normally wet areas during episodes of more intensive
seasonal dryness and often associated with lithologies near the
clastic-to-chemical rock transition; the plants associated with
these floras often are of kinds more typical of the later Pennsyl-
vanian and Permian, such as callipterids and Taeniopteris.
These latter deposits are interpreted to have formed under drier
climatic conditions (Cecil and Dulong 2003). Examples include
the 7-11 flora of Ohio (McComas 1988), the Garnett flora of
Kansas (Cridland and Morris 1963; Winston 1983, 1985), the
Hamilton Quarry flora of Kansas (Rothwell and Mapes 1988;
Cunningham et al. 1993), and the Kinney Quarry flora of New
Mexico (Mamay and Mapes 1992). Numerous other dryland
floras have been reported from North America and Europe (e.g.,
White 1912; Arnold 1941; Kerp and Fichter 1985; Broutin et al.
1990; DiMichele and Aronson 1992). Beginning in the Middle
Pennsylvanian, the seasonally dry biome (following Ziegler
1990) contained a higher percentage of evolutionarily derived
taxa than did the wetland biome. In addition, Lazarus taxa,
known previously only from seasonally dry floras of the Late
Mississippian, reappear in these Late Pennsylvanian and early
Permian drier environments after a long history of unseen exis-
tence outside of the preservational window (DiMichele and
Gastaldo 2008; Gastaldo and Demko 2011). One such taxon is
Archaeocalamites Stur (Mamay and Bateman 1991). Until re-
cently, the reappearance of Sphenopteridium Schimper (Mamay
1992) also was believed to be evidence for a Lazarus taxon.
Systematic re-evaluation, however, has concluded that
Mamay’s (1992) figured material conforms to another taxon,
Sphenopteris germanica, which is an element of Middle Penn-
sylvanian, seasonally dry floras in New Mexico (DiMichele
2014) and occurs throughout western Pangea in the Pennsylva-
nian (Pfefferkorn and Resnik 1984). Ultimately, the seasonally
dry biome rose to dominance in Permian deposits throughout
most of the tropics (Kerp and Fichter 1985; Kerp 1996; Tabor et
al. 2013a; Looy et al. 2014). The exception to this is in China,
where the proximity of microcontinents to oceanic and atmo-
spheric circulation conditions permitted the everwet biome to
persist (Rees et al. 2002; Wang and Pfefferkorn 2013). This pat-
tern of appearance of new taxa in successively more seasonally
dry environments continued into the Permian, where additional
dryland biomes have been recognized, each consisting of new
kinds of conifers and associated plants (Looy et al. 2014).
In Late Pennsylvanian and early Permian stratigraphic succes-
sions, floras ascribed to wet-climate and drier-climate conditions
often occur in alternating beds that contain sedimentological in-
dicators of wetter or drier climates (Feldmann et al. 2005;
DiMichele et al. 2013; Tabor et al. 2013b; Looy and Hotton
2014). Elias (1936; Moore, Elias and Newell 1936) was one of
the first to report this observation from the Midcontinent se-
quences in Kansas. He noted that wet-climate floras, having af-
finities with Carboniferous taxa including Neuropteris ovata
Hoffmann and Macroneuropteris scheuchzerii (Hoffmann)
Cleal et al. were found in strata that are clearly of Permian age.
In contrast, the drier-climate floras were dominated by taxa
thought to be typical of the Permian, especially the foliage of
Callipteris-type pteridosperms (gymnosperms) and conifers.
These stratigraphically alternating floras represented the over-
lap of elements that, otherwise, were dominant in lowland bas-
ins and of which, ultimately, the latter biome progressively
expanded over the landscape under an increasingly dry seasonal
climate (Montañez et al. 2007; Looy et al. 2014). This is in con-
trast to the earlier Havlena effect. The latest Pennsylvanian re-
cords a drier-climate flora, the features of which are of a
stratigraphically younger aspect. Thus, the dynamics represent a
370
Hermann W. Pfefferkorn et al.: Impact of an icehouse climate interval on tropical vegetation and plant evolution
reversal of the effect noted at the beginning of the LPIA. We
call this set of conditions the Elias effect (text-figs. 2, D and 3,
D) after Maxim Konrad Elias (1889–1982). Here, innovation
occurs in extrabasinal settings and plants from those areas ex-
pand non-competitively into the depositional lowlands during
more seasonally dry times.
CONCLUSIONS
The major patterns of paleotropical vegetational change during
icehouse conditions of the Late Paleozoic Ice Age are now ap-
parent. Distinctly different patterns and dynamics constitute the
onset, height, and waning of this particular icehouse interval.
Megafloras preserved during the onset of the LPIA exhibit evo-
lutionary innovations in those areas most affected by changes in
soil moisture (swamps and mires) under increasingly hu-
mid-to-perhumid, everwet conditions. In contrast, habitats
wherein better drained soils persisted, influenced by more sea-
sonal conditions during this time, were occupied by plant taxa
that were holdovers from a similar, previous climate state.
Broad vegetational patterns during the height of the Carbonifer-
ous icehouse are similar to those of the Pleistocene, as evi-
denced during our own icehouse interval. Wetland biomes
reorganized with the same systematic diversity and composition
following biogeographic range contraction and subsequent ex-
pansion, in response to the waxing and waning of continental
ice sheets and the effect of that on global sea level and climate.
The pattern documented at the transition out of the Pennsylva-
nian cannot be compared with anything in the Recent or
Anthropocene, because we have yet to experience this direc-
tional climate shift. We, thus, suggest the following generali-
ties.
Vegetational persistence or reoccurrence is the dominant pat-
tern during the height of an ice age. Glacial advance and retreat
and associated sea-level and climatic changes appear to have
minimal effects on vegetational structure or dominance-diver-
sity patterns. This is true as long as overall extinction levels are
low between subsequent glacial and interglacial periods and no
other thresholds exist. These findings are similar to those de-
scribed for our current ice age (Bennett 1997).
Major extinctions and vegetational restructuring occur at the
transitions between globally warm and globally cool condi-
tions. Within a species pool, weedy taxa (ruderals) have advan-
tages in capturing resource space and attaining ecological
dominance during times of significant restructuring following
extinctions. Opportunist advantage has been described follow-
ing other extinctions in both the terrestrial and marine realms
(Tschudy et al. 1984; Fraiser, Twitchett and Bottjer 2005).
However, these changes do not involve major body-plan evolu-
tionary innovations. Evolutionary innovation that results in ma-
jor new body plans appears to be confined to those times and
places where extrinsic drivers either create new habitats or
cause extinctions, thus opening major resource spaces to colo-
nization (Valentine 1980; Bateman and DiMichele 2003) and
enhancing the likelihood of pulses of niche construction (Erwin
2008). These events are generally rare, and conform to periods
of ‘escalation’ (Vermeij 1987). It appears that once such re-
source spaces are filled, innovation is suppressed (Valentine
1980; DiMichele and Bateman 1996).
In the context of the ice-age tropics, innovation is most likely to
occur in wetland habitats at the onset of glaciation. This is due
to the initial creation of everwet habitats in the tropical regions,
371
Stratigraphy, vol. 14, nos. 1–4, 2017
TEXT-FIGURE 4
Maps of the United States (current outline shown to aid in orientation) with
paleo-equator (E–E’) indicated for the Carboniferous ~310 Ma during the
height of icehouse conditions. A, North America during maximum regres-
sion (lowest sea level = glacial interval of icehouse conditions), which is
associated with the maximum extent of ice in the Southern Hemisphere.
Green indicates extent of lowland floras, blue represents marine condi-
tions, and brown indicates areas of higher elevation characterized by ero-
sion. B, North America during marine transgression associated with
maximum deglaciation in an interglacial period, when lowland floras were
restricted to marginal refugia. Epicontinental seas covered large parts of
the continent. The arrows indicate the generalized direction of the trans-
gression. These two states (A and B) alternated regularly as evidenced by
the cyclostratigraphic record. Plants and plant communities persisted
through these repeated cycles even though they were extirpated over large
areas of the continent during each sea-level highstand. This persistence of
floras was demonstrated (e.g., DiMichele, Phillips and Nelson 2002) for
11 glacial-interglacial cycles in the Illinois Basin (interval C of text-figs. 2
and 3). Transgressions did not always have the same maximum extent
shown here (see Heckel 2008). Map modified after Cecil and Dulong
(2003).
while tropical seasonally dry habitats become refugia for older
taxa (the Havlena effect). At the end of glaciation, innovation is
seen more clearly in seasonally dry habitats. This is a conse-
quence of seasonally dry areas increasing their spatial distribu-
tion (spreading) during the beginning of extensive periods of
seasonal dryness, while increasingly patchy wetlands become
refugia for the older wetland elements (the Elias effect).
ACKNOWLEDGMENTS
We would like to thank Rich Lane for his decade-long labor to
keep stratigraphical and paleontological research supported in
the United States. We also remember fondly the various private
discussions at scientific meetings and in Nanjing, and we ad-
mire his achievements, his collegiality, and his input. We lost an
illustrious and important colleague.
Data used in this synthesis were collected by the authors over
decades while they were supported by their respective institu-
tions. In addition, support was made available over the years by
the NSF (U.S. National Science Foundation), the Illinois State
Geological Survey, the German Science Foundation, the U.S.
Geological Survey, and the Charles and Mary Walcott Funds of
the Smithsonian Institution. Preparatory stages of this specific
project were funded by the University of Pennsylvania Re-
search Foundation to H. W. P and NSF under INT-9627893 to
R. A. G. The project was funded by NSF under EAR-0207848
to H. W. P and under EAR-0207359 to R. A. G. We thank spe-
cifically Adam Kotas, Anna Kotasowa, Teresa Migier,
Aleksandra Trzepierczynska, Eva Purkyová, Zbynek
Šimnek, Vitaly Shulga, Cortland Eble, Mitch Blake, Maciej
Podemski, Leszek Marks, and Tadeusz M. Peryt for their sup-
port and consultation during conceptual development. We thank
Richard Lupia and Edith Taylor for constructive comments on
an earlier version of this paper.
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