Palaeogeography, Palaeoclimatology, Palaeoecology 574 (2021) 110440

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Palaeogeography, Palaeoclimatology, Palaeoecology

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Sequence stratigraphy and carbon isotopes from the Trenton and Black
River Groups near Union Furnace, PA: Constraining the role of land plants
in the Ordovician world
Page C. Quinton a, *, Michael C. Rygel a, Megan Heins a, b
a
Department of Geology, State University of New York, College at Potsdam, 44 Pierrepont Avenue, Potsdam, New York 13676, United States of America
b
Department of Earth and Atmospheric Sciences, University of Nebraska-Lincoln, 126 Bessey Hall, Lincoln, NE 68588, United States of America

A R T I C L E I N F O A B S T R A C T

Editor:Prof. Thomas Algeo Sea-level change influences carbon isotopic trends in both modern and ancient carbonate depositional envi­
ronments. Generally, this relationship is manifested as a positive carbon isotopic excursion where the rising limb
Keywords: of the excursion is associated with transgression. These excursions exist because sea level can influence 1) local/
Carbon cycle regional/global carbon cycling, 2) basin restriction and meteoric influence, and 3) carbonate sedimentation. The
Non-vascular plants
first two processes are influenced, in part, by the large carbon reservoir represented by the terrestrial biosphere.
Appalachian Basin
Our goal is to explore the relationship between carbon isotopes and sea level prior to the evolution of vascular
Sandbian
Sea level plants at a time when the terrestrial carbon reservoir was far smaller.
Epicontinental sea This study focuses on the Upper Ordovician Union Furnace section in Pennsylvania, where a previously
documented ~1‰ increase in δ13C values in the upper Sandbian (458.4–453.0 Ma; referred to as the “baseline
shift”) has been interpreted to reflect a perturbation to the global carbon cycle due to the proliferation of non-
vascular plants. By focusing on the interval deposited before the “baseline shift”, we were able to examine the
relationship between sea level, carbon isotopes, and some of the earliest terrestrial ecosystems. We present a
high-resolution δ13C record directly tied to the sedimentological and sequence stratigraphic framework. By
sampling genetically related packages of rock we were able to document a relationship between carbon isotopes
and sea level at the sequence and parasequence level. Our results show that carbon isotopic values decreased
during transgressions and increased during regressions. This seemingly inverse relationship between δ13C values
and sea level is interpreted to reflect increased terrestrial fluxes during regression in a world devoid of vascular
plants. Our results suggest that the early colonization of land by non-vascular plants impacted regional carbon
cycling in the Appalachian Basin and eventually the global carbon cycle in the Late Ordovician.

1. Introduction carbon, either in the water column or at the sediment-water interface,

Carbon isotopic trends from marine carbonates can record spatial
and temporal changes in the global carbon cycle (Saltzman and Thomas,
2012). They do so because significant fluctuations in the average δ13C
composition of the ocean (δ13C DIC) are a function of changes in the flux
of carbon between the atmosphere-ocean system and the sedimentary
reservoir (Kump and Arthur, 1999; Saltzman and Thomas, 2012). This
flux is primarily controlled by the burial and oxidation of organic car­
bon, which is enriched in 12C (Kump and Arthur, 1999; Saltzman and
Thomas, 2012). The burial of organic carbon thus results in an increase
in the δ13CDIC value of the water. Whereas the oxidation of organic
decreases the δ13C DIC value of the ocean. These changes in ocean
chemistry are recorded in the carbon isotopic ratio of the carbonate
sediments (δ13C carb) formed in many marine settings. This basic rela­
tionship is the foundation for the use of δ13C records in paleoclimatic,
paleoredox, and chemostratigraphic studies. However, there are
complicating factors that must be considered when using δ13C records
from epicontinental carbonates.
The δ13C trends recorded in carbonates of shallow epicontinental
settings can reflect a complicated set of global, regional, and local pro­
cesses. These processes include, but are not limited to, carbonate
weathering, carbonate deposition, meteoric input, organic carbon

* Corresponding author.
E-mail address: quintopc@potsdam.edu (P.C. Quinton).

https://doi.org/10.1016/j.palaeo.2021.110440
Received 18 January 2021; Received in revised form 25 April 2021; Accepted 25 April 2021
Available online 29 April 2021
0031-0182/© 2021 Elsevier B.V. All rights reserved.
P.C. Quinton et al.
burial/oxidation, carbon sources/flux, and local anoxia (Kump and
Arthur, 1999; Kump et al., 2000; Immenhauser et al., 2003; Panchuk
et al., 2005; Melchin and Holmden, 2006; Fanton and Holmden, 2007;
Schrag et al., 2013). All of these processes can be controlled directly or
indirectly by relative sea-level change because it influences the burial/
oxidation of organic carbon, proximity to terrestrial carbon sources,
basin restriction, carbonate production vs. weathering, carbonate
mineralogy, and/or meteoric fluids. This relationship is supported by
multiple studies which have documented a correlation between carbon
isotopic trends and sea level (e.g. Weissert, 1989; Jenkyns, 1996; Bug­
gisch et al., 2003; Immenhauser et al., 2003; Swart and Eberli, 2005;
Swart et al., 2009; Eltom et al., 2018). The general pattern that emerges
is one where δ13C values tend to increase during sea level rise and
decrease during sea level fall. The result is a positive carbon isotopic
excursion with the rising limb during transgression.
While there are multiple processes that can potentially produce this
sea-level-induced positive carbon isotopic excursion, many of them are
linked to an abundance of land plants. For example, proximity to shore
and terrestrial carbon sources (typically enriched in 12C) can lead to
decreasing δ13C values when relative sea level is low (Burdige, 2005;
Jarvis et al., 2006; Fanton and Holmden, 2007). At the same time,
proximity to the shoreline and basin restriction might lead to an increase
in meteoric fluids which are typically enriched in plant respired 12C
(Allan and Matthews, 1982; Immenhauser et al., 2003). The result is
decreased δ13C values when sea level is low. This influence of land plants
can be observed in modern carbonate settings such as the Great Bahama
Banks and Florida Bay, where nearshore environments record δ13CDIC
values that are up to 2‰ lower than those close to the open ocean, in
part because of freshwater and terrestrial organic matter input (Patter­
son and Walter, 1994). The importance of land plants on the carbon
isotopic values of shallow epicontinental settings and the at least partial
dependence on land plants to produce the sea-level-induced positive
carbon isotopic excursion raises an important question: will the rela­
tionship between sea level and carbon isotopes be different and/or more
variable prior to an abundance of terrestrial land plants? The aim of this
study is to examine carbon isotopic trends in a sequence stratigraphic
framework in the absence of widespread terrestrial vegetation.
2. Geologic setting
This study focused on Black River and Trenton Group carbonates
(Late Ordovician) in a ~ 240 m thick exposure along PA Route 453 near
Union Furnace, PA (40.6230◦ N, 78.1730◦ W). These rocks were
deposited in the shallow epicontinental sea that covered Laurentia
during the Sandbian–Katian Stages (458.4–445.2 Ma; Scotese, 2004;
Torsvik and Cocks, 2013; Walker et al., 2018). Our efforts were focused
on the Sandbian-aged Black River Group in the basal 105 meters of the
section (Laughrey et al., 2017). The Union Furnace section is appro­
priate for this study because: 1) the sequence stratigraphic framework
independently established by Laughrey et al. (2017) is easily identifiable
and confirmable in the field, 2) the Late Ordovician carbon isotopic
record of Laurentia suggests that sea level might have influenced carbon
cycling in the Appalachian Basin, and 3) the Union Furnace section
spans an interval when non-vascular land plants were beginning to
impact the carbon cycle.
2.1. Late Ordovician sequence stratigraphy
A detailed sequence stratigraphic framework for the Middle to Late
Ordovician of eastern North America was established by Holland and
Patzkowsky (1996). Their efforts focused on the central to southern
Appalachians and the Cincinnati Arch where they recognized and
named a total of 14 sequences. Those sequences are composed of
transgressive systems tracts (TST) and highstand systems tracts (HST),
with lowstand systems tracts (LST) occasionally preserved (Holland and
Patzkowsky, 1996). Parts of this sequence stratigraphic framework have
2
Palaeogeography, Palaeoclimatology, Palaeoecology 574 (2021) 110440
been extended into New York and southern Ontario (Brett and Baird,
2002; Brett et al., 2004), but these studies and recent work in the Cin­
cinnati Arch (Brett et al., 2020) has led to revision of the original
framework established by Holland and Patzkowsky (1996) and the
addition of at least two sequences. Much of this work has focused on the
Katian Stage of the Upper Ordovician, with similar efforts not yet
completed for the earlier Sandbian Stage.
Detailed sedimentological and sequence stratigraphic work on the
Union Furnace section was done by Laughrey et al. (2017) as part of the
pre-conference field trip for the 82nd Annual Field Conference of
Pennsylvania Geologists and as part of a larger report (Patchen et al.,
2006). They recognized four sequences in the Black River Group (BR
1–4) and six sequences in the Trenton Group (TR 1–6). While the TR 3
and TR4 sequences have been correlated to Holland and Patzkowsky’s
(1996) M4 and M5 sequences, the rest of the section has not been
correlated in detail and we use the nomenclature that Laughrey et al.
(2017) established at this location. Broad regional age equivalence of
the Union Furnace section is possible with the age model developed by
Edwards et al., 2015a, placement of Midcontinent Conodont Biozones
(Edwards et al., 2015a, Adiatma et al., 2019), and identification of the
Deicke and Millbrig K-bentonites (altered volcanic ash beds; Laughrey
et al., 2017).
2.2. Late Ordovician carbon isotopic record
The Late Ordovician carbon isotopic record is well documented with
four globally recognized positive excursions and a relatively high-
resolution composite curve spanning the Sandbian to the Hirnantian
Stages (Bergström et al., 2009; Saltzman and Thomas, 2012). Many of
these studies have focused on the Laurentian epicontinental region (e.g.
Young et al., 2005; Fanton and Holmden, 2007; Bergström et al., 2010;
Saltzman and Edwards, 2017; Adiatma et al., 2019). The large number
of carbon isotopic records and studies that attempt to constrain regional
variability in δ13C values (Holmden et al., 1998; Panchuk et al., 2006)
have demonstrated that there was significant variation in carbon iso­
topic values (~2‰) across the Laurentian epicontinental sea during the
Late Ordovician. These spatial gradients in carbonate δ13C values have
generally been attributed to basin restriction, differences in carbon
sources, and organic carbon burial (Panchuk et al., 2005; Panchuk et al.,
2006) but could also reflect miscorrelations between sections or certain
mineralogical differences (Swart et al., 2009; Metzger and Fike, 2013;
Sell et al., 2015). Regardless of the cause, the presence of these spatial
gradients increases the chances that sea level could exert a resolvable
influence on δ13C trends.
This prediction is supported by previous studies that documented a
relationship between δ13C trends and sea level during the Late Ordovi­
cian. Fanton and Holmden (2007) used neodymium (Nd) isotopes to
estimate sea level change and compared these inferred sea level cycles to
δ13C trends during the Katian. They suggested that the correlation be­
tween positive carbon isotope excursions and Nd trends indicated that
sea level was driving changes in basin restriction and upwelling (Fanton
and Holmden, 2007). Similarly, Young et al. (2008) argued that the
globally recognized Guttenberg Isotope Carbon Excursion (GICE) in the
early Katian might have been enhanced in the Laurentian epicontinental
sea as sea level rose, driving upwelling of nutrient rich, oxygen poor
waters that fueled primary productivity and the burial of organic car­
bon. While these studies focused on Katian carbon isotopic record (and
this study is focused on the slightly older Sandbian record), these results
suggest that the Union Furnace section is an appropriate interval for
exploring the relationship between sea level and δ13C.
2.3. The Late Ordovician & the evolution of land plants
The colonization of the terrestrial realm by land plants during the
Ordovician–Devonian Periods would have had a significant and lasting
impact on the carbon cycle (Figure 1). Increased biomass would have
P.C. Quinton et al. Palaeogeography, Palaeoclimatology, Palaeoecology 574 (2021) 110440

Fig. 1. Timescale with terrestrial plant stages as identified by Davies and Gibling (2010) and Dahl and Arens (2020). Major events in lant plant evolution and
influence on the carbon isotopic record are tied to the timescale. The Union Furnace section is placed within this framework using the age model from Edwards
(2015a). 1Stein et al., 2020, 2Glasspool and Scott, 2000, 3Gerrienne et al., 2011, 4Driese and Mora, 2001, 5Kennedy et al., 2013, 6Gensel et al., 2001, 7Edwards et al.,
1983, 8Tomescu and Rothwell, 2006, 9Tomescu et al., 2009, 10Kenrick et al., 2012, 11Adiatma et al., 2019, 12Jones et al., 2015, 13Malkowski and Racki, 2009.

resulted in significant terrestrial carbon storage (Berner, 1998). The
advent of root systems would have led to an increase in silicate weath­
ering that consumes CO2 (Algeo and Scheckler, 1998; Berner, 1998;
Quirk et al., 2015). The increase in silicate weathering would also supply
nutrients to the marine realm fueling primary productivity and causing
an increase in marine organic carbon burial (Algeo et al., 1995; Algeo
and Scheckler, 1998). Traditionally, these changes were thought to
mostly be confined to the Devonian Period because that is when complex
root systems evolved and when significant organic carbon storage in the
terrestrial realm began via the earliest coal (Algeo and Scheckler, 1998;
Glasspool and Scott, 2010; Dahl and Arens, 2020). However, recent
studies looking at microfossil assemblages, molecular clock origination
dates (Kenrick et al., 2012; Morris et al., 2018), and non-vascular land
plants’ ability to influence carbon storage and silicate weathering
(Boucot and Gray, 2001; Edwards et al., 2015a, 2015b; Porada et al.,
2016), suggest that land plants impacted the carbon cycle throughout
the Ordovician and Silurian and did so in a progressive, stepwise fashion
(Lenton et al., 2012, 2016; Dahl and Arens, 2020).
Recent studies suggest that land plants’ impact on the carbon cycle
may have started in the Late Ordovician (Lenton et al., 2012, 2016;
Adiatma et al., 2019; Dahl and Arens, 2020). The first unequivocal
spores appear in the Middle Ordovician (Rubinstein et al., 2010; Kenrick
et al., 2012) and macroscopic plant material attributed to liverworts
appears by the Late Ordovician (Tomescu et al., 2009). These observa­
tions have led some to argue that significant terrestrial biomass existed
by the Late Ordovician (Tomescu et al., 2009; Dahl and Arens, 2020).
These early terrestrial ecosystems would have been dominated by
bryophytes, which have little to no subsurface component. Despite the
lack of roots, bryophytes could have exerted a significant influence on
silicate weathering rates by trapping sediments in their surface
components (Kleinhans et al., 2018; McMahon and Davies, 2018) and
secreting organic acids through their rhizoids (Boucot and Gray, 2001;
Lenton et al., 2012; Quirk et al., 2015; Porada et al., 2016).
A Late Ordovician increase in silicate weathering and nutrient supply
could have caused a fundamental shift in the global cycle (Boucot and
Gray, 2001; Lenton et al., 2016; Porada et al., 2016) marked by a ~ 2‰
increase in the background marine δ13C carbonate values in the late
Sandbian (Lenton et al., 2016). This geochemical support comes in part
from the Union Furnace section where Adiatma et al. (2019) reported an
~1‰ increase in δ13C values from 60 to 120 m in their section. They
refer to this increase as a “baseline shift” caused by the influence of land
plants (Adiatma et al., 2019). By focusing on the interval below this
“baseline shift” in this study we were able to examine the relationship
between sea level, carbon isotopes, and the earliest terrestrial
ecosystems.
2.4. Previous carbon isotope work from the Union Furnace section
As noted above, Adiatma et al. (2019) reported δ13C results at a 0.5
to 3 m sampling interval from the Union Furnace section. That study
specifically focused on documenting large-scale trends related to
changes in the global carbon reservoir (e.g. the GICE and the Sandbian
“baseline shift”). Here we focus on the generating a higher resolution
record (0.1 to 1 m intervals) for the lower 113.2 m that is directly tied to
detailed sedimentological observations and a sequence stratigraphic
framework. This new paired geochemical and sedimentological dataset
will allow us to test sea level’s role on the observed carbon isotopic
patterns.

3
P.C. Quinton et al. Palaeogeography, Palaeoclimatology, Palaeoecology 574 (2021) 110440

3. Methods deposits capping progradational parasequence sets. When preserved,

3.1. Sedimentology and sequence stratigraphy
We measured and described 220 m of the Union Furnace section
(Fig. 2 and supplementary materials file). We keyed our measurements
to bedding units marked on the outcrop for the pre-conference field trip
for the 82nd Annual Field Conference of Pennsylvania Geologists which
also allowed us to directly compare our observations to the sedimento­
logical and sequence stratigraphic work by Laughrey et al. (2017). In
addition to our field-based observations, 15 samples were collected for
petrographic analysis. Using these sedimentological observations and
building on the interpretations of Laughery et al. (2017), we developed a
simplified and standardized framework for a carbonate ramp informed
by comprehensive reviews of carbonate systems (e.g. Read, 1985; Son­
nenfeld, 1996; James and Jones, 2015). This depositional framework is
consistent with the barrier-bank type ramp that characterizes the Black
River Group in this region (Laughrey et al., 2017). We were able to place
the Union Furnace section into 10 distinct depositional environments
within that framework (Table 1, Figs. 2 and 3).
Our sequence stratigraphic framework is built on the work of
Laughrey et al. (2017), which is easily recognizable and verifiable in the
field. Sequence boundaries (SB) are placed atop the shallowest water
lowstand systems tract (LST) deposits record a basinward shift in facies
and a continuation of the progradational parasequence architecture.
Transgressive surfaces (TS) mark the start of progressive deepening;
where the LST is absent this surface is merged with the SB. The trans­
gressive systems tract (TST) is composed of deposits that record pro­
gressive deepening and a retrogradational parasequence architecture.
Given the subtle lithologic differences in distal facies and the monoto­
nous succession that result, individual parasequences were difficult to
identify. The maximum flooding surface (MFS) is placed in the middle of
the deepest water deposits; given the nature of the distal facies described
above, its placement is approximate. The transgressive systems tract
(TST) records progressive shallowing. In some intervals, marked dif­
ferences in facies result in an obvious progradational parasequence
architecture.
In a few places we made slight adjustments (<2 m) to the position of
sequence stratigraphic surfaces presented by Laughrey et al. (2017). In
sequence BR 2 (27.5–61.5 m) we were able to confidently identify 10
parasequences. This level of detail was not possible in the other se­
quences and systems tracts because of a lack of lithologic contrast. Our
expanded facies scheme and sequence stratigraphic framework is pre­
sented in Figs. 2 and 3.

Fig. 2. Graphic log for 125 m of the Union Furnace section. Lithostratigraphy and bedding units are based on Laughrey et al. (2017). Graphic log, facies associations
(Table 1), and parasequence calls are based on work from this study. Sequence stratigraphic framework is based on Laughrey et al. (2017) and confirmed with our
sedimentological analysis. Grainsize scale abbreviations are mdst (mudstone), wkst (wackestone), pkst (packstone), grnst (grainstone), rxl (recrystallized). Detailed
field notes and full measured section are included in the supplementary materials file.

4
P.C. Quinton et al.
Table 1
Facies associations and interpretated depositional environments for the Union Furnace section. The framework for depositional environments we deploy is informed by
Read (1985), Sonnenfeld (1996), and James and Jones (2015). The bioturbation index was developed by Droser and Bottjer (1986).
Facies Depositional Lithologies and sedimentary structures
Associations Environment
A Anaerobic Laminated to faintly laminated dark gray mudstone
Outer Ramp (>90%) with discrete interbeds of fissile organic-
rich(?) mudstone (<10%). Rare fossiliferous
wackstone beds. Sparse bioturbation (B.I. = 1–2).
Fossils and burrows are locally replaced by chert.
B Aerobic Outer Faintly laminated to internally massive mudstone
Ramp interbedded with dark gray nodular limestone and
rare sharp-based packstone/grainstone
(fossiliferous). Abundant bioturbation (B.I. = 3–4);
burrows commonly filled with coarser grained
material
C Middle Ramp Faintly laminated mudstone and wackestone with
subordinate sharp-based fossiliferous grainstones,
1–2 cm thick laminated zones locally. Abundant
bioturbation (B.I. = 3–4). Scattered hardgrounds
with bored surfaces and intraformational
conglomerates.
D Shoreface Grainstone with interbedded packstone; both
contain abraded fossils, intraclasts, and peloids.
Faintly laminated to internally massive. Minor
mudstone/wackestone interbeds locally.
Bioturbation is difficult to distinguish in coarse-
grained facies; rare grainstone-filled burrows in
mudstone/wackestone (B.I. = 1).
E Upper Nearly identical to D, except with abundant cross-
Shoreface beds and no mudstone/wackestone.
F Restricted Interbedded wackestone/packstone (fossiliferous)
Upper with subordinate mudstone. Faintly laminated to
Shoreface internally massive. Minor bioturbation (B.I. = 1).
Commonly grades into G.
G Back-shoal Interbedded grainstones and packstones with minor
wackestone. Grainstones may be laminated, ripple
cross-laminated, or cross-bedded. Grainstones
appear sandy, and have abraded fossils and
intraclasts. Burrows are difficult to distinguish (B.I.
= 1). Commonly grades into F.
H Lagoon Mudstone to wackestone with minor packstone/
grainstone lags (fossils and intraclasts). Mudstone
and wackstone varies from internally massive to
faintly laminated and bioturbation ranges from
scattered to abundant (B.I. = 2–5). Pyrite abundant
locally.
I Intertidal Cyclically interbedded successions of a) faintly
laminated mudstone/wackestone, b) laminated to
internally massive fossiliferous packstone, and c)
fossiliferous to intraclastic grainstones. Individual
cycles are a few tens of centimeters and not well
represented given the scale of our measured
section. Scattered bioturbation (B.I. = 1–2).
Commonly grades into J.
J Supratidal Mudstone to wackestone interbedded with
laminated to ripple cross-laminated grainstone.
Scattered fossils in mudstone/wackestone;
grainstones appear sandy and may be capped with
fossiliferous horizons. Scattered bioturbation (B.I.
= 1–2). Commonly grades into I.
3.2. Stable carbon isotopic samples and analyses
Samples for carbonate carbon isotopic analyses were collected along
with the generation of our detailed measured section. This sampling
protocol allowed us to directly tie generated carbon isotopic trends to
the sedimentological information and interpretations. We used a sliding
sampling resolution based on sedimentological context. We did not take
samples at some regular stratigraphic interval. Instead, when possible
parasequence boundaries were identified, we took a minimum of three
5
Palaeogeography, Palaeoclimatology, Palaeoecology 574 (2021) 110440
Fossils Processes
Rare transported brachiopod Below storm wave base, limited carbonate
shells. Rare Chondrites. production, low oxygen limits bioturbation.
Dominated by suspension deposition.
Brachiopods with scattered Below storm wave base, limited carbonate
gastropods and rugose coral. production but closer to carbonate sources on the
shelf, aerobic conditions allow for more intense
bioturbation and somewhat higher faunal
diversity. Combination of suspension deposition
and downslope transportation from C
Abundant fossils include Above storm wave base and below fairweather
brachiopods, bryozoans, trilobites, wave base, abundant oxygen and fully marine
rugose corals, echinoderms, conditions results in intense bioturbation and
mollusks, and gastropods. abundant/diverse fauna. Grainstones and lag
deposits represent storm beds.
Abundant abraded shell Above fairweather wave base. Although fully
fragments. marine, high energy conditions and shifting
substrate results in abraded fossils and modest
bioturbation.
Similar to D, except that energy levels are even
higher and cross-beds become abundant.
Abundant abraded shell Topographic low between the upper shoreface
fragments. and the backshoal. Wave activity still occurs, but
mud can accumulate in sheltered areas.
Abundant abraded shell Highest energy (shallowest water) environment
fragments; rare bryozoans. on the ramp; represents the transition between
the wave-dominated shoal and the tidally-
influenced lagoon.
Abundant abraded shell Shoreward of shoal and subaqueous even at low
fragments; bryozoans (some in tide. Occasional washover events bring in
situ) grainstones; intraclasts record storm events.
Variable conditions result in restricted fauna.
Abundant abraded shell Subaerially exposed at low tide and covered in
fragments; bryozoans and shallow water at high tide. Generally a sheltered,
scattered brachiopods. low energy environment behind the back-shoal/
lagoon complex; occasional high energy events.
Abraded shell fragments. Subaerially exposed, even at high tide. High
energy events can flood the area.
samples through the parasequence. This approach allowed us to
generate a δ13C record that captured changes within parasequences.
Samples were given a general lithological identification (mudstone,
wackestone.. etc) in the field and this was confirmed when sampling for
carbon isotopic values in the laboratory.
Sample powders were generated from freshly exposed surfaces using
a low speed drill. When possible, we targeted the micritic zones because
muddy lithologies have a lower susceptibility to post-depositional
alteration (due to their low permeability) than coeval coarser-grained
P.C. Quinton et al.
lithologies (e.g. Hayes et al., 1989). However, that was not always
possible because some intervals lacked carbonate mud. Sample lithology
for all carbon isotopic values are reported in supplementary data file.
Sample powders were analyzed for bulk carbonate δ13C and δ18O values
on a ThermoFinnigan Delta Plus Dual Inlet isotope ratio mass spec­
trometer connected to a Kiel III Carbonate Interface at the University of
Missouri Stable Isotope and Biogeochemistry Laboratory. Analytical
precision is ±0.04‰ (one standard deviation) for δ13C and ± 0.06‰
(one standard deviation) for δ18O and is calculated from multiple ana­
lyses of NBS-19.
4. Results
We report 170 bulk carbonate δ13C values from the lower 113 m of
the Union Furnace section. All values are plotted on Fig. 3 with a three-
point moving average; a complete data table is included in the
6
Palaeogeography, Palaeoclimatology, Palaeoecology 574 (2021) 110440
Fig. 3. Bulk carbonate carbon isotopic results from
this study are plotted with a 3-point moving average.
Lithostratigraphy (column 1) and bedding units
(column 2) are based on Laughrey et al. (2017).
Facies associations and inferred depositional envi­
ronments are based on the standardized framework
developed in this study (column 3). Systems tracts
and sequence stratigraphic surfaces (column 4) are
based on Laughrey et al. (2017) and confirmed based
on the work from this study. Black River and Trenton
Group cycles as identified by Laughrey et al. (2017)
are given in column 5. The late Sandbian “baseline
shift” previously identified by Adiatma et al. (2019)
is highlighted in green and the interval of proposed
sea level influence on carbon isotopic trends in
highlighted in blue. (For interpretation of the refer­
ences to colour in this figure legend, the reader is
referred to the web version of this article.)
supplementary data file. These values, with an average of − 0.28‰ are
similar to those reported elsewhere for this interval (Young et al., 2010;
Bergström et al., 2010; Quinton et al., 2016) and broad trends match
those reported previously by Adiatma et al. (2019). In particular, there is
a general trend of increasing values from ~0.5‰ at 41 m to 1.5‰ at 113
m. We did not sample the upper 113–220 m of the section (i.e. the GICE
interval) because a) the outer ramp deposits of those upper 107 m do not
contain the sedimentological contrast needed to confidently identify
parasequences and b) this study focused on Sandbian events.
In the lower 61.5 m, below the interval previously identified by
Adiatma et al. (2019) as the “baseline shift” in the late Sandbian, δ13C
values are correlated with the sequence stratigraphic framework. We see
decreasing values in the TST, increasing values in the HST, and stable
values in the single occurrence of a LST. Inflection points in these trends
occur at or near sequence stratigraphic surfaces. In sequence BR 1,
values decrease from –1‰ to –1.5‰ in the TST and increase from − 1.5‰
P.C. Quinton et al. Palaeogeography, Palaeoclimatology, Palaeoecology 574 (2021) 110440

to 0‰ in the HST. In sequence BR 2, values decrease from 0 to –1.5‰ in

the TST and increase from − 1.5‰ to 0‰ in the HST. We used Pearson’s
regression analyses to test the direction and significance of these visually
identified trends. Those results are reported in Table 2 and indicate that
there is a moderate to strong correlation in the negative direction be­
tween δ13C values and transgressive system tracts. For highstand sys­
tems tracts, there is a strong correlation in the positive direction. To
demonstrate that this correlation was truly unique to the identified
systems tracts, we also tested the correlation between δ13C values and
the sequences (BR 1 and BR 2) and found a weak correlation.
In sequence BR 2 we were able to confidently identify parasequences
and we report corresponding δ13C trends in Fig. 4. Averages from those
10 parasequences display a stepwise pattern consistent with their place
in the sequence stratigraphic framework (parasequence averages and
standard deviations are reported in the supplementary data file). Para­
sequence 1 (P1) represents the LST (P1 average 0.24‰). Parasequences
2 through 4 (P2 – P4) comprise the TST and show progressively lower
averages (P2 average 0.02‰, P3 average − 1.10‰, P4 average −
1.51‰). Parasequences 5 through 10 (P5 – P10) comprise the HST and
progressively higher averages (P5 average − 1.23‰, P6 and P7 average
− 0.56‰, P8 average − 0.57‰, P9 average − 0.47‰, P10 average −
0.41‰).

5. Discussion

5.1. Carbon isotopic trends at Union Furnace

Union Furnace carbon isotopic results record the same gradual in­
crease in δ13C values through the Erismodus quadridactylus to Phragmo­
dus undatus Midcontinent Conodont Biozones documented on Laurentia
(Quinton et al., 2016; Bergström et al., 2015) and Baltoscandia (Ainsaar
et al., 2010). This observation suggests that the Union Furnace carbon
isotopic record does reflect primary changes in the global carbon
reservoir as has been previously argued by Adiatma et al. (2019).
However, the correlation between δ13C trends and the sequence strati­
graphic framework observed in sequences BR 1 and BR 2 at Union
Furnace indicates that regional and/or local processes were also influ­
encing the record.
Carbon isotopic results from Union Furnace are interpreted to reflect
primary changes in the δ13C value of the local water mass (δ13CDIC)
rather than diagenetic processes. The main diagenetic concern for car­
bonate δ13C is recrystallization and exchange of carbon isotopes with
meteoric fluids (Allan and Matthews, 1982; Saltzman and Thomas,
2012). Carbon and oxygen isotopic results show no systemic relation­
ship (supplementary material file) as might be the case if the record was
altered by meteoric diagenesis. Traditionally, cross-plots between δ13C
and δ18O values of carbonates were used as a tool to test for systematic
alteration due to exchange with meteoric fluids (Marshall, 1992). The

Table 2
Results from Pearson’s regression analysis. Strength of correlation is determined
by using a scale informed from Akoglu (2018): 0.1 < |r| < 0.3 for weak corre­
lation, 0.3 < |r| < 0.5 for moderate correlation, and 0.5 < |r| for strong
correlation.
a b c
Sequence Systems Pearson’s Pearson’s n Statistical
Tract r r Significance

BR 1 TST − 0.284 − 0.512 16 Moderate

correlation
BR 2 TST − 0.820 − 0.906 14 Strong correlation
BR 1 HST 0.608 0.965 4 Strong correlation Fig. 4. Bulk carbonate carbon isotopic results for sequence BR 2 plotted by
BR 2 HST 0.539 0.687 30 Strong correlation parasequence. Each parasequence is numbered and the parasequence δ13C
BR 1 All − 0.045 − 0.062 20 Weak correlation average is indicated with a pink line. The carbon isotopic averages for para­
BR 2 All − 0.158 − 0.158 50 Weak correlation sequences in the transgressive systems tract get progressively lower and pro­
a
Calculated based on original data with no smoothing. gressively higher in the highstand systems tract. (For interpretation of the
b
Calculated based on 3-point moving average of carbon isotopic data. references to colour in this figure legend, the reader is referred to the web
c
Number of data points in analysis. version of this article.)

7
P.C. Quinton et al.
positive correlation between δ13C and δ18O values is interpreted to
reflect the varying contribution of light isotopes from meteoric fluids
(Allan and Matthews, 1982; Marshall, 1992; Banner and Hanson, 1990).
We should note that the utility of carbon and oxygen cross-plots as a test
for meteoric diagenesis has been challenged based on work in the Great
Bahama Banks. Swart and Oehlert (2018) demonstrated that positive
correlation between carbon and oxygen isotopes might also result from
varying amounts of recrystallization and development of diagenetic
cements and, more importantly, that meteoric diagenesis does not al­
ways produce a positive correlation.
Another way to constrain potential meteoric influence is to look at
the relationship between δ13C and δ18O values and lithology. Lithologies
dominated by fine grained material (e.g. mudstone and wackestone)
have lower permeability and porosities, such that it is less likely that
fluids will penetrate and alter these than their coarser grained coun­
terparts (e.g. grainstone and packstone) (Hayes et al., 1989; Marshall,
1992). We observe no systematic relationship between lithology and
isotopic values (Supplementary material file). Lastly, petrographic work
did not show significant and/or obvious meteoric cements and we
avoided any such features when sampling for carbon isotopes. Overall,
we see no evidence that the carbon isotopic values at Union Furnace
reflect significant alteration due to meteoric diagenesis.
5.2. Is sea level influencing carbon isotopes at Union Furnace?
The statistically significant correlation between carbon isotopic
trends and the sequence stratigraphic framework in sequences BR 1 and
BR 2 suggests that sea level was influencing the carbon isotopic record at
Union Furnace. There are multiple lines of evidence that support this
interpretation. The sequence stratigraphic framework was established
independently (though verified by us based on our facies identifications)
of the δ13C record. For this reason, there is no chance that the
geochemical trends influenced our placement of sequence stratigraphic
surfaces. We see the same pattern in both sequences with similar start
and end values for each systems tract (TST trend ends at ~ − 1.5‰ and
HST trend ends at ~0‰). This observation suggests that there is some
specific process that is allowing sea level to influence δ13C trends (dis­
cussed below). Furthermore, the inflection points in the carbon isotopic
trends occur at or near sequence stratigraphic surfaces. For example, in
BR 2 when flooding begins at the transgressive surface, δ13C values start
to decrease and when progressive shallowing begins following the
maximum flooding surface, δ13C values start to increase.
We also document a relationship between sea level and δ13C trends at
the parasequence level in sequence BR 2. The progressively deepening
parasequences of the TST (P2 – P4) show progressively lower δ13C
values while the progressively shallowing parasequences of the HST
show progressively higher δ13C values. While there is certainly some
scatter in δ13C values, averages of each parasequence show the same
relationship to δ13C that we observe at the systems tract level. Com­
bined, these observations provide significant evidence that sea level is
influencing δ13C values in the BR 1 and BR 2 sequences at Union
Furnace. The question is: what processes are allowing sea level to con­
trol carbon isotopes and are they different from what is traditionally
deployed to explain positive carbon isotope excursions?
5.3. How is sea level controlling carbon isotopes at Union Furnace?
The pattern of decreasing δ13C values during the TST and increasing
values during the HST in BR 1 and BR 2 is statistically significant
(Table 2) and the opposite of the pattern typically documented in most
younger carbonates (Weissert, 1989; Jenkyns, 1996; Immenhauser
et al., 2003; Swart and Eberli, 2005; Swart et al., 2009; Eltom et al.,
2018). In these studies, a rise in sea level causes an increase in carbon
isotopic values followed by a return to pre-excursion values as sea level
falls. The processes that are commonly thought to produce this sea level
induced positive carbon isotopic excursion fall into three categories 1)
8
Palaeogeography, Palaeoclimatology, Palaeoecology 574 (2021) 110440
carbon cycling, 2) meteoric influence, and 3) carbonate sedimentation.
Carbon cycling is the most common explanation for the relationship
between sea level and carbon isotopic values (e.g. Jenkyns, 1996; Bur­
dige, 2005; Jarvis et al., 2006; Fanton and Holmden, 2007; Katz et al.,
2007; Ainsaar et al., 2010). The two may be related because sea level
controls the net surface area for primary productivity and upwelling of
nutrient rich (that can potentially fuel primary productivity) and/or
oxygen poor waters (that can potentially promote the preservation of
organic carbon). The net result is that during transgressions, increased
upwelling of nutrient rich waters and increased surface area for primary
productivity results in increased organic carbon burial. Combined with
increased terrestrial organic carbon input and basin restriction during
sea level falls, the result is a classic sea-level-induced positive carbon
isotope excursion with the rising limb in the TST and the falling limb in
the HST.
Meteoric fluids can also cause a relationship between sea level and
δ13C patterns. During sea level lows, penetration of meteoric fluids
(potentially enriched in respired 12C) can lead to alteration of recorded
carbon isotopic values. The result is decreasing carbon isotopic values
up to the sequence boundary. This model, described by Allan and Mat­
thews (1982), is readily acknowledged and geochemical studies attempt
to avoid meteoric cements. However, Immenhauser et al. (2003) pro­
posed the idea that sea level’s control on meteoric fluids can potentially
have a more active role in the generation of carbon isotopic trends. Sea
level can influence the degree of basin restriction and therefore the ratio
of meteoric fluids to sea water. During sea level lows, basin restriction
can create an isotopically evolved fluid (low carbon isotopic values due
to increased meteoric influence). As sea level rises, decreased basin re­
striction and/or shifting shoreline leads to decreased meteoric influence
and higher carbon isotopic values in any carbonate sediments produced
at the time. The result is a positive carbon isotopic excursion that has the
rising limb associated with transgression (TST).
Sea level can also influence carbon isotopic patterns because it
controls the production, export, mixing, and type of carbonate sedi­
ments. Aragonite tends to form in the shallowest water settings and
typically has carbon isotopic values 2–3‰ higher than the low magne­
sium calcite (LMC) that forms on the flank of the carbonate banks (Swart
and Eberli, 2005). Based on these relationships and observations from
the Great Bahama Banks, Swart (2008) argued that sea level change
would result in positive carbon isotopic excursions that get preserved
even after the aragonite is recrystallized to LMC after burial. In this
model, as sea level rises, aragonite production increases and carbon
isotopic values increase. Again, we get a positive carbon isotope
excursion with the rising limb in the transgressive systems tract.
In all of these scenarios, the result is the inverse of the one that we
document at Union Furnace. We propose two possible explanations for
the observed relationship between carbon isotopes and sequence stra­
tigraphy in BR 1 and BR 2 at Union Furnace: 1) increased weathering
fluxes during HST fueling marine primary productivity and/or 2) a
spatial gradient in marine δ13CDIC with higher values nearshore.
Carbon isotopic values could be increasing in the HST due to
increased marine primary productivity and a corresponding increase in
organic carbon burial. Normally, primary productivity and enhanced
organic carbon burial is thought to increase during transgression as
described above. However, it is equally as plausible that nutrients (e.g.
phosphorus and nitrogen) can be sourced from the terrestrial realm.
Progressive shallowing during the HST and the basinward movement of
the shoreline could have led to increased weathering fluxes. Because
adequate shallow water would still have existed for primary producers,
the nutrient supply led to an increase in primary productivity and
eventually an increase in organic carbon burial in these shallow marine
settings. Basin restriction would have enhanced the effect by stalling
mixing with the open ocean.
The second possible explanation is that spatial gradients in δ13CDIC
were the opposite of what we observe in a fully vegetated world. In this
scenario, nearshore settings could have had higher δ13CDIC values than
P.C. Quinton et al.
distal settings. In modern settings, like the Great Bahama Banks or
Florida Bay, nearshore settings have distinctly lower δ13CDIC values due
in part to the high influx of respired 12C from terrestrial vegetation
(Patterson and Walter, 1994; Saltzman and Thomas, 2012). With smaller
amounts of biomass in the terrestrial realm, this might not have been
possible in the Ordovician world. This interpretation is supported by
results from Panchuk et al. (2006) looking at broad spatial gradients
across the Laurentian epicontinental sea during the Katian Stage. Their
study documented that δ13C values in the Pennsylvania to Virginia re­
gion generally increased toward shore and the Taconic Highlands.
There are a variety of reasons why terrestrial fluxes could have
contributed a greater percentage of 13C into the marine realm. Near­
shore settings could have received a greater proportion of 13C from the
weathering of exposed carbonates (e.g. Ainsaar et al., 2010). Meteoric
fluids, which traditionally have lots of respired 12C, would have had
much higher δ13CDIC values during the Late Ordovician. With no root
systems to supply respired 12C, the main source of carbon in meteoric
fluids would have been atmospheric CO2. Meteoric δ13CDIC values would
have on average reflected the atmospheric δ13C values at the time
(Driese and Mora, 2001), which have been estimated to be around − 6‰
to − 8‰ (Tomescu et al., 2009). The presence of high meteoric δ13CDIC
values in the Late Ordovician is supported by geochemical studies
constraining the carbon isotopic composition of meteoric carbonates (e.
g. Tobin and Walker, 1994; Driese and Mora, 2001; Jones et al., 2015).
5.4. A role for early land plants at Union Furnace and more broadly?
Regardless of which proposed mechanism is generating the inverse
relationship between sea level and carbon isotopic trends (and it could
certainly be a combination of those processes), our results do suggest a
possible role for early land plants. The progressive increase in the
amount of vegetation in nearshore environments may have exerted a
significant influence on silicate weathering and supplied important
nutrients into the marine realm (Quirk et al., 2015; Lenton et al., 2016;
Porada et al., 2016). The increasing δ13C values during the highstand
systems tract in sequences BR 1 and BR2 could record intervals of
enhanced marine primary productivity due to the influx of those plant
released nutrients. The early terrestrial biosphere could also have
contributed to an inverse spatial gradient in δ13CDIC values by suppling a
higher percentage of 13C to nearshore marine settings. Terrestrial
organic carbon in the Late Ordovician is estimated to have had higher
δ13C values than marine organic matter. The average δ13C value of
marine organic matter during the Late Ordovician is estimated to be ~
− 28‰, while estimated values for liverworts at the time range from
− 26‰ to − 27‰ (Tomescu et al., 2009). Alternations between recycling
marine and terrestrial organic matter in shallow water settings could
explain some of the 1.5‰ shift we observe between transgressions and
regression at Union Furnace.
If this relationship between early land plants, sea level, and carbon
isotopes exists, the Late Ordovician would reflect a unique interval in
Earth’s history where the limited size and extent of the terrestrial
biosphere allowed it to impact shallow carbonate settings in ways not
observed before or after. The temporary nature of the inverse relation­
ship between sea level and carbon isotopes documented at Union
Furnace is supported by Cambrian-Middle Ordovician δ13C records that
display the predicted relationship with sea level (Buggisch et al., 2003;
Schiffbauer et al., 2017). In both intervals, carbon isotopic values in­
crease during transgressions and decrease during regression. Buggisch
et al. (2003) interpreted this relationship to reflect changes in the
relative proportions of marine organic carbon remineralization verses
carbonate deposition. By the Late Ordovician, with the ever-increasing
influx of terrestrial organic carbon enriched in 13C, land plants could
have influenced nearshore settings and altered the relationship between
sea level and carbon isotopic trends.
The proposed role of early land plants is uncertain and needs to be
tested. The first step should be to determine if a similar relationship
9
Palaeogeography, Palaeoclimatology, Palaeoecology 574 (2021) 110440
between sea level and carbon isotopic trends is documented elsewhere.
The Late Ordovician carbon isotopic record has received significant
attention, but much of it focuses on intervals with positive carbon iso­
topic excursions (e.g. GICE and HICE) or on long term trends. There are a
few studies that cover the same interval (Pl. aculeata – E. quadridactylus
Midcontinent Conodont Biozones) at a high enough resolution to make
comparison of δ13C curves possible (Fig. 5). The stratigraphic interval
with the proposed relationship between sea level and carbon isotopes at
Union Furnace is approximately correlated with records from Oklahoma
(Edwards et al., 2015a, 2015b), Alabama (Quinton et al., 2016), Ten­
nessee (Kozik et al., 2019), and West Virginia (Adiatma et al., 2019)
based on the position of conodont biozones and the nature of the carbon
isotopic curves. This correlation is compelling as it suggests that a
similar pattern is present across eastern North America. However, these
comparisons cannot be made without more chronostratigraphic con­
straints and detailed sequence stratigraphic frameworks. We hope that
our work will motivate future studies to pair high resolution δ13C re­
cords with detailed sedimentological and sequence stratigraphic
frameworks to test the relationship between early land plants, sea level,
and carbon isotopes during this dynamic interval in the Late Ordovician.
5.5. Support for the late Sandbian “baseline shift”
Our results add additional support to the interpretation that the
“baseline shift” in δ13C values in the late Sandbian could reflect a
perturbation to the global carbon reservoir possibly due to early land
plants. The most compelling evidence this study can add is that the
relationship between δ13C values and sea level (in BR 1 and BR 2) ends at
the start of the “baseline shift” identified by Adiatma et al. (2019). This
observation suggests that the ~1‰ increase observed at Union Furnace
does reflect a change in the global carbon reservoir that is overriding the
interpreted regional signal in BR 1 and BR 2.
However, there is an alternative explanation for the decoupling of
sea level and δ13C trends in sequences BR 3 and BR 4. These cycles occur
in deeper water settings where changes in sea level might not be able to
exert as significant of a control on carbon isotopes. In these deep-water
settings, the shoreline (and accompanying terrestrial fluxes) is far
removed and development of restricted water masses is less likely. In
fact, identified facies and δ13C values are correlated, with inner shelf
settings on average 0.55‰ (σ = 0.51‰) and mid-shelf to outer ramp
settings on average at − 0.64‰ (σ = 0.58‰). A student t-test indicates
that the difference in δ13C values between environments is statistically
significant (supplementary materials file). It should be noted that this
increase in baseline δ13C values has been observed at the margin of the
epicontinental sea where records are thought to reflect the open ocean
signal (Quinton et al., 2016). This observation supports the interpreta­
tion that the late Sandbian “baseline shift” does reflect a change in the
global carbon cycle.
6. Conclusions
By coupling a high resolution δ13C record to the detailed sedimen­
tological and sequence stratigraphic framework, we demonstrate that
sea level and local/regional processes can influence carbon isotopic
patterns. The observed decrease in δ13C values in the TST and increase in
values in the HST suggests that sea level was controlling carbon isotopic
values during the BR 1 and BR 2 sequences. This relationship likely re­
flects a combination of factors, but we argue that during regressions,
marine organic carbon burial increased due to enhanced nutrient supply
from nearshore terrestrial environments. The ratio of meteoric fluids
and terrestrial organic carbon was much higher when sea level was low
and the shoreline closer. In the Late Ordovician both meteoric fluids and
organic carbon would have been enriched in 13C relative to the modern.
Consequently, nearshore environments would have had higher δ13CDIC
values than distal settings. This is the opposite of modern shallow car­
bonate settings where the abundance of vascular plants contributes large
P.C. Quinton et al. Palaeogeography, Palaeoclimatology, Palaeoecology 574 (2021) 110440

Fig. 5. Regional comparison of bulk carbonate carbon isotopic trends for the late Sandbian. Correlation is approximate based on the position of the Pl. aculeata and
E. quadridactylus Midcontinent Conodont Biozone boundary and the correlation of that boundary to the B. gerdae North Atlantic Conodont Biozone based on Cooper
and Sadler (2012). Sections include the Interstate-35 section in Oklahoma (Edwards et al., 2015a, 2015b), the Tidwell Hollow section in Alabama (Quinton et al.,
2016), Evans Ferry section in Tennessee (Kozik et al., 2019), the Germany Valley section in West Virginia (Adiatma et al., 2019), and the Union Furnace section in
Pennsylvania (this study). The blue shaded region represents the stratigraphic interval in the Union Furnace section where sea level is influencing δ13C trends. The
green shaded region is the baseline shift identified by Adiatma et al. (2019). These intervals are approximately identified in the other sections based on the position of
conodont biozones and the nature of the carbon isotopic curves. North Atlantic Conodont Biozones for Oklahoma and Tennessee are as identified in Edwards et al.,
2015a, 2015b) and Kozik et al. (2019) respectively. Midcontinent Conodont Biozones for Alabama and West Virginia are as identified in Quinton et al. (2016) and
Adiatma et al. (2019) respectively. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

quantities of 12C. Declaration of Competing Interest

Our results also suggest that non-vascular plants could have played a
significant role in the Ordovician carbon cycle by supplying nutrients
and organic carbon enriched in 13C. This study adds support to the idea
that the late Sandbian increase in δ13C values reflects a change in the
global carbon cycle. We demonstrate that the relationship between sea
level and carbon isotopes at Union Furnace ends when this “baseline
shift” begins. We argue that this observation could reflect the fact that
the global signal is overriding the local/regional signal observed in se­
quences BR 1 and BR 2. Furthermore, the lack of relationship between
sea level and δ13C values in this part of the section (above 61.5 m) in­
dicates that sea level is not driving the “baseline shift”. In short, there is
a perturbation to the global carbon cycle that falls at a time when land
plants were increasing in abundance and diversity. As suggested by
previous studies, there is likely a cause-and-effect relationship and we
are seeing land plants’ first impact on the global carbon cycle.
Finally, these results and interpretations are only possible because of
detailed sampling coupled with sedimentological observations. Sam­
pling at regular intervals that are not directly tied to sedimentological
observations can show general trends. Only by targeting genetically
related packages of rock and using a sliding sampling resolution can one
capture δ13C patterns within those packages and constrain the rela­
tionship between carbon isotopic trends and environmental processes.
None.
Acknowledgements
This study was funded by the SUNY Potsdam Presidential Scholars
program, the Kilmer Fund, and the Neil R. O’Brien & William T.
Kirchgasser Undergraduate Research Fund at SUNY Potsdam. This
manscript benefited from thoughtful reviews by Editor Thomas Algeo,
William McMahon, and one anonymous reviwer. We thank Emily Downs
for assistance with field work, Kenneth MacLeod for analyses at the
University of Missouri Stable Isotope and Biogeochemistry Laboratory,
and Sheetz Inc. for logistical support.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.palaeo.2021.110440.
References
Adiatma, Y.D., Saltzman, M.R., Young, S.A., Griffith, E.M., Kozik, N.P., Edwards, C.T.,
Leslie, S.A., Bancroft, A.M., 2019. Did early land plants produce a stepwise change in
atmospheric oxygen during the Late Ordovician (Sandbian ~485 Ma)? Palaeogeogr.
Palaeoclimatol. Palaeoecol. 534 https://doi.org/10.1016/j.palaeo.2019.109341.
Ainsaar, L., Kaljo, D., Martma, T., Meidla, T., Männik, P., Nõlvak, J., Tinn, O., 2010.
Middle and Upper Ordovician carbon isotope chronostratrigraphy in Baltoscandia: a

10
P.C. Quinton et al.
correlation standard and clues to environmental history. Palaeogeogr.
Palaeoclimatol. Palaeoecol. 294, 189–201. https://doi.org/10.1016/j.
palaeo.2010.01.003.
Akoglu, H., 2018. User’s guide to correlation coefficients. Turkish J. Emerg. Med. 18 (3),
91–93. https://www.sciencedirect.com/science/article/pii/S2452247318302164.
Algeo, T.J., Scheckler, S.E., 1998. Terrestrial-marine teleconnections in the Devonian:
links between the evolution of land plants, weathering processes, and marine anoxic
events. Philos. Trans. R. Soc. Lond. 353, 113–130. https://doi.org/10.1098/
rstb.1998.0195.
Algeo, T.J., Berner, R.A., Maynard, J.B., Scheckler, S.E., 1995. Late Devonian oceanic
anoxic events and biotic crises: “rooted” in the evolution of vascular land plants. GSA
Today 5 (3), 45–66.
Allan, J.R., Matthews, R.K., 1982. Isotope signatures associated with early meteoric
diagenesis. Sedimentology 29, 797–817. https://doi.org/10.1111/j.1365-
3091.1982.tb00085.x.
Banner, J.L., Hanson, G.N., 1990. Calculation of simultaneous isotopic and trace element
variations during water–rock interaction with applications to carbonate diagenesis.
Geochim. Cosmochim. Acta 54, 3123–3137. https://doi.org/10.1016/0016-7037
(90)90128-8.
Bergström, S.M., Chen, X., Gutierrez-Marco, J.C., Dronov, A., 2009. The new
chronostratigraphic classification of the Ordovician System and its relations to major
regional series and stages and the δ13C chemostratigraphy. Lethaia 42, 97–107.
https://doi.org/10.1111/j.1502-3931.2008.00136.x.
Bergström, S.M., Schmitz, B., Saltzman, M.R., Huff, W.D., 2010. The Upper Ordovician
Guttenberg δ13C excursion (GICE) in North American and Baltoscandia: occurrence,
chronostratigraphic significance, and paleoenvironmental relationships. In:
Finney, S.C., Berry, W.B.N. (Eds.), The Ordovician System, The Geological Society of
America Special Paper 466, pp. 37–67.
Bergström, S.M., Saltzman, M.R., Leslie, S.A., Ferretti, A., Young, S.A., 2015. Trans-
Atlantic application of the Baltic Middle and Upper Ordovician carbon isotope
zonation. Est. J. Earth Sci. 64, 8–12.
Berner, R.A., 1998. The carbon cycle and carbon dioxide over Phanerozoic time: the role
of land plants. Philos. Trans. R. Soc. Lond. B Biol. Sci. 353 (1365), 75–82. https://
doi.org/10.1098/rstb.1998.0192.
Boucot, A.J., Gray, J., 2001. A critique of Phanerozoic climatic models involving changes
in the CO2 content of the atmosphere. Earth Sci. Rev. 56 (1–4), 1–159. https://doi.
org/10.1016/S0012-8252(01)00066-6.
Brett, C.E., Baird, G.C., 2002. Revised stratigraphy of the Trenton Group in its type area,
central New York State: sedimentology and tectonics of a Middle Ordovician shelf-to-
basin succession. Phys. Chem. Earth, Parts A/B/C 27 (1–3), 231–263. https://doi.
org/10.1016/S1474-7065(01)00007-9.
Brett, C.E., McLaughlin, P.I., Cornell, S.R., Baird, G.C., 2004. Comparative sequence
stratigraphy of two classic Upper Ordovician successions, Trenton Shelf (New
York–Ontario) and Lexington Platform (Kentucky–Ohio): implications for eustasy
and local tectonism in eastern Laurentia. Palaeogeogr. Palaeoclimatol. Palaeoecol.
210 (2–4), 295–329. https://doi.org/10.1016/j.palaeo.2004.02.038.
Brett, C.E., Aucoin, C.D., Dattilo, B.F., Freeman, R.L., Hartshorn, K.R., McLaughlin, P.I.,
Schwalbach, C.E., 2020. Revised sequence stratigraphy of the upper Katian Stage
(Cincinnatian) strata in the Cincinnati Arch reference area: Geological and
paleontological implications. Palaeogeogr. Palaeoclimatol. Palaeoecol. 540, 109483.
https://doi.org/10.1016/j.palaeo.2019.109483.
Buggisch, W., Keller, M., Lehnert, O., 2003. Carbon isotope record of Late Cambrian to
Early Ordovician carbonates of the Argentine Precordillera. Palaeogeogr.
Palaeoclimatol. Palaeoecol. 195 (3–4), 357–373.
Burdige, D.J., 2005. Burial of terrestrial organic matter in marine sediments: A re-
assessment. Glob. Biogeochem. Cycles 19 (4). https://doi.org/10.1029/
2004GB002368.
Cooper, R.A., Sadler, P.M., 2012. The Ordovician Period. In: Gradstein, F.M., Ogg, J.G.,
Schimtz, M.D., Ogg, G.M. (Eds.), The Geologic Time Scale 2012. Elsevier,
Amsterdam, pp. 489–523.
Dahl, T.W., Arens, S.K., 2020. The impacts of land plant evolution on Earth’s climate and
oxygenation state–An interdisciplinary review. Chem. Geol. 547 https://doi.org/
10.1016/j.chemgeo.2020.119665.
Davies, N.S., Gibling, M.R., 2010. Cambrian to Devonian evolution of alluvial systems:
The sedimentological impact of earliest land plants. Earth Sci. Rev. 98 (3–4),
171–200. https://doi.org/10.1016/j.earscirev.2009.11.002.
Driese, S.G., Mora, C.I., 2001. Diversification of Siluro-Devonian plant traces in paleosols
and influence on estimates of paleoatmospheric CO2 levels. In: Gensel, P.G.,
Edwards, D. (Eds.), Plants Invade the Land: Evolutionary and Environmental
Perspectives, pp. 237–253. https://doi.org/10.7312/gens11160.
Droser, M.L., Bottjer, D.J., 1986. A semiquantitative field classification of ichnofabric.
J. Sediment. Res. 4 (1986), 558–559. https://pubs.geoscienceworld.org/sepm/
jsedres/article-abstract/56/4/558/113776/A-semiquantitative-field-classification-
of?redirectedFrom=PDF.
Edwards, D., Feehan, J., Smith, D.G., 1983. A late Wenlock flora from Co. Tipperary,
Ireland. Bot. J. Linn. Soc. 86 (1–2), 19–36. https://doi.org/10.1111/j.1095-
8339.1983.tb00715.x.
Edwards, C.T., Saltzman, M.R., Leslie, S.A., Bergström, S.M., Sedlacek, A.R., Howard, A.,
Bauer, J.A., Sweet, W.C., Young, S.A., 2015a. Strontium isotope (87Sr/86Sr)
stratigraphy of Ordovician bulk carbonate: implications for preservation of primary
seawater values. Bulletin 127 (9–10), 1275–1289. https://doi.org/10.1130/
B31149.1.
Edwards, D., Cherns, L., Raven, J.A., 2015b. Could land-based early photosynthesizing
ecosystems have bioengineered the planet in mid-Palaeozoic times? Palaeontology
58 (5), 803–837. https://doi.org/10.1111/pala.12187.
11
Palaeogeography, Palaeoclimatology, Palaeoecology 574 (2021) 110440
Eltom, H.A., Gonzalez, L.A., Hasiotis, S.T., Rankey, E.C., Cantrell, D.L., 2018.
Paleogeographic and paleo-oceanographic influences on carbon isotope signatures:
Implications for global and regional correlation, Middle-Upper Jurassic of Saudi
Arabia. Sediment. Geol. 364, 89–102. https://doi.org/10.1016/j.
sedgeo.2017.12.011.
Fanton, K.C., Holmden, C., 2007. Sea-level forcing of carbon isotope excursions in epeiric
seas: implications for chemostratigraphy. Can. J. Earth Sci. 44, 807–818. https://doi.
org/10.1139/e06-122.
Gensel, P.G., Kotyk, M.E., Basinger, J.F., 2001. Morphology of above- and below-ground
structures in Early Devonian (Pragian–Emsian) Plants. In: Gensel, P.G., Edwards, D.
(Eds.), Plants Invade the Land: Evolutionary and Environmental Perspectives.
https://doi.org/10.7312/gens11160.
Gerrienne, P., Gensel, P.G., Strullu-Derrien, C., Lardeux, H., Steemans, P., Prestianni, C.,
2011. A simple type of wood in two Early Devonian plants. Science 333 (6044), 837.
https://doi.org/10.1126/science.1208882.
Glasspool, I.J., Scott, A.C., 2010. Phanerozoic concentrations of atmospheric oxygen
reconstructed from sedimentary charcoal. Nat. Geosci. 3 (9), 627–630. https://doi.
org/10.1038/ngeo923.
Hayes, J.M., Popp, B.N., Takigiku, R., Johnson, M.W., 1989. An isotopic study of
biogeochemical relationships between carbonates and organic carbon in the
Greenhorn Formation. Geochim. Cosmochim. Acta 53, 2961–2972. https://doi.org/
10.1016/0016-7037(89)90172-5.
Holland, S.M., Patzkowsky, M.E., 1996. Sequence stratigraphy and long-term
paleoceanographic changes in the Middle and Upper Ordovician of the eastern
United States. Geol. Soc. Am. Spec. Pap. 306, 117–128.
Holmden, C., Creaser, R.A., Muehlenbachs, K., Leslie, S.A., Bergström, S.M., 1998.
Isotopic evidence for geochemical decoupling between ancient epeiric seas and
bordering oceans: Implications for secular curves. Geology 26 (6), 567–570. https://
doi.org/10.1130/0091-7613(1998)026<0567:IEFGDB>2.3.CO;2.
Immenhauser, A., Della Prorta, G., Kenter, J.A.M., Bahamonde, J.R., 2003. An alternative
model for positive shifts in shallow-marines carbonate δ13C and δ18O. Sedimentology
50, 953–959. https://doi.org/10.1046/j.1365-3091.2003.00590.x.
James, N.P., Jones, B., 2015. Origin of carbonate sedimentary rocks. Am. Geophys.
Union 464. ISBN: 978-1-118-65270-1.
Jarvis, I., Gale, A.S., Jenkyns, H.C., Pearce, M.A., 2006. Secular variation in Late
Cretaceous carbon isotopes: a new δ13C carbonate reference curve for the
Cenomanian-Campanian (99.6-70.6 Ma). Geol. Mag. 143 (5), 561–608. https://doi.
org/10.1017/S0016756806002421.
Jenkyns, H.C., 1996. Relative sea-level change and carbon isotopes: data from the Upper
Jurassic (Oxfordian) of central and Southern Europe. Terra Nova 8, 75–85. https://
doi.org/10.1111/j.1365-3121.1996.tb00727.x.
Jones, D.S., Creel, R.C., Rios, B., Santiago Ramos, D.P., 2015. Chemostratigraphy of an
Ordovician–Silurian carbonate platform: δ13C records below glacioeustatic exposure
surfaces. Geol 43 (1), 59–62.
Katz, D.A., Buoniconti, M.R., Montañez, I.P., Swart, P.K., Eberli, G.P., Smith, L.B., 2007.
Timing and local perturbations to the carbon pool in the lower Mississippian
Madison Limestone, Montana and Wyoming. Palaeogeogr. Palaeoclimatol.
Palaeoecol. 256, 231–253. https://doi.org/10.1016/j.palaeo.2007.02.048.
Kennedy, K.L., Gibling, M.R., Eble, C.F., Gastaldo, R.A., Gensel, P.G., Werner-
Zwanziger, U., Wilson, R.A., 2013. Lower Devonian coaly shales of northern New
Brunswick, Canada: plant accumulations in the early stages of Terrestrial
colonization. J. Sediment. Res. 83 (12), 1202–1215. https://doi.org/10.2110/
jsr.2013.86.
Kenrick, P., Wellman, C.H., Schneider, H., Edgecombe, G.D., 2012. A timeline for
terrestrialization: consequences for the carbon cycle in the Palaeozoic. Philosoph.
Transact. Royal Society B 367 (1588), 519–536. https://doi.org/10.1098/
rstb.2011.0271.
Kleinhans, M.G., de Vries, B., Braat, L., van Oorschot, M., 2018. Living landscapes:
muddy and vegetated floodplain effects on fluvial pattern in an incised river. Earth
Surf. Process. Landf. 43 (14), 2948–2963. https://onlinelibrary.wiley.com/doi/full/
10.1002/esp.4437.
Kozik, N.P., Young, S.A., Bowman, C.N., Saltzman, M.R., Them II, T.R., 2019.
Middle–Upper Ordovician (Darriwilian–Sandbian) paired carbon and sulfur isotope
stratigraphy from the Appalachian Basin, USA: Implications for dynamic redox
conditions spanning the peak of the Great Ordovician Biodiversification Event.
Palaeogeogr. Palaeoclimatol. Palaeoecol. 520, 188–202.
Kump, L.R., Arthur, M.A., 1999. Interpreting carbon-isotope excursions: carbonates and
organic matter. Chem. Geol. 161, 181–198. https://doi.org/10.1016/S0009-2541
(99)00086-8.
Kump, L.R., Brantley, S.L., Arthur, M.A., 2000. Chemical weathering, atmospheric CO2,
and climate. Annu. Rev. Earth Planet. Sci. 28 (1), 611–667. https://doi.org/
10.1146/annurev.earth.28.1.611.
Laughrey, C.D., Kostelnik, J., Doden, A.G., Gold, D.P., Harper, J.A., Morrow, L.,
Pedlow, G., 2017. Trenton and black river carbonates in the union furnace roadcut,
state route 453, Huntingdon County, Pennsylvania. In: 82nd Annual Field Conference
of Pennsylvania Geologists Pre-conference Field Trip Guidebook, pp. 1–56.
Lenton, T.M., Crouch, M., Johnson, M., Pires, N., Dolan, 2012. First plants cooled the
Ordovician. Nat. Geosci. 5, 86–89. https://doi.org/10.1038/ngeo1390.
Lenton, T.M., Dahl, T.W., Daines, S.J., Mills, B.J., Ozaki, K., Saltzman, M.R., Porada, P.,
2016. Earliest land plants created modern levels of atmospheric oxygen. Proc. Natl.
Acad. Sci. 113 (35), 9704–9709. https://doi.org/10.1073/pnas.1604787113.
Malkowski, K., Racki, G., 2009. A global biogeochemical perturbation across the
Silurian–Devonian boundary: Ocean–continent–biosphere feedbacks. Palaeogeogr.
Palaeoclimatol. Palaeoecol. 276, 244–254. https://doi.org/10.1016/j.
palaeo.2009.03.010.
P.C. Quinton et al.
Marshall, J.D., 1992. Climatic and oceanographic isotopic signals from the carbonate
rock record and their preservation. Geol. Mag. 129 (2), 143–160. https://doi.org/
10.1017/S0016756800008244.
McMahon, W.J., Davies, N.S., 2018. Evolution of alluvial mudrock forced by early land
plants. Science 359 (6379), 1022–1024. https://science.sciencemag.org/content
/359/6379/1022.abstract.
Melchin, M.J., Holmden, C., 2006. Carbon isotope chemostratigraphy of the llandovery
in arctic canada: implications for global correlation and sea-level change. gff 128,
173–180. https://doi.org/10.1080/11035890601282173.
metzger, g.j., fike, d.a., 2013. Techniques for assessing spatial heterogeneity of carbonate
δ13c values: implications for craton-wide isotope gradients. Sedimentology 60,
1405–1431. https://doi.org/10.1111/sed.12033.
Morris, J.L., Puttick, M.N., Clark, J.W., Edwards, D., Kenrick, P., Pressel, S., Wellman, C.
H., Yang, Z., Schneider, H., Donoghue, P.C., 2018. The timescale of early land plant
evolution. Proc. Natl. Acad. Sci. 115 (10), E2274–E2283. https://doi.org/10.1073/
pnas.1719588115.
Panchuk, K.M., Holmden, C., Kump, L.R., 2005. Sensitivity of the epeiric sea carbon
isotope record to local-scale carbon cycle processes: tales from the Mohawkian Sea.
Palaeogeography, Paleaclimatology, Palaeoecology 228, 320–337. https://doi.org/
10.1016/j.palaeo.2005.06.019.
Panchuk, K.M., Holmden, C.E., Leslie, S.A., 2006. Local controls on carbon cycling in the
Ordovician Midcontinent region of North America, with implications for carbon
isotope secular curves. J. Sediment. Res. 76, 200–211. https://doi.org/10.2110/
jsr.2006.017.
Patchen, D.G., Hickman, J.B., Harris, D.C., Drahovzal, J.A., Lake, P.D., Smith, L.B.,
Nyahay, R., Schulze, R., Riley, R.A., Baranoski, M.T., Wickstrom, L.H., Laughrey, C.
D., Kostelnik, J., Harper, J.A., Avary, K.L., Bocan, J., Hohn, M.E., McDowell, R.,
2006. A geologic play book for Trenton-Black River Appalachian Basin exploration.
Department of Energy (DOE Award Number DE-FC26-03NT41856) Report. https
://www.netl.doe.gov/sites/default/files/2018-04/NT41856-Final-Rpt.pdf.
Patterson, W.P., Walter, L.M., 1994. Depletion of 13C in seawater ΣCO2 on modern
carbonate platforms: Significance for the carbon isotopic record of carbonates.
Geology 22, 885–888. https://doi.org/10.1130/0091-7613(1994)022%3C0885:
DOCISC%3E2.3.CO;2.
Porada, P., Lenton, T.M., Pohl, A., Weber, B., Mander, L., Donnadieu, Y., Beer, C.,
Pöschl, U., Kleidon, A., 2016. High potential for weathering and climate effects of
non-vascular vegetation in the Late Ordovician. Nat. Commun. 7 (1), 1–13. https://
doi.org/10.1038/ncomms12113.
Quinton, P.C., Herrmann, A.D., Leslie, S.A., MacLeod, K.G., 2016. Carbon cycling across
the southern margin of Laurentia during the Late Ordovician. Palaeogeogr.
Palaeoclimatol. Palaeoecol. 458, 63–76. https://doi.org/10.1016/j.
palaeo.2015.08.020.
Quirk, J., Leake, J.R., Johnson, D.A., Taylor, L.L., Saccone, L., Beerling, D.J., 2015.
Constraining the role of early land plants in Palaeozoic weathering and global
cooling. Proc. R. Soc. B Biol. Sci. 282 (1813), 20151115. https://doi.org/10.1098/
rspb.2015.1115.
Read, J.F., 1985. Carbonate platform facies models. Am. Assoc. Pet. Geol. Bull. 66,
860–878. https://doi.org/10.1306/AD461B79-16F7-11D7-8645000102C1865D.
Rubinstein, C.V., Gerrienne, P., de la Puente, G.S., Astini, R.A., Steemans, P., 2010. Early
Middle Ordovician evidence for land plants in Argentina (eastern Gondwana). New
Phytol. 188 (2), 365–369. https://nph.onlinelibrary.wiley.com/doi/full/10.1111/
j.1469-8137.2010.03433.x.
Saltzman, M.R., Edwards, C.T., 2017. Gradients in the carbon isotopic composition of
Ordovician shallow water carbonates: A potential pitfall in estimates of ancient CO2.
Earth Planet. Sci. Lett. 464, 46–54. https://doi.org/10.1016/j.epsl.2017.02.011.
Saltzman, M.R., Thomas, E., 2012. Carbon isotope stratigraphy. In: Gradstein, F.M.,
Ogg, J.G., Schmitz, M., Ogg, G. (Eds.), The Geologic Time Scale, pp. 207–232.
https://doi.org/10.1016/B978-0-444-59425-9.00011-1.
Schiffbauer, J.D., Huntley, J.W., Fike, D.A., Jeffrey, M.J., Gregg, J.M., Shelton, K.L.,
2017. Decoupling biogeochemical records, extinction, and environmental change
during the Cambrian SPICE event. Sci. Advanc. 3 (3), e1602158.
12
Palaeogeography, Palaeoclimatology, Palaeoecology 574 (2021) 110440
Schrag, D.P., Higgins, J.A., Macdonald, F.A., Johnston, D.T., 2013. Authigenic carbonate
and the history of the global carbon cycle. Science 339 (February), 540–543. https://
doi.org/10.1126/science.1229578.
Scotese, C.R., 2004. A continental drift flipbook. J. Geol. 112, 729–742. https://www.
journals.uchicago.edu/doi/abs/10.1086/424867.
Sell, B.K., Samson, S.D., Mitchell, C.E., MacLaughlin, P.J., Koenig, A.E., Leslie, S.A.,
2015. Stratigraphic correlations using trace elements in apatite from Late Ordovician
(Sandbian-Katian) K-bentonites of eastern North American. Geol. Soc. Am. Bull. 127,
1259–1274. https://doi.org/10.1130/B31194.1.
Sonnenfeld, M.D., 1996. Sequence evolution and hierarchy within the lower
Mississippian Madison Limestone of Wyoming. In: Longman, M.W., Sonnenfeld, M.D.
(Eds.), Paleozoic systems of the Rocky Mountain region. Society for Economic
Paleontologist and Mineralogist (Society for Sedimentary Geology) Rocky Mountain
Section, pp. 165–192.
Stein, W.E., Berry, C.M., Morris, J.L., Hernick, L.V., Mannolini, F., Ver Straeten, C.,
Landing, E., Marshall, J.E.A., Wellman, C.H., Beerling, D., Leake, J.R., 2020. Mid-
Devonian Archaeopteris roots signal revolutionary change in earliest fossil forests.
Curr. Biol. 30, 1–11. https://doi.org/10.1016/j.cub.2019.11.067.
Swart, P.K., 2008. Global synchronous changes in the carbon isotopic composition of
carbonate sediments unrelated to changes in the global carbon cycle. Proc. Nation.
Acad. Sci. 105 (37), 13741–13745.
Swart, P.K., Eberli, G., 2005. The nature of the δ13C of periplatform sediments:
implications for stratigraphy and the global carbon cycle. Sediment. Geol. 175,
115–129. https://doi.org/10.1016/j.sedgeo.2004.12.029.
Swart, P.K., Oehlert, A.M., 2018. Revised interpretations of stable C and O patterns in
carbonate rocks resulting from meteoric diagenesis. Sediment. Geol. 364, 14–23.
https://doi.org/10.1016/j.sedgeo.2017.12.005.
Swart, P.K., Reijmer, J.J.G., Otto, T., 2009. A reevaluation of facies on Great Bahama
Bank II: variations in the δ13C, δ18O and mineralogy of surface sediments. Int. Assoc.
Sedimentol. Spec. Publ. 41, 47–59.
Tobin, J., Walker, K.R., 1994. Meteoric diagenesis below a submerged platform:
implications for δ13C compositions prior to pre-vascular plant evolution, Middle
Ordovician, Alabama, USA. Sediment. Geol. 90 (1–2), 95–111. https://doi.org/
10.1016/0037-0738(94)90019-1.
Tomescu, A.M.F., Rothwell, G.W., 2006. Wetlands before tracheophytes: Thalloid
terrestrial communities of the Early Silurian Passage Creek biota (Virginia). In:
Greb, S.F., Dimichele, W.A. (Eds.), Wetlands through time, Special Papers-Geological
Society of American, 399, p. 41.
Tomescu, A.M., Pratt, L.M., Rothwell, G.W., Strother, P.K., Nadon, G.C., 2009. Carbon
isotopes support the presence of extensive land floras pre-dating the origin of
vascular plants. Palaeogeogr. Palaeoclimatol. Palaeoecol. 283 (1–2), 46–59. https://
doi.org/10.1016/j.palaeo.2009.09.002.
Torsvik, T.H., Cocks, R.M., 2013. New global palaeogeographical reconstructions for the
Early Palaeozoic and their generation. In: Harper, D.A.T., Servais, T. (Eds.), Early
Palaeozoic Biogeography and Palaeogeography. Geological Society of London,
Memoirs 38, pp. 5–24. https://mem.lyellcollection.org/content/38/1/5.short.
Walker, J.D., Geissman, J.W., Bowring, S.A., Babcock, L.E., 2018. Geologic Time Scale v.
5.0. Geological Society of America. https://doi.org/10.1130/2018.CTS005R3C.
Weissert, H., 1989. C-isotope stratigraphy, a monitor of paleoenvironmental change: A
case study from the Early Cretaceous. Surv. Geophys. 10, 1–61. https://link.springer.
com/article/10.1007/BF01901664.
Young, S.A., Saltzman, M.R., Ausich, W.I., Desrochers, A., Kaljo, D., 2010. Did changes in
atmospheric CO2 coincide with latest Ordovician glacial–interglacial cycles?
Palaeogeogr. Palaeoclimatol. Palaeoecol. 296 (3–4), 376–388.
Young, S.A., Saltzman, M.R., Bergström, S.M., 2005. Upper Ordovician (Mohakian)
carbon isotope (δ13C) stratigraphy in eastern and central North America: regional
expression of a perturbation of the global carbon cycle. Palaeogeogr. Palaeoclimatol.
Palaeoecol. 222, 53–76. https://doi.org/10.1016/j.palaeo.2005.03.008.
Young, S.A., Saltzman, M.R., Bergström, S.M., Leslie, S.A., Xu, C., 2008. Paired δ13Ccarb
and δ13Corg records of Upper Ordovician (Sandbian-Katian) carbonates in North
America and China: implications for paleoceanographic change. Palaeogeogr.
Palaeoclimatol. Palaeoecol. 270, 166–178.