Climatic reversals related to the Central Atlantic magmatic province caused the end-Triassic biotic crisis— Evidence from continental strata in Poland

Abstract

Eight climatic events can be distinguished in the Triassic-Jurassic (ca. 201 Ma) continental strata of Poland. These events are distinguished by kaolinite/illite ratio, chemical index of alteration (CIA), color of sediments, and palynomorphs. The first transition to wetter climate, evidenced by a shift from smectite- to kaolinite-dominated mudrocks, coincides with the earlier ("precursor") Rhaetian negative δ13Corgexcursion, which means that the beginning of climate perturbations predates the oldest known Central Atlantic magmatic province flood basalts by some 100-200 k.y. The later global, late Rhaetian "initial" negative δ13Corgexcursion is divided into two subpeaks, each corresponding to hot and humid events, separated by a cooler and drier event. The upper subpeak is also associated with perturbation of the osmium isotope system (attributed to volcanic fallout), and darkened miospores, pointing to acid rains. Between the "initial" excursion and the Triassic-Jurassic boundary interval, five climatic fluctuations are inferred from the changing kaolinite/illite ratio, the last two of which are also associated with an Os isotope perturbation, polycyclic aromatic hydrocarbon (PAH) occurrences, a "spore peak," and darkened miospores. A series of periodic atmospheric loading events by CO2, CH4, or alternatively by SO2, sulfate aerosols, and toxic compounds, is inferred to have caused this series of rapid climatic reversals and resulting extinction of many less-adapted forms. Just above the palynofloral extinction level, appearance of new forms commenced Jurassic palynofloral recovery. Tetrapod evolution events in the end-Triassic-earliest Jurassic were related to the extinction of the Pseudosuchia, Dicynodontia, Capitosauroidea, Plagiosaroidea, and Rhynchosauria, while appearance of highly diversified tetrapod ichnofauna in the earliest Jurassic strata indicates a rapid recovery and refill of ecological niches by dinosaurs.

Full text

doi:10.1130/2014.2505(13) , published online August 21, 2014;Geological Society of America Special Papers
Grzegorz Pienkowski, Grzegorz Niedzwiedzki and Pawel Branski
Evidence from continental strata in Poland−−end-Triassic biotic crisis Climatic reversals related to the Central Atlantic magmatic province caused the
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263
The Geological Society of America
Special Paper 505
2014
Climatic reversals related to the Central Atlantic
magmatic province caused the end-Triassic biotic crisis—
Evidence from continental strata in Poland
Grzegorz Pieńkowski*
Polish Geological Institute–National Research Institute, Rakowiecka 4, PL-00-975 Warszawa, Poland
Grzegorz Niedźwiedzki*
Subdepartment of Evolution and Development, Department of Organismal Biology, Evolutionary Biology Centre,
Uppsala University, Norbyvägen 18A, 752 36 Uppsala, Sweden
Paweł Brański
Polish Geological Institute–National Research Institute, Rakowiecka 4, PL-00-975 Warszawa, Poland
ABSTRACT
Eight climatic events can be distinguished in the Triassic–Jurassic (ca. 201 Ma)
continental strata of Poland. These events are distinguished by kaolinite/illite ratio,
chemical index of alteration (CIA), color of sediments, and palynomorphs. The
fi rst transition to wetter climate, evidenced by a shift from smectite- to kaolinite-
dominated mudrocks, coincides with the earlier (“precursor”) Rhaetian negative
δ
13Corg excursion, which means that the beginning of climate perturbations predates
the oldest known Central Atlantic magmatic province fl ood basalts by some 100–
200 k.y. The later global, late Rhaetian “initial” negative
δ
13Corg excursion is divided
into two subpeaks, each corresponding to hot and humid events, separated by a cooler
and drier event. The upper subpeak is also associated with perturbation of the osmi-
um isotope system (attributed to volcanic fallout), and darkened miospores, point-
ing to acid rains. Between the “initial” excursion and the Triassic-Jurassic boundary
interval, fi ve climatic fl uctuations are inferred from the changing kaolinite/illite ratio,
the last two of which are also associated with an Os isotope perturbation, polycyclic
aromatic hydrocarbon (PAH) occurrences, a “spore peak,” and darkened miospores.
A series of periodic atmospheric loading events by CO2, CH4, or alternatively by SO2,
sulfate aerosols, and toxic compounds, is inferred to have caused this series of rapid
climatic reversals and resulting extinction of many less-adapted forms. Just above the
*grzegorz.pienkowski@pgi.gov.pl; grzegorz.niedzwiedzki@ebc.uu.se.
Pieńkowski, G., Niedźwiedzki, G., and Brański, P., 2014, Climatic reversals related to the Central Atlantic magmatic province caused the end-Triassic biotic
crisis—Evidence from continental strata in Poland, in Keller, G., and Kerr, A., eds., Volcanism, Impacts, and Mass Extinctions: Causes and Effects: Geological
Society of America Special Paper 505, p. 263–286, doi:10.1130/2014.2505(13). For permission to copy, contact editing@geosociety.org. © 2014 The Geological
Society of America. All rights reserved.
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264 Pieńkowski et al.
INTRODUCTION
At least four of the fi ve Phanerozoic mass extinction events
can be associated with large igneous province formation. The
eruption of continental fl ood basalts of the Central Atlantic
magmatic province at the end of the Triassic (ca. 201 Ma) was
approximately coeval with the breakup of Pangea, sea-level
changes, abrupt negative carbon isotope excursions (CIEs) trig-
gered by volcanism and/or methane hydrate dissociation, cli-
matic and environmental changes, and mass extinction (e.g.,
McHone, 1996, 2003; Hesselbo and Jenkyns, 1998; Marzoli et
al., 2004, 2011; McElwain et al., 1999, 2007; Olsen et al., 2002;
Hesselbo et al., 2002, 2004, 2007; Hounslow et al., 2004; Guex
et al., 2004; Pieńkowski, 2004; Tanner et al., 2004; Galli et al.,
2005; Jourdan et al., 2009; Ruhl et al., 2009, 2010; Korte and
Kozur, 2011; Pieńkowski et al., 2012). Central Atlantic magmatic
province ages gathered so far from the North American basins
and Morocco (Jourdan et al., 2009; Cirilli et al., 2009; Deenen et
al., 2010; Marzoli et al., 2011; Blackburn et al., 2013) suggest a
short (~1.5 m.y.) duration for the main magmatic activity, extend-
ing from the latest Triassic to the earliest Jurassic.
Mass extinctions are thought to refl ect signifi cant interac-
tions among volcanic/geochemical cycles, climate, and biota.
Recently, Central Atlantic magmatic province volcanism has
been shown to coincide with the end-Triassic extinction (Cirilli
et al., 2009; Deenen et al., 2010; Ruhl et al., 2010; Blackburn
et al., 2013), although some inconsistency of chronology occurs
between the studies of Deenen et al. (2010) and Blackburn et al.
(2013) from one side and Marzoli et al. (2011) on the other side.
Nevertheless, there remain uncertainties concerning cause and
effect, particularly in the terrestrial realm. Global anoxic events,
regression, primary productivity crash, methane hydrate disso-
ciation, and rapid global warming have all fi gured in recent pro-
posed kill mechanisms (Hallam and Wignall, 1997, 1999; Pálfy
et al., 2000, 2001; Hesselbo et al., 2000, 2002, 2007; Ward et al.,
2001, 2004; Galli et al., 2005; McElwain et al., 1999, 2007).
Series of perturbations in the global carbon cycle in par-
ticular have proved useful in evaluating the paleoenvironmental
changes during the biotic crisis, and also in discussion of the
separate issue of the placement of the Triassic-Jurassic boundary
and its global correlation. After recognition of an “initial” and a
“main” negative carbon isotope excursion (CIE), a more com-
plex picture is emerging with other excursions, both negative and
positive, prior to and following the Triassic-Jurassic boundary
(e.g., Pálfy et al., 2001; Galli et al., 2005; Hesselbo et al., 2007;
Hillebrandt et al., 2007; Korte and Kozur, 2011; Pieńkowski et
al., 2012).
The origin of the huge amount of CO2 or methane enriched
in 12C (13C depleted) is still a matter of debate; it could have
been derived directly from the Central Atlantic magmatic prov-
ince basalts, subvolcanic thermal metamorphism of subsurface
organic-rich strata, or from methane hydrates. None of these
mechanisms is mutually exclusive, and all three may have con-
tributed to the release of 12C-enriched carbon at the end of the
Triassic (Ruhl et al., 2011). Relative contributions of these end
members remain unknown, but given the very short duration and
magnitude of the observed end-Triassic 12C release, a signifi -
cant contribution from methane hydrates is likely. Nevertheless,
cumulative carbon release might have been capable of amplify-
ing an initial warming, resulting in runaway greenhouse condi-
tions. Support for this scenario comes from fossil plants through
stomatal density studies and other paleobotanical work (McEl-
wain et al., 1999, 2007; Bacon et al., 2013).
Resolution of the extinction timing and the recognition of
an ecological selectivity for extinctions of different groups of
plants and animals have also helped to clarify the nature of the
crisis (McRoberts and Newton, 1995; Ward et al., 2001, 2004;
McElwain et al., 1999, 2007; Pieńkowski et al., 2012). The direct
causal link between carbon-cycle disturbance (methane release)
and onset of terrestrial biotic disturbances has been recently chal-
lenged, on the basis that the onset of the biotic crisis predates
the initial negative δ13C excursion (Lindström et al., 2012). Fur-
thermore, it is unclear what caused the quite conspicuous (Hes-
selbo et al., 2002; Ruhl et al., 2009; Lindström et al., 2012) ear-
lier Rhaetian δ13C excursions, which predate the onset of Central
Atlantic magmatic province volcanism. There are also reports
pointing out possible cooling and/or release of toxic compounds
as a cause of extinction (Hubbard and Boutler, 2000; Schoene et
al., 2010; van de Schootbrugge et al., 2009). Furthermore, Korte
et al. (2009) provided δ18O data from fossil oysters that argue for
cool ocean temperatures immediately after the initial δ13C excur-
sion, followed by ~8 °C early Hettangian warming.
The present study is based on data sets obtained from two
boreholes (Kamień Pomorski IG-1 and Niekłań PIG-1, referred
to herein as Kamień Pomorski and Niekłań) and two outcrops,
Lisowice and Sołtyków (Fig. 1), which yielded records across
the Triassic-Jurassic transition in continental deposits (lacus-
trine–alluvial plain). Many of the sedimentological, geochemi-
cal, and palynological results from the Kamień Pomorski bore-
hole have been published (Pieńkowski et al., 2012). In this paper,
palynofl oral extinction level, appearance of new forms commenced Jurassic palyno-
fl oral recovery. Tetrapod evolution events in the end-Triassic–earliest Jurassic were
related to the extinction of the Pseudosuchia, Dicynodontia, Capitosauroidea, Plagio-
saroidea, and Rhynchosauria, while appearance of highly diversifi ed tetrapod ichno-
fauna in the earliest Jurassic strata indicates a rapid recovery and refi ll of ecological
niches by dinosaurs.
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Climatic reversals related to volcanism caused the end-Triassic biotic crisis—Evidence 265
we complement these results with clay mineral studies and dis-
cuss the results in the context of currently expanded knowledge
of the Triassic-Jurassic boundary events, including mass extinc-
tion. A temporal sequence of events at the end of the Triassic and
beginning of Jurassic is presented, and we also attempt to link
environmental/climate changes with tetrapod biota evolution,
based on data from Lisowice and Sołtyków, in order to arrive at
a more comprehensive picture of the continental end-Triassic–
earliest Jurassic climate, mass extinction of plants and tetrapods,
and their subsequent recovery.
GEOLOGICAL SETTING
Tectonics
In the late Rhaetian and early Hettangian, the Polish Basin
was located at about 40 degrees of the northern latitude, some
1200–1500 km from the nearest Central Atlantic magmatic
province volcanic extrusions in Europe (Fig. 1). The Kamień
Pomorski and Niekłań boreholes, ~500 km away of each other,
are both located in the Mid-Polish Trough, as is the Sołtyków
outcrop, situated close to the Niekłań borehole, while the Liso-
wice outcrop is located further to the south-west, outside the
Mid-Polish Trough (Fig. 1). The Mid-Polish Trough is one of
many inverted basins in Western and Central Europe (Ziegler,
1990), running generally along the Teisseyre-Tornquist zone and
the Trans-European suture zone. The Mid-Polish Trough is the
largest of these basins, with a length of more than 700 km, and
it contains an accumulated sedimentary thickness of the order of
10 km. Rapid subsidence in the trough commenced in the early
Hettangian (Pieńkowski, 2004), and in the Rhaetian subsidence
was less marked (Fig. 1). Additionally, in Rhaetian-Hettangian
times, the area around Kamień Pomorski was affected by a num-
ber of N-S–trending faults, grabens, and semigrabens (Dadlez,
1969; Dadlez et al., 1995). These synsedimentary faults and gra-
bens acted as “catchment traps” for sandy sediments delivered
from the Kaszuby Land, located eastward from the Koszalin-
Chojnice fault zone (Fig. 1).
100 km
WARSAW
p
a
t
h
r
i
a
a
n
C
-
L
e
a
r
n
o
d
F
inferred maximum range
of the Rhaetian deposits
main synsedimentary
faults
maximum thickness of the
Rhaetian deposits > 100 m
MID - POLISH TROUGH
Kaszuby
Land
Lublin Land
Sudety Land
HOLY CROSS
MOUNTAINS
BASIN
P
O
M
A
I
E
N
R
A
POLISH
BASIN
500 km
boreholes and outcrops
Lisowice
800 km
sills and
lava flows
CAMP
boundary
Polish Basin
main directions
of sediment delivery
Koszalin - Chojnice
fault zone
Figure 1. Late Rhaetian Polish Basin, its location in the European Basin, and its position on Earth ca. 201 Ma, in relation to the Central Atlantic
magmatic province (CAMP).
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266 Pieńkowski et al.
Figure 2. Kamień Pomorski IG-1 profi le showing integrated stratigraphy, lithology, kaolinite-illite ratio in the <2 µm fraction, chemical index of alteration (CIA) in bulk rock, iridium
content, carbon and osmium isotopes, total organic carbon (TOC), organic geochemistry (pyrolytic polycyclic aromatic hydrocarbons [PAH] and total organic carbon [TOC]), and
miospore characteristics (color, diversity, and abundance). Inferred climate changes are based on clay mineralogy, and to lesser extent on lithology (sediment color), palynology,
and other indices. Note increasing frequency of these changes above the “initial” carbon isotope excursion (CIE) toward the Triassic-Jurassic boundary, established on the basis of
biostratigraphy. Osmium isotope ratios 187Os/186Os and 187Os/188Os show almost identical changes; therefore, they are presented as one plot with two scales, respectively. Stratigraphy,
lithology, C and Os isotopes, and miospore characteristics are after Pieńkowski et al. (2012); pyrolytic PAH is after Marynowski and Simoneit (2009). FAD—fi rst appearance datum;
LAD—last appearance datum; VPDB—Vienna Pee Dee Belemnite standard.
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Climatic reversals related to volcanism caused the end-Triassic biotic crisis—Evidence 267
Stratigraphy and Sedimentology
The Triassic-Jurassic boundary is located between the last
appearance datum (LAD) of megaspore Trileites pinguis (Mar-
cinkiewicz, 1962, 1971) and fi rst appearance datum (FAD) of
Cerebropollenites thiergartii in the Kamień Pomorski profi le
(Fig. 2; Pieńkowski et al., 2012). This placement of the Triassic-
Jurassic boundary in Kamień Pomorski is further supported by
major palynofl oral turnover from the Cingulizonathes rhaeticus–
Limbosporites lundblandii association, which corresponds to the
Rhaetipollis-Ricciisporites (= Rhaetipollis-Limbosporites) zone
of Rhaetian age, and the typically Hettangian Conbaculatisporites
mesozoicus–Dictyophyllidites mortoni– Cerebropollenites thier-
gartii association (with the age-diagnostic pollen C. thiergar-
tii), which corresponds to the Pinuspollenites-Trachysporites (=
Trachysporites-Heliosporites) zone (Pieńkowski et al., 2012).
This boundary is correlated with the base-Jurassic global stra-
totype section and point (GSSP) profi le in Kuhjoch (Kürschner
et al., 2007; Hillebrandt et al., 2007). A factor of crucial signifi -
cance for stratigraphy is chemostratigraphic correlation between
Kamień Pomorski and other profi les, based on carbon isotopes
(Pieńkowski et al., 2012, their fi g. 7).
Due to local tectonic conditions, the majority of coarser
sediments were deposited east of Kamień Pomorski, and further
to the west, the channel energy is inferred to have decreased on
a “sand-starved” alluvial plain (Pieńkowski et al., 2012). Thus,
muddy overbank deposits dominate in the Kamień Pomorski pro-
fi le (Fig. 2). Fault activity in Rhaetian times periodically created
additional accommodation space, allowing temporarily enhanced
deposition of thicker packages of sediments.
Subsequent Early Jurassic sedimentation in Poland, includ-
ing Pomerania and the Holy Cross Mountains, was characterized
in detail by Pieńkowski (2004). Earliest Hettangian alluvial and
lacustrine deposition (although much steadier and faster) was a
continuation of Rhaetian deposition (Figs. 2 and 3). Therefore,
both upper Rhaetian and lowermost Hettangian alluvial and
lacustrine deposits are assigned to the same lithoformation—
Zagaje Formation (Pieńkowski, 2004).
Preliminary geological and sedimentological observations
from Lisowice outcrop (spanning the Triassic-Jurassic boundary)
were presented by Szulc et al. (2006), Dzik et al. (2008a, 2008b),
Niedźwiedzki and Sulej (2008, 2010), and Niedźwiedzki et al.
(2012). We interpret these sediments as having been deposited
on an alluvial plain (see Fig. 4 and the “Results” section). The
whole 12-m-thick section at Lisowice is assigned herein to the
informal lithostratigraphic units of the Zbąszynek Beds (lower
red beds) and Wielichowo Beds (gray sandstone-mudstone bone-
bearing interval plus heterolith unit). In previous work, the sec-
tion in Lisowice was included in the other informal lithological
unit, Woźniki Limestones or Woźniki Formation (Szulc et al.,
2006). However, this diachronous red bed–carbonate unit does
not fi t into the general lithological character of the beds exposed
in Lisowice, particularly in regard to the middle, gray-colored
part of the outcrop (Fig. 4). Presumably, the Woźniki Limestones
can be correlated laterally only with palustrine carbonate depos-
its (crenogenic in origin; see Szulc et al., 2006). These carbonates
are also exposed in a small, abandoned quarry located ~600 m to
the north of the clay pit.
Based on the palynomorph assemblages (Staneczko, 2007;
Dzik et al., 2008a, 2008b; Świło et al., 2013), these strata would rep-
resent the subzone IVc (defi ned by the appearance of Rhaetipollis
germanicus in Orłowska-Zwolińska 1983, 1985) of the Corollina
meyeriana zone, and the Ricciisporites tuberculatus zone, which
was later incorporated into the Rhaetian Rhaetipollis germanicus
zone (Lund, 1977; Herngreen, 2005; Kürschner and Herngreen,
2010). According to the existing stratigraphical interpretation (Fig.
4), the Lisowice strata correlate with latest Norian Zbąszynek
Beds and Rhaetian Wielichowo Beds (Deczkowski, 1997; Franz et
al., 2007a, 2007b; Franz, 2008). The presence of Rhaetian strata
is further supported by cuticle fragments of the typical Rhaetian
seed-fern Lepidopteris ottonis (Goeppert, 1836) (Staneczko, 2007;
Ociepa et al., 2008; Dzik et al., 2008a, 2008b; Wawrzyniak and
Ziaja, 2009; Wawrzyniak, 2010a, 2010b, 2010c, 2011), the conifer
Stachyotaxus septentrionalis (this genus has a distribution strongly
restricted to the Rhaetian deposits of Scania and Greenland; see
Taylor et al., 2009), isoëtalean macrospores Trileites cf. pinguis
(Harris, 1935), and Horstisporites bertelseni Fuglewicz, 1977
(Fuglewicz and Śnieżek, 1980). At Lisowice, two horizons with
conchostracans were identifi ed, the lower with numerous Shipingia
sp. and rare small Euestheria sp., and the upper with Gregoriusella
polonica Kozur et Weems, 2010. These two conchostracan zones
identify the presence of the latest Norian and early (possible also
younger) Rhaetian deposits (Kozur and Weems, 2005, 2007, 2010),
consistent with palynological data.
An outcrop of the Hettangian alluvial-plain deposits in
Sołtyków, located close to the Niekłań borehole (Fig. 1), was
described by Pieńkowski (2004). The strata there represent a
siliciclastic coal-bearing lithofacies association of an alluvial
wetland, with deposition controlled by high-sinuosity, anasto-
mosing streams. The climate was warm to temperate, with wet-
ter and drier seasons. An earliest Hettangian age is indicated by
sequence stratigraphic correlation (Pieńkowski, 2004), macro-
fl ora dominated by the conifer Hirmeriella muensteri (Schenk)
Jung and the benettite Pterophyllum alinae Barbacka, 2010 (Rey-
manówna, 1992; Wcisło-Luraniec, 1991; Barbacka et al., 2010),
palynomorphs (Ziaja, 2006), and conchostracans Bulbilimnadia
kilianorum Kozur, Weems, et Lucas, 2010).
MATERIAL AND METHODS
Recovery of core in the Kamień Pomorski borehole was
~42% in the Rhaetian and Hettangian, but, fortunately, a more
complete core was recovered from the most important Triassic-
Jurassic boundary interval. Moreover, the Kamień Pomorski
profi le represents a fairly continuous succession through the
Triassic-Jurassic transition, which is unique in the Polish Basin.
The new Niekłań core is 100% complete, but the Rhaetian pro-
fi le is much more fragmentary due to hiatuses. However, despite
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268 Pieńkowski et al.
Figure 3. Niekłań PIG 1 borehole Triassic-Jurassic continental transition section showing stratigraphy, lithology, clay mineralogy, and weathering indices established in conti-
nental mudrocks (s.b.—sequence boundary). Note the mineralogical and geochemical record of a major climatic shift in the Rhaetian and series of frequent and abrupt changes
in the clay mineral composition and weathering indices values below and at the Rhaetian-Hettangian (= Triassic-Jurassic) boundary, which correspond to inferred climate rever-
sals. Explanations: Clay minerals: K—kaolinite, I—illite, Ch—chlorite, Sm—smectite; weathering indices: Al/K—aluminum/potash ratio; chemical index of alteration (CIA) =
(Al2O3/[Al2O3 + CaO* + Na2O + K2O]) × 100 (Nesbitt and Young, 1982); chemical index of weathering (CIW) = (Al2O3/[Al2O3 + CaO* + Na2O]) × 100 (Harnois, 1988); plagioclase
index of weathering (PIA) = {(Al2O3 – K2O)/([Al2O3 – K2O] + CaO* + Na2O)} × 100 (Fedo et al., 1995; for lithology column legend see Fig. 2). Stratigraphical position of the nearby
(~4 km) Sołtyków outcrop is shown.
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Climatic reversals related to volcanism caused the end-Triassic biotic crisis—Evidence 269
Figure 4. Lithologic, stratigraphic, and sedimentologic profi le of the main exposure in the Lisowice (= “Lipie Śląskie” clay pit at Lisowice,
Silesia) with position of the bone-bearing interval, tetrapod trace fossils, other faunal and fl oral remains, and intervals with red-bed deposits.
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270 Pieńkowski et al.
fragmentary preservation of the Rhaetian section, this core
yielded a valuable, continuous profi le through the Hettangian,
together with clay mineral data refl ecting important Rhaetian cli-
matic changes, and giving insight into the mineralogy of under-
lying upper Norian red beds, deposited under dry climate condi-
tions. Moreover, the Niekłań borehole is located very close to the
Sołtyków outcrop, allowing integrated interpretation with very
rich tetrapod ichnofossils (Gierliński et al., 2004; Niedźwiedzki,
2006, 2011b).
We acknowledge that the analysis of 24 samples (Kamień
Pomorski) for δ13Corg, 15 samples for palynology, and even fewer
for Os isotopes, spanning >60 m of section, is a relatively small
number of data on which to report. However, 35 m of those 60
m section were not cored at all. We sampled the remaining 25 m
for δ13Corg as densely as possible (depending on the core preser-
vation), and the average density of sampling in cored profi le is
close to 1 sample per meter, although sampling density is irregu-
lar, again due to core preservation and lithology (only mudstones
yielded suffi cient palynomaceral material).
Palynology, Carbon and Osmium Isotopes, Iridium
Content, and Organic Geochemistry in the Kamień
Pomorski Profi le
Palynological results, carbon isotopes, osmium isotopes,
and organic geochemistry in the Kamień Pomorski profi le are
taken from the previous paper by Pieńkowski et al. (2012),
and a detailed description of material and methods is included
therein. Fifteen rock samples from the Kamień Pomorski core
were selected for quantitative palynological analysis, 12 of which
yielded palynomorphs (Fig. 2).
Samples for δ13Corg from the Kamień Pomorski borehole
(Fig. 2) were separated manually under a binocular microscope
from palynomorphs, and only woody phytoclast separates were
taken for the carbon isotope analysis (homogenized samples).
The carbon isotope ratios were measured using an elemental ana-
lyzer Carlo-Erba 1110 connected online to a Thermo Finnigan
Delta Plus mass spectrometer (see Pieńkowski et al., 2012).
Analyses of osmium, iridium, and rhenium content, as well
as analyses of the following isotopes, 186Os, 187Os, 188Os, 192Os,
and 187Re, were carried out from nine samples (Fig. 2). The ana-
lytical procedure used here followed that described in detail by
Brauns (2001). The isotopic composition of Os in the blank is
very close to the natural composition and has a ratio of 0.112
for 187Os/188Os. All data were blank corrected on the basis of
these measurements in combination with a yield of 90%, and
an Os blank of 0.10 pg. For more information, see Pieńkowski
et al. (2012).
The total organic carbon (TOC) and polycyclic aromatic
hydrocarbons (PAH) were determined in 24 mudstone samples
using a chromatographic, coulometric method (procedure PB-23)
in an automated LECO analyzer. The gas chromatography–mass
spectrometry (GC-MS) analyses of the aliphatic and aromatic
fractions were performed with an Agilent 6890 Series Gas Chro-
matograph interfaced to an Agilent 5973 Network Mass Selective
Detector and Agilent 7683 Series Injector (Agilent Technologies,
Palo Alto, California) by Marynowski and Simoneit (2009).
Clay Mineralogy
The bulk-rock mineralogy, clay mineralogy, and major-ele-
ment geochemistry were determined from 40 claystone and mud-
stone samples from Kamień Pomorski and the lower part of the
Niekłań borehole. Bulk-rock mineralogy and clay minerals (in
the <2 µm fraction) were identifi ed by X-ray diffraction (XRD)
using a Phillips PW 3020 X’Pert diffractometer with CuKα radi-
ation. Major-element amounts were measured by X-ray fl uores-
cence using a Phillips PW 2400 spectrometer. All samples were
analyzed at the Polish Geological Institute–National Research
Institute laboratories. For more details, see Brański (2014).
Tetrapods
The vertebrate remains and trace fossils were described
from the uppermost Norian–Rhaetian strata of Lisowice, while
a rich assemblage of the earliest Hettangian tetrapod trace fossils
comes from the Sołtyków outcrop. In order to obtain the sedi-
mentological and stratigraphical framework (Fig. 4) for the most
important part of the profi le in Lisowice with spectacular bone-
bearing deposits, a few short trenches were excavated through
the site (2007–2013). A detailed description of the taphonomic
processes recognized for all bone-bearing horizons at this site is
presented in Niedźwiedzki (2014).
RESULTS
Lithology
The Upper Norian deposits in the Niekłań borehole (Fig. 3)
and uppermost Norian Lisowice outcrop (Fig. 4, the lower red
bed) represent a red-bed association (mostly mudstones and silt-
stones) with rare calcrete, other carbonate nodules, and oxidized
plant roots, and are assigned to the Zbąszynek Beds. The super-
posed Rhaetian sediments generally show bipartite development
(Figs. 2 and 3). The lower part (belonging to the Wielichowo
Beds) is represented by red-brownish, yellow-greenish, or var-
iegated mudstone, with poorly preserved (oxidized) rootlets and
scattered calcium carbonate concretions or carbonate calcretes
in its lower part. In the Kamień Pomorski and Niekłań bore-
holes, the Upper Rhaetian (belonging together with the overlying
Lower Hettangian strata to the Zagaje Formation) commences
with sandstone resting on an erosional boundary, identifi ed with
the sequence boundary (depth of 698 m in Kamień Pomorski
[Fig. 2]; 180 m in Niekłań [Fig. 3]). Upper Rhaetian mudrock of
the Zagaje Formation (678.4–693 m [Fig. 2]; 162–180 m [Fig.
3]) shows a conspicuous change in color into gray/dark gray in
Kamień Pomorski. However, there are thin intervals showing an
intermittent return to a variegated or reddish gray color (Fig. 2).
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Climatic reversals related to volcanism caused the end-Triassic biotic crisis—Evidence 271
In Niekłań, the color change from red to variegated colors occurs
in the bottom of the 162–180 m interval, and gray sediments
appear in the top of this interval (Fig. 3).
This change of color into gray/dark gray is associated with
the appearance of frequent paleosols with coalifi ed rootlets,
occasionally topped with thin lignite beds, which further point
to a general humidifi cation of climate. At the top of the Rhae-
tian deposits, the second sequence boundary occurs (678.4 m
in Kamień Pomorski; 161.9 m in Niekłań), slightly below or
at the Triassic-Jurassic boundary. This boundary, linked to
another sea-level lowstand at the Triassic-Jurassic boundary
(Pieńkowski, 2004), is usually marked by regional erosion in the
whole sedimentary basin of Poland (quite often, the Rhaetian
deposits are missing in the whole basin or for a signifi cant part).
Only in Kamień Pomorski, due to a local tectonic regime, is this
erosion less signifi cant (fi ne-grained sandstone above the ero-
sional boundary does not contain coarser grains or mud clasts,
and bedding at the bottom of pyritic sandstone is characteristic
of a rhythmic, lower-fl ow regime; see Pieńkowski et al., 2012).
On the other hand, the Niekłań profi le (Fig. 3) shows signifi cant
erosion and hiatuses at the inter-Rhaetian and Triassic-Jurassic
sequence boundaries. Judging from clay mineral composition
and its comparison to the more complete Kamień Pomorski
profi le, the whole uppermost Rhaetian section is missing in
Niekłań. The Sołtyków outcrop exposes only the lowermost
Hettangian strata (Fig. 3; for detailed profi le of the outcrop, see
Pieńkowski, 2004).
Above the Norian red beds of the clay pit at Lisowice (Fig.
4), the major part of the section is represented by gray mud-
stones and cross-bedded sandstones (graywackes), mainly with
trough cross-bedding (general transport of material was to NE;
Fig. 1) and ripple-drift cross-lamination, including climbing rip-
ples. The epsilon cross-bedding produced by lateral migration
of channels is common. Both sandstone and mudstone lithofa-
cies contain numerous drifted plant fossils and pedogenic hori-
zons. The mudstones and sandstones are slightly calcareous,
likely because of redeposition and destruction of older carbon-
ate rocks. Layers of palustrine carbonates or dolomitic mud-
stones also occur (called regionally the “Woźniki Limestone,”
a diachronous and genetically inhomogeneous lithological
unit). Conglomerates and breccia-like coarse sediments appear
mostly at the bottoms of sandstone layers (= bottoms of fl uvial
cycles) and are composed of reworked carbonate nodules from
older Upper Triassic sediments and rock fragments (quartzites
of shales, probably of Paleozoic age). These conglomerates are
traditionally known as the “Lisów breccia” bed (diachronous
lithological unit). A factor of note is a layer of disturbed bed-
ding in the middle part of the profi le (Fig. 4), possibly repre-
senting a tectonic shake effect. A red-greenish heterolith unit
(interbedded mudstone, siltstone, and sandstone beds) occurs in
the top of the outcrop, and it contains carbonate nodules, often
infi lling plant roots (Fig. 5).
There are two major intervals with bones in Lisowice (Fig.
4), and the upper (main) one has four discrete bone-bearing
Figure 5. Top 3 m of the Lisowice outcrop (see Fig. 4), showing the
gray mudstones (the upper part of the interval with bones), capped
by the red-greenish heterolithic unit with pedogenic horizons. Note
numerous plant roots associated with carbonate nodules (inset). The
heterolithic unit probably refl ects climatic fl uctuation toward aridity.
Such fl uctuations might have caused biotic crises, including extinction
of vertebrates. Scale bar is 1 m.
beds, separated by bone-barren mudstone-siltstone levels with
numerous plant remains and casts of bivalve shells. Lithologi-
cal characteristics, geometry of the lithic units, and sedimentary
structures (Fig. 4) suggest that the gray (middle) strata in Liso-
wice were formed in a mosaic of alluvial-plain environments—
swamps with ephemeral ponds and laterally shifting river chan-
nels. In particular, texturally immature sediments (graywackes),
lack of stable channels, and presence of climbing ripples point to
an increased rainfall and runoff, combined with fast deposition.
This interpretation is supported by macrofl oral and microfl oral
remains (Fuglewicz and Śnieżek, 1980; Wawrzyniak, 2010a,
2010b, 2010c, 2011). Both red bed units at the bottom and at the
top of the outcrop were also deposited on an alluvial plain, but in
semiarid climate conditions.
Clay Mineralogy
The examined bulk-rock samples of claystones and mud-
stones from the Zbąszynek Beds (Norian), Wielichowo Beds, and
Zagaje Formation (late Rhaetian–early Hettangian) of Kamień
Pomorski (Fig. 2) and Niekłań borehole (Fig. 3) are mainly com-
posed of phyllosilicates (50%–90%). Quartz is less abundant, but
occurs in variable amounts (7%–43%). In the Wielichowo Beds
(early-middle Rhaetian), hematite, goethite, feldspars, calcite,
and locally dolomite were observed with proportions of 0%–8%.
In contrast, in the Zagaje Formation, calcite and dolomite are
absent, but instead siderite is present in some samples with pro-
portions of 0%–7%. Feldspar, hematite, and goethite appear spo-
radically, mostly in accessory amounts.
with paleosols and carbonate nodules
with paleosols and carbonate nodules
Interval with bones
Interval with bones
Red and green/gray heteroliths
Red and green/gray heteroliths
(uppermost part)
(uppermost part)
with paleosols and carbonate nodules
Interval with bones
Red and green/gray heteroliths
(uppermost part)
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272 Pieńkowski et al.
The clay mineral composition is very diverse in the <2 µm
fraction, which is refl ected in kaolinite/illite, and kaolinite/illite +
chlorite, and in places kaolinite/smectite ratios (Figs. 2 and 3).
In three samples taken from the Upper Norian strata (Zbąszynek
Beds; Fig. 3), illite (with subsidiary chlorite) totally predomi-
nates, and kaolinite content is insignifi cant. In the overlying
Wielichowo Beds (Rhaetian), the composition of clay mineral
assemblage is different (Figs. 2 and 3). With exception of a few
samples (see following), an abundance of smectite is observed;
kaolinite appears mostly in lesser amounts (Fig. 3) or is absent
(Fig. 2). In Kamień Pomorski, smectite makes up 100% of clay
minerals in claystones of the Wielichowo Beds (Fig. 2). Small
amounts of illite and chlorite were detected only in a single
sample in Niekłań. In the superposed Zagaje Formation, kaolin-
ite generally predominates (with an average content of 54% in
Kamień and 49% in Niekłań) over illite (average content of 46%
and 33%, respectively). Chlorite is still absent in the late Rhae-
tian part of the Zagaje Formation. It should be noted that in the
lowest part of this formation in Niekłań section, a signifi cant con-
tent of smectite (up to 70%) was still observed (Fig. 3). However,
smectite is absent in the Zagaje Formation in Kamień Pomorski,
while the kaolinite phase is accompanied by signifi cant amounts
of berthierine or serpentine. In the Hettangian part of Zagaje For-
mation (Fig. 3), the clay mineral assemblage consists of kaolin-
ite and illite, with signifi cant subsidiary chlorite, and smectite is
almost absent.
Clay mineral data are refl ected in the major-element geo-
chemistry and in the values of alteration indices, which are high
in Upper Rhaetian and Lower Hettangian strata. The chemical
index of alteration (CIA) ranges from 79 to 96, but in the vast
majority of cases, it exceeds 85, and these are due to residual
clays (Nesbitt and Young, 1982). CIA shows the loss of Ca2+, K+,
and Na+ in relation to Al2+ (Nesbitt and Young, 1982; see caption
of Fig. 2). Therefore, CIA data are very useful in recognizing
climate fl uctuations.
Clay minerals in the studied sections are largely detrital and
show generally insignifi cant diagenetic overprint (Brański, 2009),
due to moderate burial and to the fairly closed hydrologic sys-
tem. There is no distinct and systematic evolution of clay- mineral
assemblages from top to bottom of the studied boreholes that
would be indicative of burial diagenesis. Importantly, smectite
is abundant in Lower-Middle Rhaetian Wielichowo Beds despite
its sensitivity to diagenetic alteration. Low thermal alteration
of the Rhaetian-Hettangian strata in the Polish Basin was con-
fi rmed by palynomorph color (Pieńkowski and Waksmundzka,
2009) and biomarkers (Marynowski and Simoneit, 2009). More-
over, a focus of this study is kaolinite content, which is resistant
under moderate diagenetic conditions. Moreover, the scanning
electron microscope (SEM) observations show that clay miner-
als are mostly detrital, and authigenic kaolinite appears rarely.
In the studied <2 µm fraction, most diagenetic kaolinite should
be excluded, because this has a typical particle size of 5–10 µm
(when it has replaced feldspar) and an order of magnitude greater
when it pseudomorphs mica (e.g., Hesselbo et al., 2009). On the
other hand, breaking of kaolinite crystallites may occur during
sample preparation (Hesselbo et al., 2009).
The kaolinite peaks in some intervals in the Rhaetian and
at the beginning of the Hettangian on clay mineral ratio curves
are of particular interest (Figs. 2 and 3). It should be noted that
the fi rst distinct rise of kaolinite content already appeared ear-
lier in the Rhaetian Wielichowo Beds (Figs. 2 and 3; see the 1
hh climatic step at 700 m in Fig. 2, where the sequences of fre-
quent and marked climatic reversals are named as 1hh, 2hh, 3
cd, 4 hh, 5 cd, 6 h, 7 cd, and 8 hh events [hh—hot and humid,
cd—cold and dry, h—humid]; Fig. 2; Table 1). Similar peaks are
observed in the Zagaje Formation, and they correspond in the
Kamień Pomorski profi le to the three rapid increases of kaolinite
through the late Rhaetian section of the Zagaje Formation, and
the last one at the beginning of Hettangian section of this forma-
tion. These peaks are described (Fig. 2; Table 1) as 2 hh, 4 hh,
6 hh, and 8 hh climatic steps.
Higher in the section, the kaolinite content signifi cantly
decreases and stabilizes (Fig. 3). The kaolinite peaks usually
correlate with the highest values of weathering indices (in the
most cases, CIA values are greater than 90). A few discrepancies
may relate to the fact that clay minerals were analyzed in the
<2 µm fraction, but the major-element content was measured on
bulk-rock samples, and most probably some of them (in Kamień
Pomorski borehole; Fig. 2) may have been somewhat infl uenced
by Na metaso matism (Brański, 2014).
Interestingly, recent studies focused on the weathering prox-
ies (CIA, chemical index of weathering [CIW] = [Al2O3/{Al2O3
+ CaO* + Na2O} × 100 [Harnois, 1988]; plagioclase index of
weathering [PIA] = [(Al2O3 – K2O)/{(Al2O3 – K2O) + CaO* +
Na2O}] × 100 [Fedo et al., 1995]) are indicative of intense con-
tinental weathering associated with pulsed end-Cretaceous Dec-
can eruptions and abundant rainfall (acid rains), which resulted in
mesotrophic waters (Gertsch et al., 2013).
Carbon Isotopes, Osmium Isotopes, Organic
Geochemistry, and Miospores
The values of δ13Corg (see Pieńkowski et al., 2012; Fig. 2;
Table 1) show signifi cant fl uctuations through the Rhaetian. The
most conspicuous negative δ13Corg excursion is identifi ed with the
Rhaetian “precursor” and double-peaked “initial” CIE, followed
by a positive excursion and again by slightly more negative val-
ues, probably representing subordinate fl uctuations within a lon-
ger positive excursion, where the Triassic-Jurassic boundary is
now inferred (Korte et al., 2009; Korte and Kozur, 2011). The
double-peaked initial CIE shows negative values of -29.38‰
δ13Corg at 691 m and -28.85‰ δ13Corg at 686 m, separated by a
more positive value (~-26‰ at 687 m). However, the interpreta-
tion of “subpeaks” should be treated with some caution, because
there are only a few carbon-isotope data, and each feature is rep-
resented only by a single data point, with signifi cant data gaps
separating each inferred excursion. The amplitude of negative
values indicates that these disturbances can be compared only to
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Climatic reversals related to volcanism caused the end-Triassic biotic crisis—Evidence 273
TABLE 1. SEQUENCE OF δ13C EXCURSIONS, LITHOLOGICAL, CLAY MINERAL, AND MINERALOGICAL CHANGES, OSMIUM ISOTOPE AND POLYCYCLIC AROMATIC
HYDROCARBON (PAH) EVENTS, CHARACTERISTICS OF MIOSPORE APPEARANCE, AND INFERRED SUCCESSION OF MAIN CLIMATE AND EVENTS IN THE KAMIE
POMORSKI BOREHOLE SECTION ACROSS THE TRIASSIC-JURASSIC (T-J) BOUNDARY
Depth (m),
event no. (Fig. 2)
δ13C excursions Sediment color, kaolinite/illite/
smectite, CIA
Os isotopes PAH and
TOC
Color of miospores Environment, climate
676.9
8hh
Slightly more positive Gray, kaolinite >> illite PAH
increase
Darkened
miospores
Intermittent drop in palynofl oral
diversity, fi nal disappearance of
Triassic miospores
677.5
7 cd–8hh
Slightly more negative Gray Increase of 192Os Appearance of new miospores, high
diversity, onset of Jurassic palynofl oral
recovery
678.6
(T-J boundary)
7cd
Slightly more positive Sequence boundary, dark-
gray, illite > kaolinite, drop in
CIA, pyrite
Drop in 187Os/186Os PAH and
TOC
increase
Strongly darkened
miospores, fern
peak
CAMP volcanic fallout, acid rains,
cool episode, extirpation of Triassic
palynofl ora
679.5
6 h
Slightly more negative Dark gray-olive gray, illite >
kaolinite, kaolinite rise
Intermittent warming
680–682.5
5 cd
Positive excursion Dark gray–reddish, illite >
kaolinite (rapid kaolinite drop)
Pollen grain>spores Drier and cooler
684.5, 4hh Return to background
values
Dark gray, rapid rise of
kaolinite, kaolinite >> illite
Hot and humid
686.0
3 cd–4 hh
“Initial” negative
excursion (upper
subpeak of the INE)
Dark gray, kaolinite = illite Os isotope disturbance,
decrease in 187Os/186Os,
187Os/188Os and 192Os rise
Darkened
miospores
CAMP volcanic fallout, acid rains,
beginning of fl oral turnover
687.0
3 cd
Positive subpeak
within the INE
Gray, illite > kaolinite Stabilization High PAH,
high TOC
Cooler and drier, forest fi res and/or
volcanogenic PAH release
689–692
2 hh
“Initial” negative
excursion (lower
subpeak of the INE)
Dark gray, k aolinite >> illite Slight disturbance in
187Os/186Os, 187Os/188Os, and
192Os
Hot and humid, ? oldest known CAMP
volcanism
699.3–700
1 hh
“Precursor” negative
excursion
Variegated, kaolinite >> illite;
698 m - sequence boundary
Slight decrease in 187Os/186Os
and 187Os/188Os
Spores>pollen
grains
Major climate turnover into more
humid, intensifi ed hydrolysis
>700 Reddish, 100% smectite,
carbonate nodules
Semidry, seasonal climate
Note: INE—“initial” negative excursion; CIA—chemical index of alteration; TOC—total organic carbon; CAMP—Central Atlantic magmatic province.
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274 Pieńkowski et al.
the “initial” CIE, because there are not such light values known
in the Rhaetian terrestrial organic matter (Hesselbo et al., 2002;
Whiteside et al., 2010; Ruhl et al., 2010, 2011).
A subsequent positive excursion follows (-23.67‰ δ13Corg
at 681.5 m, although this is again based on a single data point),
which is in turn followed by more negative values with subor-
dinate fl uctuations. At the Triassic-Jurassic boundary (677.5 m),
there is a slight trend toward more positive values at the interval
676.9–677.5 m, followed by an uncored section between 653.4
and 676.5 m for which there are no data. Noticeably, the sedi-
ments spanning the “initial” double-peaked δ13Corg excursion are
markedly expanded in comparison to other known profi les (~6 m
thick; Fig. 2), and assuming that the “initial” CIE lasted for only
some 10,000–20,000 yr (Ruhl et al., 2011) and the precompaction
thickness of this mud-sandy interval could be some 8–9 m (assum-
ing compaction factor of 1.4), we obtain an average sedimentation
rate of some 80–90 cm/1000 yr (taking 10,000 yr duration, or half
of this fi gure in case of 20,000 yr). Such a sedimentation rate is
reasonable during an intensifi ed hydrologic cycle for an alluvial-
plain environment, under steady subsidence rate.
The fi rst suites of osmium/rhenium isotope data from
the continental deposits across the Triassic-Jurassic boundary
were obtained by Pieńkowski et al. (2012); here, we use these
results to further investigate geochemical events occurring at the
Triassic-Jurassic boundary. The data set, obtained from nine sam-
ples (Fig. 2), represents corrected, initial Re/Os isotopic ratios
of 187Os/186Os and 187Os/188Os, as well as content of unradiogenic
osmium isotope 192Os (Fig. 2; Table 1).
In the lowermost part of the section, 187Os/188Os and
187Os/186Os ratios slightly decrease upward, along with the sta-
ble 192Os, up to the sequence boundary at 678.4 m. Conspicu-
ous decreases of 187Os/188Os and 187Os/186Os, linked with a sharp
increase in 192Os, are associated with the CIE (higher “subpeak”
of the “initial” CIE at 686 m) and are interpreted as the result
of a volcanic fallout event (Pieńkowski et al., 2012). Another
decrease of 187Os/188Os and 187Os/186Os at 678.6 m, just below
the Triassic-Jurassic boundary (though linked with a decrease
in 192Os), could also refl ect volcanic fallout (association with
darkened miospores). Higher in the profi le, the 187Os/188Os and
187Os/186Os ratios in the Hettangian section increase. The con-
tent of 192Os drops back to Lower/Middle Rhaetian values (one
sample at depth 652.1 m). All measured values of the initial ratio
187Os/186Os are around 3 or higher (Fig. 2), which is indicative
of a crustal origin (Koeberl and Shirey, 1997; Koeberl, 1998).
Additionally, iridium content was measured, and all the values
were very low, below 10 ppt (Fig. 2). The disturbances in the Os
isotopic system are coeval or almost coeval with two levels show-
ing elevated polycyclic aromatic hydrocarbon (PAH) contents
(Marynowski and Simoneit, 2009). Their fi rst increase of PAH
content slightly predates (is 1 m below) the major Os excursions
at the depth of 686 m, the second PAH abundance peak coincides
with the less-marked excursions at the Triassic-Jurassic bound-
ary (677.5–678.6 m), and the third at 652 m (Hettangian) is not
related to any Os system disturbance (Fig. 2).
Systematic description of miospores was given by
Pieńkowski et al. (2012). There are three levels of darkened mio-
spores (Fig. 2; Table 1; 686 m, 678.6 m [Rhaetian], and 676.6 m
[Hettangian] section), indicating acid rains (van Schootbrugge et
al., 2009; Pieńkowski et al., 2012). Darkened miospores at 686 m,
within the “initial” negative isotope excursion, coincide with
carbon and osmium isotope as well as with PAH disturbances.
Between 682.5 and 700 m in Kamień Pomorski (Fig. 2), the mio-
spores are very scarce, 3–5 specimens per sample. Above 682.5 m,
there is an increase in abundance and diversity of spores, and
the changes of spore/pollen grain proportions generally fol-
low the changes in kaolinite/illite ratio (Fig. 2). Just below the
Triassic-Jurassic boundary, the abundance of miospores is very
high (1355 specimens), linked with their marked darkening,
187Os/186Os and 187Os/188Os drop, as well as a signifi cant drop in
kaolinite and CIA (Fig. 2; Table 1). Both diversity and abundance
of miospores are generally much higher in the Lower Hettangian
strata, and spores dominate over pollen grains (Pieńkowski et al.,
2012; Fig. 2). Another spore peak occurs higher up, within the
Hettangian section (Fig. 2, depth 641.8 m), but within this peak,
we observe much more diversifi ed palynomorphs (i.e., number
of palynofl oral taxa), and also many more pollen grains produced
by gymnosperm plants.
Tetrapod and Other Vertebrate Fossils
The recovered accumulation of bones in Lisowice includes
over 400 large tetrapod bones and over 500 small vertebrate
remains (fi sh teeth and scales, small tetrapod bones and teeth),
representing up to 15 species (Niedźwiedzki, 2014). The verte-
brate assemblage contains typical Late Triassic components and is
dominated by bones of a giant dicynodont (Fig. 6D) and remains
of actinopterygian fi shes. Other vertebrate skeletal remains are
rare and include small to large archosauromorphs, a large capi-
tosaur, and a small plagiosaur. Well-preserved bones occur only
in the lenticular body of gray mudstone and claystone deposits
(Figs. 4 and 5). The Lisowice clay pit is dominated by terrestrial
rather than amphibious or aquatic tetrapods. In two layers with
large wood fragments, remains of huge dicynodonts are associ-
ated with remains of Smok wawelski (a basal predatory dinosaur),
small- to medium-size archosaurs (pterosaur, dinosauromorph,
dinosaur, and poposaurid), and other more non-archosaur archo-
sauromorphs (a choristodere-like animal and rhynchosaur). Iden-
tifi cation of many of the disarticulated, often fragmentary bones
of other tetrapods is diffi cult and usually problematic (e.g., Dzik
et al., 2008a, 2008b; Niedźwiedzki et al., 2011, 2012).
The temnospondyl amphibians (Cyclotosaurus sp. and Ger-
rothorax sp.) are known from an isolated, partially preserved
skull (Fig. 6B), skull bones, jaw bones, and numerous long bones
collected in a layer less than a meter above the principal bone-
bearing bed of the clay-pit exposure. A few isolated long bones
of capitosaur and plagiosaur were also found in the main bone-
bearing horizon. Such “amphibian assemblages” are typical of a
frequently fl ooded alluvial fl oodplain. Numerous macroremains
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Climatic reversals related to volcanism caused the end-Triassic biotic crisis—Evidence 275
of coelacanth and dipnoan fi shes, hybodont sharks (Świło,
2010a, 2010b, 2010c; Świło and Kowalski, 2011), and palae-
onisciform fi shes were also found. A very rare mammaliaform
or mammal tooth (Morganucodontidae indet.) was also recorded
(Świło et al., 2013).
The tetrapod ichnofauna of the Zagaje Formation (lower
Hettangian) exposed at the Sołtyków clay pit and also other
sites (Figs. 7A–7F) shows a high ichnotaxonomic diversity
(Gierliński and Pieńkowski, 1999; Gierliński et al., 2004;
Niedźwiedzki, 2011b). Rare tetrapod bones have also been
found (Fig. 7G). After re-examination of the collected material,
specimens observed in the fi eld, and the study of extended, as-
yet-undescribed material from more recent collecting activities
(Niedźwiedzki, 2011b), the ichnotaxa assemblage includes 12
ichnotaxa of dinosaurs representing: predatory dinosaur foot-
prints; trackways and tracks of herbivorous sauropodomorphs;
and isolated prints of early ornithischia (Stenonyx isp., Grallator
isp., Anchisauripus isp., Kayentapus soltykovensis, Eubrontes
isp., cf. Megalosauripus isp., Anomoepus isp., Delatorrichnus
isp., Parabrontopodus isp., cf. Otozoum isp., cf. Tetrasauropus
isp.). Dinosaur swimming traces are represented by Charac-
ichnos isp. The assemblage refl ects notable dinosaur diversity,
thus far not known from other Hettangian or pre–Early Juras-
sic localities. By the morphology of imprints, the most abun-
dant large, tridactyl theropod dinosaur ichnotaxa (Kayentapus
soltykovensis, Eubrontes isp., cf. Megalosauripus isp.) can be
attributed to three species of early predatory dinosaurs. Par a-
brontopodus isp., cf. Otozoum isp., and cf. Tetrasauropus isp.
might possibly represent footprints of early sauropodomorphs
(“prosauropods” and early sauropods). The numerous gigan-
tic and large theropod dinosaur footprints were discovered in
the Sołtyków track site. These fi nds provide evidence for the
occurrence of gigantic and large predatory dinosaurs in the ear-
liest Jurassic (early Hettangian) times (Gierliński et al., 2001,
Figure 6. Characteristic tetrapod fossils from the Late Triassic of Poland: (A) Metoposaurus diagnosticus krasiejovensis Sulej, 2002, skull,
Krasiejów; (B) part of mandible of Cyclotosaurus sp., Lisowice; (C) maxilla of Polonosuchus silesiacus (Sulej, 2005), Krasiejów (Silesia);
(D) femur, tibia, and fi bula of the new species of large dicynodont, Lisowice (Silesia); (E) cf. Grallator isp., tridactyl dinosaur footprint, Liso-
wice (Silesia); (F) Chirotheriidae indet., pentadactyl pseudosuchian footprint, Woźniki (Silesia); (G) cf. Pseudotetrasauropus isp., tetradactyl
pseudosuchian footprint, Skarszyny (Holy Cross Mountains). Scale bars: A and B = 10 cm; C, E–G = 5 cm; D = geological hammer 32 cm long.
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276 Pieńkowski et al.
Figure 7. Characteristic tetrapod fossils from the Early Jurassic of Poland: (A) cf. Megalosauripus isp., footprint of large theropod dinosaur,
Sołtyków; (B) Parabrontopodus isp., manus and pes imprints of the early sauropod, Sołtyków; (C) Moyenisauropus cf. natator Ellenberger,
1974, footprint of the early ornithischian dinosaur, Gromadzice; (D) Anomoepus isp., footprint of the early ornithischian dinosaur, Sołtyków;
(E) Kayentapus isp., footprint of the large theropod dinosaur, Sołtyków; (F) Eubrontes isp., footprint of the large theropod dinosaur, Gromadzice;
(G) dinosaur bones, Sołtyków. Scale bars: A, B, D, and G = 5 cm; C, E, and F = 10 cm.
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Climatic reversals related to volcanism caused the end-Triassic biotic crisis—Evidence 277
2004). The largest footprints were described as cf. Megalo-
sauripus isp. (50–65 cm long). Another large theropod foot-
prints (30–45 cm) identifi ed at Sołtyków resemble Kayentapus
-like ichnites (Kayentapus soltykovensis) and classic ichnotaxa
of the Newark Supergroup, i.e., Eubrontes giganteus (Olsen et
al., 2002). Of note are also the fi nds of nondinosaur footprints:
Ameghinichnus isp., left by a mammal-like animal, and small
tetra- or pentadactyl footprints left by small reptiles (pterosaurs
and small diapsids) and basal crocodylomorphs.
CLIMATIC AND ENVIRONMENTAL CHANGES
VERSUS TERRESTRIAL EXTINCTION AND
RECOVERY—DISCUSSION
Climatic Changes and Palynofl oral Crisis
Results presented herein allow characterization of climatic
and other environmental changes at the end of the Triassic and
beginning of the Jurassic and link them with the end-Triassic
terrestrial extinction and ensuing Early Jurassic recovery. The
most continuous (though not completely recovered) profi le from
Kamień Pomorski shows conspicuous fl uctuations in clay min-
eral composition, carbon and osmium isotopes, PAHs, and paly-
nomorph frequency and preservation (Fig. 2; Table 1).
Our climatic interpretation is mostly based on clay mineral-
ogy and to a lesser extent on color of sediments and fl oral remains
(Figs. 2, 3, 4, and 5; Table 1). Much of clay deposits in sedimen-
tary basins represent a fi nal product of continental weathering
processes, and they may reveal climatic fl uctuations on con-
tinents, if diagenetic transformations were not very signifi cant
(e.g., Singer, 1984; Chamley, 1989; Ruffell et al., 2002; Ahlberg
et al., 2003; Deconinck et al., 2003; Raucsik and Varga, 2008;
Hesselbo et al., 2009; Dera et al., 2009; Brański, 2009, 2010,
2012). In particular, variations in the detrital kaolinite content of
the clay fraction are considered as reliable proxy for humidity.
Abundance of major elements is intimately related to min-
eralogy of the mudrocks. A distinct domination of illite (with
chlorite) in the Zbąszynek Beds and comparatively low values
of chemical indices (Fig. 3) are clearly related to the dominance
of physical weathering due to the (semi-) arid climate in Norian
time. The abundance of smectite in part of the Wielichowo Beds
(Figs. 2 and 3) indicates some increase in precipitation and a
distinct seasonality in the early-middle Rhaetian compared with
the Norian. The crucial change in the clay mineral assemblage is
observed between the Wielichowo Beds and Zagaje Formation
(Fig. 2, event 1 hh; Fig. 3), which refl ects the climate humidifi ca-
tion during Rhaetian time, with a shift from smectite- to kaolinite-
dominated mudrocks. Moreover, the clay mineral change was
associated with the earlier Rhaetian “precursor” δ13C excursion
of Ruhl and Kürschner (2011), which, according to Lindström et
al. (2012), commenced the series of environmental perturbations.
In the upper part of the Wielichowo Beds and in the Zagaje
Formation, the kaolinite-illite association prevails and signifi es
the predominance of warm climate with high year-round rain-
fall, even though initially some seasonality was still possible,
as indicated by the appearance of smectite in the Niekłań sec-
tion (Fig. 3) but not in Kamień Pomorski (Fig. 2). The values
of weathering indices are similar to modern residual clays or
muds from rivers draining strongly weathered tropical areas like
central Uganda, where the CIA values from 87 to 96 have been
recorded (Nyakairu and Koeberl, 2001). In general, progressive
weathering caused the loss of Ca, Na, and Mg and fi nally, K,
which refl ect the decrease in both nonclay silicate minerals (pla-
gioclases, K-feldspars) and compositionally immature clay min-
erals such as smectite (cf. Weaver, 1989).
A signifi cant admixture of berthierine and serpentine in
the Kamień section is probably linked to the co-occurrence of
weathered mafi c rocks in the source area (Kaszuby Land of the
Baltic Shield; Fig. 1). Importantly, some beds with very high
kaolinite/illite ratio and highly elevated values of chemical indi-
ces were observed in the sections studied (Figs. 2 and 3). The
sharp increase in kaolinite content records the abrupt change in
weathering regime. These layers, particularly rich in kaolinite,
with usually very high CIA value, were developed as a result of
extreme chemical weathering in the aftermath of rapid warming
and abundant rainfall. Strong leaching has almost removed the
mobile alkali and alkaline earth elements while immobile ele-
ments such as Al and Ti are enriched. As noted already, the fi rst
such greenhouse event was recorded in the earlier Rhaetian Wieli-
chowo Beds. The last distinct kaolinite episode was detected in
the lowermost Hettangian. High amounts of kaolinite are also
reported from the Rhaetian-Hettangian sections in southern Swe-
den (Ahlberg et al., 2002, 2003), from the Tatra Mountains in Slo-
vakia (Michalik et al., 2010), and from topmost Triassic Triletes
beds in the German Basin (van de Schootbrugge et al., 2009).
Recently, distinct kaolinite enrichment was also documented
from the topmost Triassic Kossen Formation (Eiberg Basin) in
Austria (Pálfy and Zajzon, 2012; Zajzon et al., 2012). Regardless
of this, the periods of less intense chemical weathering during
cooler and less humid conditions were recorded in some layers
characterized by lesser kaolinite content and lower CIA values.
Rapid and episodic fl uctuations in the composition of clay
minerals and in the values of weathering indices point to the
sequence of frequent and marked climatic reversals, named as
1 hh, 2 hh, 3 cd, 4 hh, 5 cd, 6 h, 7 cd, and 8 hh events (Fig. 2;
Table 1). These changes above the “initial” CIE become con-
spicuously more frequent as the Triassic-Jurassic boundary is
approached, assuming a relatively constant average sedimenta-
tion rate (Fig. 2).
The data provided by the palynomorphs from Triassic-
Jurassic transitional beds in Poland show conspicuous changes in
composition and character of plant biodiversity (see Pieńkowski
et al., 2012). The extinction processes and replacement in the
latest Triassic fl oras in Poland probably began along with an
increase in humidity in the middle to late Rhaetian time, as it
is associated with a marked clay mineral and CIA shift (Fig. 2;
Table 1; depth ~700 m, the 1 hh climate event). This is the shift
that likely correlates (at least partly) with a series of negative
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278 Pieńkowski et al.
δ13Corg excursions (so-called “precursor” negative δ13Corg excur-
sion [Ruhl and Kürschner, 2011], or Neg-1 excursion [Lindström
et al., 2012]), also observed in Kamień Pomorski δ13Corg excur-
sions. This somewhat neglected CIE event is also associated with
spore peak and bivalve extinction, as well as foraminiferal and
marine plankton change (Deenen et al., 2010; Ruhl et al., 2010;
Lindström et al., 2012). Interestingly, this CIE seems to be unre-
lated to any known magmatic/volcanic event, as it predates the
oldest known volcanic events of the Central Atlantic magmatic
province (Deenen et al., 2010; Lindström et al., 2012; Blackburn
et al., 2013).
Increased rainfall did not result in an increase of miospore
diversity and abundance, although dominance of spores at 700 m
contrasts with the lower, pollen grain–dominated sample (703 m),
which seems to support humidifi cation. The scarcity of mio-
spores between 682.5 and 700 m in Kamień Pomorski (Fig. 2)
is most probably caused by a low preservational potential, given
the lithology, which is unfavorable to organic matter preservation
(although a fl oral crisis might have also been involved). Dark-
ened miospores appear at 686 m, coincident within the “initial”
CIE and osmium isotope as well as with PAH disturbances. This
coincidence of indices points to marked environmental distur-
bances, which could be a refl ection of the disturbances that are
generally regarded to mark the onset of the end-Triassic extinc-
tion (Cirilli et al., 2009; Deenen et al., 2010; Ruhl et al., 2010;
Blackburn et al., 2013). Above 682.5 m, there is an increase in
abundance and diversity of miospores, and the changes of spore/
pollen grain proportions follow the changes in kaolinite/illite
ratio (Fig. 2). The super-abundances of spores just below the
Triassic-Jurassic boundary, linked with their marked darkening,
decreases in 187Os/186Os and 187Os/188Os, as well as signifi cant
drops in kaolinite and CIA (Fig. 2; Table 1), point to a marked
cooling, acid rains, and domination of “plant disaster taxa” asso-
ciated with this “spore peak” (Fig. 2; event 7 cd at 678.4 m). The
spore spike could be explained by a short but severe crisis for
land plants, generated by an eruption (or an impact), in which
all adult photosynthesis organs died off for lack of light, or in
a prolonged frost, or in acid rain, or other toxic compounds, or
all of them. Abundance of spores might also be partly related
to better preservational conditions (i.e., diminishing rate of bio-
genic decay of miospores, associated with acidifi cation of soil).
Perhaps ferns were the fi rst plants to recolonize the debris, and
higher plants returned later.
This event has been noticed in many parts of the world at the
Triassic-Jurassic boundary (Deenen et al., 2010; Whiteside et al.,
2010; Blackburn et al., 2013). In North America, high-diversity
pollen assemblages composed of monosulcates and monosac-
cates give way to lower-diversity assemblages dominated by
Classopollis, a pollen type normally associated with hot and/or
arid climate conditions, and palynofl oral diversity loss is esti-
mated at ~60% (Fowell and Olsen, 1993). It should be noted,
that Cirilli et al. (2009) cast some doubts on the existence of both
widespread palynological turnover and a “fern peak” in North
America (particularly in the Newark Basin), based on palyno-
logical results from the Fundy Basin (Nova Scotia, Canada); they
attributed abundance of spores to rather diachronous and short-
term ecological perturbations that were regional in scope (see
also Lucas and Tanner, 2007b). However, it should be noted that
Cirilli et al. (2009) were dealing with entirely Triassic material.
Some current studies from Europe (Kürschner and Herngreen,
2010; Cirilli, 2010) show that palynofl oral composition between
the Late Triassic and Hettangian was relatively steady and with-
out abrupt changes and consequently claim that the end-Triassic
biotic crisis appears to have little affected palynofl oral species
diversity, at least in Europe. Also estimates of diversity loss
based on macrofossils are typically much higher than estimates
of diversity loss based on miospores (Mander et al., 2010).
As the Triassic-Jurassic palynofl oral turnover in Poland is
more conspicuous than changes observed in other regions in
Europe, it may also be interpreted that the extinction rate was
related to the original paleolatitudinal position, implying that
the associated plant migration could also have been involved in
the Triassic-Jurassic palynofl oral turnover (Pieńkowski et al.,
2012). In that respect, is also noticeable that that the Rhaetian
palynofl oral assemblage from Pomerania (Pieńkowski et al.,
2012) shows differences compared to Lisowice (Staneczko,
2007; Dzik et al., 2008a, 2008b; Świło et al., 2013). The fl o-
ral crisis observed in Poland was relatively short, and the next
humid event sparked the re-composition of the fl ora, as indi-
cated by the increased proportion of hygrophilous plants (and
spores) in the earliest Hettangian ecosystems. Another “spore
peak” (Fig. 2, depth 641.8 m), much more diversifi ed in terms of
miospore taxa and containing many gymnosperm pollen grains,
could be related to a similar event (i.e., volcanic eruption) or to
localized change of environment associated with very favorable
hydrologic and preservational conditions.
Osmium Isotopic System, Iridium Content, and Organic
Geochemistry—Impact versus Volcanic Scenario
The 187Os/186Os ratio and very low iridium content lend no sup-
port for a role for asteroid impact at the Triassic-Jurassic boundary
(Pieńkowski et al., 2012). Furthermore, the recently redated 201 ±
2 Ma impact structure in Rochechouart in France (Schmieder et al.,
2010) seems to be too small (some tens of kilometers) to have caused
global extinction, despite its original size being still a matter of debate
(Smith, 2011). A hypothetical comet impact or “comet shower,”
leaving behind no geochemical traces, is still a theoretically pos-
sible explanation, but it seems that volcanic volatiles from the Cen-
tral Atlantic magmatic province continental fl ood basalts remain the
most plausible explanation for end-Triassic continental mass extinc-
tion. Noteworthy factors are increased abundances of 192Os, observed
twice in the Kamień Pomorski profi le (Fig. 2). Content of 192Os, the
most common unradiogenic osmium isotope, is thought to be mainly
derived from igneous activity (Cohen and Coe, 2002, 2007; Kuroda
et al., 2010). Also, the negative shift in 187Os/188Os values suggests
input of unradiogenic Os of mantle (or extraterrestrial) origin or a
reduction of continentally derived Os or both (Cohen and Coe, 2002;
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Climatic reversals related to volcanism caused the end-Triassic biotic crisis—Evidence 279
Kuroda et al., 2010). Observed changes in osmium isotopes in the
Kamień Pomorski profi le (the higher value of 187Os/188Os at the
bottom) resemble, to some extent, trends noted by Cohen and Coe
(2002). Higher in the profi le, values of this ratio are stable, followed
by the marked drop concomitant with the upper “subpeak” of the
“initial” CIE, and lower values continue until the top of Triassic strata
(Fig. 2). Because these are terrestrial deposits, the higher 187Os/188Os
values can be tentatively linked to volcanic fallout coming from the
Central Atlantic magmatic province (Fig. 1). Such fallout could be
associated with other volcanic impacts, as acid rain and subvolcanic
PAH releases (triggered when hot magma interacts with organic-rich
rocks), which could lead both to observed darkening of spores in sev-
eral horizons (Fig. 2) and defoliation, which in turn might have led to
intensifi cation of forest fi res.
The PAHs were produced also by local wildfi res
(Marynowski and Simoneit, 2009; Pieńkowski and Waksmun-
dzka, 2009), which should also be associated with the presence
of charcoal in palynomacerals. In some samples, this is the case
(i.e., Fig. 2, 687 m, 3 cd event; many Hettangian samples in
Kamień Pomorski and Sołtyków), but elsewhere it is not. Lack of
charcoal linked with elevated PAHs would rather support a sub-
volcanic origin of these compounds (Pieńkowski et al., 2012; see
also van Schootbrugge et al., 2009). Moreover, charcoal, even if
present, must be taken with some caution as an “in situ” indicator
of wildfi re frequency because it can be widely redeposited due to
its resistance to biogenic degradation and buoyancy (Pieńkowski
and Waksmundzka, 2009).
Anyhow, release of toxic pollutants such as SO2, sulfate aero-
sols, and PAHs certainly led to defoliation, which increased for-
est fl ammability and resulting fi re activity (see also Pieńkowski
et al., 2012), similar to the climate-driven shift from broad-leaved
to narrow-leaved taxa at the Triassic-Jurassic boundary (McEl-
wain et al., 2009; Belcher et al., 2010).
Tetrapods across the Triassic-Jurassic Boundary in Poland
The Lisowice and Sołtyków localities, linked with other
data from older or younger sites (Figs. 6, 7, 8, and 9), provide
paleontological data valuable for determining biodiversity and
evolutionary changes of the terrestrial tetrapod fauna across the
Mammaliaformes
Figure 8. Distribution of terrestrial tetrapods and tetrapod assemblages (based on both body and trace fossils record)
across the Late Triassic–Early Jurassic time interval in Poland. Although extinction period cannot be precisely indicated
due to the lack of continuous outcrops spanning the Triassic-Jurassic boundary, the dramatic faunal turnover following
the extinction period between the middle and end Rhaetian is visible, as is rapid tetrapod recovery in earliest Jurassic time
(Sołtyków-Gromadzice assemblage). Figure is based on data from Gierliński et al. (2004), Dzik and Sulej (2007), Sulej
(2009), Sulej et al. (2011, 2012), Dzik et al. (2008a, 2008b), Niedźwiedzki (2006, 2011a, 2011b, 2014), Niedźwiedzki
et al. (2011, 2012, 2014).
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280 Pieńkowski et al.
Figure 9. Reconstructed vertebrate fauna and simplifi ed food web for the Lisowice community (Late Triassic: a—
hybodont sharks; b—coelacanth fi sh; c—dipnoan fi sh; d—palaeonisciform fi shes; e—capitosaur; f—plagiosaur;
g—choristodere-like archosauromorph; h—rhynchosaur; i—poposaur; j—dicynodont; k—small predatory dinosaur;
l—sphenodontid; m—small dinosauromorph; n—large predatory dinosaur; o—pterosaur; p—mammal or mamma-
liaform) and Sołtyków community (Early Jurassic: a—large theropod dinosaur; b—medium theropod dinosaur; c—
sauropod dinosaur; d—small theropod dinosaur; e—medium ornithischian dinosaur; f—small ornithischian dinosaur;
g—small crocodylomorph; h—palaeonisciform fi shes; i—mammal or mammaliaform). Solid lines with arrows show
feeding pathways.
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Climatic reversals related to volcanism caused the end-Triassic biotic crisis—Evidence 281
Triassic-Jurassic boundary in Poland (Fig. 8). Reconstructions of
assemblages from both localities reveal a rather complex trophic
array, which today is characterized as stable and not disturbed
ecosystems (Fig. 9).
The two well-characterized assemblages of tetrapod fossils
(Lisowice and Sołtyków assemblages) from the southern Poland
bracket the Triassic-Jurassic boundary. Those two assemblages
encompass body fossils and footprints from a variety of litho-
facies that represent similar depositional systems. This makes it
easiest to simply compare each assemblage to the other because
the differences between the assemblages in large part arose from
taphonomic and not paleoenvironmental factors. In addition, the
similar ages and paleogeographical positions of both sites are a
good indicator of temporal succession and evolution of tetrapods
across Triassic-Jurassic boundary in Poland.
In the Lisowice site, both body and trace fossil records of tet-
rapods are rich, and both are comparable from a taxonomic point
of view. The Sołtyków site shows only an ichnological record
(bones are extremely rare), but it is very diverse. Despite these
differences, clear events in tetrapod evolution in the end of Trias-
sic (or across Triassic-Jurassic boundary) in Poland are visible.
The fi rst event is the extinction of the pseudosuchians in the end
of Triassic. This extinction, usually referred to as the extinction
of “thecodonts” (or pseudosuchians), was identifi ed as the prin-
cipal tetrapod extinction at the end of the Triassic (see Colbert,
1958; Olsen et al., 2002; Lucas and Tanner, 2007a; Brusatte et
al., 2008; Langer et al., 2010; Sues and Fraser, 2010). Pseudo-
suchian body fossils and footprints (“Rauisuchia”) are pres-
ent in the Lisowice sites, and also older Triassic sites in Poland
(Fig. 6C), but are absent in Sołtyków. We take this to indicate
pseudosuchian extinction somewhere between the Lisowice and
Sołtyków assemblages, thus very close to the Triassic-Jurassic
boundary. The second event is the extinction of typical Triassic
fauna elements such as Dicynodontia, Capitosauroidea, Plagio-
sauroidea, and Rhynchosauria (Fig. 8). All these tetrapods dis-
appear from the fossil record between the middle Rhaetian and
earliest Hettangian. The third trend in tetrapod evolution across
the Triassic-Jurassic boundary in Poland worth additional com-
ment is the dramatic latest Triassic change in dinosaur diversity.
The tetrapod assemblage from the earliest Hettangian of Poland
shows that a sudden increase in numbers, diversity, and body
sizes of dinosaurs took place during the latest Triassic (probably
late Rhaetian) or just after the Triassic-Jurassic boundary. Thus,
earliest Hettangian tetrapod assemblage of Poland is dominated
by dinosaur tracks, with relatively rare tracks of other small tet-
rapods (basal crocodylomorphs, early mammals, small reptiles,
and pterosaurs). The large-sized theropod dinosaurs (~7–8 m in
length) might have been the top predators hunting the sauropodo-
morphs (probably early sauropods or last “prosauropods”; Fig.
9). The tetrapod ichnofauna from the Zagaje Formation supports
the presence of a locally distributed theropod-sauropodomorph
assemblage in the earliest Jurassic, preceding characteristic asso-
ciations with numerous tracks of early ornithischia occurring in
the late Hettangian and Sinemurian.
Numerous workers (e.g., Benton, 1986; Hunt, 1991; Olsen
et al., 2002; Brusatte et al., 2008, 2010) have drawn attention
to a relatively sudden increase in dinosaur abundance, diversity,
and body size during the latest Triassic–earliest Jurassic, and this
event is geographically widespread and not lithofacies correlated,
so probably it is a record of evolutionary events connected with
terminal Triassic extinction of therapsid-pseudosuchian faunas
on the land. Additionally, according to paleontological records
from Poland (and also from Italy, United States, Argentina, South
Africa), a few evolutionary lines of dinosaurs were already wide-
spread in the latest Triassic and earliest Jurassic. The relatively
high diversifi ed dinosaur ichnofauna from Sołtyków indicates to
us a rapid recovery (or radiation) and refi ll of ecological niches
after the end-Triassic extinction and within ~0.5–1.5 m.y. during
the latest Rhaetian and earliest Hettangian.
The fi rst Central Atlantic magmatic province pulse (latest
Rhaetian in age; see Blackburn et al., 2013) probably coincided
with the major extinction event of plants and other organisms in
the continental realm. As recognized in the Newark and Fundy
Basins (North America), the extinction of pseudosuchians and
the rise of dinosaur-dominated fauna (Olsen et al., 2002) were
also linked to Central Atlantic magmatic province volcanism
(Blackburn et al., 2013). The record of extinction of the Tri-
assic elements in the tetrapod fauna across Triassic-Jurassic
boundary in Poland is probably connected with initial Central
Atlantic magmatic province volcanism (i.e., according to the
existing knowledge, also with the “initial” CIE). Some of the
later (post–“initial” CIE, pre–Triassic-Jurassic boundary; 5 cd
and 7 cd events; Fig. 2) releases of toxic pollutants (SO2, sul-
fate aerosols, and other gases) were most likely the fi nal blow
to the end-Triassic pseudosuchian-therapsid– dominated eco-
systems. We agree with the conclusion of Langer et al. (2010)
that the Rhaetian stage was probably associated also with a
crisis in dinosaurs.
The end-Triassic extinction event also affected many groups
of large aquatic tetrapods and was perhaps an ecological event
linked with changes in water chemistry of lakes and ponds (poi-
soning of surface water). Successive Central Atlantic magmatic
province eruptions contributed both CO2 (and CH4 and PAHs) and
sulfur injections (of note is the presence of highly pyritic sand-
stone just below the Triassic-Jurassic boundary in the Kamień
Pomorski profi le, different in its high intensity from other forms
of pyritic mineralization in the profi les studied; see Fig. 2) with
cooling lasting years or decades. Relatively longer periods of
greenhouse events (2 hh, 4 hh, 8 hh; Fig. 2) should have allowed
some migration of “vulnerable” taxa (temnospondyls, pseudosu-
chians, therapsids) to the lower latitudes, but abrupt cooling led
to less time for those animals to survive (either by migration or
accommodation). In contrast, dinosaurs, early mammals, small
crocodylomorphs, pterosaurs, and also their insulated relatives
could better withstand the cold challenge. Although still specula-
tive (because of insuffi cient data), this scenario is consistent with
the post-Triassic recovery in which dinosaurs took over the world
(Figs. 8 and 9).
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282 Pieńkowski et al.
INTEGRATED STRATIGRAPHIC FRAMEWORK
The existing chronostratigraphic framework based on zircon
U-Pb geochronologic and astronomical constraints from marine
and continental sections (Deenen et al., 2010; Ruhl et al., 2010;
Blackburn et al., 2013) can be matched to our profi les based on
carbon isotope stratigraphy and palynofl ora. All the aforemen-
tioned authors correlate the end-Triassic extinction with the onset
of Central Atlantic magmatic province volcanism and the marked
“initial” CIE (see also Ruhl et al., 2011). The following recovery
at the Triassic-Jurassic boundary occurred just ~120 k.y. after-
wards, spanning six precession cycles (Ruhl et al., 2010). We
inferred four climatic events (reversals) in this time interval, but
some part of sedimentary record may be missing at the sequence
boundary, close to the Triassic-Jurassic boundary (Fig. 2).
Interestingly, Bonis et al. (2010) recognized four pronounced
spore peaks in the St. Audrie’s Bay section of the UK, in the
end-Rhaetian Lilstock Formation. They attributed these peaks
to precession-induced increased runoff, which would require the
duration of the initial CIE to have been at least 20,000 yr, which is
just on the maximum limit indicated by Ruhl et al. (2011). Bonis
et al. (2010) also linked spore peaks to climate change rather than
to any catastrophic events. Linking our climatic steps inferred
from clay mineralogy with astronomical cycles (precession
cycles) is not impossible. However, our study and other reports
(i.e., Korte and Kozur, 2011; Ruhl et al., 2011) show that the ini-
tial CIE is bipartite, with two negative subpeaks. If the duration
of the whole initial CIE was between 10,000 and 20,000 yr, these
changes (refl ected also in climate; Fig. 2; Table 1) are of too high
frequency to be linked with astronomical forcing. Instead, they
could be associated with episodes of Central Atlantic magmatic
province eruption, which is further supported by differences of
certain geochemical and palynological properties (Os isotope sys-
tem disturbances as well as darkening of miospores and, circum-
stantially, PAH content) of these two peaks (Fig. 2; Table 1). It is
possible that those independent mechanisms (astronomical forc-
ing and volcanic activity) could amplify or oppositely, alleviate
adverse effects, but Central Atlantic magmatic province–related
processes seem to be the most probable explanation for the biotic
crisis at the end of the Triassic. A series of periodic atmospheric
loadings by CO2, CH4, or alternatively by SO2, sulfate aerosols,
and toxic compounds, is inferred to have caused this series of
rapid climatic reversals and resulting biota crisis.
Our study confi rms the results of Lindström et al. (2012),
which indicated that the climatic and biotic changes commenced
earlier than the initial CIE, somewhere at the level of the “precur-
sor” CIE (Ruhl and Kürschner, 2011), judging from the astrochro-
nological scale of Ruhl et al. (2010), some 100–200 k.y. earlier
than the initial CIE and hitherto known onset of Central Atlantic
magmatic province volcanism (Blackburn et al., 2013). The direct
cause of this CIE is still unclear; perhaps an earlier phase of Central
Atlantic magmatic province volcanism is still to be discovered.
The only other event that seems to be approximately cor-
relatable with this shift is a marked erosional surface (698 m in
Kamień Pomorski; Fig. 2), identifi ed with the sequence bound-
ary, which is probably concomitant with the Rhaetian lowstand,
inferred to be one of the lowest in Phanerozoic (Hallam, 2001).
This sequence boundary can be identifi ed with emergence sur-
faces within the Lilstock Formation, occurring both in the St.
Audrie’s (Hesselbo et al., 2002) and Larne (Simms and Jeram,
2006) sections and a lowstand (correlative with a sequence
boundary) at Kuhjoch, located at the top of the Koessen Forma-
tion (Hillebrandt et al., 2007) and in Csövár (Pálfy et al., 2001,
2007). However, the lowstand and exposure of vast areas would
rather lead to steepened latitudinal temperature gradients and
increased environmental extremes due to continentality, rather
than to marked humidifi cation.
Our conclusion is that the extinction period was more pro-
longed than the 20,000 yr duration of the initial CIE. Slight paly-
nofl oral change commenced at the initial δ13C excursion, corre-
lated with the main onset of Central Atlantic magmatic province
volcanism, but the following positive carbon isotope excursion,
accompanied by a rapid drop in kaolinite content and intermit-
tent reappearance of a thin layer of red beds (5 cd event; Fig. 2;
Table 1; possibly also red-greenish strata in Lisowice [Fig. 5],
although existing stratigraphical resolution in Lisowice does not
allow precise correlation with the Kamień Pomorski profi le), was
of equal, if not greater signifi cance, causing the deepening crisis
for Triassic fl ora and tetrapod fauna. Increased aridity might have
been unfavorable, particularly for amphibians (i.e., Plagiosauri-
dae). The next three steps, particularly 7 cd (Fig. 2; Table 1) at
the Triassic-Jurassic boundary, associated with acid rains, “spore
peaks,” and darkened miospores, probably dealt the “fi nal blow”
to the otherwise weakened Triassic ecosystem, which is refl ected
by the disappearance of most of the Triassic palynomorphs and a
number of characteristic tetrapod taxa. Obtained values of initial
187Os/186Os between 2.905 and 4.873 and very low iridium con-
tent (Fig. 2) lend no support for a role of an asteroid impact at
the Triassic-Jurassic boundary event. However, Central Atlantic
magmatic province–related volcanic volatiles causing climatic
disturbances and infl uencing the ecosystem in many ways (Self
et al., 2006) seem to be a much better substantiated cause for the
end-Triassic extinction.
ACKNOWLEDGMENTS
We are grateful to Stephen Hesselbo and Simonetta Cirilli for
their constructive reviews. Gerta Keller, Andrew Kerr, and Gina
Harlow are thanked for their editorial assistance. This paper
is a part of a project fi nanced by the Polish National Science
Centre, granted on the basis of decision no. DEC-2012/06/M/
ST10/00478. Niedźwiedzki is currently funded by a Wallenberg
scholarship grant awarded to P.E. Ahlberg (Uppsala University).
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