1 s2.0 S0012825220303780 Main

Earth-Science Reviews 210 (2020) 103332
Contents lists available at ScienceDirect
Earth-Science Reviews
journal homepage: www.elsevier.com/locate/earscirev
Review Article
Catastrophic soil loss associated with end-Triassic deforestation T

a,⁎ a a,1 b c
B. van de Schootbrugge , C.M.H. van der Weijst , T.P. Hollaar , M. Vecoli , P.K. Strother ,
N. Kuhlmannd, J. Theind, H. Visschera, H. van Konijnenburg-van Citterta, M.A.N. Schobbena,
A. Sluijsa, S. Lindströme
a
Department of Earth Sciences, Utrecht University, Marine Palynology & Paleoceanography, Princetonlaan 8a, 3584, CB, Utrecht, the Netherlands
b
Exploration Technical Services Department, Saudi Aramco, Dhahran, Saudi Arabia
c
Weston Observatory, Boston College, United States
d
Institute for Geology, University of Bonn, Nussallee 8, 53115 Bonn, Germany
e
GEUS–Geological Survey of Denmark and Greenland, Øster Voldgade 10, DK-1350 Copenhagen, Denmark
ABSTRACT
Soils are a crucial link between the atmosphere, biosphere, and geosphere. Any disturbance to the health of soils will severely impact plants as well as a multitude of
organisms living in or on soils, such as fungi, bacteria, and insects. Catastrophic soil loss is thought to have played a pivotal role during mass-extinction events as a
result of major deforestation, but the exact feedbacks remain elusive. Here, we assess the role of soil loss during the end-Triassic mass-extinction event based on proxy
data obtained from four sediment sections recovered from France, Germany, and Denmark. Clay mineral and palynological data indicate a strong increase in erosion
during the latest Rhaetian with the influx of kaolinite and abundantly reworked Palaeozoic and Neoproterozoic organic matter. Based on a new timeline, these
changes were coeval with intense volcanic activity in the Central Atlantic Magmatic Province (CAMP). In addition to vegetation dieback, repeated forest fires, as well
as widespread seismic activity related to CAMP emplacement led to landscape destruction triggering removal of soils. The biological degradation of fern spore walls
by fungi and bacteria, a process coupled to organic matter decay in soils, strongly decreased across the T/J boundary. We interpret this counter-intuitive result as
evidence for rapid and widespread removal of soils. Taken together, CAMP induced environmental changes led to profound changes in erosion and removal of soils,
while soil resilience during the Hettangian appears to have proceeded hand in hand with recovery in Jurassic seas.
1. Introduction as well as prevailing climatic conditions, mainly temperature and

runoff, and the type of vegetation cover (Nesbitt and Young, 1984;
Soils are a key factor in the health of global ecosystems. Acting as an Drever, 1994). Weathering of continental landmasses is thus a pre-
interface between the atmosphere, geosphere, and biosphere, soil bio- requisite to form soils in the first place and should be clearly dis-
diversity is likely higher than diversity in above-ground ecosystems tinguished from erosion, which is the physical removal and transport of
(Freckman et al., 1997; Orgiazzi et al., 2016). Not only do soils play a regolith and soil by gravity, wind, water or ice.
vital role in influencing plant diversity, but healthy soils are also Soil loss via erosion is the most extreme form of soil disturbance,
characterized by diverse communities of among others fungi, bacteria, which can trigger profound changes in ecosystems. Catastrophic soil
arthropods, and nematodes that together control nutrient fluxes in and loss can be due to several mechanisms, including earthquakes and forest
out of soils. Many of these interactions reinforce each other with plants fires, but the foremost disturbance is deforestation. Loss of vegetation
actively shaping soil formation, chemistry, and retention of nutrients cover leads to rapid removal of topsoils during rainfall. Today, elevated
(Drever, 1994; Van Nuland et al., 2016). In addition, soils store very rates of erosion are, for example, particularly evident on Madagascar
large quantities of carbon that upon loss can be released to the atmo- where anthropogenic destruction of tropical rainforests leads to ex-
sphere and oceans (Scharlemann et al., 2014). treme soil loss (Cox et al., 2010). Extensive sediment plumes can be
Soil formation is not only a biological process, but it is closely tied seen entering the Indian Ocean using satellite imagery; the landscape of
to the chemical weathering of crystalline rock at the Earth's surface, Madagascar proverbally “bleeds to death” as astronauts on the ISS re-
which drives large-scale patterns in mineralogy of clay particle as- marked in 2004 while observing the Betsiboka estuary (see also https://
semblages (Nesbitt and Young, 1984). Chemical weathering depends on earthobservatory.nasa.gov/images/4388/betsiboka-estuary-
bedrock mineralogy, texture, permeability, and the age of the substrate, madagascar).
⁎
Corresponding author.
E-mail address: [email protected] (B. van de Schootbrugge).
1
Now at the University of Exeter, Exeter, United Kingdom.
https://doi.org/10.1016/j.earscirev.2020.103332
Received 9 December 2019; Received in revised form 25 August 2020; Accepted 26 August 2020
Available online 01 September 2020
0012-8252/ © 2020 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY license
(http://creativecommons.org/licenses/by/4.0/).
B. van de Schootbrugge, et al. Earth-Science Reviews 210 (2020) 103332
In currently accepted scenarios for the end-Permian and end- driving ecosystem change.
Triassic extinctions, deforestation and soil loss are key triggers driving Here, we use palynological data for the end-Triassic mass-extinction
the extinction of species on land and likely played a role in the eu- event (ETME; 201.6 Ma) from four different cores from Denmark
trophication and spread of shallow marine anoxia (Visscher et al., 1996; (Stenlille), France (Boust), and Germany (Schandelah and
Looy et al., 2001; Algeo and Twitchett, 2010; Algeo et al., 2011a). Mingolsheim) as a proxy to examine intense soil erosion and bedrock
Hence, soil loss is regarded as a crucial link between ecosystem dis- weathering related to forest dieback across the Triassic-Jurassic
turbance on land and in the oceans. Elevated erosion rates associated boundary. In addition, we develop a new proxy-record for soil erosion
with the end-Permian extinction have been inferred from a wealth of based on the intensity of biological degradation of palynomorphs by
information on changes in river morphology (Smith, 1995; Retallack soil-borne organisms. Our long-term palynological records allow us to
et al., 2006; Newell et al., 2010; Zhu et al., 2019), as well as strong also trace the resilience of soils following the end-Triassic extinction.
increases in sediment fluxes (Algeo and Twitchett, 2010; Algeo et al.,
2011b). Influx of soil-derived organic debris and organic biomarkers 2. Material and methods
has been attributed to increased soil erosion (Sephton et al., 2005a,
2005b; Wang and Visscher, 2007; French et al., 2012; Sawada et al., 2.1. Material
2012; Kaiho et al., 2016). Eutrophication and widespread anoxia, in-
cluding photic zone euxinia, were likely important proximate kill me- The presented work is based on new palynological analyses from the
chanisms that contributed to the extinction of Permian marine in- Schandelah-1 core (northern Germany) and Boust core (northern
vertebrates (Isozaki, 1997; Grice et al., 2005; Isozaki et al., 2007; Shen France), combined with previously published data from the Stenlille
et al., 2007; Wignall et al., 2010; Kaiho and Koga, 2013). core (Denmark; Lindström et al., 2012) and Mingolsheim core
For the end-Triassic extinction, many of the general triggers and (southern Germany; van de Schootbrugge et al., 2009). The four cores
feedback mechanisms that contributed to extinction (van de form a north to south transect and represent different basins within the
Schootbrugge and Wignall, 2016) have been implicated in recently European Epicontinental Seaway, with the Stenlille core representing
published detailed studies, including work on country-rock basalt in- the Danish Basin, the Schandelah-1 core representing the North German
teractions in the CAMP province (Heimdal et al., 2018), carbon cycle Basin, the Mingolsheim representing the South German Basin and the
perturbations (Bacon et al., 2011; Ruhl et al., 2011; Ruhl and Boust core representing the Paris Basin (Fig. 1).
Kürschner, 2011), terrestrial devastation (Lindström et al., 2012; 2016), The Stenlille structure is a gentle anticline on top of a salt pillar.
possible effects of sulfuric acid on leaf roundness and plant cuticles More than 20 wells were drilled through the T/J-transition encom-
(Bacon et al., 2011; Steinthorsdottir et al., 2017), increases in pCO2 passing the sandstone-dominated Gassum Formation and succeeding
(Steinthorsdottir et al., 2011), and photic zone euxinia (Richoz et al., mudstone-dominated Fjerritslev Formation to assess the reservoir and
2012; Jaraula et al., 2013; Kasprak et al., 2015) as well as widespread cap rock quality, respectively, as the structure is used for storage of
anoxia (Jost et al., 2017; Luo et al., 2018). Based on a marked change in natural gas. The Stenlille-1 well was drilled in 1980 and cored over the
fluvial style in East Greenland, Steinthorsdottir et al. (2012) argued for T/J-transition. The litho-, bio-, and chemostratigraphy of the Stenlille
an increase in run-off, which was perhaps driven by a global decrease in succession in the Danish Basin has been presented in Nielsen (2003)
evapotranspiration and an associated increase in humidity, as indicated and Lindström et al. (2012, 2015, 2017). The lower to middle Rhaetian
by lower stomatal densities on plant leaves. A dominance of C29 ster- Gassum Formation consists of interbedded fine- to medium-grained,
anes in sedimentary organic matter from the marine Kennecott Point occasionally coarse-grained and pebbly sandstones, heteroliths, mud-
section on the Haida Gwaii Islands, Canada, suggests that the main stones and a few thin coaly beds, formed in a shallow marine to coastal
carbon source was from land (Jiao et al., 2008). Deforestation is also environment. It is succeeded by the lowermost part of the Fjerritslev
implicated in the end-Triassic mass-extinction event (van de Formation which consists of dark grey shale. This shale unit was de-
Schootbrugge et al., 2009; van de Schootbrugge and Wignall, 2016), posited during the late Rhaetian transgression and a regionally re-
however its effect on soil erosion remains largely unaddressed and cognized maximum flooding surface (MFS 7 of Nielsen, 2003) is located
difficult to quantify. in the middle of the ca. 3 m thick shale. Palynological assemblages of
Analyses of micro- and macrofloral abundance and diversity in- the Gassum Formation and the dark grey shale at the base of the
dicate severe changes in land plant cover during the latest Rhaetian in Fjerritslev Formation are assigned to the Granuloperculatipollis-Classo-
NW Europe (van de Schootbrugge et al., 2009; van de Schootbrugge pollis-Perinopollenites (GCP) zone, which is dominated by conifer pollen,
and Wignall, 2016), Greenland (McElwain et al., 1999), and North primarily the nominate taxa (Lindström, 2016). These pollen groups
America (Cirilli et al., 2009; Fowell et al., 1994). Detailed palynological decrease in abundance upwards within the dark grey shale and are
studies of cores and outcrops in Denmark (Lindström et al., 2012), virtually absent in the succeeding zone. Dinoflagellate cysts also de-
Sweden (Lindström and Erlström, 2006), Germany (Heunisch et al., crease dramatically in abundance and Rhaetogonyaulax rhaetica has its
2010), England (Bonis et al., 2010), Hungary (Götz et al., 2009), and last common occurrence (LCO) in the middle of the dark grey shale.
Austria (Bonis et al., 2009) show simultaneous changes in floral as- The succeeding “grey siltstone” interval consists of greenish grey
semblages over a very broad area. Early Rhaetian vegetation in NW siltstones and fine-grained sandstones. Here, primary bedding features
Europe consisted of a diverse community of pollen-producing gym- are undisturbed by bioturbation, allowing for the observation of mul-
nospermous trees and shrubs, including seed ferns, cycads, ginkgo- tiple levels of soft-sediment deformations structures, seismites, which
phytes, gnetaleans, bennettitaleans, and conifers with an understory of can be correlated with similar disturbed intervals across Europe
ferns and fern allies (mosses, horsetails, lycopsids). Palynological as- (Lindström et al., 2015). The seismites are interbedded by intervals of
semblages from the uppermost Rhaetian Triletes Beds in Germany, and undisturbed strata. Wave-ripple cross-lamination indicates deposition
equivalent beds in Denmark and Sweden, show a strong decline in tree at depths above storm wave base (Lindström et al., 2015). The lower c.
pollen abundance and a surge in fern spores dominated by Poly- 3 m of the “grey siltstone” interval contains abundant clay clasts
podiisporites polymicroforatus and various morphotypes of the Deltoi- (Lindström et al., 2017). The palynoflora from the grey siltstone in-
dospora-Concavisporites complex. Across NW Europe the predominantly terval belongs to the Polypodiisporites-Ricciisporites-Deltoidospora (PRD)
arborescent (tree-dominated) flora was replaced during the latest zone, which differs significantly from the previous zone. This zone
Rhaetian by a pioneer vegetation consisting of ferns and fern allies, correlates with the Ricciisporites-Polypodiisporites Zone of Lund (1977)
being dominated by only a few groups of ground ferns (van de and is dominated by Polypodiisporites polymicroforatus, Ricciisporites tu-
Schootbrugge et al., 2009). Taken together these observations suggest berculatus and Deltoidospora spp., while conifer pollen are virtually
that deforestation during the end-Triassic extinction was a key factor in absent (Lindström, 2016). Ricciisporites tuberculatus is common to
2
Fig. 1. Geological base map and location map for the Triassic-Jurassic transition in NW Europe. This reconstruction is based on combining two maps originally
published by Ziegler (1990). The reconstruction uses the Permian base map as an overlay on the Jurassic palaeogeographic marine facies map. The arrows indicate
directions of reworking and probable source areas. (1) Stenlille, Demark. (2) Schandelah, Germany. (3) Mingolsheim, Germany. (4) Boust, France.
abundant in the GCP zone and remains relatively abundant throughout grey sand- and siltstones characterized by pronounced and varied se-
the grey siltstone interval (Lindström, 2016). High levels of Poly- dimentary structures. Liquefaction horizons, described as seismites in
podiisporites polymicroforatus, are restricted to the PRD zone after which Lindström et al. (2015), are common in this interval. Mud-chip con-
it decreases markedly. Cerebropollenites thiergartii is first recorded in the glomerate beds occur throughout the Triletes Beds, with clasts typically
uppermost part of the interval (Lindström et al., 2017). highly rounded and up to 1 cm in diameter. Also, sandstone beds with
The grey siltstone is followed by marine claystones, mudstones and cross-stratification include climbing ripples, suggesting high sediment
fine-grained sand intervals, deposited during another transgressive influx. Although fossils are largely absent, there is evidence of bio-
event. The palynofloras are interpreted to reflect floral recovery, and turbation between 330.35 and 330.00 mbs. The Rhaetian-Hettangian
are assigned to the Calamospora-Conbaculatisporites-Monosulcites (CCM) boundary is placed at a sharp surface (318.6 mbs) at the base of a 70 cm
and Perinopollenites-Deltoidospora-Stereisporites (PDS) zones (Lindström, thick shale which lies within a bioturbated sandstone unit characterized
2016). Kraeuselisporites reissingerii and Ischyosporites variegatus have by increasingly better developed laminations towards the top. The
their FOs in the lowermost part of the PDS zone, and in its upper part Hettangian (318.60–233.30 mbs) consists of a heterolithic succession of
the dinoflagellate cyst R. rhaetica occurs for the last time. The FCO of alternating silt- and sandstone lenses interspersed with organic-rich
Pinuspollenites minimus marks the lower boundary of the Deltoidospora- mudstones and shales, typical of mudflats under tidal influences. Gutter
Perinopollenites-Pinuspollenites (DPPi) zone, which is dominated by Del- casts are common. Several intervals contain thick sandstone beds that
toidospora spp., Perinopollenites elatoides and P. minimus. Lindström are recognized as regional marker beds. The lower part of the Hettan-
(2016) correlated the DPPi zone and the succeeding Perinopollenites- gian from 318.00 to 298.00 consists of laminated dark-grey mudstones
Pinuspollenites (PPi) zone with the Pinuspollenites-Trachysporites Zone of with intervals that contain paper shales. Macrofossil content is of low
Lund (1977), and placed these zones within the post-extinction phase. abundance throughout the Hettangian, with the exception of trace
The Schandelah-1 core was drilled close to the town of fossils, crinoid remains, and sparse ammonite remains. Noticeable in-
Braunschweig in northern Germany and detailed bio-, chemo-, and li- tervals contain the gradual transition into distinctly red-coloured la-
thostratigraphy can be found in van de Schootbrugge et al. (2019). The minated claystones at 258.65–254.80 mbs and 238.00–234.00 mbs.
upper Rhaetian (338.00–318.60 mbs) Exter Formation includes the Apart from the colour, the lithology is not strikingly different from
‘Contorta Beds’ (334.80–332.00 mbs) and the overlying ‘Triletes Beds’ under- and overlying dark-grey laminated shales. It remains unclear
(332.00–318.60 mbs). The former consists of current-rippled sand- whether these colour changes are primary or whether they are diage-
stones with intercalations of thin mud beds. Charcoal and plant mate- netic in origin.
rial are abundant in this unit. The ‘Triletes Beds’ consists of grey to dark The Mingolsheim core was drilled in 1968 (Hettich, 1974; von
3
Hildebrandt and Schweizer, 1992) in the vicinity of Bad Mingolsheim, dried sample was crushed to ~1–3 mm3 fragments. Carbonates and
renamed Bad Schönborn in 1971 (Baden-Württemberg, Southern Ger- silicates were removed by washing the samples twice with 30% HCl and
many). The litho,- bio-, and chemostratigraphic framework for the 40% HF. Where needed, oxidative maceration was performed with by
Mingolsheim core, including trace element and bulk organic carbon KClO3 and 65% HNO3 to remove the bulk of the amorphous organic
isotopes, was presented by Quan et al. (2008) and van de Schootbrugge matter. After neutralizing, the samples were sieved over a 250 μm mesh
et al. (2009). The sampled succession starts with the coarse grained to remove coarse material and subsequently over a 10 μm mesh to re-
continental ‘Rhätsandstein’, a massive sandstone that contains abun- move the finest, amorphous material. The heavy mineral fraction was
dant plant remains. This is overlain by the Contorta Beds, black shales removed from the lighter organic fraction by bringing the light fraction
rich in marine dinoflagellate cysts, such as Rhaetogonyaulax rhaetica, a in suspension in an ultrasonic tub. The remaining light fraction was
marker species for the Rhaetian. The Contorta Beds are overlain by 7 m mounted on glass microscopic slides with glycerin jelly. The surplus
of uniform grey claystones known as the Triletes Beds of latest Rhaetian material was collected in a small glass vial and is stored at the De-
age. These claystones are devoid of any macrofossil remains. The partment of Earth Sciences, Utrecht University for future reference.
transition to the Jurassic is marked by an indurated sandy limestone Samples were analyzed under 1000× magnification with an optical
bed known as the Psilonoten Limestone, named after the first occur- microscope (Leica) and relative abundances were determined by
rence of Jurassic psiloceratid ammonites. The lowermost part of the counting approximately 300 palynomorphs per slide. A subset of sam-
Hettangian appears to be either missing or is highly condensed in the ples was spiked with Lycopodium and these were counted quantita-
Mingolsheim core. The Hettangian and base of the Sinemurian in the tively.
Mingolsheim core consist of organic rich shales. Compared to other
successions in this report, the Hettangian is extremely condensed, 2.3. Biodegradation
suggesting deposition in deeper marine conditions.
The Boust core was drilled in 1986 close to the Hettangian strato- There is no single method for quantifying palynomorph preserva-
type in Grande Hettange, northeastern Lorraine, near the Luxembourg tion and none tailored for geological deep time. Therefore, the meth-
and German borders. The investigated and sampled part of the core odologies we use here are based on a pollen preservation system de-
measures 37.30 m from 69.90 m below surface (mbs) in the uppermost veloped for Quaternary sporomorphs (Havinga, 1967). Typically 5
Norian to 32.20 mbs in the Hettangian. The Rhaetian sequence ranges classes of degradation are distinguished: (1) corrosion, (2) degradation,
from 67.00 m to 38.81 m, representing by far the thickest and most (3) crumpling, (4) physical damage, and (5) intact preservation
complete Rhaetian from all cores analyzed recently in the northeastern (Havinga, 1967; Twiddle and Bunting, 2010). As the aim of this study
Paris Basin (Kuhlmann, 2019). The uppermost Norian (‘Steinmergelk- was to quantify biodegradation across the Rhaetian-Hettangian
euper’) consists generally of light greenish-grey dolomitic marls. It is boundary, the focus was solely on corrosion. The other categories are
truncated by a thin conglomerate, an indicator of the Rhaetian trans- related to transport as opposed to biological degradation in soils. Cor-
gression that is found in all drillcores in the northeastern Paris Basin. rosion of palynomorphs is described as etching or pitting of the exines,
This is overlain by the Contorta Beds of the lower Rhaetian up to whereby the circular to rectangular holes often with scalloped edges are
47.16 m, representing in northern Lorraine the facies of the so called randomly scattered on the surface of the sporomorphs. Generally, this
“Sables de Mortinsart”. Its lower part consists of exceptionally thick form of corrosion only affects the ektexine, the outer layer of the wall of
laminated silty clay, with lenses and layers of silt. From 57.00 to the sporomorph grains. In extreme cases, however, the ektexine may be
55.00 m the proper Sables de Mortinsart occurs. It forms the top of a almost completely eaten away, or leaving only fragments of the ek-
sandy coarsening-up cycle that ends with conglomeratic layers with texine on the intact endexine (Cushing, 1967). Corrosion also includes
evidence for desiccation. Above 50.80 m clay reappears. Its planar la- deterioration a material undergoes as a result of its interaction with its
mination is perturbed by convolute structures of soft sediment de- surroundings, including physical, chemical or biological processes.
formation (seismites). The lower Rhaetian ends with some irregular Microbial attack is one of the more important biological processes
conglomerate beds with intercalated, microfolded layers of silty clays, grouped in this class (Havinga, 1967). Specific, repetitive patterns of
topped by structureless grey clay with Rhaetavicula contorta and Iso- corrosion shown on pollen, spores and other plant debris, can be of
cyprina sp. This sequence has been suggested to represent tsunami de- fungal or bacterial origin (Goldstein, 1960), but it is the primitive
posits triggered by the Rochechouart impact, which can be recognized fungal group, the chytridiomycetes (“chytrids”), that are the most well-
in the end-Triassic sediment archive throughout the Paris Basin know pollen pathogens today. These saprophagous microfungi pene-
(Kuhlmann, 2019). A rapid change to thick, reddish-brown clays starts trate the pollen wall and consume the cell contents of pollen floating in
at a depth of 45.16 m. They reach up to 38.81 m and represent the lakes or retained in soils (Skvarla and Anderegg, 1972; Czeczuga and
upper Rhaetian, stratigraphically belonging to the ‘Argiles de Levallois’. Muszynska, 2001). Chytrids, which first occur in the Devonian Rhynie
Near 45.00 m a microfolded slumping horizon (seismite) can be ob- Chert, have a long association with soil ecosystems throughout the
served. The clays grade sharply into the dark grey, bioturbated marls of geological record of terrestrial deposits.
the lower Hettangian (‘Psiloceras beds’) with intercalated biodetritic For the purpose of this study, the broadly defined category “corro-
limestone beds in the lower part. The typical ammonite Psiloceras pla- sion” was split into two subcategories: (1) corrosion as a result of the
norbis was found at 36.23 m. At the top of the section (32.20 m) more physical and chemical alteration of the sporomorphs. (2) biodegrada-
frequent occurrence of plant debris is observed. A detailed sedimento- tion due the activity of micro-organisms. We quantified the level of
logical, geochemical and mineralogical description of Boust and other corrosion attributable to micro-organisms using only smooth, trilete
sections of the northeastern Paris Basin is presented in detail in fern spores belonging to Deltoidospora and Concavisporites
Kuhlmann (2019). (Concavisporites-Deltoidospora-complex; Fig. 2). These genera were
chosen as proxy taxa because of their high abundance and continuous
2.2. Palynology occurrence from the Rhaetian to upper Hettangian. Furthermore, these
taxa are characteristic for the fern peaks during the Late Triassic – Early
A description of sample preparation methods used for Mingolsheim Jurassic in the Schandelah-1 core. Fluctuations in abundance of Con-
and Stenlille is provided in van de Schootbrugge et al. (2009) and cavisporites-Deltoidospora led to the exclusion of ~30 samples in which a
Lindström et al. (2012), respectively. The drillcore Boust was described minimum of 50 specimens per sample could not be attained. In samples
and sampled by N. Kuhlmann and J. Thein at the University of Lorraine with abundant proxy taxa, counts are based on 100 specimens of trilete
in Nancy (France). Samples from the Schandelah-1 and Boust cores spores. A distinction was made between differing levels of biode-
were processed at Utrecht University. Approximately 10 g of freeze gradation observed in each specimen. We partitioned these degrees of
4
Fig. 2. Examples of Triassic-Jurassic smooth trilete spores of the Concavisporites-Deltoidospora complex. The specimens show an increase in the degree of biode-
gradation from (a) no biodegradation to (b) severe biodegradation of the spore wall.
biodegradation into three classes: (1) none – minor pitting, (2) high plants, including cycads, conifers, and peltasperms, make up on average
amount of pitting and/or etching but no distinct biological pattern, (3) 30 to 50% of the total palynomorph fraction. A particularly prominent
distinct biological patterns. In total, 91 slides were counted over the victim of the end-Triassic extinction was the enigmatic gymnospermous
depth interval 228–338 mbs, spanning the Rhaetian to lowermost Si- pollen taxon Ricciisporites tuberculatus (Mander et al., 2012a). In the
nemurian. Schandelah-1 core, R. tuberculatus gradually decreases in abundance
and finally disappears at 318.65 mbs around the Triassic-Jurassic
boundary. The extinction of R. tuberculatus is part of a general trend
3. Results
towards an assemblage that is nearly devoid of tree-pollen, differing
starkly from underlying Rhaetian and overlying Hettangian assem-
3.1. Reworking and vegetation change
blages (compare for example van Eldijk et al. (2018) for a plot of long-
term trends in pollen taxa). The step-wise disappearance and eventual
In all four cores the loss of four of the most common taxa of late
extinction of these four taxa can be observed in all four areas (Fig. 3),
Triassic pollen, Ovalipollis ovalis, Lunatisporites rhaeticus, Rhaetipollis
although spikes in abundance in the Stenlille core are somewhat ob-
germanicus, and Ricciisporites tuberculatus occurs during the latest
scuring this trend.
Rhaetian (Fig. 3), reaching a nadir at the Triassic-Jurassic boundary.
In all four cores the loss of pollen is accompanied by substantial
These four pollen taxa, representing a diverse mix of gymnospermous
Fig. 3. Correlation of pollen abundance for the four investigated cores. Plotted here is the sum of the “big four” Triassic taxa that go extinct at the Triassic-Jurassic
boundary, including Ovalipollis ovalis, Rhaetipollis germanicus, Lunatisporites rhaeticus, and Ricciisporites tuberculatus.
5
Fig. 4. Palynological diagram showing total pollen abundance and reworking as a percentage of the total fraction. Reworking strongly increases within the Triletes
Beds in Germany and in correlatable units in Denmark and France.
increases in fern spore abundance together with increases in bryophyte examples of Aulisporites astigmosus, signaling reworking of Middle-
spores and spores from horsetails. For the Germanic and Danish Basins, Upper Triassic (upper Ladinian to Carnian) sediments.
these data have been discussed extensively in a set of papers on fern Both red shale intervals are characterized by declines in both re-
proliferation during the end-Triassic extinction (Lindström et al., 2012, lative and absolute pollen abundance (Fig. 7), although the absolute
Lindström, 2016; van de Schootbrugge et al., 2009, 2019; Barth et al., quantitative data set is somewhat patchy due to low counts of Lycopo-
2018a;). Here, we focus on the link between pollen loss indicating de- dium in several samples. Minima in pollen total abundance are pre-
forestation, and the reworking of palynomorphs. dominantly related to decreased abundance of Classopollis and Perino-
All four studied cores provide a consistent record of reworking of pollenites. Their relative decline is compensated by increases in fern
palynomorphs that is highly correlated with the main phase of defor- spores of the Concavisporites-Deltoidospora complex. Reworking appears
estation (Fig. 4) within the Triletes Beds in Germany, and its equiva- to have ceased at the Hettangian-Sinemurian boundary, however, the
lents in Denmark (‘grey siltstone’ interval) and France (‘Argiles de Le- reworking signal could possibly be masked by a very pronounced in-
vallois’). In the Schandelah-1 core, the exact same interval within the crease in pollen abundance (Fig. 7), mainly of Perinopollenites elatoides.
upper part of the Triletes Beds that contains evidence for forest dieback, The Sinemurian shows a dominance of tree pollen and very reduced
also contains a remarkable and unusual record of abundant reworked numbers of spores overall.
Palaeozoic palynomorphs. Within the top part of the Triletes Beds In the Mingolsheim core (Fig. 8) the Triletes Beds contain numerous
(327.00 to 318.80 mbs) large quantities of Palaeozoic acritarchs and specimens of reworked Veryhachium and several specimens of Multi-
prasinophyte cysts (phycomata) were encountered (Figs. 4, 5 and 6) plicisphaeridium, together with a suite of other unidentified Palaeozoic
comprising up to 35% of the total palynomorph assemblage. In several acritarchs and prasinophytes. Perhaps, much more significant is an
samples around 321.00 mbs, reworked acritarchs/prasinophytes con- abundance of peculiar palynomorphs that bear similarity to Neopro-
stitute the second-most dominant palynomorph group. Prasinophyte terozoic palynomorph assemblages known from localities worldwide.
phycomata (e.g. Tasmanites and leiosphaerids) are mostly long-ranging These reworked palynomorphs share a common grey-brown preserved
undiagnostic taxa, and many are poorly preserved. In contrast, the appearance that is distinctly different from authochthonous material
quality of preservation of the acritarch specimens is exceptional and also differs from reworked Palaeozoic acritarchs. The genus Ger-
(Figs. 5, 6), enabling species-level taxonomic identifications. The minosphaera (Fig. 8a–c) is present in multiple samples from the Triletes
identified acritarchs are exclusively Silurian and Ordovician in age Beds, occurring together with undetermined acanthomorph acritarchs
(Table 1). Except for Acanthodiacrodium ubui, and some Polygonium and (Fig. 8d–e). Germinosphaera has recently been recovered from the
Baltisphaeridium species, all of the encountered acritarch specimens are Shaler Group of Meso- to Neoproterozoic age (Loron et al., 2019), but it
singletons (Table 1). This pattern of recovery is typical of reworked has a very wide distribution and also occurs in the Tonian of the East
assemblages. Overall, 20 different Palaeozoic acritarch species were European Platform in Russia (Vorob'eva et al., 2009). Ostiana micro-
recognized. cystis is present in sample MING-39 (Fig. 8f). In addition, we find many
Reworking reached a climax just before the Triassic-Jurassic differently shaped palynomorphs that are reminiscent of the Neopro-
boundary in the Schandelah-1 core and declined sharply into the terozoic form taxon Jacutianemia solubila (Fig. 8i, h and k) as recently
Hettangian. However, reworking did continue at low levels for much of described from the Tonian of North China (Li et al., 2019), but also
the Hettangian (Fig. 7). In addition to minor occurrences of additional present in the East European Platform (Vorob'eva et al., 2009). We also
Palaeozoic acritarchs, reworking during the Hettangian is mainly find numerous distinctly branched tubes (Fig. 8g) in different samples,
characterized by reworking of Triassic palynomorphs, marked by single which resemble the recently described Neoproterozoic form genus
occurrences of Ovalipollis and Ricciisporites. There are two two red shale Ouraisphaera giraldae (Loron et al., 2019). Also present are coiled flat-
intervals within the upper Hettangian Angulata Zone (Fig. 7) that show tened filaments (Fig. 8j) that can be assigned to Obruchevella, a pre-
increased numbers of reworked palynomorphs, including numerous sumed filamentous cyanobacteria that has been reported from the
6
(caption on next page)
7
Fig. 5. Identified Ordovician and Silurian acritarchs from the Triassic-Jurassic boundary interval in the Schandelah core (see also Table 1). Scale bars are 20 μm. (a)
Cymatiogalea cf. C. deunfii - Sample Sch 324.10. (b) Acanthodiacrodium sp. - Sample Sch 324.10. (c) Multiplicisphaeridium sp. - Sample 322.70. (d) Oppilatala
eoplanktonica - Sample Sch 322.70. (e) Ammonidium cf. A. microcladum - Sample Sch 323.40. (f) Multiplicisphaeridium sp. - Sample Sch 322.50. (g) Evittia sp. - Sample
Sch 322.70. (h) Ankyrotrochus crispum - Sample Sch 322.50. (i) Multiplicisphaeridium cf. M. raspa - Sample Sch 320.20. (j) Stellechinatum cf. S. uncinatum - Sample Sch
321.10. (k) Baltisphaeridium cf. B. dasos - Sample Sch 321.10. (l) Baltisphaeridium cf. B. dasos - Sample Sch 320.20. (m) Stelliferidium sp. - Sample Sch 320.20.
Neoproterozoic of the East European Platform by Vorob'eva et al. to chemical or physical corrosion as compared to percentages of the
(2009) and elsewhere, including the Doushanto. Taken together there is observed biodegradation pattern. Throughout the studied interval va-
clear evidence for abundantly reworked Neoproterozoic organic-walled lues of pitting vary between 0 and 30% and no clear pattern or trend is
palynomorphs in the Triletes Beds in the Mingolsheim core. evident (not shown). For the entire interval, the mean percentage of
In the Boust core (Fig. 9) reworking increases dramatically within pitting is 6.8%. Also, despite a few exceptions, the majority of the
the latest Rhaetian Argiles de Levallois, which is the Paris Basin samples (~80%) show less than 10% pitting.
equivalent to the Triletes Beds in Germany. Reworked palynomorphs In the upper Rhaetian (318.6–338.0 mbs) biodegradation levels are
are mostly comprised of Carboniferous spores that stand out from au- low with a mean of ~26.4% (Fig. 7). Throughout the upper Rhaetian to
thochtonous spores by being darker and generally more poorly pre- the Hettangian-Sinemurian boundary the mean percentage of biological
served. Also undetermined Palaeozoic acritarchs and prasinophyte patterns of degradation on fern spores is 43.5%. A minimum of 8%
phycomata are present although they are less abundant than in biodegradation occurs in the uppermost Rhaetian, just before the
Schandelah-1. We note the presence of spores assignable to Lycospora, boundary at 318.65 mbs. Across the Triassic-Jurassic boundary, at
Spencerisporites, and Cingulizonates, all of which have a late Carboni- 318.6 mbs, the percentage of biodegraded fern spores increases sharply.
ferous age. Some of these taxa have in the past been incorporated in A small drop occurs at 304.0 mbs of approximately 20%, which is
quantitative analyses of Rhaetian floras, which suggests that there coeval with the black and grey laminated mudstones in the lower
could be a serious problem with vegetation reconstructions. Hettangian, just after the paper shales at a depth between 305.0 and
In the Stenlille cores (Fig. 9), reworked early Palaeozoic, Carboni- 306.0 mbs. The number of spores with biological degradation patterns
ferous and Middle to Late Triassic palynomorphs (up to 10% of the total increases hereafter and values modestly fluctuate between ~45–85%.
fraction) are common within the latest Rhaetian “grey siltstone” in- Following this steady increase, the biodegradation record rapidly de-
terval of the Fjerritslev Formation (Lindström et al., 2012). Lower Pa- creases showing a sudden drop from 78% (259.0 mbs) to 26% (257.0 12
laeozoic (Ordovician–Silurian) reworked palynomorphs include Evittia mbs). Low values of biodegradation continue to 254.0 mbs with only
denticulata, Nexosarium leherrisei, Oppilatala sp., Cymbosphaeridium sp., 16.7% biodegradation at this depth. This interval, 254.0–259.0 mbs,
Multiplicisphaeridium spp. and Neoveryhachium carminae. The cryptos- corresponds to the red-coloured laminated claystone deposited at
pore Tetrahedraletes medinensis is also found reworked within this in- 254.8–258.7 mbs. Hereafter, the amount of biodegraded spores rises
terval. It is known from Upper Silurian to Lower Devonian strata in again reaching a maximum value of 84% at 252.0 mbs. This high value
Scania in southern Sweden (Mehlqvist et al., 2015). A single specimen is followed by a moderate low value of 31.5%, after which the biode-
of Emphanisporites micrornatus present in the “grey siltstone” interval in gradation record fluctuates between approximately 32–51%. Following
the Stenlille-4 core, also occurs in the Lower Devonian (Lockhovian) of these relatively minor oscillations, two spikes show up in the record at
Scania (Mehlqvist et al., 2015). Reworked Carboniferous spores and 242.3 mbs and 240.0 mbs of 69.3% and 71.4%, respectively. This is
pollen include Cingulizonates spp., Crassispora kosankei, Densosporites followed by a second drop in the record that continues from 239.0 mbs
spp., Lycospora pusilla, Florinites sp., Schulzospora rara and Tripartites to 234.0 mbs with biodegradation values between ~18–28%. This de-
vetustus, derived from Visean and Namurian to Westphalian palyno- cline in the biodegradation record is coeval with the second red-co-
floras (Clayton et al., 1977). Carboniferous spores and pollen have loured laminated claystone interval from 234.0–238.0 mbs. Next, to-
previously been encountered in Jurassic and Cretaceous strata of wards the Hettangian – Sinemurian boundary (~230.2 mbs), the
southern Sweden and the Danish island of Bornholm (Nielsen and biodegradation values increase once more reaching 70% at 228.0 mbs.
Koppelhus, 1991; Lindström et al., 2017). Late Middle to Late Triassic A minimum characterizes the Hettangian-Sinemurian boundary at
pollen are also encountered, including Retisulcites perforates, Echinitos- 331.0 mbs (13.6%).
porites iliacoides, Staurosaccites quadrifidus, Illinites chitonoides, Proto-
diploxypinus gracilis and Striatoabieites spp. Several of these Silurian to
mid-Triassic taxa have also been found reworked within the “grey 4. Discussion
siltstone” in southern Sweden, along with rare chitinozoans (Lindström
et al., 2017). Although initially not considered to be reworked, abun- 4.1. Widespread soil erosion associated with the end-Triassic extinction
dant sphaeromorphs (Fig. 9), likely representing the vegetative cysts of
prasinophyte algae, may constitute up to 50% of the total palynomorph When exposed bedrock, in any terrain, contains organic-rich sedi-
assemblage in some beds (Lindström et al., 2012). These sphaer- mentary rocks, those rocks may release palynomorphs as weathering
omorphs vary significantly in appearance within a sample, both in and erosion take place. Such eroded and subsequently transported pa-
colour and preservation, which may be explained by reworking from lynomorphs behave like silt-sized organic particles that are readily re-
the Lower Palaeozoic along the Sorgenfrei-Tornquist Zone. In Scania deposited together with siliciclastic particles as sediments come to fill a
(southern Sweden) the Lower Palaeozoic is intruded by numerous dykes depositional basin. Although the recognition of this reworked organic
of Carboniferous–Permian age. These dykes caused significant contact component presents a basic, but well-known challenge in stratigraphic
metamorphism in aureoles around the intrusives (Olsson-Borell and palynology (Wilson, 1964), it is not a topic that is often addressed in the
Ahlberg, 2003) and may explain the variations in colour observed in the general literature, because, under normal conditions of sediment ac-
sphaeromorphs and associated prasinophycean phycomata. cumulation, the ratio of reworked to autochthonous palynomorphs is
overwhelmingly diluted by input from contemporary material, and may
become vanishingly small. Reworked components can be further di-
3.2. Corrosion of palynomorphs luted due to taphonomic loss during extended transport from more
distant source areas, and, of course, much source bedrock itself (e.g.
In the Schandelah core, the biodegradation record (Fig. 7) is based metamorphic and igneous rocks) does not act as a source of reworked
solely on the percentage of smooth trilete spores showing clear biolo- palynomorphs at all. The recognition of reworking, therefore, becomes
gical patterns. Only a few spore specimens contain pitting marks related a potential tool for interpreting former sedimentary dynamics in ways
8
Fig. 6. Identified Ordovician and Silurian acritarchs from the Triassic-Jurassic boundary interval in the Schandelah core (see also Table 1). Scale bars are 20 μm. (a)
Elektroriskos cf. E. pogonius - Sample Sch 320.20; (b) Baltisphaeridium klabavense - Sample Sch 320.20; (c) Arbusculidium cf. A. ramusculosum – Sample Sch 319.50; (d)
Evittia remota - Sample Sch 320.20; (e) Gorgonisphaeridium sp. – Sample Sch 319.50; (f) Vogtlandia sp. - Sample Sch 319.50; (g) Polygonium cf. P. fragilis - Sample Sch
319.66; (h) Polygonium sp. – Sample Sch 319.66; (i) Gorgonisphaeridium sp. - Sample 319.50; (j) Acanthodiacrodium ubui – Sample Sch 319.66; (k) Gorgonisphaeridium
sp. – Sample Sch 319.80; (l) Cymbosphaeridium aniae – Sample Sch 331.60.
9
Table 1
List of identified Ordovician and Silurian acritarchs reworked into the uppermost Rhaetian of the Schandelah-1 core together with their stratigraphic and absolute
age range.
Depth (m) Species Age range Stages Top (Ma) Base (Ma) Average Sample average SD
319.5 Arbusculidium cf. A. ramusculosum Lower Ordovician 471.80 488.30 480.05 461.62 17.3
319.5 Gorgonisphaeridium sp. Upper Ordovician - Silurian 416.00 460.90 438.45
319.5 Vogllandia sp. Middle Ordovician 460.90 471.80 466.35
319.66 Polygonium cf. P. fragilis Middle Ordovician 460.90 471.80 466.35 468.39 10.0
319.66 Polygonium spp. Ordovician 443.70 471.80 457.75
319.66 Polygonium spp. Ordovician 443.70 471.80 457.75
319.66 Acanthodiacrodium ubui Lower Ordovician 471.80 488.30 480.05
319.66 Acanthodiacrodium ubui Lower Ordovician 471.80 488.30 480.05
319.80 Ordovicidium heteromorphicum Middle - Upper Ordovician 471.80 443.70 457.75 457.75 14.1
320.20 Baltisphaeridium cf. B. dasos Upper Ordovician 443.70 460.90 452.30 453.48 16.1
320.20 Elektoriskos cf. E. pogonius Lower - Middle Silurian 426.20 443.70 434.95
320.20 Evittia remota Upper Ordovician 443.70 460.90 452.30
320.20 Stelliferidium sp. Lower Ordovician 471.80 488.30 480.05
320.20 Multiplicisphaeridium cf. M. raspa Lower - Middle Silurian 426.20 443.70 434.95
320.20 Baltisphaeridium klabavense Middle Ordovician 460.90 471.80 466.35
320.80 Acanthodiacrodium ubui Lower Ordovician 471.80 488.30 480.05 480.05 8.3
321.10 Baltisphaeridium cf. B. dasos Upper Ordovician 443.70 460.90 452.30 463.45 11.2
321.10 Stellechinatum cf. S. unctinatum Lower - Middle Ordovician 460.90 488.30 474.60
322.50 Multiplicisphaeridium sp. Middle Ordovician - Silurian 471.80 416.00 443.90 455.13 11.2
322.50 Ankyrotrochus crispum Middle Ordovician 460.90 471.80 466.35
322.70 Evittia sp. Lower Silurian 443.70 428.70 436.20 438.77 3.6
322.70 Multiplicisphaeridium sp. Middle Ordovician - Silurian 471.80 416.00 443.90
322.70 Oppilatala eoplanktonica Lower Silurian Rhuddanian - Wenlock 443.70 428.70 436.20
323.40 Ammonidium cf. A. microcladum Middle Silurian Wenlock 422.90 428.90 425.90 425.90 0.0
324.10 Acanthodiacrodium sp. Lower Ordovician 471.80 488.30 480.05 480.50 0.0
324.10 Cymatiogalea cf. C. deunfii Lower Ordovician Tremadocian - Floian 471.80 488.30 480.05
325.50 Ordovicidium cf. O. elegantulum Upper Ordovician 443.70 460.90 452.30 452.30 0.0
331.6 Cymbosphaeridium aniae Middle - Upper Silurian Wenlock - Pridoli 428.90 416.00 422.45 422.45 0.0
Fig. 7. Long term palynological records for the Schandelah-1 core. (a) Total pollen (% of total palynomorph fraction) showing clearly dramatic disappearance of
pollen grains within the upper part of the Triletes Beds and declines associated with the red shale intervals. (b) Reworking as percentage of total palynomorph
fraction. Both the Rhaetian and upper Hettangian show high numbers of reworked palynomorphs. (c) Biodegradation based on corrosion of 50 counted trilete spores
per sample. (d) Absolute pollen abundance (pollen per gram sediment) showing decreased pollen abundance in red shale intervals. (e) Absolute pollen abundance
(pollen per gram sediment) showing strong decline in absolute pollen abundance in the Triletes Beds marking deforestation during the end-Triassic mass-extinction.
(For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)
10
Fig. 8. Selected Neoproterozoic reworked palynomorphs from the Triassic-Jurassic boundary in the Mingolsheim core. All scale bars 20 μm. (a-c) Germinosphaera cf.
G. bispinosa; (d, e) Undetermined acanthomorph acritarchs; (f) Ostiania microcystis Ming-39, Triletes Beds; (g) Ouraisphaera giraldae; (h, i, k) Jakutianemia spp.; (j)
Obruchevella valdaica.
11
Fig. 9. Selected reworked palynomorphs from the Boust core (northern France) and Stenlille cores (Denmark). All scale bars 20 μm. (a–h) Reworked Carboniferous
spores in the Boust core. (i–l) Reworked Palaeozoic acritarchs from Stenlille. (a) Lycospora noctuina; (b) Spencerisporites spp.; (d–h): Lycospora – Cingulizonates.; (i)
Multiplicisphaeridium sp. (cf. Circinatisphaera aenigma) Stenlille-4, 1508.83 m. (j) Evittia sp. cf. E. denticulata. Stenlille-4, 1507.80 m; (k) Neoveryhachium carminae.
Stenlille-4, 1502.15 m; (l) Cymatiosphaera sp. and large sphaeromorph. Stenlille-4, 1504.30 m.
that enhance the value of sedimentary provenance. And, given that re- constrained in time and space.
working has not been generally observed in sedimentary sequences in Reworking was not limited to the investigated cores. For example,
the Triassic and Jurassic of NW Europe, the reworked material from the sections in the Haida Gwaii Islands (Canada) contain a record of re-
extinction interval in multiple locations across NW Europe represents a worked Triassic conodonts that have been redeposited within the
unique signal that can be linked to environmental changes that are lowermost Jurassic (Greene et al., 2012). In the Mariental-1 core from
12
North Germany, reworked acritarchs are not as abundant as in Schan- with an increase in fern spore abundance (Ruckwied and Götz, 2009).
delah, but the presence of the Silurian acritarch Multiplicisphaeridium In Pakistan, kaolinite contents increases across the Rhaetian-Hettangian
and Carboniferous spores was noted by Heunisch et al. (2010). These boundary together with sedimentological and petrographical evidence
authors also described the reworking of typical Rhaetian pollen into for a marked change in weathering and erosion (Iqbal et al., 2019). As
Hettangian assemblages. Palynological analyses of the Kössen Forma- discussed by Iqbal et al. (2019) the chemical index of alteration sharply
tion and overlying Schattwald Beds in Austria by Bonis et al. (2009) increases also at the Rhaetian-Hettangian boundary. Towards the East,
show a remarkable increase in the diversity of trilete spores within the clay mineral analyses of a suite of cores from Poland show marked
Schattwald Beds. Many of these taxa are however extremely rare, and changes in clay mineraology starting with the precursor carbon isotope
most constitute single occurrences. It is not excluded that many of these excursion in the Rhaetian (Pienkowski et al., 2014; Brański, 2014). In
taxa, including Densosporites, Lycospora, and Cingulizonates are in fact addition to an increase in kaolinite contents within the Triassic-Jurassic
reworked Carboniferous taxa. The reworking of Carboniferous spores boundary beds, these authors also note an increase in osmium isotope
has been observed in several other previous studies on Hettangian to values.
Sinemurian palynological assemblages, for example on Bornholm Although elevated proportions of kaolinite in Rhaetian sediments
(Nielsen and Koppelhus, 1991), in the Paris Basin (Hagemann, 1967), in are usually related to enhanced chemical bedrock weathering, it is
Poland (Muir, 1967) and in Northeast Scotland (Windle, 1979). important to realize that the presence of clay in shallow-marine de-
posits could well be derived through physical erosion of soils and, more
4.2. Clay minerals as indicators of soil erosion particularly, older clay-rich sediments (Thiry, 2000). In Europe, notably
the extensive late Carboniferous clastic deposits are a plausible source
Clay particles are aluminosilicate minerals, typically with a size of of reworked kaolinite. Tethyan sections are illustrative for increased
2 μm or less, and they are the by-products of both physical weathering erosion rather than chemical weathering associated with the end-
(e.g. mechanically weathered mica) and chemical weathering (so-called Triassic extinction, because in many Alpine and southern European
secondary minerals). The chemical pathway of clay mineral formation sections carbonate deposition ceased prior to the Triassic-Jurassic
is a primary driver of soil formation (pedogenesis), and soils can store boundary and was replaced by mudstone or clay deposition. The
organic carbon, as clay particles have a high sorptive capacity (Hedges Schattwald Beds in the Northen Calcareous Alps (Austria) are a case-in-
and Keil, 1995), thereby stabilizing soils and improving nutrient and point, because this clay-rich unit overlaying the carbonate marl-lime-
water retention times (Ekwue, 1990). Clay formation occurs as poorly stone alternations of the Kössen Formation is the start of more gen-
soluble ions, such as aluminum, form immobile hydrolyzates resistant eralized clay-rich sedimentation during the Early Jurassic (McRoberts
to further weathering. This process is dictated largely by the large ionic et al., 1997). Although representing the southern margin of the Tethys,
radius and the high ionic charge of Al, and therefore Al has a tendency the sedimentological expression is remarkably similar in Italy, where
to form strong bonds with oxygen. On the other hand, chemical species the carbonate-rich Zu Formation of Rhaetian age is abruptly overlain by
with a small ionic radius and low ionic charge (Na, Ca, Mg and Li) are a distinct claystone at the base of the Malanotte Formation that marks
more readily weathered from the parent primary minerals, and are the end-Triassic extinction event (Bachan et al., 2014; Galli et al.,
covered by a hydration shell once liberated, making them highly so- 2007). Although the cessation of carbonate deposition has been at-
luble. As such weathering intensity, which is described as a function of tributed to ocean acidification and a crisis in biocalcification in corals
weathering rate and time of exposure, dictates the predominant mi- and other carbonate producing organisms (Hautmann et al., 2008; van
neralogy of the clay assemblage. Illites and montmorillonites are low de Schootbrugge et al., 2007), a widespread increase in erosion could
intensity secondary products of weathering, while kaolinites are their also have led to smothering and eutrophication of shallow marine
intensely weathered counterparts (Nesbitt and Young, 1984). evironments, similar to what has been proposed for the end-Permian
A change in chemical weathering during the latest Triassic has been extinction (Algeo et al., 2011b; Algeo and Twitchett, 2010).
inferred from clay mineralogical studies. A sudden change in clay mi-
neralogy in the upper Rhaetian is very obvious in sections of the 4.3. Provenance of the reworked material
northeastern Paris Basin. High kaolinite contents have been de de-
scribed in the Argiles de Levallois in the Grouft drillcore (Luxembourg) Because of their taxonomic value and their general resistance to
by Lieder-Wolf (2012). Most of the other sections in this region (in- degradation, reworked palynomorphs are excellent tools to constrain
cluding Boust) show similar stratigraphic trends in clay mineralogy. sediment provenance. However, as pointed out by Streel and Bless
Through the Rhaetian, the weathering became increasingly lateritic in (1980) and Batten (1991), all too often reworking is either ignored or
nature, with an enrichment of iron-oxyhydroxides. The Argiles de Le- reported only as accessory data, leaving its full potential unexplored.
vallois therefore show a very characteristic reddish-brown colour, due Nonetheless, several authors have successfully applied reworking to
to a high content of goethite and hematite. Additionally, the con- geological problems extending beyond glitches in biostratigraphy. For
centrations of iron and other siderophile trace elements are also rela- example, Truswell and Drewry (1984) used abundantly reworked Cre-
tively high (Kuhlmann, 2019). taceous palynomorphs to determine the provenance of sub-Recent se-
Increased amounts of kaolinite encountered in boundary beds across diments on the Antarctic shelf. Eshet et al. (1988) showed that regional
NW Europe have been taken as evidence for extremely humid condi- sedimentary cycles throughout the Permian-Triassic subsurface of Israel
tions (van de Schootbrugge et al., 2009; Zajzon et al., 2012; Lintnerová corresponded to decreased (transgressive) and increased (regressive)
et al., 2013). In Scania (Sweden), high amounts of kaolinite were levels of reworked palynomorphs. They noted specifically that re-
identified in the uppermost Rhaetian Bjuv Member (Lindström and working increased substantially as bedrock sourcing Palaeozoic paly-
Erlström, 2006). Whereas the entire Late Triassic to Early Jurassic nomorphs became more exposed during regressive phases. In a recent
succession in Scania is characterized by deltaic and flood plain deposits study on the Carboniferous of Portugal it was found that more than 90%
that show evidence for intense chemical weathering of the Fennos- of the encountered palynomorphs were reworked from Cambrian to
candian margin under humid climate conditions (Ahlberg et al., 2002), Devonian sediments that were part of the Variscan Orogeny. Under-
the Bjuv Member vertisols clearly stand-out compared to Lower Jurassic standing this regional reworking signal contributed signficantly to un-
paleosoils (Ahlberg et al., 2003). The vertisol underclays constitute derstanding basin geodynamical processes (Lopes et al., 2014). Re-
excellent kaolinitic clays for the ceramic industry (Ahlberg et al., 2003). worked palynomorphs may even be used to showcase the former
Further south, sections in the Tatra Mountains in Slovakia show a presence of sediments that are now long gone. Batten (1991) described
strong increase in kaolinite content across the Rhaetian-Hettangian reworking of Triassic and Jurassic palynomorphs in Tertiary sediments
boundary (Michalik et al., 2010; Lintnerová et al., 2013) coinciding in Scotland, in an area that contains very few remnants of these
13
Mesozoic deposits. Palynological work by several authors (e.g. Vavrdová, 2008) show ra-
We use the geological map of Fig. 1 to make predictions of probable ther poorly preserved Neoproterozoic palynomorphs in those units,
source areas for the reworked assemblages encountered in the upper- which could be due to later thermal overprinting. It seems we have an
most Rhaetian in our four studied cores. Palaeozoic acritarchs and interesting situtation, whereby the reworked palynomorphs that are
chitinozoa present in the Stenlille palynological fraction were likely now present in Mingolsheim are better preserved than their original
sourced directly from the Fennoscandia border zone towards the north. source material.
Previously, reworking of Carboniferous spores in the Lower Jurassic The reworked palynomorphs present in the Boust core were almost
(Hettangian-Sinemurian) Rønne Formation on nearby Bornholm was certainly sourced from the London-Brabant Massif towards the north. In
reported by Nielsen and Koppelhus (1991) and attributed to erosion of addition to Devonian limestones and siliciclastics, this ancient land
uplifted blocks of basement and Palaeozoic cover along the southern mass is characterized by numerous outcrops of Carboniferous sedi-
limit of the Ringkøbing-Fyn high. However, it remains unclear whether mentary rocks even today. During much of the Mesozoic it must have
these blocks were above water during much of the Early Jurassic. Al- been shedding Palaeozoic material into shallow Mesozoic basins. The
ternatively, uplifted blocks in the Colonus Shale on the southern border map in Fig. 1 can also be used to predict the predominant reworking
of the Fennoscandian Shield could have been the source. The Palaeo- signal observed, for example, in British Mesozoic basins, such as the
zoic cover in the area includes mostly Silurian, Ordovician, and Cam- Cardigan Bay and Cleveland basins. These basins were receiving mainly
brian. Another potential source area could have been the area around sediments derived from Carboniferous rocks, including the palyno-
Oslo, where the Oslo Confacies Belt comprises a thick series of Pa- morphs released from those sediments (Windle, 1979).
laeozoic and Neoproterozoic sediments (Lassen and Thybo, 2012).
Upper Silurian–lower Devonian strata are also present within the Sor- 4.4. Mechanisms driving elevated soil erosion
genfrei-Tornquist Zone in Scania, but Carboniferous–Permian sedi-
mentary rocks are missing. Thus, the reworked Carboniferous spores Absolute quantitative data (pollen per gram of sediment) from the
and pollen found in strata in the Danish Basin may either have had a Mingolsheim core indicates that pollen accumulation decreased 1000-
different source area, or alternatively the Carboniferous succession was fold in the latest Rhaetian and did not recover until the middle
completely removed at this event and later events during the Jurassic. Hettangian (Fig. 7e). This suggests that the drop in the abundance of
The rich and diverse assemblage of Ordovician and Silurian acri- tree pollen in the Triletes Beds across Germany, and in equivalent beds
tarchs encountered in the Triletes Beds in the Schandelah-1 core were across NW Europe, reflects a true decrease in vegetation cover related
most likely derived from outcrops in the Baltic Basin towards the East to the end-Triassic crisis and, therefore, can not be ascribed to relative
where much of the Palaeozoic strata must have been at the surface changes in organic matter fluxes. The widespread loss of arborescent
during the Triassic and Jurassic (Fig. 1). Paleogeographical re- vegetation, which can be traced from Sweden all the way south to
constructions indicate the presence of a large deltaic system connecting Austria, can be regarded as the main cause for increased weathering
the shallow marine areas in the Germanic Basin via Poland with the and erosion during the latest Rhaetian. However, changes in vegetation,
East European Platform in the hinterland (Barth et al., 2018a). In such as the shift to fern-dominated landscape during the Toarcian OAE
contrast, Franz and Wolfgramm (2008) proposed a Scandinavian source (Slater et al., 2019), are not uncommon during the Mesozoic. While the
for the thick sandstone units within the Rhaetian Exter Formation that TOAE has also been linked to elevated weathering and erosion (Dera
were transported via distributary channels reaching as far south as et al., 2009; Them et al., 2017), there are no other reports of a similarly
Lower Saxony and as far East as Mecklenburg-Vorpommern (Germany). enhanced increase in the reworking of palynomorphs. Hence, it is likely
Perhaps a more local source is also possible, for example with soil that other additional mechanisms were at play during the end-Triassic
erosion taking place in the Rhenish Massif towards the south of extinction event.
Schandelah. Enhanced erosion during the latest Rhaetian was exacerbated by a
Two arguments for a local source can be made. The first is related to number of reinforcing mechanisms that acted alongside deforestation.
the presence of beds rich in clay clasts within the Triletes Beds in the Large-scale tectonic events, such as large earthquakes, can trigger cat-
Schandelah-1 core. Test analyses did not provide clear evidence that the astrophic landscape devastation as was observed in the aftermath of the
clasts are the source of the reworked acritarchs, but the start of the clay 7.9 Mw Wenchuan earthquake that struck Sichuan province (China) in
clast beds does coincide with the onset in reworking (Fig. 10). The 2008. Seismic shaking led to catastrophic landslides, causing net to-
abundance of these clay clasts might suggest a local source. Further- pographic denudation (Ouimet, 2011). The latest Triassic in NW Europe
more, the ages of the reworked assemblages in the Schandelah-1 core contains unique evidence for repeated seismic activity not only in the
appears to increase in age up section. Hence, there is clear evidence for Schandelah-1 core (Fig. 10; van de Schootbrugge et al., 2019), but also
an inverted stratigraphy with the younger Silurian material being in Denmark, Sweden, Luxemburg and the United Kingdom (Lindström
eroded first and progressively older material (Ordovician and older) et al., 2015; Simms, 2003). Seismites, sedimentary layers that show
later. It appears unlikely that such a signal would be preserved if the typical signs of in situ shaking, such as slump folds, water-escape
material had been transported over hundreds of kilometers from Poland structures, and micro-faulting, occur over a wide area within the latest
or the East European Platform. A local source from the Rhenish Massif Triassic (Lindström et al., 2015). Whereas the pulses in reworking in the
towards the southwest or a Fennoscandian source is therefore likely. Schandelah-1 core are not directly coupled to the occurrences of the
The Neoproterozoic palynomorphs present in the Mingolsheim core seismite beds (see also Fig. 10), the initiation of seismic activity does
were likely sourced from the Bohemian Massif towards the east of coincide with the start of influx of reworked material. The same ob-
Mingolsheim (Fig. 1), even though the best preserved and most similar servation can be made for the Boust core.
specimens in the literature have been described from the East European Enhanced erosion and run-off events can be triggered by loss of
Platform. Neoproterozoic sediments with palynomorphs are known vegetation through burning (Sankey et al., 2017). Removal of vegeta-
from the Czech Republic in the so-called Moravo-Silesian Unit or Bru- tion due to burning not only leads to an increase in the impact of rain
novistulicum. Now buried beneath Carpathian Nappes, those Variscan drops, but heating of soil may increase water repellency, stimulating
units still reside at shallow depths (around 1000 m) and have remained run-off rather than uptake (Ferreira et al., 2008). There is good evi-
relatively undeformed. Neoproterozoic sediments represent a former dence for widespread fire activity during the Rhaetian from charcoal
siliciclastic passive margin of Gondwana. Proterozoic sediments were records from Germany (Uhl and Montenari, 2011), Greenland (Belcher
folded and thrusted during the Variscan Orogeny and were covered by et al., 2010), Sweden (Petersen and Lindstrom, 2012), and Poland
Silurian and Devonian siliciclastics and later by Carboniferous coal (Marynowski and Simoneit, 2009). The increase in forest fire activity,
basins during foreland basin evolution (Kalvoda et al., 2008). despite overall lower oxygen concentrations in the atmosphere (Belcher
14
Fig. 10. Triassic-Jurassic boundary interval in the Schandelah-1 core. Reworking increases concommittantly with the onset of seismic activity and the reduction in
tree cover. The age of the reworked assemblages shows an overall increase in age through the latest Rhaetian from mostly Silurian to predominantly Ordovician.
and McElwain, 2008), has been explained as a result of warming cli- 4.5. Biodegradation of palynomorphs and soil loss and resilience
mates with a higher incidence of lightning strikes affecting plants that
were increasingly prone to burning because of their finer leaves In Quaternary studies the corrosion (biological and chemical al-
(Belcher et al., 2010; van de Schootbrugge, 2010). Importantly, all re- teration) of sporomorphs has been used to indicate erosion of lake
cords of forest fires derive from the same geographical areas that also catchments. In those studies, higher numbers of reworked palyno-
show elevated reworking, so it seems likely that landscape devastation morphs correlate to a larger number of corroded specimens, indicating
due to burning also contributed to the observed phenomena. a positive correlation between erosion and corrosion (Wilmshurst and
Ultimately, weathering and erosion were also exacerbated by the McGlone, 2005). Mander et al. (2012b) have used corrosion of paly-
release of large amounts of volatiles to the atmosphere, including CO2 nomorphs in Triassic-Jurassic deposits in Greenland to examine the role
and SO2, the latter of which would have had the potential via acid rain of taphonomy in apparent mismatches between macro- and microfloral
to increase weathering and erosion on short time scales. Basalts within records. Although these authors did not distinguish between biological
the Central Atlantic Magmatic Province likely interacted with evapor- and chemical degradation, their plant bed 5, which represents the ex-
ites and coal in the subsurface, increasing their volatile load sub- tinction interval in Greenland, did indeed show a higher incidence of
stantially (Callegaro et al., 2013, 2014; Heimdal et al., 2018). Sulfur corrosion. Microbial activity in mesophiles is known to be temperature
aerosols would presumably not enter the stratosphere but instead dependent (Biederbeck and Campbell, 1973), hence metabolic rates of
would remain in the troposphere and rain out regionally. This could fungi and bacteria, likely increased during periods of global warming
explain why the weathering signal is so unique especially in NW Eur- such as that seen across the Triassic-Jurassic boundary. We therefore
opean basins, which would be downwind from the CAMP with the fully expected to observe an increase in biodegradation of spores within
development of a permanent Inter-Tropical Convergence Zone (ITCZ) the Triletes Beds.
over Laurasia. In contrast to SO2, carbon dioxide remains in the at- The three intervals within the Schandelah core characterized by
mosphere much longer, and its effects therefore are very long lived. The high numbers of reworked palynomorphs, however, clearly show much
duration of the Triletes Beds is estimated at perhaps 100 kyr or less, lower numbers of biodegraded spores (Fig. 7). Although seemingly
based on the latest timeline for CAMP eruptions and carbon isotope counter-intuitive, in our recorded decrease in biodegradation is com-
excursions presented by (Lindström et al., 2017). Whereas the long- patible with records of increased reworking in the same intervals. As
term continuation of reworking during the Hettangian could be due to our records of intense reworking of Palaeozoic and older bedrock ma-
elevated atmospheric CO2, it is unlikely that the massive increase in terial clearly show, it is entirely plausible that soils were already mostly
reworking in the latest Rhaetian would have been directly driven by removed during deposition of the Triletes Beds as a result of defor-
carbon injection. estation and erosion. Instead of being retained in soils, eroded paly-
nomorphs would have bypassed soils and have been washed out di-
rectly to shallow marine settings where they were subsequently
15
preserved without retaining evidence of microbial degradation. and prasinophytes, along with lesser numbers of Devonian,
Additional factors that could stymie biodegradation include high Carboniferous and Middle–early Late Triassic palynomorphs that were
runoff from intensified precipitation, flushing organic matter more ra- washed in from Fennoscandia and the Baltic Platform. In the South
pidly into estuaries and coastal seas. High runoff has been inferred by German Basin, Neoproterozoic material was likely derived from the
several authors using palynological data. Bonis and Kürschner (2012) Bohemian Massif. The Paris Basin retained mostly Carboniferous re-
interpreted elevated counts of fern spores within the Schattwald beds, worked palynomorphs, which were derived from the London-Brabant
the Austrian equivalent of the German Triletes Beds, mainly as an in- Massif. Increased erosion corresponded with the disappearance of
dication of wet conditions. In their scenario, climate change induced by dominant Triassic pollen taxa and a proliferation in fern spores.
CAMP eruptions led to stronger monsoonal rainfall and increased Deforestation, therefore, was the driving mechanism behind this major
runoff. A similar conclusion was reached by Ruckwied and Götz (2009) increase in erosion. Denudation of the Triassic landscape is evident
based on an increase in fern spores within the transition from the up- from an inverted stratigraphy, whereby the age of the reworked ma-
permost Rhaetian Fatra Formation to the lower Hettangian Kopieniec terial increases upsection. This indication of bedrock erosion further
Formation in Slovakia. In addition, based on changes in the chemical implies a general loss of soil cover. In addition to deforestation as a
alteration index of their palynomorphs, Iqbal et al. (2019) proposed a direct result of volcanism, earthquakes and forest fires all contributed to
significant increase in humidity across the Rhaetian-Hettangian soil loss across the Triassic-Jurassic boundary.
boundary.
The increase in the amount of biodegraded spores in the lower Declaration of Competing Interest
Hettangian of the Schandelah core is interpreted to indicate terrestrial
ecosystem recovery (Fig. 7). High soil microbial diversity and abun- None.
dance reflects a healthy ecosystem (Coleman et al., 1992). Following
the initial recovery of biodegradation, reaching plateau values within Acknowledgments
the Liasicus Zone, two sharp declines in the abundance of biodegraded
spores are evident associated with the two enigmatic red beds in the BvdS acknowledges financial support from the Goethe University
upper Hettangian (Fig. 7). In accordance with high levels of biode- Frankfurt (Frankfurt am Main, Germany) allowing to drill the
gradation representing a healthy ecosystem, these drops in biode- Schandelah core. BvdS and PKS acknowledge financial support from the
gradation are here interpreted as renewed environmental perturba- German Science Foundation (DFG) grant# SCH1216/5-1. SL acknowl-
tions. This would have resulted in decreased microbial activity and edges support from GEUS. Alexander Houben (TNO) is thanked for help
diversity, hence a decrease in sporomorph biodegradation. with the identification of Carboniferous spores. Natasja Welters is
Much lower rates of reworking during the Hettangian may have thanked for her help with processing the palynological samples at
been driven by recovery of arborescent vegetation, as is clear from the Utrecht University.
recovery of tree pollen abundance in our long-term record from the
Schandelah-1 core (Fig. 7). This may have slowed down the most in- References
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Earth-Science Reviews 210 (2020) 103332

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Catastrophic soil loss associated with end-Triassic deforestation T

1. Introduction as well as prevailing climatic conditions, mainly temperature and

(caption on next page)

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