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Microbially induced wrinkle structures in Middle Devonian siliciclastics from

the Prague Basin, Czech Republic

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DOI: 10.1111/let.12280

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Microbially induced wrinkle structures in Middle Devonian
siliciclastics from the Prague Basin, Czech Republic
STANISLAVA VODRA 

ZKOV
, RADEK VODRA
A 

ZKA, AXEL MUNNECKE, JURAJ FRANC
U,

AND PETRA TONAROVA
KHALDOUN AL-BASSAM, PATRICIE HALODOVA

Vodr
azkov
a, S., Vodr
azka, R., Munnecke, A., Franc

u, J., Al-Bassam, K., Halodov
a, P.
& Tonarov
a, P. 2018: Microbially induced wrinkle structures in Middle Devonian
siliciclastics from the Prague Basin, Czech Republic. Lethaia, https://doi.org/10.1111/le
t.12280.

The occurrence of wrinkle structures in Middle Devonian deep-water siliciclastic

sequences of the Prague Basin (Robl
ın Member, Srbsko Formation, Givetian) is
reported and interpreted to be microbially induced. Current and/or gravitational
forces are considered the simplest explanation of the origin of these structures. Taking
into account the tectonic activity linked with ongoing Variscan orogeny, the wrinkle
structures could also be interpreted as soft sediment deformation structures originat-
ing due to exposure of mat-bearing sediment to seismic shocks. The distribution of n-
alkanes and isoprenoids suggests two types of prevailing biological sources, namely
phytoplankton, representing organic material from the water column, and benthic
bacteria. Judging from the sedimentary facies and lack of petrographic characteristics
suggestive of light competition, the microbial mats are interpreted to be formed by
non-autotrophic bacteria. □ Deep-water siliciclastics, Devonian, microbial mats,
Prague Basin, wrinkle structures.

Stanislava Vodr
az kov
a ✉ [stana.vodrazkova@seznam.cz], Radek Vodr
az ka [radek.vo

drazka@geology.cz], Khaldoun Al-Bassam [albassam703@gmail.com], Patricie Halodov
a
[patricie.halodova@geology.cz], and Petra Tonarov
a [petra.tonarova@geology.cz], Czech
Geological Survey, Kl
arov 3, 118 21 Prague 1, Czech Republic; Axel Munnecke [axel.mun-
necke@fau.de], GeoZentrum Nordbayern, FG Pal€aoumwelt, Friedrich-Alexander Univer-
sit€at Erlangen-N€urnberg, Loewenichstraße 28, Erlangen, Germany; Juraj Franc u
[juraj.francu@geology.cz], Czech Geological Survey, Leitnerova 22, 658 69 Brno, Czech
Republic; manuscript received on 11/12/2017; manuscript accepted on 12/04/2018.

Since the paper of Black (1933), who studied micro-
bial mats of Andros Island in the Bahamas, the sig-
nificant role microbes play in recent carbonate
environments has become increasingly apparent.
Considering the low preservation potential of similar
structures in siliciclastics due to the lack of early
cementation (Logan 1961), it is not surprising that
siliciclastic sediments have not received much atten-
tion in this respect. This has changed after the paper
of Davis (1968), who was the first to report ancient
microbial mats in siliciclastics (‘algal stromatolites
composed of quartz sandstone’), followed by numer-
ous examples both from recent tidal flats (e.g.
Gunatilaka 1975; Schwarz et al. 1975; Gerdes et al.
1985; Witkowski 1990; Noffke et al. 1996) and from
ancient environments. Most of the fossil microbial
mats (or structures induced by microbial activity)
reported from siliciclastics are Proterozoic (e.g.
Gehling 1999; Hagadorn & Bottjer 1999; Noffke
et al. 2002; Banerjee & Jeevankumar 2005) and Early
Palaeozoic in age (e.g. Noffke 2000; Brett et al. 2003;
Buatois & M
angano 2003; Bailey et al. 2006) – thus
from times with less effective benthic grazing and
low or no infaunal activity (e.g. Droser & Bottjer
1989). Only a few examples of post-Ordovician
microbial mats have been recorded from siliciclastic
settings (e.g. Pfl€
uger 1999; Dragantis & Noffke 2004;
Pruss et al. 2004; Calner 2005; Kremer 2006; Calner
& Eriksson 2012; Marriott et al. 2013) mostly having
in common the occurrence in stressed and restricted
environments (Mata & Bottjer 2009).
Most modern microbial mats have been reported
from shallow water settings, with cyanobacteria
being the main constituents of the biofilms (e.g.
Noffke 2010 and references therein). Similarly, as
implied from Table 1 in Davies et al. (2016), most
of the fossil microbial mats have been reported from
tidal and storm influenced settings, and only a few
examples of fossil microbial mats from deeper water
facies were recorded. The relatively recent discovery
of modern microbial mats formed by the bacterium
Thioploca inhabiting vast areas of the oxygen mini-
mum zone (OMZ) off the coasts of Chile and Peru
(Gallardo 1977) represented a breakthrough in
understanding of microbial mat-forming communi-
ties. More examples followed (e.g. Fossing et al.
1995; Buck & Barry 1998; Schulz et al. 1999; Gal-
lardo & Espinoza 2007) showing that biofilms

DOI 10.1111/let.12280 © 2018 Lethaia Foundation. Published by John Wiley & Sons Ltd
2 Vodr
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a et al. LETHAIA 10.1111/let.12280

Table 1. Molecular ratios of the n-alkanes and isoprenoids in the rock extract: saturated fraction. Positions of the samples Hlu-1, Hlu-2,
Hlu-4 and Hlu-5 are marked in the Figure 2A. Sample Hlu-3 comes from the Hlubocepy-railroad cut, 3.5 m above the base of the Robl
ın
Member. Explanations of the abbreviations and ratios: Pri/Phy – pristane/phytane ratio; Pr/n-C17 – pristane/n-C17 alkane; Ph/n-C18 –
phytane/n-C18 alkane; % Norpr – relative amount of norpristane = Norpr/(Norpr+Pr+Ph); %Pr – relative amount of pristane = Pr/
(Norpr+Pr+Ph); %Ph – relative amount of phytane = Ph/(Norpr+Pr+Ph); Sum (n-C17–20) % – relative amount of the saturated hydro-
carbons in the selected range; Sum (n-C25–28) % – relative amount of the saturated hydrocarbons in the selected range; Sum (n-CC31–34)
% – relative amount of the saturated hydrocarbons in the selected range.

Sample Pri/Phy Pr/n-C17 Phy/n-C18 % Norpr %Pr %Ph Sum (n-C17–20) % Sum (n-C25–28) % Sum (n-C31–34) %

Hlu-1 0.92 0.57 0.52 20.60 38.14 41.26 0.879 0.117 0.004
Hlu-2 0.99 0.59 0.61 21.30 39.14 39.56 0.859 0.137 0.004
Hlu-3 1.46 0.59 0.50 26.75 43.48 29.77 0.943 0.054 0.002
Hlu-4 1.11 0.61 0.55 21.55 41.28 37.17 0.925 0.058 0.017
Hlu-5 0.83 0.74 0.73 18.28 37.11 44.61 0.888 0.109 0.003

formed by sulphur-oxidizing bacteria are character-
istic components of sediments in the oxygen mini-
mum zones of modern oceans.
Unfortunately, there has been much confusion in
the literature with respect to terminology of the sedi-
mentary structures interpreted as microbially
induced. Various distinctive types of microbial mat
features are termed by Noffke et al. (1996) as Micro-
bially Induced Sedimentary Structures (MISS). The
most common forms of MISS are wrinkle structures
(Hagadorn & Bottjer 1997), palimpsest ripples (Sei-
lacher 1999), roll-up structures (Simonson & Carney
1999) and laminar structures (Noffke et al. 1996).
Wrinkle structures are among the best-documented
MISS (Noffke et al. 2002; Fernandez & Pazos 2013;
Zhong et al. 2013). Hagadorn & Bottjer (1997)
introduced the term ‘wrinkle structures’ as a collec-
tive term for structures described as wrinkle marks
and Kinneyia ripples. According to these authors,
although Kinneyia structures differ from wrinkle
marks by flat-topped crests, they both may represent
‘preservational end members of similar structures’
(Hagadorn & Bottjer 1997; p. 1047). As stressed by
Eriksson et al. (2007) and Porada & Bouougri
(2007), both terms – wrinkle structures and Kin-
neyia – are often treated as synonyms in the
literature.
Origin of wrinkle structures
There has been much speculation about wrinkle
structure formation, involving both biotic and abi-
otic processes. Many hypotheses have been put for-
ward to explain the biotic formation of wrinkle
structures, or ‘Kinneyia’, with nearly all proposed
methods requiring the presence of cohesive micro-
bial mats at the sediment’s surface (Hagadorn &
Bottjer 1997; Noffke et al. 2003; Parizot et al. 2005;
Shi et al. 2008; Xing et al. 2011; Zhong et al. 2013).
Noffke (2000) and Bottjer & Hagadorn (2007)
regarded wrinkle structures as fossilized microbial
mats. In contrast, Noffke et al. (2002) interpreted
wrinkle structures as a result of loading pressure
originated by deposition of sediment on fluid-rich
microbial mat-bearing sediment. In this case, the
casts and moulds are seen as water-escape structures.
Pfl€
uger (1999) considered some wrinkles in
Silurian turbidites to result from gas, produced by
decay of organic mats, trapped underneath the mat.
Mariotti et al. (2014) performed wave-tank experi-
ments and interpreted the pits and ridges to be
formed by physical interaction of mat aggregates,
sediment grains and waves, implying formation of
the wrinkle structure at the sediment–water inter-
face. Thomas et al. (2013) and Herminghaus et al.
(2016) ascribed the formation of wrinkle structures
to flaw-induced hydrodynamic instability. In their
model, the hydrodynamic instability caused by the
presence of microbial film gave rise to ripple pattern
on the surface of the mat, which subsequently led to
sediment deposition in the ripple depressions and
preservation even after the original mat was disinte-
grated. Hagadorn & Bottjer (1997) and Bouougri &
Porada (2002) regarded wrinkle structures as fos-
silized mats reflecting original mat morphology
influenced by physical action of currents, waves and
wind. Similarly, Bose & Chafetz (2009), who
observed modern tidal siliciclastic sediments with
microbial mats, ascribe formation of wrinkles to mat
deformation due to current shear. Brett et al. (2003)
observed wrinkle structures (Kinneyia) in associa-
tion with seismites and suggested that such struc-
tures could be seismically induced.
Although all of the above-mentioned models dif-
fer, they do have one necessary prerequisite in com-
mon – the presence of microbial mats. Some
authors, however, consider purely abiotic processes
as responsible for the formation of wrinkled sedi-
ment surfaces. For example, ‘wrinkle marks’
LETHAIA 10.1111/let.12280
originating solely by abiotic loading processes were
described by Allen (1985). Wind action on the sur-
faces of wet and fine-grained sediment was supposed
as a main trigger of ‘runzel-marken’ formation by,
for example Reineck (1969) and Teichert (1970).
Dzulynski & Simpson (1966) ascribed formation of
wrinkles to the shearing effect of turbidite currents.
As implied from this brief outline, undisputed
evidence of the origin of wrinkle structures is still
missing, which is also due to the fact that exam-
ples of modern shallow water wrinkle structures
are only rarely described (Hagadorn & Bottjer
1997, 1999; Bose & Chafetz 2009). Modern deep-
water wrinkle structures are unknown but can be
expected as modern deep-water microbial mats
composed of sulphur-oxidizing bacteria are due to
extracellular polymeric substances (EPS, Wingender
et al. 1999) capable of producing sedimentary
structures that are comparable to those produced
by cyanobacteria in shallow water settings (Flood
et al. 2014). Some types of sulphur-oxidizing bac-
teria were found able to trap and bind sediments
in much the same way as phototrophic cyanobac-
teria, giving rise to many of the same wrinkle
structures that are commonly attributed to
cyanobacteria. Sulphate-reducing bacteria have also
been found to construct significant microbial mats
(Noffke et al. 2006a,b).
Evidence of biogenic nature
Seeking evidence for the biogenic nature of wrinkle
structures, thus demonstrating that microbial com-
munities were involved in their formation, must be
the key task – however, what are the chances of
preservation of microbial mat signatures in the sedi-
mentary record? As stressed by Davies et al. (2016,
p. 241), ‘smoking guns for microbial involvement
are the exception rather than a rule’. Such rare direct
evidence was provided by Kremer (2006), who
reported cellular preservation of Silurian benthic
cyanobacteria, but these were preserved in cherts
and no surface structures were described. Several
microscale features of siliciclastic rocks were sug-
gested as criteria for recognizing former microbial
mats and thus could serve as indirect evidence of
biogenicity. Those criteria include the following: (1)
trapping and binding of grains in a more or less lam-
inar matrix and authigenic mineral precipitation
(e.g. Noffke et al. 1997; Gerdes et al. 2000; Elster
et al. 2016); (2) early diagenetic pyrite formation
(Schieber 2002); (3) wavy-crinkly laminae (Schieber
1986, 1989); (4) replacement of organic matter by
minerals (‘destructive biomineralization’ sensu Nof-
fke 2010); and (5) accumulation of translucent
Wrinkle structures in Middle Devonian siliciclastics 3
quartz grains within organic filamentous material
(Gerdes et al. 2000; Noffke 2009).
In this paper, we report the occurrence of wrinkle
structures, interpreted to be microbially induced,
from the Middle Devonian (Givetian) deep-water
siliciclastic sequences of the Robl
ın Member (Srbsko
Formation) from the Prague Basin, Czech Republic.
We suggest possible modes of origin of the struc-
tures and discuss environmental parameters that
facilitated formation and preservation of microbial
mats.
Geological setting
The Prague Basin is situated between the cities of
Prague and Pilsen (Fig. 1) and represents the cen-
tral part of the Tepl
a-Barrandian unit (TBU) of the
Bohemian Massif, which is widely accepted as a
Gondwana-derived crustal block characterized by
Neoproterozoic (Cadomian) basement uncon-
formably overlain by Lower Palaeozoic sequences
(see e.g. Chlup
ac et al. 1998; Kachl
ık 1999; Franke
2000). After opening of the Rheic Ocean in the
Early Ordovician (e.g. Linneman et al. 2007; and
references therein), the TBU drifted towards lower
latitudes and was incorporated into the Variscan
orogen during the Late Devonian-Early Carbonifer-
ous (e.g. Franke 2000). Evidence of tectonic insta-
bility caused by initial collision of TBU with
adjacent terranes can be observed in the Prague
Basin already in the Lower Devonian (lower
Emsian) at the base of the Zl
ıchov Formation (sedi-
mentation of intraformational breccias; Chlup
ac
1957; Chlup
ac & Kukal 1986) and especially in the
Middle Devonian (Givetian), when sedimentation
of the Robl
ın Member of the Srbsko Formation
took place (Schulmann et al. 2009). The influence
of Variscan tectonic movements on Robl
ın Member
development was stressed already by Chlup
ac
(1958, p. 183), followed by Kukal & J€ager (1988),
Strnad & Hladil (2001) and Schulmann et al.
(2009).
Robl
ın member of the Srbsko formation
(Middle Devonian, Givetian)
The succession of the Robl
ın Member, almost 260 m
thick (preserved thickness), is developed as an irreg-
ular rhythmic alternation of grey, grey-green and
brownish siltstones, clay-rich siltstones, and fine-
grained sandstones with characteristic features, such
as graded bedding, lamination, cross-bedding and
flute marks (Chlup
ac 1958, 1960; Kukal & J€ager
1988).
4 Vodr
az kov
a et al.
Fig. 1. Schematic map showing the geographical distribution of the Srbsko Formation in the Prague Basin, with the position of the sam-
pling site marked.
The unit was interpreted as ‘very distal turbidites
deposited in a basin of moderate depth with gentle
paleoslopes’ (Kukal & J€ager 1988, p. 79). The latter
authors also suggested lower salinities of the water
masses due to river water inflow. Marine fauna was
recorded only at the very base and upper part of the
unit by Chlup
ac (1958, 1960).
Clay-rich beds of the Robl
ın Member are rich in
remnants of the land flora (especially in the lower
parts, e.g. Obrhel 1961; Jurina et al. 2009). Mikul
as
(1995) and Mikul
as & Pek (1996), who studied ich-
nofossils from the uppermost preserved parts of the
Robl
ın Member, documented rather rare and a low-
diversity mixture of elements belonging to Arenicol-
ites, Cruziana and Nereites ichnofacies and compared
it to assemblages known from flysch deposits.
Material and methods
Wrinkle structures were observed on several bedding
planes of sandy siltstones of the Robl
ın Member,
which crop out sporadically in a gorge in Praha-
LETHAIA 10.1111/let.12280
Hlubocepy, along the northeast slope directed
towards Daleje-Prokop valley (50.0387672N,
14.3826744E). The entire gorge exposes several tens
of metres thick succession of the Robl
ın Member.
For this study, we exposed and cleaned 3.5 m of the
sequence (Fig. 2A, C). Except for plant remains the
beds are barren, which prevents biostratigraphical
assignment. Based on local settings, we can only esti-
mate that the sampled interval represents approxi-
mately the middle part of the member. The cleaned
outcrop was documented for sedimentary structures
and sampled for wrinkle structures, thin sections,
palynomorph and biomarker analyses. It is notewor-
thy that this locality remained unnoticed until
recently. Previous studies focused on the classic out-
crop known as the Hlubocepy- railroad cut, which is
situated ca 1 km east of the outcrop described herein
(e.g. Chlup
ac 1960; Jurina et al. 2009).
In total, 21 thin sections with dimensions of
45 9 27, 50 9 50, 75 9 75 and 75 9 100 mm were
studied using the Carl Zeiss Stemi 2000C and Nikon
Eclipse E600 microscopes. Photographs were taken
with a Nikon DS-Fi1 camera mounted onto the
LETHAIA 10.1111/let.12280
latter. A scanning electron microscope FEG-SEM
Tescan Mira 3GMU fitted with EDS SDD X-Max
80 mm2 analyzer (Oxford Instruments) was used to
investigate thin sections in detail. The analytical con-
ditions used were 15 kV accelerating voltage, 1.5 nA
beam current and 15 mm working distance. BSE
detector was used for imaging. The surface of the
samples was covered by a 15 nm thick layer of amor-
phous carbon to avoid charging by the SEM.
For biomarker analysis, organic compounds were
extracted from pulverized and homogenized rocks
using 93:7 DCM: methanol for 20 min in a Dionex
accelerated solvent extractor at 80 °C and 15 MPa.
The extract was evaporated under nitrogen flow in a
Zymark TurboVap instrument, asphaltenes were
precipitated, and saturated, aromatic and polar frac-
tions were separated by column chromatography.
The saturated and aromatic hydrocarbons were
analysed by gas chromatograph-mass spectrometer
Agilent Technologies 7890 and 5973N in full-scan
and SIM modes.
For palynomorph analysis, three pilot samples
were treated with 10% HF. The insoluble residue
was sieved into fractions (20–50, 50–80, >80 lm).
The 20–50 lm fraction was used to prepare perma-
nent micro-palaeontological slides. As all samples
were barren, they will not be discussed herein.
Results
Field observations
The section exposed exhibits an alternation of thin-
to medium-bedded siltstones, sandy siltstones and
fine-grained sandstones (Fig. 2A, C). Cross-bedding
was occasionally observed in sandy siltstone beds.
Common cross-bedding, planar lamination, grada-
tion and erosional bases with flute marks were
recorded in the sandstone beds. The wrinkle struc-
tures were observed almost exclusively on the bed-
ding planes of those sandy siltstones that are
overlain by clayey siltstones with land plant remains,
and they always occur on the upper bedding planes.
In total, 38 slabs with wrinkle structures were pho-
tographed and measured. The structures observed
are formed of small mounds and ridges, with charac-
teristic patchy occurrence on the bedding planes
(Fig. 2B). Two different types of wrinkle structures
were observed. The first type is represented by deli-
cate, well-distinguished mounds, 2–5 mm in length
and 1.5–2 mm in width, joined with an adjacent
mound with a low ridge, forming a depression in the
centre (Figs 2B, 3A–D, 4B, C). The spacing between
the mounds is 2–3 mm. The second type (Fig. 4A)
Wrinkle structures in Middle Devonian siliciclastics 5
is represented by slightly wider (3–4 mm), less well-
distinguished mounds, with a rather diffused
appearance. However, the overall pattern of these
two types is the same – mounds joined with a low
ridge, forming a depression in the central area. The
length of the mound can be shorter in both of the
groups, so they are rather isometric, showing a hon-
eycomb-like pattern (Fig. 4B, C). It is important to
mention that wrinkle structures were observed by us
also in other parts of the Prague Basin in the silt-
stones of the Robl
ın Member, namely in the Hlu-
bocepy-railroad cut locality and in the Srbsko and
Host
ım vicinity (ca 20 km SW from Hlubocepy).
Petrographic observations
The wrinkle structure-bearing sandy siltstones are
moderately sorted, dominated by sub-angular to
sub-rounded, corroded, monocrystalline quartz
grains, mica flakes (biotite, muscovite), minerals of
the chlorite group and a minor amount of feldspars.
Petrographic observations, SEM and EDX analyses
(performed both on polished and on non-polished
thin sections) revealed the presence of filamentous
wavy structures (wavy-crinkly texture sensu Schieber
1986) composed of organic matter (Figs 5–8) and its
diagenetic products, such as pyrite and iron oxides
(Fig. 6C). Individual filaments, which are ca 5–
10 lm thick (Fig. 6E), tend to stack and form
thicker bundles. Those bundles have an appearance
of discontinuous wavy laminae, which are ca 10–
30 lm thick (Figs 5, 6A, B, D, F). Euhedral pyrite
replacing organic matter (6C) and collapsed pyrite
framboids within the organic, crinkle laminae, were
observed (Figs 7, 8B). Grains overgrown by organic
filaments were also recorded (Figs 6F, 8C). The EDX
spectrum shows the presence of carbon as a main
component in the filament (Figs 7, 8C). The minor
amount of other elements represents a contribution
from the surrounding matrix, coming from the sub-
surface interaction volume of the analysed feature.
Biomarkers and n-alkanes
The molecular characteristics of the organic matter
are summarized in Tables 1 and 2 and presented in
Figure 9 as the labelled chromatograms or ‘finger-
prints’ of the saturated hydrocarbons. Each sample
has a unique molecular composition, which provides
details on the biological origin of the organic matter
and of the changes in the depositional conditions.
The full-scan GC-MS analysis shows a strong preva-
lence of alkanes (m/z 57) over steranes (217) and
terpanes (191) by several orders of magnitude. All
fingerprints have a notable UCM hump (unresolved
6 Vodr
az kov
a et al. LETHAIA 10.1111/let.12280

Fig. 2. A, lithological log of the studied Hlubocepy gorge section showing alternation of fine-grained sandstones, sandy siltstones and
siltstones of the Robl
ın Member. B, examples of sandy siltstones of the Robl
ın Member with wrinkle structures on bedding planes. C,
field photo of the exposed and cleaned part of the section. [Colour figure can be viewed at wileyonlinelibrary.com]
LETHAIA 10.1111/let.12280 Wrinkle structures in Middle Devonian siliciclastics 7

A B

C D

Fig. 3. Examples of the wrinkle structures preserved on bedding planes of sandy siltstones of the Robl
ın Member from the Hlubocepy
section. A, B, D, correspond to 85–90 cm above the base of the section, C corresponds to the level of 260 cm above the base. All samples
belong to the first type of wrinkle structures described herein. Scale bar = 1 cm.

compound mixture). Aromatic hydrocarbons con-
sist of well-preserved phenanthrene series while
naphthalenes occur in low quantities.
Discussion
Petrographic observations
The petrographic observations and SEM and EDX
analyses of the siltstones of the Robl
ın Member
bearing wrinkle structures revealed features that
are suggestive of the former presence of microbial
mats, that is discontinuous wavy laminae formed
by organic matter and its diagenetic products
(Figs 5–8). Filament-like micro-structures appear
in thin sections in a tangled mass. The structures
preserved appear to be the sheaths and the fila-
ments typical of bacteria. As such, the structures
were most likely remnants of mucilaginous sheaths
of ensheathed forms, one of the most common
morphotypes of benthic bacteria involved in mat
and biofilm formation (Gerdes et al. 2000). Col-
lapsed pyrite framboids documented within the
organic filaments (Fig. 7) can be regarded as sup-
portive evidence for the microbial nature of the
observed filamentous structures (e.g. Schieber
2002). We can reasonably suppose that the mats
were formed by micro-organisms that produced
EPS (extracellular polymeric substance), given that
it is EPS, which is responsible for biofilm cohesion
and adhesion to substrates. It was shown by Flem-
ming (1995) that EPS has a high cation-binding
capacity. Thus, microbial mats themselves could
have provided the iron needed for sulphide
(pyrite) formation. Interestingly, sheaths of the
modern bacterium Thioploca coated by iron sul-
phides were reported by Schulz et al. (2000).
Kez dzierski et al. (2015) described pyrite framboids
in Trichichnus, formerly regarded as ichnofossil,
and interpreted it to represent a remnant of pyri-
tized Thioploca-related microbial mats. The sulphur
needed for pyrite precipitation could form during
bacterial decomposition of the mat, but there is
another plausible sulphur source, which should be
taken into consideration. The modern sulphur-oxi-
dizing bacterium Thioploca is capable of storing
elementary sulphur and therefore can act as a
potential site for pyrite precipitation. Thus, the
pyrite framboids reported herein could also be
related to the metabolic pathway of the mat-form-
ing micro-organism. On the other hand, one
would expect more common occurrence of the
framboids within the filaments, if framboid forma-
tion was related to metabolism of the mat-forming
bacteria. We can only speculate about the strategy
of the mat-forming micro-organisms; however,
there are a few other hints, which point to non-
8 Vodr
az kov
a et al. LETHAIA 10.1111/let.12280

A B

Fig. 4. Examples of the wrinkle structures preserved on bedding planes of sandy siltstones of the Robl
ın Member from the Hlubocepy
section. A, (260 cm above the base of the section) correspond to the second type of wrinkle structures described herein (less distinctive
and slightly wider mounds). B, C, (100 cm above the base) shows honeycomb pattern of the wrinkle structures. Scale bar = 1 cm.

A B

C D

E F

Fig. 5. Petrographic thin sections of wrinkle structures bearing sandy siltstones of the Robl
ın Member. A-F, crinkled laminae formed by
organic matter and its diagenetic products (iron oxides) interpreted herein to represent remnants of microbial mats. Note the grain coat-
ing and enclosing (e.g. B-arrowed) and several episodes of mat colonization of the sediment (e.g. E-arrowed). A–D, 9–15 cm above the
base of the section; E, F, 85–90 cm above the base of the section. [Colour figure can be viewed at wileyonlinelibrary.com]
LETHAIA 10.1111/let.12280 Wrinkle structures in Middle Devonian siliciclastics 9

A B

C D

E F

Fig. 6. Petrographic thin sections of wrinkle structures bearing sandy siltstones of the Robl
ın Member. A-F, crinkled laminae formed by
organic matter and its diagenetic products (iron oxides) interpreted herein to represent remnants of microbial mats. Note the grain coat-
ing and enclosing (A, B, D, F arrowed), aligned euhedral pyrite crystals replacing organic matter of the microbial mat (C) and the single
filament of the microbial mat (E). A, B, D–F = 85–90 cm above the base of the section, C = 260 cm above the base of the section.
[Colour figure can be viewed at wileyonlinelibrary.com]

autotrophic metabolic pathways below the photic

zone. This evidence includes the sedimentary
facies, which are thin-bedded distal turbidites with-
out sedimentary structures implying shallow water
deposition and also the lack of biogenic features
resulting from the need of light, such as domal
structures, quartz accumulations or authigenic car-
bonate mineral precipitation as described by, for
example Noffke (2009) and Gerdes et al. (2000).
Given the ubiquity of microbes in the deep ocean
today, often forming huge areas covered by bacte-
rial films (e.g. Gallardo 1977), such a record of
deep-water Palaeozoic microbial sediments (or
microbially induced sedimentary structures) is not
surprising. In addition, the geometry of biofilms
formed by sulphur-oxidizing bacterium Beggiatoa
shown by Thar & Kh€ ul (2002; Fig. 7D) resembles
the pattern of the wrinkle structures observed
herein (Fig. 4B, C). Given the analogy with micro-
bial mats, which thrive in oxygen minimum zones
of modern oceans, a reasonable assumption could
be made, that the Robl
ın Member was deposited in
a similar environment.
Molecular fossils
Two types of molecular fingerprints occur in the
sample set (Fig. 9). The first type has odd > even n-
alkane homologues in the 15–19 range, OEP(17) > 1,
and maximum at n-C17. The second type (Hlu-1 and
Hlu-2) shows even>odd predominance, that is OEP
(17) < 1, with maximum at n-C18 and n-C16. Such a
record indicates that two possible biological sources
contributed to the n-alkane distribution in the sedi-
ments. The first type of fingerprint with n-C17 as the
dominant alkane is here interpreted in agreement
with the earlier observations made by, for example
Or
o et al. (1967), Han & Calvin (1969) or Blumer
et al. (1971) as evidence of organic material originat-
ing from phytoplankton living in the water column.
The even carbon predominance of n-C16 and n-C18
over n-C17 alkanes recorded at the second type is
suggestive of benthic bacteria (Han & Calvin 1969;
Nishimura & Baker 1986). It can be reasonably spec-
ulated that the benthic mucilaginous biofilms
enhanced preservation of the organic matter pro-
duced by phytoplankton derived from the water
10 Vodr
az kov
a et al.
36
38
25 µm
Fig. 7. SEM image of microbial mat filament with collapsed pyr-
ite framboids embedded within and corresponding spectrum
diagrams obtained from EDX analysis showing that the filament
is formed by organic matter (Spectra 36 and 38).
column. Similar observations, especially in carbon-
ates, are noted by Logan et al. (1999), Killops & Kil-
lops (2005), Peters et al. (1999, 2005) with some
minor differences, such as a lower abundance of
mid-range alkanes (n-C20–25) in our case. There is a
negligible or no contribution by terrestrial organic
matter in the studied samples evidenced by rather
high n-C17/(n-C17 + n-C27) ratio. The low pristane/
n-C17 and phytane/n-C18 ratios provide evidence,
that the biodegradation of the organic matter was
low and that the fingerprints are diagnostic of their
LETHAIA 10.1111/let.12280
200 µm
B
50 µm
C
6
10 µm
Fig. 8. SEM images of microbial mat filaments (A, B) and a
grain aggregate coated by filament (C) of organic composition,
as inferred from the spectrum diagram (Spectrum 6).
biological origin. The low pristane/phytane ratios of
the sample series suggest rather reducing conditions,
except for sample Hlu-3, which points to more oxic
ones. Thermal maturity of all the samples is very
stable and is equivalent to a vitrinite reflectance Rc of
0.67–0.70%, and a burial temperature of 110–
115 °C, that is early oil-generation window calcu-
lated from the aromatic hydrocarbons (MPI-1) using
the equation by Radke et al. (1986) and Cassani et al.
LETHAIA 10.1111/let.12280
(1988). This inferred temperature is further sup-
ported by the CPI-2 ratio of n-alkanes in the n-C15-19
range (Table 2 and Fig. 9). The features mentioned
above suggest that the fingerprints preserve the signa-
tures of biological input and that their alteration due
to thermal maturation and biodegradation is low.
Environmental controls on the origin of the
wrinkle structures in the Robl
ın member
As noted by Noffke et al. (2002), the interplay of an
ecological window (mat development) and a tapho-
nomic window (mat preservation) is necessary for
the development of fossilized microbial mats. Biotur-
bation is widespread in oxygenated environments in
the Phanerozoic, and the continuous reworking of
the sediment does not allow microbial mats to
develop. In addition, microbial mats can be
destroyed by metazoan grazers within days (Flenchel
Fig. 9. Fingerprints of the n-alkanes and isoprenoids in the saturated hydrocarbon fraction of the rock extracts. Positions of the samples
Hlu-1, Hlu-2, Hlu-4 and Hlu-5 are marked in Figure 2A. Sample Hlu-3 comes from the Hlubocepy-railroad cut locality, 3.5 m above
the base of the Robl
ın Member.
Wrinkle structures in Middle Devonian siliciclastics 11
1998). Therefore, harsh environmental conditions,
such as abnormal salinity, oxygenation or tempera-
ture as well as episodic sedimentation, are required
(e.g. Farmer 1992). The importance of benthic graz-
ers is also illustrated by a number of papers reporting
microbial sediments from stratigraphical intervals in
which evidence for benthic grazing is limited or miss-
ing (see references above). The requirements outlined
above were obviously fulfilled during sedimentation
of the Robl
ın Member. The pristane/phytane ratio
(Table 1) points to the frequent occurrence of reduc-
ing conditions. In addition, considering the short
time needed for recent cyanobacteria to form a mat,
which is on the order of hours to days (Shepard &
Sumner 2010) and long time needed for seafloor re-
colonization by benthic organisms after depositional
events (hundreds and thousands of years, Young
et al. 2001), we suppose that low sedimentation rates
between episodes of turbidity current deposition
12 Vodr
az kov
a et al. LETHAIA 10.1111/let.12280

Table 2. Extracted saturated hydrocarbon yield (Sat) in mg per kg of rock and molecular ratios of the n-alkanes and aromatic hydrocar-
bons in the rock extract. Positions of the samples Hlu-1, Hlu-2, Hlu-4 and Hlu-5 are marked in the Figure 2A. Sample Hlu-3 comes from
the Hlubocepy-railroad cut, 3.5 m above the base of the Robl
ın Member. Explanations of the abbreviations and ratios: Sat (mg/kg) –
absolute amount of saturated hydrocarbons in 1 kg of rock; OEP(17) – odd/even predominance index OEP(17) =
(C13+2*C15+2*C17+C19)/(2*(C14+C16+C18)); CPI-2 (n-C25–33) – carbon preference index =(C25 + 2*C27 + 2*C29 + 2*C31 + C33)/
(2*(C26 + C28 + C30 + C32)), Marzi et al. (1993); n-C17/(n-C17 + n-C27) – ratio of the numbered n-alkane homologues; MPI-1 –
methylphenanthrene index = 1.89*(3MP+2MP)/(P + 1.26*(9MP+1MP)), Cassani et al. (1988); MPDF – methylphenanthrene distribu-
tion factor MPDF = (3MP+2MP)/(3MP+2MP+9MP+1MP); Rc(MPI-1) % – calculated vitrinite reflectance based on MPI-1: Rc =
0.38 + 0.61*MPI1.

Sample Sat (mg/kg) OEP (17) CPI-2 (n-C25–33) n-C17/(n-C17 + n-C27) MPI-1 MPDF Rc% (MPI1)

Hlu-1 859 0.92 1.11 0.90 0.51 0.46 0.70

Hlu-2 684 0.89 1.05 0.90 0.45 0.52 0.67
Hlu-3 503 1.04 1.09 0.97 0.48 0.54 0.69
Hlu-4 921 1.07 1.27 0.96 0.49 0.54 0.69
Hlu-5 638 0.92 1.06 0.91 0.46 0.53 0.67

represent enough time for mats to be formed, but
not enough time for benthic assemblages to colonize
the seafloor. Benthic invertebrate fossils are rare in
the Robl
ın Member and recorded only at the very
base and upper part of the unit (Chlup
ac 1958). The
episodic depositional events possibly coupled with
lower salinities (Kukal & J€ager 1988) and reducing
environment obviously represented harsh environ-
mental conditions. Also, the low-diversity trace fossil
assemblages described by Mikul
as & Pek (1996) are
formed by r-strategic forms of filter and sediment
feeders, which could cope with periodic disturbances
represented by turbidite flows.
Origin of the wrinkle structures in the Robl
ın
member
Although reports of deep-water wrinkle structures
are under-represented in comparison with those
from shallow water, they do show comparable mor-
phologic features. Modern examples of wrinkle
structure from tidal flats as shown by Hagadorn &
Bottjer (1997, 1999) and Bose & Chafetz (2009) and
fossil deep-water examples shown by Buatois &
M
angano (2003) and Pazos et al. (2015) show very
similar patterns. One would therefore expect action
of the same or similar forces leading to formation of
the wrinkles. Whereas wind and/or wave and/or cur-
rent action can explain wrinkle formation in shallow
water, it fails to explain their occurrence in the deep
sea as concerns the wind and waves. Bearing in mind
the supposed viscoelastic nature of microbial mats
(Klapper et al. 2002), it seems reasonable that
wrinkle structures could develop simply due to
gravitational forces, even when occurring on very
gentle slopes. That can easily explain their published
occurrences both in shallow and in deep water.
Interestingly, we observed very similar wrinkle pat-
terns on pavements made of asphalt concrete
(Fig. 10A,B). Asphalt, similarly to microbial mats, is
a viscoelastic material (Anderson & Goetz 1973),
and as such responds according to physical and ther-
mal stress.
There is another interesting feature, which should
be taken into consideration. Fossil deep sea wrinkle
structures, described by Dzulynski & Walton (1965),
Buatois & M
angano (2003) and Pazos et al. (2015),
have one thing in common, namely occurrence in
flysch-like deposits. Also, Brett et al. (2003)
recorded wrinkle structures (Kinneyia in his descrip-
tion) in the Ordovician Kope Formation in associa-
tion with seismites and suggested that they may have

A B

Fig. 10. (A) Example of pavement made of asphalt concrete with deformation resembling wrinkle structures. (B) Detail of example (A).
1-euro coin was used to indicate the scale.
LETHAIA 10.1111/let.12280
originated during seismic events. Similarly, Seilacher
(2008) noted an association of Kinneyia with earth-
quake triggered sedimentary structures in the
Silurian Tanezzuft Shales in Libya.
The deposition of the Robl
ın Member of the Srb-
sko Formation correlates with Middle Devonian
continental underthrusting of the Saxothuringian
plate underneath the TBU plate (Schulmann et al.
2009). Besides gravity instability or current shear, we
should therefore consider other possible physical
disturbances, which the microbial mats responded
to, namely seismic shocks. The sediment with micro-
bial mats behaved as thixotropic matter, and the
wrinkle structure observed could represent a defor-
mation triggered by seismic waves. Detailed facies
and petrographic study of the entire succession of
the Robl
ın Member is necessary to confirm such an
interpretation. The influence of Variscan tectonic
movements on Robl
ın Member development was
stressed already by Chlup
ac (1958; p. 183) followed
by others (see above).
Conclusions
Assessing all the characteristics of the sediments
observed, that is direct association of wrinkle struc-
tures on bedding planes and organic wavy laminae
in thin sections, the patchy occurrences of the wrin-
kles on the bedding planes of sandy siltstones and
sediment cohesive behaviour demonstrated by the
shapes of wrinkles, we interpret the observed wrinkle
structures as being microbially induced. We con-
clude that specific ecological and taphonomic condi-
tions were met during the sedimentation of the
Robl
ın Member, facilitating microbial mat forma-
tion and preservation, that is the absence of biotur-
bation (stressed environment for benthic grazers)
and/or insufficient time between sedimentation
events for colonization of benthic bioturbators, suit-
able facies (heterolithic strata of turbidites), quick
burial and moderate hydrodynamics. We interpret
the wrinkle structure as a sedimentary structure
induced by the presence of cohesive microbial mats,
which also enhanced preservation of the structure.
The mats responded to various physical stresses by
deformation, which is now preserved as a wrinkle
structure. We consider gravitational forces and/or
current as the simplest explanation of its origin,
which would also explain the origin of such struc-
tures in both shallow and deep water as reported
from the literature. However, we also consider
another alternative as probable, namely that wrinkle
structures represent soft sediment deformation
structures originating due to exposure of
Wrinkle structures in Middle Devonian siliciclastics 13
mat-bearing sediment to seismic shocks. Such an
explanation sounds reasonable especially considering
the published reports on wrinkle structure from
deep-water flysch-like deposits.
The nature of the micro-organisms forming the
mat is not known to us. The distribution of n-
alkanes and isoprenoids described herein suggests
two types of prevailing biological sources, phyto-
plankton with odd > even n-alkanes, representing
organic material from the water column and ben-
thic bacteria with even > odd homologues. Judg-
ing from the sedimentary facies (deep-water
turbidites) and lack of features suggestive of light
competition (such as translucent quartz-grain
accumulation and authigenic carbonate precipita-
tion), we interpret the mats to have been formed
by non-autotrophic bacteria.
Acknowledgements. – This research was supported by the Czech
Science Foundation through the Project No. P210/12/2018 and
the project of the Czech Geological Survey No. 339900. This
study is a contribution to IGCP 652. SV wishes to acknowledge
the support of Alexander von Humboldt foundation as part of
the study was carried out during her AvH Research Fellowship in
the University Erlangen-Nuremberg. Gilbert Klapper is acknowl-
edged for English corrections. We thank to Radek Mikul
as
(Czech Academy of Sciences, Czech Republic) for his advice on
some problematic trace fossils from the Robl
ın Member and
Zdenek Kukal (Czech Geological Survey, Prague) for discussion
about the sedimentology of the Robl
ın Member. The authors are
indebted to Carlton E. Brett (University of Cincinnati, Ohio) for
his meticulous review and very helpful comments, which
improved the manuscript. The authors declare no conflict of
interest regarding the publication of this article.
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