Journal of Sedimentary Environments (2023) 8:261–282

https://doi.org/10.1007/s43217-023-00132-y

RESEARCH

Climate‑induced changes in fluvial ichnofossil assemblages

of the Pennsylvanian–Permian Appalachian Basin
Jennifer K. Crowell1 · Daniel I. Hembree2

Received: 31 January 2023 / Revised: 28 March 2023 / Accepted: 1 April 2023 / Published online: 21 April 2023
© The Author(s), under exclusive licence to Springer Nature Switzerland AG 2023

Abstract
The long-term response of riparian communities to shifting climatic conditions can be addressed by the ichnofossil record,
because organism behavior is typically altered in response to changes in environmental factors. During the late Paleozoic,
the Appalachian Basin experienced a shift from an ever-wet to wet–dry climate. Changes in the abundance, diversity, density,
and composition of ichnofossil assemblages were investigated in fluvial point bar sandstones from five roadside outcrops of
the Middle Pennsylvanian-to-early Permian Allegheny, Conemaugh, Monongahela, and Dunkard groups located in southeast
Ohio and northwest West Virginia. Ichnofossil data were collected using a 0.5 × 0.5-m grid placed on bedding plane surfaces
and from vertically oriented thin sections. Abundance, density, diversity, and burrow widths increased through the study
interval. Behaviors changed from stationary- to mobile-deposit feeding, while community composition shifted toward more
established, permanent generalists. These changes in ichnofossil assemblages suggest that the shift to a drier, more pro-
nounced seasonal climate made short- to long-term occupation of the point bar sands more advantageous as surface conditions
were more unfavorable and resources limited. This study helps us understand how terrestrial community composition and
ecosystem dynamics shift over long time intervals in response to environmental perturbations. By assessing these changes,
we can better predict what future impacts climatic shifts will have on continental ecosystems and terrestrial communities.

Keywords Continental · Point bar · Trace fossil · Behavior · Paleoecology

1 Introduction fundamental shifts in ecosystems due to significant environ-

Ichnofossils are vital to understanding ancient ecosystems
and environments as they are often the only evidence of life
in some settings (Bromley, 1996). Ichnofossils have been
utilized to interpret various aspects of continental environ-
ments, including temperature, oxygenation, salinity, nutrient
content, and turbidity, because organism behavior is typi-
cally altered in response to changes in environmental fac-
tors (Gingras et al., 2007; Hasiotis, 2002; Hasiotis et al.,
2007; Hembree, 2018; MacEachern et al., 2007; Minter
et al., 2016). Additionally, ichnofossils can help interpret
Communicated by M. V. Alves Martins
* Daniel I. Hembree
dhembre2@utk.edu
1
Department of Geology and Geological Engineering,
University of Mississippi, Oxford, MS, USA
2
Department of Earth and Planetary Sciences, University
of Tennessee Knoxville, Knoxville, TN, USA
mental changes including climate change (Bromley, 1996;
Buatois et al., 1998).
The terrestrial climate of the late Paleozoic has been
characterized by the development of strong seasonality in
the continental interior of Pangea (Cecil, 2013; Gastaldo
et al., 1996; Powell et al., 2009; Tabor & Poulsen, 2008;
Tabor et al., 2008). Several studies have been conducted on
the effect of the late Paleozoic climatic shift on paleosols of
the Appalachian Basin (Hembree, 2022; Hembree & Blair,
2016; Hembree & Bowen, 2017; Hembree & Carnes, 2018;
Hembree & McFadden, 2020; Hembree & Nadon, 2011);
however, few have focused on changes in riparian ichno-
fossil assemblages. In this study, ichnofossil assemblages
preserved in point bar sandstones were studied through the
strata deposited during the Pennsylvanian–Permian climatic
transition in the Appalachian Basin to understand the long-
term impacts of climatic shifts on riparian ecosystems. Sec-
tions of the Middle Pennsylvanian (Moscovian)-to-early
Permian (Asselian) Allegheny, Conemaugh, Monongahela,
and Dunkard groups in southeast Ohio and northwest West

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262
Virginia were selected, because paleosols within these
groups indicate an up-section shift from a wet to a drier,
more seasonal, climate (Hembree, 2022).
Investigating changes in terrestrial ichnofossil assem-
blages provides a broader understanding of the impact that
climatic shifts have on environmentally sensitive compo-
nents of ancient ecosystems. By assessing these changes,
we can also better predict what future impacts climatic shifts
will have on continental ecosystems and, specifically, ripar-
ian invertebrate communities (LaRoux and McGeoch, 2008;
Walther, 2010; Capon et al., 2013).
2 Geologic setting
2.1 Middle Pennsylvanian‑to‑early Permian climate
transition
Widespread Late Devonian through Early Mississippian
glacial deposits are some of the earliest indicators of the
initiation of the Late Paleozoic Ice Age (LPIA) (Montañez
& Poulsen, 2013). By the Late Mississippian-to-Middle
Pennsylvanian, continental ice centers extended into areas
of southern Gondwana and the Earth was in an icehouse
state (Crowell, 1995; DiMichele, 2014; DiMichele et al.,
2009; Montañez & Poulsen, 2013). The final assembly and
northward drift of the supercontinent Pangea from the Late
Pennsylvanian to the early Permian had a long-lasting effect
on many Earth systems, including global atmospheric circu-
lation patterns and glacial states (Blakey, 2008; Montañez
& Poulsen, 2013; Tabor & Poulsen, 2008). Ice sheets waxed
and waned during this time, creating changes in the distri-
bution of land and ocean as sea-level oscillated (Tabor &
Poulsen, 2008). Low-latitude environmental changes, such
as withdrawal of epeiric seas, increasing continentality, and
decreasing precipitation, likely resulted in the drying of
near-equatorial regions of Pangea (Tabor & Poulsen, 2008).
In the Appalachian Basin, this resulted in a major shift from
a humid, nonseasonal climate to a drier, subhumid, strongly
seasonal climate (Montañez & Cecil, 2013). Seasonality was
especially stronger in the interior of Pangea as it was far
from the moderating effects of the ocean.
Many studies have utilized paleosols and, to a lesser
extent, ichnofossils, to document this climate transition in
the Appalachian Basin (Hembree, 2022; Hembree & Blair,
2016; Hembree & Bowen, 2017; Hembree & Carnes, 2018;
Hembree & McFadden, 2020). Appalachian Basin paleosols
of the Upper Pennsylvanian exhibit redox features, little-to-
no carbonate nodules, and minor vertic features in contrast to
paleosols from the Lower Permian which exhibit oxidizing
features, abundant carbonate nodules, and prominent ver-
tic features (Hembree & Blair, 2016; Hembree & Carnes,
2018; Hembree & McFadden, 2020; Hembree & Nadon,
13
J. K. Crowell, D. I. Hembree
2011). These paleosols record a regional shift to more vari-
able climatic conditions with fluctuations in paleoprecipita-
tion indicating a change from a humid ever-wet climate to
a drier, more seasonal climate (Hembree & Carnes, 2018;
Hembree & McFadden, 2020). A decrease in precipitation
and drier seasons resulted in a shift away from plant com-
munities dominated by floodplain wetland flora assemblages
with poorly rooted groundcover (Hembree, 2022). Abundant
and pervasive rhizoliths of the Middle-to-early Late Penn-
sylvanian indicate widespread surface vegetation of a high
moisture environment, whereas uncommon and clumped rhi-
zoliths of the Late Pennsylvanian to Early Permian indicate
sparse surface vegetation of a seasonally dry environment
(Hembree, 2022).
2.2 Allegheny, Conemaugh, Monongahela,
and Dunkard groups
This study involved the investigation of fluvial facies asso-
ciations in the Middle Pennsylvanian (Moscovian)-to-early
Permian (Asselian) Allegheny, Conemaugh, Monongahela,
and Dunkard groups in southeast Ohio and northwest West
Virginia (Fig. 1A). The Middle Pennsylvanian Allegheny
Group crops out in Ohio and Pennsylvania, is up to 300 m
thick, and consists of shales, limestones, and sandstones as
well as six major coal zones (Ferm, 1970; Sturgeon & Mer-
rill, 1949; Williams, 1960). During the deposition of the
Allegheny, a narrow marine embayment existed in the region
and experienced periodic flooding during times of sea-level
rise (Fig. 1B) (Martin, 1998). The Late Pennsylvanian Cone-
maugh Group of Ohio and Pennsylvania is between 325 and
375 m thick and consists of fluvial and marine lithofacies,
including shales, sandstones, limestones, and some coals
divided into two formations, the Glenshaw and Casselman
(Condit, 1912; Hembree & Nadon, 2011; Sturgeon, 1958).
The upper Allegheny and lower Conemaugh groups were
deposited by a massive northwestwardly prograding flu-
vial–deltaic system (DiMichele et al., 1996).
Outcrops of the Monongahela and Dunkard groups can be
found in Ohio, Pennsylvania, and West Virginia (Hembree &
McFadden, 2020; Martin, 1998). Both groups were deposited
close to the equator within the central to northern portions of
the Appalachian Basin and consist of fluvial and lacustrine
sediments (Fig. 1C) (Hembree & McFadden, 2020). The Mon-
ongahela Group is Late Pennsylvanian in age, 80–125 m thick,
and is regionally divided into the Pittsburgh and Uniontown
formations (Hembree & McFadden, 2020; Martin, 1998; Stur-
geon, 1958). These formations are composed of shales, mud-
stones, sandstones, and limestones that were deposited in a
river-dominated deltaic system during a regression (Hembree
& McFadden, 2020; Martin, 1998). The Dunkard Group is
Early Permian, up to 360 m thick, and regionally divided into
the Waynesburg, Washington, and Greene formations which
Climate‑induced changes in fluvial ichnofossil assemblages of the Pennsylvanian–Permian…
Fig. 1  Stratigraphy and paleogeography of the study area. A Gen-
eral stratigraphic column showing the geologic age of the Allegheny,
Conemaugh, Monongahela, and Dunkard groups. B, C Paleogeo-
graphic reconstructions of North America during the Middle Pennsyl-
are mainly composed of shale, mudstone, and sandstone and
minor coal and limestone (Hembree & Carnes, 2018; Hembree
& McFadden, 2020; Lucas, 2013; Martin, 1998; Schneider
et al., 2013). Dunkard Group sediments were derived from
the weathering of the Allegheny Mountains and deposited on
a broad fluvial plain (Martin, 1998).
3 Materials and methods
The study area consisted of five roadside outcrops in south-
east Ohio and northwest West Virginia. Field sites were
located in Athens County, Ohio and Wood County, West
263
vanian (B) and early Permian (C). The location of the study areas is
indicated by the white star. (Images by R. Blakey. Used with permis-
sion)
Virginia, USA (Fig. 2). The studied sections included por-
tions of the Middle Pennsylvanian Allegheny Group, Late
Pennsylvanian Conemaugh and Monongahela groups, and
the early Permian Dunkard Group. At each locality one to
two, 3–8 m-thick, fluvial facies associations comprised of
sandstones, shales, mudstones, siltstones, and coal were
studied. General stratigraphic sections of the facies associa-
tions were constructed to document major lithologies, sedi-
mentary structures, and ichnofossil occurrences.
A 0.5 × 0.5-m grid was used to assess the ichnofossil
assemblages along exposed bedding planes of point bar
sandstones at the top of each fluvial facies association. For
each facies association, eight grids were placed on bedding
13
264
Fig. 2  A Location of the study area within Ohio and West Virginia
(within white square) relative to the Appalachian and Dunkard basins.
The inset map shows the location of the study region within the
plane surfaces with at least 0.5 m of exposure, spaced up
to 45 m apart. Ichnofossil data collected within the grids
included abundance, density, diversity, and composition.
Abundance was determined by counting the number of indi-
vidual ichnofossils identified within the grid. Density was
assessed by recording the concentration of ichnofossils in
the grid and the degree of sediment reworking using ich-
nofabric indices (ranked 0–6) (Buatois & Mángano, 2011).
Diversity was assessed by counting the number of unique
ichnogenera. Composition was assessed by (1) evaluating
the mean ichnofossil complexity by counting segments,
branches, chambers, and openings to the surface of each
ichnofossil (Hembree, 2018); (2) measuring the widths of
each ichnofossil to determine a mean value; and (3) evalu-
ating the number of different behaviors represented within
the grid based on ichnofossil morphology (Bromley, 1996;
Buatois & Mángano, 2011).
Sandstone samples were collected for thin-section prepa-
ration from each studied grid at the five field sites. Thin
sections were vertically oriented to assess the vertical pen-
etration of the substrate by ichnofossils observed on bedding
planes, total levels of bioturbation, and to identify additional
ichnofossils in cross section that were not visible in the field.
A total of 43 thin sections (35 1.5″ × 3.0″, 8 1.0″ × 2.0″
slides) were prepared by Texas Petrographic (Houston, TX,
USA). Thin sections were examined using a Motic BA300
polarizing petrographic microscope. Mineralogy, grain size,
grain sorting and rounding, sedimentary structures, and ich-
nofossil content (i.e., burrows and rhizoliths) were described
for each thin section.
The quantitative ichnofossil data from each grid and bed-
ding plane were combined from each outcrop to compare
the ichnofossil assemblages between each unit. An analysis
of variance (ANOVA) was used to compare the different
populations of abundance, density, diversity, and composi-
tion data between each of the five localities to test whether
13
J. K. Crowell, D. I. Hembree
United States (at star). B Map showing the study sites (Sites 1–5) and
the location of the stratigraphic sections
differences between the various quantitative properties of
the ichnofossil assemblages existed. A Dunn’s post hoc
comparison was used to indicate where these differences
occurred stratigraphically.
4 Results
4.1 Site descriptions
4.1.1 Allegheny Group—Site 1
The section at Site 1 consisted of (base to top) (1) carbo-
naceous, silty shale (240 cm thick), (2) green-gray massive
claystone (30 cm thick) with root traces, (3) bituminous coal
(60 cm thick) with coarse plant fragments, (4) green-gray
micaceous shale (15 cm), and (5) medium- to coarse-grained
(coarsening upward), massive, micaceous sandstone (wacke)
(80 cm thick) (Figs. 3A, 4A).
4.1.2 Conemaugh Group—Site 2
The section at Site 2 consisted of (base to top) (1) gray,
fine-grained, micaceous sandstone (wacke) (115 cm thick),
(2) gray, medium-grained, micaceous sandstone (wacke)
(90 cm thick), and (3) fine- to medium-grained (coarsen-
ing upward), finely laminated to cross-laminated sandstone
(arenite) (150 cm thick) with iron concretions occurring near
the base (Figs. 3B, 4B).
4.1.3 Monongahela Group—Sites 3A, 3B
The section at Site 3A consisted of (base to top) (1) green-
gray, blocky claystone (30 cm thick), (2) red blocky mud-
stone (40 cm thick) with cutans and mottling, (3) green-gray
mudstone (50 cm thick) with cutans and mottling, (4) dark
Climate‑induced changes in fluvial ichnofossil assemblages of the Pennsylvanian–Permian…
Fig. 3  Stratigraphic sections from Sites 1–5. A Site 1 Allegheny
Group. B Site 2 Conemaugh Group. C Site 3A Monongahela Group
(upper). D Site 3B Monongahela Group (lower). E Site 4A Lower
gray shale (30 cm thick) with carbonized plant fragments,
(5) green-gray blocky mudstone (90 cm thick) with cutans,
slickensides, mottling, and iron nodules, (6) green-gray,
platy mudstone (80 cm thick), (7) green-gray, laminated to
cross-laminated shale (90 cm thick), and (8) four fining-
upward successions (40, 70, 70, 80 cm thick) of fine-grained,
cross-laminated to laminated, sandstone (wacke) (Figs. 3C).
265
Dunkard Group (east). F Site 4B Lower Dunkard Group (west). G
Site 5 Upper Dunkard Group
The section at Site 3B consisted of (base to top) (1)
red, silty mudstone (60 cm thick) with rhizoliths and mot-
tling, (2) green-gray, calcareous, silty mudstone (70 cm
thick), (3) red, blocky, silty mudstone (130 cm thick) with
rhizoliths and cutans, (4) variegated shale (180 cm thick),
and (5) fine- to medium-grained (coarsening upward),
cross-laminated, micaceous sandstone (108 cm thick)
13
266
Fig. 4  Outcrop images and grid examples. A Allegheny Group (Site
1). B Conemaugh Group (Site 2). C Monongahela Group (Site 3b).
D Lower Dunkard Group (Site 4a). E Upper Dunkard Group (Site 5).
F Point bar sandstone surface with grid from Allegheny Group (Site
1). G Point bar sandstone surface with grid from Conemaugh Group
with interbedded sandstone, siltstone, and mudstone lenses
(Figs. 3D,4C).
4.1.4 Lower Dunkard Group—Sites 4A, 4B
The section at Site 4A consisted of (base to top) (1) cross-
bedded, fine-grained sandstone (40 cm thick), (2) gray,
13
J. K. Crowell, D. I. Hembree
(Site 2). H Point bar sandstone surface with grid from Monongahela
Group (Site 3b). I Point bar sandstone surface with grid from Lower
Dunkard Group (Site 4a). J Point bar sandstone surface with grid
from Upper Dunkard Group (Site 5)
micaceous sandy shale (60 cm thick), (3) green-gray platy
shale (65 cm thick), (4) fine-grained massive sandstone
(20 cm thick), (5) argillaceous shale (90 cm thick), (6) green,
blocky argillaceous mudstone (20 cm thick), (7) red argil-
laceous mudstone (140 cm thick) with rhizoliths and cutans,
(8) green-gray shale (90 cm thick), and (9) fine-grained
sandstone (150 cm thick) (Figs. 3E,4D).
Climate‑induced changes in fluvial ichnofossil assemblages of the Pennsylvanian–Permian…
The section at Site 4B consisted of (base to top) (1) tabu-
lar, cross-bedded, fine-grained, micaceous sandstone (70 cm
thick), (2) green-gray sandy shale (60 cm thick), (3) green-
gray shale (60 cm thick), (4) light green-gray shale (10 cm
thick), (5) red blocky mudstone (130 cm thick), (6) gray
shale (175 cm thick), and (7) fine-grained sandstone (160 cm
thick) (Figs. 3F).
4.1.5 Upper Dunkard Group—Site 5
The section at Site 5 consisted of (base to top) (1) medium-
grained sandstone (70 cm thick) with trough crossbedding
at the base and ripple cross lamination at the top, (2) thin
sandstone (20 cm thick) with interbedded siltstone and mud-
stone and iron nodules, medium-grained partially laminated
sandstone (35 cm thick), (3) reddish-brown, blocky, fining
upward, sandy shale (110 cm thick) with iron nodules,
(4) green-gray argillaceous mudstone (30 cm thick), (5)
red argillaceous mudstone (180 cm thick) with rhizoliths,
(6) gray argillaceous shale (40 cm thick), and (7) fine- to
medium-grained, bioturbated sandstone (180 cm thick)
(Figs. 3G,4E).
4.2 Sandstone petrography
4.2.1 Allegheny Group—Site 1
Sandstones (n = 8) were medium–fine-grained, moderately
sorted, and composed of abundant angular quartz, common
micas, uncommon plagioclase feldspar and rock fragments,
and a fine matrix (quartz-wacke) (Fig. 5A).
4.2.2 Conemaugh Group—Site 2
Sandstones (n = 7) were fine-grained, well-sorted, and com-
posed of abundant angular quartz, uncommon mica, rock
fragments, and plagioclase feldspar, a fine matrix (quartz-
wacke), and iron cement (Fig. 5B).
4.2.3 Monongahela Group—Site 3
Sandstones (n = 14) were very-fine-grained, well-sorted, and
composed of abundant angular quartz, common mica and
rock fragments, uncommon iron-bearing minerals, rare pla-
gioclase feldspar, and a fine matrix (quartz-wacke) (Fig. 5C).
Cross lamination was observed in all samples. Bioturbation
was noted in several thin sections, including mixing of sedi-
ment grains and disturbance of primary sedimentary struc-
tures (Fig. 6A-D).
267
4.2.4 Lower Dunkard Group—Site 4
Sandstones (n = 11) were medium–fine-grained to very-fine-
grained, moderately sorted to well sorted, and composed
of angular quartz, micas, iron-bearing minerals, rock frag-
ments, rare plagioclase feldspar, and a green matrix (quartz-
wacke) (Fig. 5). Lamination and bioturbation occurred in
several samples (Fig. 6E-H). Bioturbation included general
mixing of sediment grains as well as distinct burrows.
4.2.5 Upper Dunkard Group—Site 5
Sandstones (n = 3) were extremely fine-grained to
medium–fine-grained, well sorted to moderately sorted,
and composed of abundant angular quartz, common mica,
rock fragments, and iron-bearing minerals, and a fine matrix
(quartz-wacke) (Fig. 5). Lamination and bioturbation were
uncommon.
4.3 Ichnofossils of the Allegheny, Conemaugh,
Monongahela, and Dunkard groups
Ichnofossils were described from sandstones at the top of
each stratigraphic sequence. Ichnofossils with distinct mor-
phologies were identified to ichnogenus level, as preserva-
tion prohibited confident identification to the ichnospecies
level. General bioturbation of some sites resulted in a rec-
ognizable ichnofabric, however, it was not possible to assign
these to specific ichnotaxa. Occurrence is described as rare
(1–5), common (5–10), and abundant (> 10).
4.3.1 
Skolithos
These were single, circular, horizontal cross sections of
vertical shafts found on sandstone bedding plane surfaces
(Fig. 7A, B). Shaft diameter ranged from 0.3 to 1.5 cm.
The shafts were unlined and had a fill similar to the matrix.
These burrows were attributed to Skolithos, which is diag-
nosed as a subcylindrical, unbranched, unlined-to-lined,
straight-to-curved structureless tube, perpendicular to the
bedding plane surface (Häntzschel, 1975). The burrows were
similar to Cylindrichum, but rounded terminations were not
observed, and Monocraterion, but lacked a funnel-shaped
opening (Häntzschel, 1975). Skolithos were rare in the Cone-
maugh, common in the Monongahela, and abundant in the
Upper and Lower Dunkard groups (Fig. 7C–D).
4.3.2 
Arenicolites
These were paired, circular, horizontal cross sec-
tions of vertical, unlined shafts with a fill similar to the
matrix occurring on sandstone bedding plane surfaces
(Fig. 7E–F). Individual shaft cross sections were similar
13
268 J. K. Crowell, D. I. Hembree

Fig. 5  Thin sections of point

bar sandstones. A Allegheny
Group (Site 1) sandstone con-
taining angular medium-grained
quartz, plagioclase feldspar, and
rock fragments. B Conemaugh
Group (Site 2) sandstone with
angular, fine-grained quartz,
mica, rock fragments, plagio-
clase feldspar, and iron cement.
C Monongahela Group (Site 3)
sandstone containing very-
fine-grained quartz, mica, rock
fragments, and a fine matrix. D
Lower Dunkard Group (Site 4)
sandstone containing very-fine-
grained quartz, mica, rock frag-
ments, and a fine green matrix.
E Upper Dunkard Group (Site
5) sandstone containing fine- to
medium-grained angular quartz,
mica, and rock fragments with a
fine matrix

in size and appearance to those assigned to Skolithos, 4.3.3 

however, the regular proximity (< 1 cm) of the two shafts
suggests they were likely connected in the subsurface. In
addition, samples from the Monongahela Group show the
presence of the shafts on both the top and bottom surfaces
of the sample. Shafts were unlined and contained a fill
like that of the surrounding matrix. These burrows were
attributed to Arenicolites which is diagnosed as a simple,
vertical, cylindrical U-shaped tube with smooth sides,
found perpendicular to the bedding plane (Häntzschel,
1975). Tube openings can be flared or funnel-shaped
(Häntzschel, 1975). These burrows are also similar to Dip-
locraterion but do not possess spreite between the two
shafts (Häntzschel, 1975). Arenicolites were rare in the
Monongahela and Dunkard groups.
Cruziana
These were straight, horizontal, bilobed trails, 1.0–1.5 cm
wide and 10–20 cm in length, with central furrows (Fig. 7G).
Surficial features (i.e., bioglyphs) could not be assessed due
to poor preservation. These trails were attributed to Cruzi-
ana which is diagnosed as a trail with elongate, bilobed fur-
rows covered with herringbone-shaped, longitudinal stria-
tions (Häntzschel, 1975). While the striations were missing
from the observed specimens, the bilobed furrows are char-
acteristic of Cruziana. Sizes and appearance of these trails
vary greatly due to the behavior of the trace makers and
substrate consistency (Häntzschel, 1975). Cruziana were
typically found in assemblages with Skolithos and were rare
in the Dunkard Group.

13
Climate‑induced changes in fluvial ichnofossil assemblages of the Pennsylvanian–Permian… 269

Fig. 6  Thin sections from point

bar sandstones showing primary
sedimentary structures and
bioturbation. A Lamination and
general bioturbation (sediment
mixing) from the Monongahela
Group (Site 3B). B Cross lami-
nation and general bioturbation
(sediment mixing) from the
Monongahela Group (Site 3B).
C Root traces and sediment
mixing from the Monongahela
Group (Site 3B). D Disrupted
and deflected lamination from
the Monongahela Group (Site
3B). E General bioturbation
(sediment mixing) of the Lower
Dunkard Group (Site 4A).
F Burrows (Skolithos) and
disrupted clay laminae from the
Lower Dunkard Group (Site
4B). G Passively filled burrows
along the upper sandstone sur-
face from the Lower Dunkard
Group (Stie 4B). H General
bioturbation (sediment mixing
and disrupted laminae) from the
Lower Dunkard Group (Site 4B)

4.3.4 
Planolites
These were simple, unlined, and actively filled horizontal
burrows (Fig. 8A, B). The burrows were generally straight,
less than 1 cm in width, and did not branch. The burrow
fill was distinct from the matrix and was raised above the
bedding plane. The tunnel surfaces were generally smooth
with no visible scratch marks or internal structure. These
burrows were attribute to Planolites which is diagnosed as
an unlined, subcylindrical to cylindrical, straight to curved,
unbranched but often overlapping, actively filled tunnel
(Häntzschel, 1975). They are also similar to Palaeophy-
cus, but lack a lining and do not branch (Häntzschel, 1975).
Planolites were common in the Lower Dunkard Group and
rare in the Upper Dunkard Group and found in assemblages
with Skolithos and Scoyenia.
4.3.5 
Scoyenia
These were horizontal, straight to curved tunnels with a
wrinkly surface texture (Fig. 8). The tunnels were non-
branching but overlapping, 1 cm wide, and no longer than
5 cm. The tunnels were raised above the bedding plane
surface with a fill darker than the matrix. The tunnel fill
consisted of irregular meniscate backfilling. Scratch marks
were observed along the sides of the burrows but were not
well preserved. These burrows were attributed to Scoyenia
which is diagnosed as a slender, straight to curved tunnel
with rope-like sculpture produced by paired striations that
cover the tunnel surface and active backfill (Häntzschel,
1975). The tunnels commonly cross each other but do not
branch (Häntzschel, 1975). These burrows are also similar to
Taenidium and Beaconites but lack the well-defined menis-
cate backfill of both and the lining of Beaconites, in addition
to exhibiting striations on the tunnel surfaces which is not
present in either ichnogenus (Häntzschel, 1975). Scoyenia
were found in assemblages with Skolithos and were common
in the Lower Dunkard and Upper Dunkard groups.
4.3.6 
Treptichnus
These were a series of short, horizontal, oval-shaped, con-
nected tunnel segments (Fig. 9). The segments were typi-
cally connected in a linear pattern, although some were
slightly curved. Segments within a series were usually
equal in size, however, some segments were elongated. The

13
270 J. K. Crowell, D. I. Hembree

Fig. 7  Ichnofossils. A, B Bed-

ding plane views of Skolithos
from the Monongahela Group
(Site 3). C, D Bedding plane
views of Arenicolites from the
Monongahela Group (Site 3). E,
F Bedding plane views of dense
concentrations of Skolithos and
Arenicolites from the Lower
Dunkard Group (Site 4). G Bed-
ding plane view of Cruziana
from the Upper Dunkard Group
(Site 5)

13
Climate‑induced changes in fluvial ichnofossil assemblages of the Pennsylvanian–Permian…
Fig. 8  Ichnofossils. A Bed-
ding plane view of Planolites
from the Upper Dunkard Group
(Site 5). B Bedding plane view
of Planolites from the Lower
Dunkard Group (Site 4). C
Bedding plane view of Scoyenia
from the Upper Dunkard Group
(Site 5). D Bedding plane view
of Scoyenia from the Lower
Dunkard Group (Site 4)
average segment length was 0.3–1.0 cm. Most commonly
each series consisted of three segments. Some bioglyphs,
including scratch marks, were present along the base of
some segments. Burrow fill was similar to the matrix. These
burrows were attributed to Treptichnus which is diagnosed
as a series of shallow, subhorizontal, J- or U-shaped tun-
nel segments joined together in irregular patterns near the
tunnel ends (Häntzschel, 1975). They have low slopes and
a passive fill (Häntzschel, 1975). While individual tunnel
segments were similar to the basal expression of Arenicolites
or Diplocraterion, their occurrence in repeated series make
these diagnoses unlikely (Häntzschel, 1975). Treptichnus
were abundant in the Upper Dunkard Group.
4.3.7 Analysis of ichnofossil assemblages
No ichnofossils were observed in point bar sandstones of the
Middle Pennsylvanian Allegheny Group. Ichnofossil assem-
blages in point bar sandstones of the Conemaugh Group
were composed of Skolithos and had a mean abundance of
3.6, diversity of 1, density of 1, complexity of 1, mean width
of 5.2 mm, and an ichnofabric index of 1 (low bioturbation)
(Table 1). The mean values from the Conemaugh were the
lowest in the study interval. Ichnofossil assemblages of the
Monongahela Group also consisted mostly of Skolithos, but
also included rare Arenicolites. The mean abundance was
11.6, diversity was 1.2, density was 1, complexity was 1,
mean width was 6.3 mm, and ichnofabric index was 1.5 (low
271
bioturbation) (Table 1). Ichnofossil assemblages of the Lower
Dunkard Group were dominated by Skolithos but also included
Arenicolites, Cruziana, Scoyenia, and Planolites. The mean
abundance was 188, diversity was 2.8, density was 3.6, com-
plexity 1.1, mean width was 5.3 mm, and ichnofabric index
was 4 (moderate to high bioturbation) (Table 1). Most quanti-
tative assemblage properties increased from the Monongahela
Group, except for mean width. Ichnofossil assemblages of the
Upper Dunkard Group included abundant Treptichnus as well
as Skolithos, Arenicolites, Cruziana, Scoyenia, and Planolites.
The mean abundance was 73, diversity was 2.4, density was
1.9, complexity was 2, mean width was 5.7 mm, and ichnofab-
ric index was 2 (moderate bioturbation) (Table 1). Abundance,
density, and ichnofabric index of the Upper Dunkard decreased
relative to the Lower Dunkard, while the diversity, complexity,
and width increased.
Quantitative data was combined from the bedding planes
of each outcrop to compare assemblages between the Alle-
gheny, Conemaugh, Monongahela, and Dunkard groups. An
analysis of variance (ANOVA) was performed to compare the
different populations of abundance, density, diversity, com-
plexity, and mean width data between each of the five groups.
P values for abundance, diversity, density, and complexity
were all less than 0.05 indicating that significant differences
existed between these aspects of the ichnofossil assemblages
(Table 2). Dunn’s post hoc comparison showed that the assem-
blages of the Allegheny and Conemaugh groups were distinct
from those of the Monongahela and Dunkard groups (Table 2).
13
272
Fig. 9  Specimens of Treptichnus from the underside of sandstone
blocks of the Upper Dunkard Group (Site 5). A, B Cluster of long-to-
short, linked segments of Treptichus. C Close up of a single specimen
of Treptichnus consisting of at least three, linked tunnel segments
5 Discussion
5.1 Unit interpretations
5.1.1 Allegheny Group—Site 1
The section at Site 1 is interpreted as a transition from flood-
plain pond (Unit 1), proximal floodplain and mire (Units 2,
3), levee (Unit 4), to point bar (Unit 5) (Collinson, 1996;
Brierly et al., 1997; Aslan & Autin, 1998; Stow, 2005;
Miall, 2010). The features of the paleosol (Unit 2) indicate
poor development and drainage as well as water saturation
13
J. K. Crowell, D. I. Hembree
(Retallack et al., 2001). The upper sandstone coarsens
upwards indicating a change to a higher energy condition
(Miall, 2010).
5.1.2 Conemaugh Group—Site 2
Units 1–3 of Site 2 are interpreted as a series of stacked
point bar and channel deposits (Collinson, 1996; Miall,
2010). Units 1 and 2 coarsen upward and appear massive,
suggesting an upward increase in energy from the levee to
point bar (Collinson, 1996; Miall, 2010). Unit 3 contains a
succession of fine-grained sandstones with multiple, coarser-
grained, erosively based channel forms suggesting rapid lat-
eral movement of the channel (Miall, 2010).
5.1.3 Monongahela Group—Sites 3A, B
The section at Site 3A is interpreted as a transition from a
distal to proximal floodplain (Units 1–3, 5), floodplain pond
(Unit 4), levee (Units 6–7), to point bar (Units 8–11) (Collin-
son, 1996; Brierly et al., 1997; Aslan & Autin, 1998; Tabor
& Montañez, 2004; Stow, 2005; Miall, 2010). The prop-
erties of the paleosols indicate shifts from poorly drained
(Units 1, 3), well-drained (Unit 2), to moderately drained
(Unit 5) conditions, likely related to changes in proximity
to the active channel (Collinson, 1996; Tabor & Montañez,
2004). The properties of the succession of upper sandstones
(Units 8–11) indicate changes in energy from high to low as
the point bars migrated (Miall, 2010).
The section at Site 3B is interpreted as a transition from
moderately drained, distal to proximal floodplain (Units
1–3), levee (Unit 4), to point bar (Unit 5) (Collinson, 1996;
Brierly et al., 1997; Retallack et al., 2001; Tabor & Mon-
tañez, 2004; Stow, 2005; Miall, 2010). The interbedded
sands, silts, and clays of Unit 5 indicate fluctuating energy
conditions from high to low during periods of high and low
discharge, respectively (Collinson, 1996; Miall, 2010).
5.1.4 Lower Dunkard Group—Sites 4A, B
The section at Site 4A is interpreted as a transition from
point bar (Unit 1), levee (Units 2–3), point bar (Unit 4),
levee (Unit 5), proximal to distal floodplain (Units 6–7),
levee (Unit 8), to point bar (Unit 9) (Collinson, 1996; Bri-
erly et al., 1997; Aslan & Autin, 1998; Tabor & Montañez,
2004; Stow, 2005; Miall, 2010).
The section at Site 4B is interpreted as a transition from
point bar (Unit 1), levee (Units 2–4), distal floodplain (Unit
5), levee (Unit 6), to point bar (Unit 7) (Collinson, 1996; Bri-
erly et al., 1997; Retallack et al., 2001; Tabor & Montañez,
2004; Stow, 2005; Miall, 2010).
Climate‑induced changes in fluvial ichnofossil assemblages of the Pennsylvanian–Permian… 273

Table 1  Quantitative properties of ichnofossil assemblages from Sites 2–5

Location Grid Abundance Diversity Density Complexity MW: Sk MW: Ar MW: Pl MW: Sc MW: Cr MW: Tr Behavior

Site 2 1
2 2 1 1 1 10 D, F
3 3 1 1 1 4 D, F
4 2 1 1 1 3 D, F
5
6 3 1 1 1 5 D, F
7
8 3 1 1 1 4 D, F
Site 3A 1 8 1 1 1 10 D, F
2 7 2 1 1 10 10 D, F
3 15 1 1 1 5 D, F
4 3 1 1 1 5 D, F
5 11 1 1 1 10 D, F
6 3 1 1 1 10 D, F
7 7 1 1 1 5 D, F
8 23 2 1 1 5 D, F
Site 3B 9 18 2 1 1 5 5 D, F
10 28 1 1 1 5 D, F
11 6 1 1 1 3 D, F
12 6 1 1 1 4 D, F
13 9 1 1 1 5 D, F
14 12 1 1 1 4 D, F
15 14 1 1 1 7 D, F
16 15 1 1 1 5 D, F
Site 4A 1 60 4 2 1 10 10 5 3 F
2 110 4 3 1 3 3 20 5 F
3 135 2 3 1 5 5 F
4 200 2 3 1 3 3 F
5 200 3 4 1 3 3 4 F
6 400 3 5 1 6 6 3 F
7 120 3 3 1 5 5 5 F
8 140 3 3 1 4 4 5 F
Site 4B 9 120 2 3 1 5 5 F
10 200 3 3 1 5 5 3 F
11 100 4 3 2 5 5 4 10 F, L
12 300 2 5 1 3 3 F
13 200 2 5 1 5 5 F
14 75 2 2 1 5 5 F
15 300 2 6 1 5 5 F
16 350 3 5 1 3 3 6 F
Site 5 1 150 1 2 3 3 F
2 130 1 2 2 3 F
3 110 2 2 4 4 3 D, F
4 75 2 2 3 2 4 F
5 30 3 1 1 15 5 10 F, L
6 20 3 1 1 10 5 3 F, L
7 50 4 4 1 3 3 10 5 F, L
8 15 3 1 1 10 5 5 F

MW mean width (measured in mm), Sk Skolithos, Ar Arenicolites, Pl Planolites, Cr Cruziana, Sc Scoyenia, Tr Treptichnus. In the Behavior col-
umn, D dwelling, F feeding, L locomotion

13
274 J. K. Crowell, D. I. Hembree

Table 2  ANOVA and Dunn’s post hoc results comparing abundance, diversity, density, complexity, and mean width values from the Cone-
maugh, Monongahela, and Dunkard group ichnofossil assemblages
ANOVA
Property p value

Abundance 9.917 x 10E-8

Diversity 0.0134
Density 1.246 x 10E-9
Complexity 0.0024
Width 0.2354
Dunn's Post Hoc

Abundance Complexity
Site 2 Site 3A Site 3B Site 4A Site 4B Site 5 Site 2 Site 3A Site 3B Site 4A Site 4B Site 5
Site 2 0.25 0.12 1.99E-05 7.34E-06 1.65E-03 Site 2 0.77 0.77 0.77 0.83 0.11
Site 3A 0.25 0.63 3.65E-04 1.38E-04 0.02 Site 3A 0.77 1 1 0.93 0.03
Site 3B 0.12 0.63 2.02E-03 8.51E-04 0.07 Site 3B 0.77 1 1 0.93 0.03
Site 4A 1.99E-05 3.65E-04 2.02E-03 0.8 0.2 Site 4A 0.77 1 1 0.93 0.03
Site 4B 7.34E-06 1.38E-04 8.51E-04 0.8 0.13 Site 4B 0.83 0.93 0.93 0.93 0.04
Site 5 1.65E-03 0.02 0.07 0.2 0.13 Site 5 0.11 0.03 0.03 0.03 0.04
Diversity Width
Site 2 Site 3A Site 3B Site 4A Site 4B Site 5 Site 2 Site 3A Site 3B Site 4A Site 4B Site 5
Site 2 0.76 0.84 0.04 0.33 0.03 Site 2 0.54 0.42 0.24 0.31 0.29
Site 3A 0.76 0.56 0.04 0.45 0.03 Site 3A 0.54 0.09 0.02 0.03 0.03
Site 3B 0.84 0.56 0.01 0.18 0.01 Site 3B 0.42 0.09 0.76 0.89 0.84
Site 4A 0.04 0.04 0.01 0.21 0.92 Site 4A 0.24 0.02 0.76 0.82 0.9
Site 4B 0.33 0.45 0.18 0.21 0.17 Site 4B 0.31 0.03 0.89 0.82 0.93
Site 5 0.03 0.03 0.01 0.92 0.17 Site 5 0.29 0.03 0.84 0.9 0.93
Density
Site 2 Site 3A Site 3B Site 4A Site 4B Site 5
Site 2 0.9 0.9 1.98E-03 5.12E-04 0.18
Site 3A 0.9 1 2.40E-04 4.02E-05 0.1
Site 3B 0.9 1 2.40E-04 4.02E-05 0.1
Site 4A 1.98E-03 2.40E-04 2.40E-04 0.66 0.04
Site 4B 5.12E-04 4.02E-05 4.02E-05 0.66 0.01
Site 5 0.18 0.1 0.1 0.04 0.01

Bold indicates significance

5.1.5 Upper Dunkard Group—Site 5 2004, 2011). In continental environments, Skolithos are

The section at Site 5 is interpreted as a transition from point
bar (Units 1–3), levee (Unit 4), proximal to distal floodplain
(Units 5–6), levee (Unit 7), to point bar (Unit 8) (Collinson,
1996; Brierly et al., 1997; Aslan & Autin, 1998; Tabor &
Montañez, 2004; Stow, 2005; Miall, 2010).
5.2 Ichnofossil interpretation
5.2.1 
Skolithos
Given the depositional setting, Skolithos most likely rep-
resents dwelling or deposit feeding (Buatois & Mángano,
usually associated with active channels and sandbar depos-
its (Buatois & Mángano, 2004, 2011; Hembree & Blair,
2016). Fluvial channels are high-energy environments
with rapid fluctuations in sedimentation and erosion rates,
resulting in very stressful and unstable conditions (Buat-
ois & Mángano, 2004, 2011). As a result, Skolithos typi-
cally occur in low diversity assemblages and tracemakers
were likely opportunistic generalists that rapidly colonized
exposed surfaces (Buatois & Mángano, 2004; Miller &
Collinson, 1994). Potential trace makers include various
arthropods, such as spiders, beetles, and numerous insect
larvae, however, the unstable nature of the environment

13
Climate‑induced changes in fluvial ichnofossil assemblages of the Pennsylvanian–Permian…
would be unsuitable for the permanent dwellings of spi-
ders (Hembree, 2017; Hils & Hembree, 2015).
5.2.2 
Arenicolites
In this study area Arenicolites likely represent dwelling or
deposit feeding activities since point bar deposits are suba-
erial environments (Hasiotis, 2002; Rindsberg & Kopaska-
Merkel, 2005). Potential producers of Arenicolites include
myriapods (millipedes and centipedes) and insect larvae
such as those of chironamids, mayflies, and beetles (Hasio-
tis, 2002; Hembree, 2009, 2019; Mikuś & Uchman, 2013;
Rindsberg & Kopaska-Merkel, 2005; Thacker & Hembree,
2021).
5.2.3 Cruziana
Behaviors likely associated with Cruziana include loco-
motion and deposit feeding (Buatois & Mángano, 2004;
Häntzschel, 1975; Zonneveld et al., 2002). Cruziana have
been previously described from fluvial settings, specifically
in desiccated overbank deposits (Buatois & Mángano, 2011).
Scratch marks associated with Cruziana are attributed to
active digging in a firm substrate (Buatois & Mángano,
2004, 2011; Zonneveld et al., 2002). Potential tracemak-
ers are isopods or beetles given the age and riparian setting
(Lavelle & Spain, 2005; Lawrence & Newton, 1982).
5.2.4 
Planolites
Activities associated with Planolites include mobile-deposit
feeding and are suggestive of sediment with high organic
content (Häntzschel, 1975; Hasiotis, 2002). Many differ-
ent invertebrate tracemakers may produce Planolites, but in
continental settings are most typically attributed to annelids
and insects (Buatois & Mángano, 2004; Hasiotis, 2002; Kale
et al., 1997).
5.2.5 Scoyenia
Deposit feeding and dwelling are behaviors associated with
Scoyenia (Häntzschel, 1975). Producers of Scoyenia are
indicative of moist to wet, compact, silty clay substrates
and are typically associated with point bar, levee, and flood-
plain deposits (Buatois & Mángano, 2011; Frey et al., 1984;
Hasiotis, 2004; Hasiotis & Dubiel, 1993). Scratch marks on
Scoyenia are indicative of an organism with appendages that
gripped the burrow walls including larvae of beetles or other
insects (Hasiotis, 2004).
275
5.2.6 
Treptichnus
Treptichnus is associated with a variety of feeding activities
including detritus and deposit feeding and predatory activity
(Getty et al., 2016). In the Upper Dunkard Group, deposit
feeding is the most probable given their depth of occur-
rence and abundance. Treptichnus is typically associated
with relatively diverse ichnofossil assemblages (Buatois &
Mángano, 2011). Treptichnus is widely distributed and has
a broad stratigraphic range, with an equally broad range of
potential tracemakers (Getty et al., 2016). Continental Trep-
tichnus are usually considered to be made by insect larvae
(Getty et al., 2016).
5.3 Changes in ichnofossil assemblages
from the Allegheny to Dunkard groups
Ichnofossils can be used to reconstruct paleoenvironmen-
tal conditions at the time of trace production because they
represent direct interactions of an organism with their envi-
ronment. Environmental conditions that can result in differ-
ent organism responses in continental settings may include
temperature, precipitation, nutrient levels, water-table depth,
substrate moisture, and environmental stability (Hembree,
2018). Changes in these conditions may be reflected in ich-
nofossils abundance, density, diversity, and composition.
5.3.1 Allegheny Group
There are several explanations for the absence of ichnofos-
sils in the Allegheny Group. Destruction of traces can occur
during high-energy events that mobilize the upper layer of
sediment (Gingras et al., 2007), high sedimentation rate
may limit time for the colonization of the surface (Gingras
et al., 2007), or harsh environmental conditions within the
substrate may inhibit its colonization (Beatty et al., 2008;
Morrissey & Braddy, 2004). None of these scenarios are
supported by the sedimentological data from Site 1. Instead,
the absence of ichnofossils in the point bar sandstones of
the Allegheny Group may have been related to the favorable
conditions above the substrate. The Allegheny Group was
deposited under an ever-wet climate (Garcés et al., 1997;
Hembree & Nadon, 2011; Milici, 2005) defined by high
annual rainfall, high humidity, and available surface water
throughout the year (Hasiotis et al., 2007; Underwood et al.,
2014). High soil moisture and water-table levels of ever-
wet climates result in low ichnodiversity because epigeal
and arboreal organism lifestyles are more dominant due to
water-saturated substrate conditions (Hasiotis et al., 2007).
The abundance of land plants during this time would have
contributed to a significant amount of detritus in continental
settings (DiMichele, 2014). Detritus is a common source
of energy and nutrients for soil faunal communities; it can
13
276
also serve as a habitat (i.e., shelter, refugia), making it an
integral part of ecosystems (Condron et al., 2010; Moore
et al., 2004). The favorable surface conditions of an ever-
wet environment would have limited the need to colonize
point bar sands in search of resources or shelter. Readily
available detritus on the surface would have been selected
over resources found within the substrate that would have
required extra energy to acquire by burrowing (Jumars &
Wheatcroft, 1989).
5.3.2 Conemaugh Group
The appearance of Skolithos in the point bar deposits of the
Conemaugh Group suggests colonization by opportunistic
arthropods engaged in deposit feeding and dwelling activi-
ties (Buatois & Mángano, 2004, 2011; Miller & Collin-
son, 1994). Modern burrows similar to Skolithos are often
associated with opportunistic organisms that display a high
tolerance to physiologically stressful conditions (Buatois &
Mángano, 2011). Burrowing is an energy intensive form of
locomotion and is typically only utilized when there is a
strong selective pressure to do so (Jumars & Wheatcroft,
1989). Likewise, soil fauna, such as arthropods, resort to
deposit feeding only if there is a significant selective pres-
sure to feed on nutrient-poor sediments (Jumars & Wheat-
croft, 1989). The Conemaugh Skolithos may have also been
produced as temporary refuges in response to seasonally
stressed or unstable surface conditions, such as rapidly shift-
ing water-table levels (Buatois & Mángano, 2004; Hembree
& Nadon, 2011). Overall, the transition from an absence of
ichnofossils in the Allegheny Group to their occurrence in
the Conemaugh Group suggests the onset of less optimal
surface conditions as the region began its shift to a drier,
more seasonal climate (Cecil et al., 1985; Joeckel, 1995;
Hembree & Nadon, 2011; Catena & Hembree, 2012; Dze-
nowski and Hembree, 2012), resulting in the need for vari-
ous animals to occasionally engage in deposit feeding or cre-
ate temporary dwelling spaces to avoid stressful conditions.
5.3.3 Monongahela Group
Climate conditions continued to shift from the tropical-
seasonal, wet–dry climate of the Conemaugh to the drier,
subhumid climate of the Monongahela (Brezinski & Kollar,
2011; Cecil et al., 2004). Across this interval, most quantita-
tive aspects of the ichnofossil assemblages in the point bar
sandstone increased. The increased abundance of vertical
burrows such as Skolithos and Arenicolites in low diver-
sity assemblages suggests that this environment contained
soil arthropods engaging in deposit feeding and dwelling
behaviors in a stressed environment (Buatois & Mángano,
2004; Vossler & Pemberton, 1988). Deposit feeders survive
on food sources with relatively poor nutritional value as
13
J. K. Crowell, D. I. Hembree
ingested sediment usually consists of 95% inorganic material
(Lopez & Levinton, 1987). Increased dependence on deposit
feeding, which is also energetically expensive, suggests that
surface conditions were becoming less favorable, with an
overall trend of decreasing precipitation and increasing sea-
sonality (Hembree & Bowen, 2017; Hembree & Carnes,
2018; Hembree & McFadden, 2020).
5.3.4 Lower Dunkard Group
The ichnofossil assemblages of the Lower Dunkard Group
show a significant increase in their quantitative properties,
especially in abundance and diversity, from the Mononga-
hela Group. Multiple studies of the Lower Dunkard Group
have indicated that it was a dry-seasonal climate (Cecil,
2013; Fedorko & Skema, 2013; Hembree & Carnes, 2018;
Hembree & McFadden, 2020; Montañez & Cecil, 2013).
Drier wet–dry climates like that of the Lower Dunkard have
a strongly seasonal distribution and lower amounts of pre-
cipitation (Hasiotis et al., 2007). These aspects of wet–dry
climates result in an increase in ichnodiversity in conti-
nental settings due to lower soil moisture levels (Hasiotis
et al., 2007). The major increase in abundance, diversity,
and density of ichnofossils in the Lower Dunkard shows a
substantial increase in mobile-deposit feeding activities of
animals to obtain buried resources from within the sediment.
Opportunistic animals are noted for their ability to vary their
feeding styles and sources based on food availability in their
environment (Cadée, 1984; Vossler & Pemberton, 1988). If
surface conditions were not optimal and contained minimal
resources, then these animals would utilize the best feeding
strategy to obtain sustenance, in this instance, deposit feed-
ing. A diversity of arthropods like beetles, spiders, insect
larvae, myriapods, and isopods were the probable trace
makers of the ichnofossil assemblage of the Lower Dunkard
(Buatois & Mángano, 2004; Hasiotis, 2002, 2004; Hembree
& Nadon, 2011).
5.3.5 Upper Dunkard Group
The Upper Dunkard Group represents a continuation of
stressed environments in a dry-seasonal climate (Cecil,
2013; Fedorko & Skema, 2013; Hembree & Carnes, 2018;
Hembree & McFadden, 2020; Montañez & Cecil, 2013).
The ichnofossils of the Upper Dunkard point bar sandstones
are predominantly associated with mobile-deposit feeding
behaviors. The mean complexity of the Upper Dunkard
assemblages was the highest of the studied units, due to the
appearance of Treptichnus. Although not necessarily a new
behavior, Treptichnus is significant, because it represents
an animal systemically moving through the substrate in a
complex way not seen in the other studied units (e.g., Getty
et al., 2016; Miller, 2003).
Climate‑induced changes in fluvial ichnofossil assemblages of the Pennsylvanian–Permian…
5.4 Riparian community composition change
Modern riparian habitats are the most dynamic, diverse,
complex, and ecologically productive terrestrial environ-
ments on Earth (Kondolf et al., 1996; Naiman et al., 1993).
For these reasons, it is particularly important to understand
how communities adapt to changing climatic and environ-
mental conditions in these settings. Ancient riparian com-
munities are beneficial to study, because they record changes
that occur through prolonged periods of time providing con-
text to evaluate the long-term responses of modern riparian
communities to climatic shifts now and in the future.
This study compared differences in the ichnofossil assem-
blages from the tropical (ever-wet) riparian environments
of the Allegheny Group to the semi-arid (wet–dry) ripar-
ian environments of the Upper and Lower Dunkard groups.
Modern ever-wet tropical climates have abundant precipita-
tion with a short dry season, constant high temperatures,
and consistently wet soil conditions (Hasiotis et al., 2007;
Walsh et al., 2009). This type of climate is most common
in dense rainforests and swamps that receive over 240 rainy
days a year such as parts of the Amazon Basin, Indonesia,
and Malaysia (Boyce & Lee, 2010; Hasiotis et al., 2007).
Arthropod assemblages of ever-wet climates in modern set-
tings typically require moist habitats with abundant leaf litter
and include groups, such as millipedes, centipedes, amphi-
pods, and isopods (Nakamura et al., 2003). Point bar, levee,
and floodplain deposits commonly contain terrestrial beetle
traces (Hasiotis et al., 1998).
Modern semi-arid (wet–dry) climates experience high
seasonality and a range of precipitation where the amount of
precipitation can be less, equal, or greater than evaporation
rates (Hasiotis et al., 2007). Savannahs, grasslands, steppes,
and mixed savannah woodlands are typical of wet–dry cli-
mates (Hasiotis et al., 2007). In modern wet–dry climates,
riparian zones are noted for their ecological importance as
habitats, because they contain high biodiversity in compara-
tively small areas (Chan et al., 2008; Dimmitt, 2000; Kon-
dolf et al., 1996). In wet–dry regions that are exposed to
seasonal environmental extremes such as high temperatures
and low humidity such as the subtropical Sonoran Desert, it
is common for surface dwelling animals to construct subsur-
face shelters to avoid extreme surface conditions (Cloudsley-
Thompson, 1975; Dimmitt, 2000; Hembree et al., 2017).
Ancient semi-arid riparian environments also likely con-
tained a high biodiversity reflected in the ichnofossil record.
Population strategies of trace-making organisms can be
interpreted using ichnofossil abundance, density, and diver-
sity (Ekdale, 1985). The ichnofossil assemblages of the
Conemaugh and Monongahela groups were produced by
generalists within opportunistic, temporary communities.
Opportunistic, or r-selected, organisms are those that exhibit
high reproductive rates, rapid growth rates, small body sizes,
277
wide environmental tolerances, short lifespans, high density
of individuals, low species diversity, and generalized feeding
habits (Pianka, 1970; Ekdale, 1985; Buatois and Mangano,
2011). Opportunistic organisms are known for their ability
to quickly colonize a habitat after a major environmental
change and thrive in highly stressful and unstable environ-
ments with low resources (Ekdale, 1985; Vossler & Pem-
berton, 1988; Zhao et al., 2020). Many insects fit into this
life strategy and are the most likely producers of the ichno-
fossils found at the study sites (Pianka, 1970). Ichnofauna
produced by opportunistic organisms usually display low
ichnodiversity, localized distributions, high abundance and
density, and simple morphologies (Ekdale, 1985; Buatois
and Mangano, 2011; Zhao et al., 2020). Based on the sub-
tropical environments of the Conemaugh and Monongahela
groups, it is likely that organisms in these groups only fed
occasionally in point bar sands if the opportunity arose and
likely tended to stay in the deposits of floodplains containing
plentiful organic matter.
The ichnofossil assemblages of the Monongahela and
Dunkard groups were also produced by generalists; how-
ever, the ichnofossils were indicative of a more permanent,
established community. Such communities are mostly com-
posed of equilibrium, or K-selected, organisms (Ekdale,
1985). Equilibrium organisms are much slower to colonize
a new environment compared to opportunistic organisms;
however, once established, they tend to be better adapted to
the new environment (Ekdale, 1985; Pianka, 1970). Equilib-
rium organisms are typically specialized feeders that occupy
specific niches in a habitat and exhibit narrow environmental
tolerances (Ekdale, 1985). They are usually part of high-
diversity, persistent climax communities with maximized
resource utilization occurring where the environment is sta-
ble and steady (Ekdale, 1985; Whittaker, 1953). The ichno-
fossil assemblages of the Monongahela and Dunkard groups
exhibited traits of a community that was more complex than
an r-selected community, but not a complex as a K-selected
community. For instance, the Dunkard Group had ichnofos-
sil assemblages with high diversity, abundance, and density,
but most ichnogenera were produced by simple dwelling,
locomotion, and feeding activities. Despite the simplicity
of the behaviors exhibited, the shift to an established com-
munity provides evidence that the long-term occupation of
point bar sands had become more advantageous for perma-
nent occupation by generalists as surface conditions became
drier and more seasonal.
The shift from no ichnofossils in the Allegheny Group
to hundreds of ichnofossils within the Dunkard Group sup-
ports the interpretation that as the climate changed, animals
expanded into less-suitable substrates to find available,
concentrated resources. Selective pressures on changing
food quality and quantity greatly affect organism forag-
ing behavior (Jumars & Wheatcroft, 1989). The energy
13
278
spent searching, assessing, and exploiting a resource must
be gained back by the obtained resource (Koy & Plotnick,
2007). The transition to a drier, more seasonal climate
from the Late Pennsylvanian-to-early Permian would have
resulted in changes in resource availability and, conse-
quently, a shift in organism behavior to acquire food. Most
generalist organisms can alter their feeding habits depending
on resource availability in their environment, such as shift-
ing from detritus to deposit feeding if it was the most viable
option to obtain nutrients (Cadée, 1984; Vossler & Pember-
ton, 1988). Deposit feeding is an energy intensive form of
foraging and is engaged in when there is a selective pressure
to consume nutrient-poor particles (Jumars & Wheatcroft,
1989). A shift to obligate rather than opportunistic deposit
feeding in the Dunkard Group was likely driven by increas-
ingly stressful surface conditions requiring the expenditure
of more energy to gain nutrients within the less ideal point
bar sands.
Organisms respond to climatic changes in various ways
depending mainly on the nature, rate, and length of the
change as well as the range of biological responses available
to organisms (Erwin, 2009). Some modern biotic responses
to either short- or long-term climatic events suggest flex-
ible community composition structure that allows many
organisms to respond to rapid climatic shifts by migration
(Roy et al., 1996). In the case of the Dunkard Group, arthro-
pods may have responded to shifting climatic conditions by
migrating into sandy, nutrient-poor substrates and adjust-
ing feeding behaviors to manage the demands of increas-
ingly more arid and highly seasonal environments.
6 Conclusions
Continental ichnofossils provide important insight into
riparian ecosystems not revealed from body fossils alone
and help provide a broader understanding of how long-term
climatic changes impact terrestrial communities and eco-
system dynamics. This study involved the investigation of
changes in the abundance, density, diversity, and complexity
of riparian ichnofossil assemblages in point bar sandstones
from the Middle Pennsylvanian-to-early Permian Allegheny,
Conemaugh, Monongahela, and Dunkard groups of Ohio
and West Virginia. Sedimentological evidence from the
Allegheny to Dunkard groups record a climatic shift from
the mid-Pennsylvanian to the early Permian from a tropi-
cal, ever-wet climate to a wet–dry, semi-arid, and highly
seasonal climate, respectively. The riparian trace-making
communities of the Appalachian Basin were comprised
mostly of arthropods, such as isopods, beetles, and myri-
apods whose interactions with the subaerial substrate pro-
vide salient information about how terrestrial invertebrate
communities responded to these climatic changes.
13
J. K. Crowell, D. I. Hembree
The absence of ichnofossils in the Allegheny Group
sandstone was characteristic of an ever-wet climate in
which the environment was well vegetated with abundant
resources resulting in no need for organisms to expend
energy on burrowing through point bar sands to deposit
feed. During the deposition of the Conemaugh Group,
there was a shift toward a more seasonal, subtropical cli-
mate. Surface conditions started to become less favorable
resulting in the appearance of ichnofossils produced by
dwelling and deposit feeding activities in the point bar
sandstones. Surface conditions progressively declined as
the climate shifted to a drier, subhumid one during the
deposition of the Monongahela Group. The ichnofossils
of the Conemaugh and Monongahela groups consisted
of Skolithos and Arenicolites suggesting a community
composed of opportunistic generalists that temporarily
occupied point bar sands. By the Lower Permian Dunkard
Group, sedimentary units were representative of a dry-
seasonal climate. Ichnofossil abundance, diversity, and
density increased significantly and more ichnofossil types
were observed including Cruziana, Planolites, Scoyenia,
and Treptichnus. The community composition transitioned
to an established, permanent community of generalists that
inhabited the substrate for long durations. Another trend
in community composition observed was a shift from sta-
tionary detritus feeding and dwelling to mobile detritus
and deposit feeding. This was indicated by an increase in
the complexity of ichnofossil morphology as organisms
engaged in slightly more sophisticated feeding styles to
maximize resources gained from the sediment.
Examining how continental trace-making commu-
nity composition and ecosystem dynamics shift over time
is important, because it provides crucial details about
how communities responded to environmental perturbations
in the past. With modern changes in climate and environ-
ment becoming increasingly of great concern, particularly
aridification, it is important to understand how ancient
organisms and environments responded to climatic shifts to
more arid conditions (Capon et al., 2013; LeRoux & McGe-
och, 2008; Walther, 2010). The study revealed that drier,
highly seasonal climate conditions resulted in organisms
needing to colonize harsher, nutrient-poor substrates and
engage in energy-expensive feeding styles to have enough
resources to sustain themselves. By assessing how ancient
communities adapted to changing environmental conditions,
more informed predictions can be made about what future
impacts climatic shifts will have on continental ecosystems
and terrestrial communities.
Acknowledgements We would like to thank Joseph Wislocki for help
with field work. This project would not have been possible without
funding by the Ohio University Geological Sciences Graduate Student
Alumni Grant (to JKC), the Geological Society of America Student
Research Grant (to JKC), the Society for Sedimentary Geology Student
Climate‑induced changes in fluvial ichnofossil assemblages of the Pennsylvanian–Permian…
Research Grant (to JKC), and the Paleontological Society Student
Research Grant (to JKC).
Author contributions J.C. and D.H. wrote the manuscript text and
prepared the figures and tables. All authors reviewed the manuscript.
Funding Funding for this research was provided by the Ohio University
Geological Sciences Graduate Student Alumni Grant (to JKC), the
Geological Society of America Student Research Grant (to JKC), the
Society for Sedimentary Geology Student Research Grant (to JKC), and
the Paleontological Society Student Research Grant (to JKC).
Availability of data and materials All data collected are available in
the tables and appendices.
Code availability Statistical analyses were performed using PAST ver-
sion 3, free software available at https://​folk.​uio.​no/​ohamm​er/​past/.
Declarations
Conflict of interest The authors have no competing interests to declare
that are relevant to the content of this article.
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