GeolCarp_Vol60_No3_233

www.geologicacarpathica.sk

GEOLOGICA CARPATHICA, JUNE 2009, 60, 3, 233—250 doi: 10.2478/v10096-009-0016-1

Introduction

Red-coloured rocks displaying all signs of deep marine sedi-
mentation  (turbidites,  hemipelagites)  are  called  Oceanic  Red
Beds. This relatively rare facies is associated with specific en-
vironmental  parameters.  Although  their  presence  in  the  geo-
logical  record  is  not  exclusively  restricted  to  the  Cretaceous
Period,  these  sediments  are  common  in  Upper  Cretaceous
oceanic basins, where they are referred to as Cretaceous Oce-
anic  Red  Beds  (CORB;  see  Hu  et  al.  2005).  In  the  last  15
years,  their  significance  for  the  reconstruction  of  paleoenvi-
ronmental conditions in Late Cretaceous oceans has been ac-
knowledged (cf. Skupien et al. 2009 and references therein).

Upper Cretaceous CORB are present in several tectonic

units in the Outer Western Carpathians. Identification of more
or  less  complete  sections  as  well  as  mutual  correlation  be-
tween  isolated  outcrops  are  complicated  by  the  nappe  struc-
ture, minor tectonic deformations and locally also by the high
thicknesses of the red beds. The correlation is also hampered
by the considerable lateral diversity of the CORBs and the ex-
istence  of  transitional  facies  (Skupien  et  al.  2009).  Field  and
laboratory  studies  were  conducted  in  years  2005—2007  with
the aim to solve the persisting correlation problems and to lay
the basis for a more detailed interpretation. These studies were
focused  on  integrated  biostratigraphy  (foraminifers,  di-
noflagellates,  calcareous  nannoplankton),  sedimentology,
mineralogy and ichnology (Skupien et al. 2009).

The ichnological characteristics of sequences containing

CORB and some transitional facies is the subject of the
present paper. The following aims were outlined: 1. provide
information on substrate colonization and its fluctuations, and

Ichnology of the Cretaceous Oceanic Red Beds
(Outer Western Carpathians, Czech Republic)

RADEK MIKULÁŠ

, PETR SKUPIEN

, MIROSLAV BUBÍK

and ZDENĚK VAŠÍČEK

Institute of Geology, Academy of Sciences of the Czech Republic, Rozvojová 269, Praha 6, Czech Republic; mikulas@gli.cas.cz

Institute of Geological Engineering, VŠB-Technical University, 17. listopadu, Ostrava-Poruba, Czech Republic

Czech Geological Survey, Leitnerova 22, 65 869 Brno, Czech Republic

(Manuscript received December 18, 2007; accepted in revised form December 18, 2008)

Abstract: Large differences in the intensity and overall character of bioturbational structures were found in five facies
containing hemipelagic red beds. Red beds (CORB) of the Godula facies of the Silesian Unit and their equivalents
(mostly not red) in the Kelč facies of the Silesian Unit and the CORB in the non-calcareous sediments of the Rača Unit
display a very low degree of bioturbation. The CORB facies of the Rača Unit, containing calcareous intercalations,
displays a very high degree of bioturbation as expressed by a high ichnofabric index. They contain trace fossils Chon-
drites, Zoophycos, Planolites, Thalassinoides, Palaeophycus, Teichichnus and Phycosiphon. The supply of food obvi-
ously acted as the controlling factor. The “calcareous” facies of the CORB of the Rača Unit has a considerably higher
proportion of sand-dominated interbeds and also carbonates than the non-calcareous facies. This (especially the pres-
ence of carbonates) suggests a relative proximity of food-rich environments and an easy transport of nutrition-rich
substrate by turbidite currents into the basin directly, not only by periodical fall-out of dead plankton (which is probably
responsible for the rhythmicity of poor colonization horizons in weakly bioturbated units).

Key words: Upper Cretaceous, Western Carpathians, hemipelagic, ichnofossils, ichnofabric, red beds.

on feeding strategies of the benthos (thus contributing to re-
gional paleoenvironmental and paleogeographic conclusions);
2. define the Oceanic Red Beds phenomenon against the tran-
sitional facies; 3. provide comparative information for other ar-
eas with occurrences of the Oceanic Red Beds, notably CORB.

Previous ichnological studies in the Oceanic Red Beds

No synoptic ichnological study of the CORB from a large

area  comprising  several  units  and  lithofacies  has  been  com-
pleted yet. Ichnological research before the early 1990s, when
the  ichnofabric  concept  was  widely  developed  (ichnofabric;
Ekdale et al. 1991), gave preference to facies and sites yield-
ing  well  preserved  biogenic  sedimentary  structures  of  recur-
rent morphology, namely trace fossils. This, however, was not
the  case  with  distal  turbidites  or  hemipelagic/pelagic  clays.
Detailed  studies,  dealing  not  only  with  lists  of  determinable
ichnotaxa, but also with the degree of reworking (ichnofabric
index), separate phases (sequences) of substrate colonization,
ichnofabric  rhythmicity,  hence  also  with  the  rhythmicity  of
colonization  windows,  and  with  the  relationship  between  the
ichnocoenosis  and  the  substrate  colour,  were  published  after
the early 1990s (Leszczyński 1993; Wetzel & Uchman 1998a,
2001). These studies mostly concentrated on muddy turbidites
as  a  whole,  without  any  particular  reference  to  their  colour
(which is the essential indicator of substrate oxidation, cf. Pot-
ter et al. 1980). Exceptional in this respect are the papers by
Leszczyński (1993; a report of sections in the Cretaceous and
Tertiary  turbidite  sequences  from  Spain,  with  generally  very
low or no intensity of bioturbation in red clays/claystones) and

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MIKULÁŠ, SKUPIEN, BUBÍK and VAŠÍČEK

Bąk (1995; Cretaceous marls of the Polish Carpathians). There
are also other papers, which concern mostly the Paleogene red
shales, namely Leszczyński & Uchman (1991, 1993).

Wetzel & Uchman (1998a) stated that red to brown clay-

stones deposited in the oceans are characterized by slow sedi-
mentation, limited food sources – at least inside the sediment
(a temporary accumulation of dead phytoplankton occasional-
ly rests on the surface) – and by generally complete bioturba-
tion. The number of tiers is, however, limited, and the depth
range  of  the  bioturbation  is  only  a  few  centimeters.  An  in-
crease  in  the  sedimentation  rate  may  result  in  a  considerable
increase in food content, hence also in the depth of burrowing
(and in the diameter of burrows). The common ichnotaxa are
Chondrites,  Zoophycos,  Planolites,  Thalassinoides,  Palaeo-
phycus,  Teichichnus,  Phycosiphon,  Lophoctenium  and  Nere-
ites.  Most  of  them  represent  burrows  permanently  connected
to the sea floor (Chondrites is a typical example); such adapta-
tion may be favoured due to the low permeability of the bottom.

In the Outer Western Carpathians, more general attention

has been given to the bioturbation structures of the CORB and
the  surrounding  units.  Intensively    bioturbated  rocks  have
been reported from some units underlying the CORB (Sku-
pien  &  Vašíček  2003),  and  simple  characteristics  of  the
CORB  ichnology  have  been  provided  from  some  sections  of

the Silesian and Rača Units (Skupien et al. 2009). However,
detailed studies have not been presented.

Geological settings

The Outer Carpathians represent the northernmost zone of

the Alpine-Carpathian orogen. They consist of two groups of
nappes (from top to bottom): Magura Group of Nappes (Bílé
Karpaty,  Bystrica,  Rača  Units)  and  Outer  Group  of  Nappes
(Fore-Magura,  Silesian  and  Subsilesian  Units;  Fig. 1);  each
of  the  units  may  have  its  specific  local  lithofacies.  During
the  Mesozoic,  the  Outer  Western  Carpathians  domain
rimmed  the  continental  foreland  in  the  SE  continuation  of
Paleo-Europe.  This  domain  was  separated  by  the  Penninic
Oceanic Branch from the Central Carpathian—Alpine micro-
continent,  associated  with  the  Adriatic  microcontinental  as-
semblage.  The  Penninic  rifting  resulted  in  tensional  stress
accelerating both the subsidence and tilting of the Outer Car-
pathian  intraplatform  basins,  accompanied  by  local  volcan-
ism (e.g. Mišík 1992; Michalík 1994).

Locally more than 6 km thick flysch deposits are typical of

the Outer Carpathian sedimentary sequences. They were de-
posited in several troughs separated by ridges; from South to

Fig. 1.  Location  map  showing  the
main  tectonic  units  of  the  Outer
Western  Carpathians  in  the  Czech
Republic. After Skupien et al. (2009).

North,  these  are  called:  1  –
Magura  Basin;  2  –  Silesian  Ba-
sin; 3 – Subsilesian Basin. Dep-
osition started in the Late Jurassic
as  a  consequence  of  enhanced
subsidence.

Sedimentation

black shales with local submarine
clastic  fans  embraced  almost  the
whole  Outer  Carpathian  basin  in
the  Early  Cretaceous.  Slow  and
uniform  sedimentation  of  green
and  black  shales  took  place  dur-
ing  the  Albian  and  Cenomanian,
followed by sedimentation of red
and  variegated  shales  (CORB)
under  well-oxygenated  condi-
tions  (Fig. 2).  The  CORB  forma-
tion  was  terminated  by  the
progradation  of  sand-dominated
turbidites  deposited  in  the  axial
parts  of  the  subbasins  and  along
the  continental  slope  during  the
Late Senonian.

The tectonic structure of the

Outer  Carpathians  is  a  result  of
several  tectonic  events  (Neo-Al-
pine  orogenic  processes).  Initial
folding took place during the Oli-
gocene in the southern part of the
Magura  Subbasin  and  then  pro-

235

ICHNOLOGY OF THE CRETACEOUS OCEANIC RED BEDS (CZECH REPUBLIC)

gressed northeastwards. The formation of the thrusts and up-
lifting of this area was completed during the Early Miocene
(e.g. Mišík 1992; Michalík 1994).

Material and methods

Ichnological study at all documented sections followed ear-

lier lithological description and integrated biostratigraphic
study (Skupien et al. 2009). The thickness of the sections from
tens of meters to 300 m at Bystrý potok Stream did not allow a
detailed, layer-by-layer ichnofabric documentation using an

abbrasive  paper  (in  mm-resolution).  The  essential  resolution
was on the order of tens of centimeters, with particular atten-
tion  given  to  lithological  boundaries  and  colonization  hori-
zons:  here,  documentation  works  were  designed  to  achieve
cm-resolution.  Despite  all  the  effort,  it  can  be  assumed  that
some of the weak colonization horizons have not been encoun-
tered. Information on their typical vertical spacing in the section
and their overall character is, however, well substantiated.

A principal problem in the study of ichnofabrics of litholog-

ically monotonous pelitic sediments is the differentiation be-
tween completely bioturbated facies and facies with no
bioturbation. The absence of lamination is usually taken as ev-

Fig. 2. Stratigraphy, lithology and ichnology of the Bystrý potok section (lower part).

236

MIKULÁŠ, SKUPIEN, BUBÍK and VAŠÍČEK

idence for total sediment reworking. Nevertheless, no distinct
laminae may be visible in pelites with a very low proportion
of detrital mica or other material subject to planar arrangement
during the deposition. The most effective tool for a correct so-
lution  of  this  dilemma  is  an  approximation  by  lithological
boundaries. Such approximation should, however, always in-
volve a consideration on the origin of the respective boundary:
it  may  be  connected  with  previous  sea-floor  erosion  or  a
swing  in  environmental  parameters  potentially  affecting  the
benthic biocoenosis. In any case, we are aware of the fact that
the ichnofabric index (abbreviated ii further in the text; Droser
&  Bottjer  1986)  itself  in  some  portions  of  the  studied  sedi-
ments is a matter of interpretation rather than a mere mechani-
cal determination.

Descriptive part

Silesian Unit – Godula facies

CORB and adjacent or similar rocks were studied in two fa-

cies of the Silesian Unit, namely the Godula and Kelč facies

(or  Godula  and  Kelč  developments  by  Menčík  et  al.  1983).
Different sedimentological settings resulted in separate lithos-
tratigraphies  of  the  facies.  The  Godula  facies  was  deposited
below  the  CCD  in  the  continental  rise  setting  in  the  form  of
thick turbidite fans. Upper Cretaceous strata have the charac-
ter of shale-sandstone flysch up to 3500 m thick. The Kelč de-
velopment  represents  mostly  slope  shales.  The  thickness  of
the Upper Cretaceous strata reaches several hundred meters.

The Godula facies, comprising turbidite fan facies, was

studied  in  a  continuous  section  through  the  CORB  at  the
Bystrý  potok  locality  (Skupien  &  Vašíček  2003).  The  mea-
sured section starts with the Lhoty Formation (Albian, Lower
to  low-Middle  Cenomanian).  The  CORB  are  present  within
the Mazák Formation (Middle to Upper Cenomanian and Tu-
ronian) and the lower member of the Godula Formation (Co-
niacian, Santonian and Lower Campanian; Figs. 2 and 3).

Lhoty Formation in its typical development.

The Lhoty For-

mation underlying the CORB-bearing complex is composed of
medium grey to green-grey bioturbated shales, occasionally in-
tercalated with non-bioturbated, thin sandstone turbidite beds.
No visible ichnofabrics are displayed by most mudstones in the
lowermost part (ca. 15 m) of the section. This can be explained

Fig. 3. Stratigraphy, lithology and ichnology of the Bystrý potok section (upper part).

238

MIKULÁŠ, SKUPIEN, BUBÍK and VAŠÍČEK

primarily by two facts: (1) no colonization of substrate ever oc-
curred, as indicated by the preservation of the primary lamina-
tion  (especially  in  siltstones  with  sand  admixture);  (2)
homogenization by bioturbation took place, as shown by lighter
grey claystones with no preserved lamination; homogenization
was followed by an episode of rapid sedimentation, which pre-
vented  colonization  by  deep-tier  burrowers  (Chondrites).
Where neither of the above mentioned possibilities is valid, an
ichnofabric  Planolites—Thalassinoides—Chondrites  is  present;
see  Fig. 4B,D,H.  Some  intervals  manifest  a  very  contrasting
preservation of the deepest tier, represented by “large forms” of
Chodrites  on  a  bioturbated  background  (sometimes  a  com-
pletely  bioturbated  backround,  i.e.  ichnofabric  index = 5 ;
Fig. 4A,C,E,F).  The  given  order  of  ichnotaxa  corresponds  to
their order of appearance in the substrate and also to the com-
pleteness of preservation of the biogenic structures. The ichno-
fabric index therefore oscillates between the values of 1 and 4 or
5  over  more  or  less  irregular  intervals,  with  a  prevalence  of
higher  values  (almost  complete  or  complete  bioturbation),  see
Fig. 2.  Such  ichnofabric  documents  normal  oxygen  and  food
conditions  and  has  many  parallels  in  other  Mesozoic  and
younger flysch sediments (e.g. Wetzel & Uchman 2001).

Uppermost part of the Lhoty Formation.

The uppermost

part of the Lhoty Formation consists of light grey shale with
few thin intercalations of dark grey shale with fish remains, si-
licified  siltstones  and  cherts  (Fig. 5B).  Several  centimeters
thick intercalations of grey, fine-grained siliceous glauconitic
sandstone  occur  in  the  higher  part  of  this  unit.  A  round  bio-
genic structure resembling ichnogenus Bergaueria (interpret-
ed as dwelling burrows of sea anemones; Fig. 4G) was found
at the base of silicified siltstones. Vertical shafts are occasion-
ally  present,  indicating  a  period  of  relatively  stable,  strong
flow. The overlying part of the section reaching to the base of
the Mazák Formation is dominated by light grey-green to dark
grey  claystones.  The  ichnofabric  Planolites—Thalassinoides—
Chondrites is present at only a few places. The ichnofabric in-
dex in this part of the Lhoty Formation is considerably lower
than  in  intervals  with  usual  development  of  the  same  forma-
tion: it generally oscillates between values of 1 and 2 over cm-
intervals while values of 3 or 4 were recognized only rarely in
irregular intervals tens of centimeters thick.

Mazák Formation – variegated strata.

The Mazák Forma-

tion is a sequence of variegated (red, red-brown and greyish-
green)  non-calcareous  shales,  occasionally  interbedded  with
thin  beds  of  greenish  grey  quartzose  sandstones.  Red  shales
intercalated  with  grey  or  greenish  grey  shales  represent  the
most frequent lithotype. At the Bystrý potok section, the for-
mation is 94 m thick.

A preliminary study of the section revealed only a few bio-

turbated intervals (i.e. records of colonization windows) in the
red claystones of the Mazák Formation. Small Chondrites was
mostly the only ichnofossil encountered (e.g. at 68 m, Fig. 6F,
and  at  94.5 m,  Fig. 6C—E).  Exceptional  is  the  interval  with
clearly  visible  Chondrites—Planolites  ichnofabric  (95 m,
Fig. 6G,J), indicating a short-lived increase in nutrient supply.
This  is,  after  all,  also  documented  by  grey  fillings  of  the
Planolites tunnels. The overlying claystone bed is much light-
er than the above mentioned fillings of ichnofossils and con-
tains abundant, very minute Chondrites.

A repeated investigation of this interval (which is of key im-

portance for the understanding of the ichnological contents of
CORB  in  the  Outer  Western  Carpathians  due  to  its  context,
completeness of the section, and good exposure) brought finds
of  other  weak  colonization  horizons,  generally  comprising
solitary or very sparse minute individuals of Chondrites isp. It
can be stated that records of poorly colonized bottom – diffi-
cult to document due to its low intensity – occur irregularly
throughout  the  CORB  section  of  the  Mazák  Formation,  with
spacings of tens of centimeters. Somewhat more compact red
beds with macroscopically visible flakes of detrital mica were
found  at  35 m.  Vertical  sections  in  the  rock  show  no  visible
lamination. Samples prepared by splitting of rock along bed-
ding planes, however, show more or less developed lustre (ori-
entation  of  mica  flakes  and  other  clasts),  which  is  not  fully
homogeneous and suggests a relatively intensive bioturbation
(Fig. 6A,B;  ii = 3).  Rather  coarse  (siltstone)  beds  with  trace-
able lustrous bedding surfaces can be equally found elsewhere
in the section; with the exception of the above mentioned red
beds at 35 m, however, they show no (ii = 1) or a very weak
(ii = 2) degree of bioturbation.

A sample of similar rock with notable greyish-green lami-

nae was found in a streambed below the interval at 35 m. The
greyish-green  colour  of  “variegated”  hemipelagic  pelites  and
aleurites  is  either  primary  (iron  oxides  and  hydroxides  are
never  present  in  large  proportions  in  the  rock)  or  secondary
(“depigmentation” by weathering processes); in the case of the
mentioned  find  it  is  difficult  to  decide  which  possibility  is
more  probable.  Nevertheless,  the  sample  found  also  shows
“mottling” on a lithological boundary analogous to that from
35 m, moreover, with contrasting colours (Fig. 6K).

Therefore, the primary absence of bioturbation (ii = 1) from

a  large  part  of  red  beds  is  much  more  likely  than  their  com-
plete  reworking  (ii = 5):  red,  definitely  non-bioturbated  beds,
much like bioturbated beds filled with rock of the same grain
size and composition as the substrate for burrowing, were en-
countered.  On  the  other  hand,  the  find  from  35 m  indicates
that many colonization horizons of shallow tier with relatively
large  tunnel  diameters  (maximum  10 mm)  were  probably
overlooked.  One  of  the  reasons  may  be  the  primary  extreme
substrate homogeneity, preventing the identification of coloni-
zation  windows  by  any  routine  method  of  study.  We  should
also consider that the coarser portions with detrital mica could
have been deposited at a higher rate than the surrounding clay,
which may have resulted in a higher frequency of colonization
windows in the clay beds (conversely, pore volume in slowly
deposited clay substrates may be very low, thus restricting the
bioturbation).  At  the  present  stage  of  field  and  laboratory
study, the variegated strata of the Mazák Formation should be
interpreted as rather weakly bioturbated, with prevailing ii = 1.

Mazák Formation – sand intercalations and rhythmic

sand-dominated flysch.

An interesting interval was encoun-

tered in situ around 59 m: two thin (ca. 8 cm) beds of glauco-
nitic  sandstone.  Its  base  shows  hypichnial  tunnels  of
Thalassinoides/Ophiomorpha,  Phycodes  and  Planolites.
Similarly, tabular beds of sandy flysch, which are relatively
abundant  at  48—66 m,  contain  a  rich  ichnoassemblage:
?Palaeophycus  cf.  sulcatus,  ?Pilichnus,  Helminthopsis  and
Arthrophycus (Fig. 7G,H). This suggests that episodes of in-

241

ICHNOLOGY OF THE CRETACEOUS OCEANIC RED BEDS (CZECH REPUBLIC)

put of coarse detrital material implied a short-lived influx of
nutrients necessary for the development of rich benthic com-
munities. Moreover, episodes of sand deposition (probably
of turbidite-flow origin) are favourable for the preservation
of biogenic structures in flysch sediments and, as such, rep-
resent only a weakly filtered taphonomic record.

Lower member of the Godula Formation.

It consists of

thin- to medium-bedded flysch dominated by grey shales; var-
iegated intervals still occur, but their colours are less intense:
red grey, brown grey, reddish brown grey. A 15 m thick body
of grey claystones with frequent clayey ironstones and no var-
iegated intercalations (160—175 m of the section) is present in
the  lower  part  of  the  member.  Higher  up  in  the  section,  the
lower  member  of  the  Godula  Formation  contains  two  bodies
of sandstone flysch, which are 38 and 13 m thick. Green-grey
colours  of  shale  prevail  in  the  uppermost  part  of  the  lower
member (Figs. 2 and 3).

Greyish-green or grey mudstones contain relatively frequent

intervals with Planolites—Thalassinoides—Chodrites ichnofab-
ric (Fig. 7A,D). Intervals with Chondrites only are even more
frequent (Fig. 7B). According to the above postulated criteria,
however, the lower member of the Godula Formation is gener-
ally dominated by no or very weakly bioturbated rocks (ii = 1).
Non-bioturbated intervals alternate with intervals of weakly to
moderately bioturbated rocks every several centimeters.

Clearly visible colonization horizons in red and brownish-

red shales are somewhat more frequent than in the same rocks
of the underlying Mazák Formation. They contain Chondrites
as well as Planolites. The topmost variegated strata host typi-
cal, markedly developed “large” Chondrites isp. (Fig. 7C).

Sandstone intercalations in the lower part of the lower

member  of  the  Godula  Formation  contain  Arthrophycus,
usually preserved in hyporelief. Intercalations of sand-domi-
nated  flysch  in  the  middle  part  of  the  lower  member  of  the
Godula  Formation  contain  Helminthopsis,  Arthrophycus
(Figs. 7E    and  8B),  Ophiomorpha  and  Planolites,  rare Zoo-
phycos  (Fig. 8C)  and  occasional  Nereites  isp.  (Fig. 8A).
Megagrapton  and  Zoophycos  occur  in  the  uppermost  strata
of  the  lower  member  of  the  Godula  Formation  (mostly
coarsely rhythmic alternation of sandstone and siltstone, sel-
dom claystone). The basal bed of the sand-dominated flysch
of the middle member of the Godula Formation (Fig. 5E) is
marked by an exceptional, giant biogenic structure ascribed
to the ichnogenus Treptichnus (Fig. 8E).

Silesian Unit, Godula facies – a summary.

Four types of

strongly bioturbated sediments were repeatedly identified: (1)
grey hemipelagic to pelagic mudstones completely bioturbat-
ed  at  levels  of  the  colonization  horizons,  with  two  well-de-
fined  tiers  of  biogenic  activity;  (2)  red  claystones  with
sporadic  presence  of  clearly  visible  colonization  horizons
mostly  represented  solely  by  Chondrites  with  low  density  of
individuals,  more  rarely  by  the  Planolites—Chondrites
succession with a low density of individuals. The presence of
additional  colonization  horizons  (probably  outnumbering
those clearly identifiable by several times, probably a shallow-
er tier) can be assumed on the basis of an analogy with occa-
sional  beds  with  higher  silt  content  in  the  studied  sequence;
(3)  moderately  to  coarsely  rhythmic  sand-dominated  flysch
with  Thalassinoides/Ophiomorpha,  Arthrophycus,  Phycodes

and  others,  which  roughly  corresponds  to  a  modification  of
the  “seilacherian”  Cruziana  ichnofacies;  (4)  moderately  to
coarsely  rhythmic  sand-dominated  flysch  with  Zoophycos,
Megagrapton  and Treptichnus  referring  to  the  “seilacherian”
Zoophycos ichnofacies with elements of the Nereites ichnofa-
cies.  Finely  to  coarsely  rhythmic  flysch  with  regular  alterna-
tion of pelites, siltstones and sandstones and with a suite of the
Nereites ichnofacies is missing (Paleodictyon, Nereites, Uro-
helminthoida,  Glockerichus,  Lorenzinia  and  other  gra-
phoglyptids).

Silesian Unit – Kelč facies

In the Kelč facies, the CORB are underlain by grey and

greenish-grey “mottled”, usually calcareous shales with vari-
able sand content. These are placed to the Jasenice Formation
(Eliáš 1979), which is roughly analogous in age and character
of sediments to the Lhoty Formation of the Silesian Unit (Sku-
pien  et  al.  2009).  In  the  Kelč  facies,  the  CORB  occur  in  the
Němetice  Formation,  which  was  defined  by  Eliáš  (1979)  as
green-grey  and  grey  shale  with  sporadic  red  and  brown-red
beds;  recently,  several  black-grey  shale  horizons,  grey  marl-
stones  to  clayey  limestones,  grey-green  siltstones,  and  thin
banks  and  lenses  of  fine-grained  calcareous  subgreywackes
were  also  encountered  in  the  type  area  near  Němetice  (Sku-
pien  et  al.  2009).  The  overlying  Milotice  Formation  (Eliáš
1979) consists of grey clays with variable contents of carbon-
ate,  silt  and  sand  admixture.  Red-brown  intercalations  occur
rarely. Sandstones are also rare and occur in thin isolated beds
(Fig. 9).

Jasenice Formation.

Several outcrops of this formation in-

cluding a larger one with the overlying red beds are exposed
in  a  gorge  of  a  stream  flowing  north,  from  the  easternmost
edge of Němetice to the area SE of the Pod Doubravou settle-
ment.  Greyish-green  shales  are  “mottled”,  completely  or  al-
most completely bioturbated (ii = 3—5); the separate “mottles”
can  be  interpreted  as  deeper  tier  trace  fossils  Planolites  and
Thalassinoides  (Fig. 10H,J).  Smaller  outcrops  further  down-
stream  and  talus  accumulations  yielded  fragments  of  tabular
sandstone  beds  with  hypichnial  Ophiomorpha  (Fig. 10G).  A
continuous mixing of substrates indicates favourable nutrition
and oxygenation conditions on the bottom.

Němetice Formation.

Approximately 4 m of mottled grey-

ish-green shales and ca. 3 m of red beds of the lower part of
the Němetice Formation crop out in a slope on the right bank
gorge of a stream flowing towards the north from Němetice
(ca. 1500 m SE from the Pod Doubravou settlement). These
red beds show no signs of bioturbation.

Milotice Formation

. It was studied in outcrops on both

banks  of  the  Bečva  River  downstream  from  its  confluence
with the Juhyně River at the Choryně village. Outcrops on the
left bank are currently deformed by landsliding and the section
cannot  be  fully  studied.  The  section  starts  with  a  sill  of
amygdule-rich lava of the teschenite association. The overly-
ing silty shales are baked, mostly dark grey in colour, with no
bioturbation.  Greyish-green  claystones  with  tabular  beds  of
sandstone  are  exposed  some  20 m  downstream.  The  clay-
stones are strongly compressed, which hampers the identifica-
tion of ichnofabrics. Ichnofabric in the proximity of sandstone

244

MIKULÁŠ, SKUPIEN, BUBÍK and VAŠÍČEK

Fig. 9. Supposed stratigraphy, lithology and ichnology of the Němetice (Cenomanian to Turonian) and Choryně (Coniacian to Santonian)
sections. Overall thickness of the depicted interval is hundred(s) of meters; not to scale.

beds is, however, distinct (“pressure shadow”): it can be char-
acterized as Planolites-dominated with ii = 2—3 (low-intensity
substrate  feeding,  probably  in  a  single  episode).  The  sand-
stone  beds  show  hypichnial  ?Palaeophycus  cf.  P.  sulcatus
(Fig. 10A)  and  more  rarely  Ophiomorpha  isp.,  which  indi-
cates episodes of higher physical energy of the environment.
Epichnial idiomorphic Chondrites isp. suggests an episode of
low physical energy after sand deposition.

Further up in the section (not further than 50 m from the

confluence), claystones with giant (

≥1 m in diameter) con-

cretions of micritic siderite ironstone are present. One of the
concretions  revealed  ichnofabric  (Fig. 10C)  composed  of  a
horizontal  tunnel  network  (Planolites/Megagrapton)  and  a
bundle of probable feeding tunnels (Phycodes isp.). The lat-
ter trace fossil is filled with glauconitic sandstone not docu-
mented  elsewhere  in  the  section.  This  is  followed  by  an
interval  that  is  strongly  deformed  by  landsliding,  with  first
continuous outcrops of Cretaceous rocks in situ present 100—
150 m  from  the  confluence.  These  outcrops  show  greyish-
black  clays  to  claystones  with  sporadic  thin  (a  few
centimeters)  tabular  beds  of  sandstone.  The  sandstone  bed
revealed a perfectly preserved hypichnial, radial structure of
the ichnogenus Gyrophyllites Glocker, 1841 (Fig. 10B). The
base  of  the  bed  also  contained  Palaeophycus  cf.  tubularis
Hall,  1847.  The  surrounding  shales  also  contain  sandstone-
filled  tunnels  of  Palaeophycus  isp.  or  Ophiomorpha  isp.
(difficult  to  distinguish  due  to  the  mode  of  preservation).
Such  an  assemblage  documents  that  this  part  of  the  section
probably originated under ecologically less restricted condi-
tions than the rest of the section, with more sophisticated and

long-lasting food exploitation of the substrate and rather per-
manent dwelling burrows of filtrators or predators. The re-
maining part of the section is hidden beneath ?colluvia with
blocks of algal limestone.

On the right bank, a more or less continuous outcrop starts

downstream of the weir on the Bečva River. It features grey
claystones and siltstones, occasionally with a greenish tinge,
containing very rare lenticular and tabular sandstone beds. It
can  probably  be  correlated  with  the  middle  part  of  the  out-
crop on the opposite bank. The ichnofabric is weak and colo-
nization horizons are generally several meters apart. The true
thickness  and  spacing  of  the  separate  colonization  horizons
mostly  cannot  be  determined  due  to  tectonic  deformation.
The  first  encountered  colonization  horizon  is  characterized
by  Planolites-dominated  ichnofabric  and  ii = 2—3  (substrate
feeding  of  low  intensity  and  low  efficiency,  probably  in  a
single episode). Several meters up in the section, fragments
of tabular sandstone beds and of sandy lenses can be found
with  sporadic  hypichnial  and  epichnial  Ophiomorpha  isp.
and Planolites isp. Greenish claystones with perfectly devel-
oped  colonization  horizon  of  the  ichnogenus  Chondrites
(Fig. 10E)  are  visible  another  several  meters  up  in  the  sec-
tion. After several tens of centimeters, these are followed by
a  sandstone  bed  with  a  suite  of  Planolites—Ophiomorpha  at
its  base.  Greyish-black  claystones  above  the  sandstone  bed
contain  a  ca.  2 cm  thick  interval  with  Planolites  cf.  monta-
nus  (ii = 3).  The  claystones  are  followed  by  a  long  non-bio-
turbated  interval  with  a  single  visible  red  bed  –  the  only
equivalent of the CORB found in this section. Some 130 m
downstream  from  the  weir,  a  small  but  notable  outcrop  of

246

MIKULÁŠ, SKUPIEN, BUBÍK and VAŠÍČEK

laminated siltstones contains a trace fossil of Phycosiphon
isp. (Fig. 10F; very small loop-shaped spreiten-structures),
that is traces of substrate feeding.

Silesian Unit, Kelč facies – a summary.

The onset of

CORB or their equivalents in this facies resulted in consider-
ably  restricted  conditions  for  the  development  of  benthic  or-
ganisms.  Colonization  horizons  with Chondrites  isp.,
Planolites/Ophiomorpha  and  Phycosiphon  indicate  short  in-
cursions of conditions favourable for infauna. Lower bedding
planes  of  rare  sandstone  intercalations  yielded  a  more  com-
plex assemblage of trace fossils (Gyrophyllites, Palaeophycus,
Phycodes,  Ophiomorpha)  showing  less  restricted  conditions
and more diversified feeding strategies.

Rača Unit

Unlike in the Silesian Unit, tectonic deformation and poor

outcrops did not allow the construction of a composite sec-
tion  for  the  Rača  Unit.  A  generalized  stratigraphic  chart  of
the  Upper  Cretaceous  in  the  Rača  Unit  (Fig. 11)  was  based
on biostratigraphic data from isolated outcrops and short sec-
tions. The CORB form the Kaumberg Formation as brown-
red  and  greenish-grey  shales  with  variable  silt  content.  The
Kaumberg  Formation  is  300—400 m  thick;  it  is  nearly  com-
pletely  non-calcareous  at  the  Smradlavá  site  (Fig. 1)  but  it
contains  numerous  calcareous  beds  ca.  20 km  west  of  this
site – at the Bučkový Stream site S of Horní Bečva. Nota-
bly,  the  sites  also  considerably  differ  in  their  ichnological
contents.

The shale-sandstone flysch overlying the CORB of the Rača

Unit is called the Soláň Formation, which contains several ho-
rizons of red-brown shales, reflecting short recovery episodes
of oligotrophic setting controlling the CORB formation.

Kaumberg Formation at the Smradlavá site.

At Smrad-

lavá, the lower part of the CORB shows the sequence of
grey-green and brown-red silty shales containing 0.5 to 4 cm

thick  hypoxic  black-grey  shale  horizons  with  rare  fish  re-
mains.  Fine-  to  medium-grained  sandstones  rarely  compose
thin-bedded  flysch.  Thick  banks  of  sandstones  and  slump
bodies  of  poorly  sorted  muddy  sandstones  occur  locally.
Rhythmic  thin-  to  thick-bedded  turbidite  sandstones  appear
in the upper part of the CORB. This turbidite member of the
formation contains rare grey to red calcareous mudstone ho-
rizons. Most of the CORB are not bioturbated. Weak coloni-
zation horizons marked by sparse Chondrites and Planolites
can  be  traced  at  intervals  of  approximately  2 m.  A  single
sandstone bed provided the ichnologic record of a short-time
colonization  by  sediment-feeders,  suspension-feeders  and
chemichnia  (ichnofossils  Planolites,  ?Trichichnus  and
?Arenicolites).

The succession of red, locally greenish claystones, occa-

sionally with secondary light grey colouration, is rather mo-
notonous from the ichnological point of view. Colonization
horizons  with  idiomorphic  Chondrites  isp.  are  developed
(Fig. 8H,J).  Horizons  with  very  weak  Planolites-dominated
bioturbation  are  also  present,  accentuated  by  stenomorphic
Chondrites isp. in the fillings of Planolites isp. (Fig. 8D,G).
The average spacing of the colonization horizons is 2 m but
any rhythmicity of the phenomenon is difficult to assess be-
cause  the  intensity  of  reworking  is  low  and  most  horizons
with  trace  fossils  cannot  be  identified  by  routine  collection
techniques.  It  is  a  very  restricted  environment  for  in-fauna
(oligotrophy), however, permitting episodic, highly econom-
ic life strategies with a low density of tracemakers. A bed of
laminated  sandstone  ca.  5 cm  thick  immediately  below  the
bed  marked  MB16C  (micropaleontological  sampling;  Sku-
pien et al. 2009) bears Planolites isp., ?Trichichnus isp. and
?Arenicolites isp. (Fig. 8F). The suite of these generally sim-
ple traces of opportunistic tracemakers and the low intensity
of  bioturbation  point  to  a  colonization  after  a  short  episode
of elevated physical energy (probably related to the deposi-
tion of the sandstone bed).

Fig. 11. Supposed stratigraphy, lithology and ichnology of the Smradlavá section. Overall thickness of the section is hundred(s) of meters;
not to scale.

248

MIKULÁŠ, SKUPIEN, BUBÍK and VAŠÍČEK

Kaumberg Formation at the Bučkový Stream site.

This

site exposes the upper part of the Kaumberg Formation and
the  lower  part  of  the  Soláň  Formation.  Exposure  of  the
former unit in the stream bed is incomplete, strata are mostly
overturned and monoclinally dipping. The tectonic deforma-
tion is relatively low, as revealed by tentative biostratigraph-
ic analyses. Reddish-brown shales are the prevailing (but not
absolutely  prevailing)  lithology.  With  very  rare  exceptions
(Fig. 12C),  they  mostly  lack  ichnofabrics  with  Chondrites
isp., characteristic for the preceding site. These shales are in-
tercalated  with  greenish  beds  (cm  thicknesses)  to  laminae
(mm thicknesses) containing large flakes of detrital mica and
sand with traces of shallow reworking (Fig. 12F). One of the
thin  green  laminae  yielded  minute  ?Phycosiphon  isp.  The
red  shales  themselves  show  indications  of  mottled  lustre  in
some  portions  (indicative  of  intensive  reworking  by  in-fau-
na).  Relatively  common  tabular  intercalations  of  sandstone,
centimeters to tens of centimeters in thickness, contain hyp-
ichnial  Arthrophycus,  Helminthopsis,  Thalassinoides  and
Planolites  (Fig. 12B)  and  bases  of  Diplocraterion.  Helmin-
thopsis  is  mostly  the  only  epichnion.  Thinner  intercalations
are richer in ichnological content: their deposition was prob-
ably not connected with such intensive erosion of the previ-
ous substrate.

Beds of strongly calcareous, light grey claystone

(Fig. 12D), several centimeters thick, appear some 10—15 m
below  the  onset  of  coarse-grained  rhythmic  flysch  of  the
Soláň Formation. They contain only sporadic “large” Chon-
drites isp. (Fig. 12H). The ambient red shales have a variable
but locally considerable proportion of carbonate, which can
be  seen  macroscopically  by  their  consistency  and  elevated
resistance  to  mechanical  weathering.  At  least  two  of  these
beds show intensive bioturbation: minute spreiten-structures
generally attributable to small forms of the ichnogenus Zoo-
phycos  by  the  presence  of  helicoidal  elements.  A  smaller
proportion  of  these  structures,  however,  show  elements  of
bilateral, “feather-like” spreite symmetry, thereby represent-
ing  “composites”  of  the  ichnogenera  Zoophycos  and  Lo-
phoctenium.

Soláň Formation.

The Soláň Formation, overlying the

Kaumberg Formation, is only poorly  exposed at the Smrad-
lavá site. The degree of its exposure at the Bučkový Stream
site is considerably higher. The streambed reveals thinly tab-
ular (first tens of centimeters in thickness) to thickly tabular
(maximum  0.6 m)  beds  of  coarse-grained  sandstone.  The
bases of these beds bear current marks and less common tun-
nels of deep tier trace fossils (Ophiomorpha, Thalassinoides)
exhumed by erosion. The proportion of hemipelagites is very
low.  They  are  green  or  grey  in  colour  and  are  either  com-
pletely bioturbated with no visualization of any ichnotaxa or
show the Thalassinoides—Planolites—Chondrites ichnofabric
(Fig. 12G) in which  the trace fossils are filled with colour-
contrasting material as the last stage of bioturbation.

Discussion

As has been stated above, ichnological study of the CORB

and the Tertiary oceanic red beds from a larger area with a

high facies variability has not been carried out yet. A partial
exception is the study of Leszczyński (1993) on Cretaceous
and Tertiary turbidite sequences in Spain with generally very
low-intensity  or  no  bioturbation  of  red  clays/claystones.
Other above-mentioned studies (i.e. Leszczyński & Uchman
1991,  1993;  Bąk  1995)  focus  on  less  variable  geological
units. A more general characteristic based on various small-
scale studies and unpublished observations was provided by
Wetzel & Uchman (1998a). These authors stated that red to
brown  claystones  accumulated  in  the  oceans  are  usually
characterized  by  complete  bioturbation;  the  number  of  tiers
is  limited  and  the  typical  depth  of  bioturbation  is  several
centimeters. An increase in the rate of sedimentation may re-
sult in a considerable increase in the food content, hence also
in  the  depth  of  burrow  penetration  and  in  the  diameter  of
tunnels and shafts.

The studied units provide examples confirming the validity

of both the above cited studies. A very low degree of bioturba-
tion is displayed by the CORB of the Godula facies of the
Silesian Unit, by their equivalents (mostly not red) in the Kelč
facies of the Silesian Unit, and by the CORB in non-calcare-
ous sediments of the Rača Unit. In contrast, a high degree of
bioturbation was observed in the CORB with calcareous inter-
calations in the Rača Unit: this facies provides an almost com-
plete list of ichnotaxa given for the CORB by Wetzel &
Uchman (1998a), namely Chondrites, Zoophycos, Planolites,
Thalassinoides, Palaeophycus, Teichichnus and Phycosiphon.

The above facts imply that the range of bioturbation of the

CORB may be extremely broad, with the supply of food obvi-
ously  acting  as  the  controlling  factor.  The  “carbonate-rich”
portion  of  the  CORB  of  the  Rača  Unit  has  a  considerably
higher  proportion  of  sand-dominated  interbeds  and  also  car-
bonates than the other described facies. This suggests a rela-
tively easy transport of nutrition-rich substrate into the basin
directly by turbidite currents, not only by periodical fall-out of
dead plankton.

The correlation between high diversity of ichnotaxa/strong

bioturbation/food  rich  environments,  however,  cannot  work
outside  a  narrow  range  of  parameters.  In  general,  eutrophy
favours  opportunistic  strategies;  trace  fossils  resulting  from
them  tend  to  be  present  with  low  diversity  and  high  abun-
dance.  Higher  productivity,  however,  may  increase  the
amount of organic particles on or in the sediment but also low-
er  oxygen  contents.  Considering  these  relations,  we  have  to
conclude  that  the  nutrition  richness  of  the  “carbonate-rich”
CORB  was  only  relative  in  comparison  with  the  “carbonate-
poor” CORB facies.

Conclusions

1. In the study area, CORB in the facies of reddish non-cal-

careous shales display a very low degree of bioturbation with
sparse  colonization  horizons  (mostly  with  Chondrites  as  the
only  trace  fossil,  less  often  with  the  Chondrites—Planolites
ichnofabric.  The  colonization  horizons  are  roughly  rhythmi-
cally distributed, several decimeters.

2. CORB facies displaying calcareous intercalations show a

very high degree of bioturbation as expressed by a high ichno-

249

ICHNOLOGY OF THE CRETACEOUS OCEANIC RED BEDS (CZECH REPUBLIC)

fabric index. They contain trace fossils Chondrites, Zoophy-
cos, Planolites, Thalassinoides, Palaeophycus, Teichichnus
and Phycosiphon.

3. The appearance of green layers in the carbonate-poor

CORB usually leads to the increasing of density of coloniza-
tion horizons and higher ichnofabric indices to few meters
from each other. The Chondrites—Planolites ichnofabric re-
mains the most frequent result of the colonization.

4. Lateral equivalents of CORB, namely monotonous green-

ish and grey shales, resulted in considerably restricted condi-
tions for the development of benthic organisms. Sparse
colonization horizons with Chondrites isp., Planolites/Ophio-
morpha and Phycosiphon indicate short incursions of condi-
tions more favourable for in-fauna.

Synopsis of ichnotaxa

The ichnotaxa responsible for the bioturbation of the CORB

and adjacent facies are listed and briefly characterized below.
With  the  notable  exception  of  Zoophycos—Lophoctenium
“composites”, which could be a valuable topic for the future,
the  material  does  not  bring  important  ichnotaxonomic  data,
therefore,  the  standard  systematic  ichnology  and  synonymy
are not given here.

Arenicolites isp. is a simple U-shaped burrow without a

spreite, oriented perpendicular to bedding. Arenicolites is
generally interpreted as dwelling burrow (e.g. Fillion &
Pickerill 1990).

Arthrophycus tenuis (Książkiewicz 1977) occurs gregari-

ously in the form of subhorizontal, hypichnial, convex ridg-
es. They are straight, usually transversely striated, rarely
branched and oriented in different directions (e.g. Uchman
1998).

Bergaueria is a large, typically very regular, hemispherical

pit; the “hemisphere” may represent only the bottom part of a
shallow vertical cylinder. Some burrows show simple radial or
concentric (central knob, rugged surface) ornamentation (e.g.
Häntzschel 1975).

Chondrites is a regularly branching tunnel system, typically

with radial arrangement of branchings at angles of 30—60°.
Three to four orders of branches can be observed. For a more
extensive discussion of the ichnogenus Chondrites see Fu
(1991) and Uchman (1999).

Gyrophyllites is a vertically or obliquely oriented burrow

that has a number of projections extending radially from the
central shaft. Gyrophyllites is interpreted as a feeding burrow
made by an animal that made repeated probes into the sedi-
ment in a radial pattern (e.g. Häntzschel 1975).

Helminthopsis is represented by smooth, unbranched, ir-

regularly winding, strictly horizontal burrows, interpreted as
repichnia and/or fodinichnia (e.g. Fillion & Pickerill 1990).

Megagrapton isp. is a system of usually lined tubular bur-

rows, branching at obtuse angles, thus forming scarce open
networks. For a detailed description of the morphology of the
ichnogenus see Książkiewicz (1977).

Nereites is a tightly to loosely meandering, rarely spirally

coiled endichnial trace fossil, typically 2—5 mm wide, com-
posed of a central, light – coloured faecal string, and a dark
envelope zone (e.g. Uchman 1995).

Ophiomorpha isp. is composed of cylindrical tubes lined

with pellets, forming deep, three-dimensional boxworks. Ver-
tical (at openings) and horizontal components prevail over ob-
lique ones (e.g. Häntzschel 1975).

Ophiomorpha annulata (Książkiewicz 1977) is an exich-

nial,  hypichnial  or  rarely  epichnial,  straight  to  slightly  wind-
ing,  vertical,  oblique  to  horizontal,  cylindrical,  walled  trace
fossil preserved in full-relief, 3—9 mm in diameter. It is filled
mostly with sand-sized material. In flysch deposits this trace
fossil  rarely  displays  a  wall  covered  with  small  oval  knobs,
which  are  characteristic  of  the  ichnogenus  Ophiomorpha
(Uchman 1995).

Paleophycus is an endichnial horizontal, unbranched to

non-systematically branched, lined cylindrical burrow. For
discussion of Palaeophycus see Pemberton & Frey (1982) and
Keighley & Pickerill (1995).

Phycodes is a horizontally to obliquely oriented burrow sys-

tem showing a “broomlike“ branching from a central burrow.
Phycodes is interpreted as a feeding trace made by repeated
probes by an animal into the sediment (e.g. Fillion & Pickerill
1990).

Planolites – variably oriented, but mostly horizontal, cy-

lindrical trace fossils without wall lining (Pemberton & Frey
1982; Keighley & Pickerill 1995).

Phycosiphon is observed in horizontal polished slabs as

curved endichnial lobes, 2 mm wide and up to 20 mm long,
encircled by a marginal tunnel, which is 0.7—1.0 mm wide. In
vertical cross-sections it has the character of patches of dark,
elongated spots, about 1 mm in diameter, surrounded by a
light mantle (e.g. Wetzel & Bromley 1994).

Thalassinoides is a walled or unwalled trace fossil com-

posed of wide cylindrical, mostly horizontal branched tunnels
(e.g. Ekdale 1992).

Trichichnus is a vertical to oblique, rarely horizontal,

straight to curved, simple or rarely branched, very thin, cylin-
drical trace fossil, typically less than 1 mm in diameter (Fil-
lion & Pickerill 1990; Uchman 1995, 1999).

Zoophycos is a concave funnel structure with radially ar-

ranged spreite. The spreite laminae are straight to slightly ar-
cuate. Zoophycos s.l. is generally assumed to be the trace of
an as yet undiscovered deposit-feeder. For discussion of this
ichnogenus see Ekdale (1992) and Bromley & Hanken
(2003). Very few finds of hybrids of Zoophycos and other
trace fossils have been reported; the Zoophycos-Lophocteni-
um hybrid structures as mentioned above deserve a more de-
tailed report.

Acknowledgments: The research was supported by the Czech
Science  Foundation  project  No. 205/05/0917  “Upper  Creta-
ceous oceanic red beds in the Czech part of the Outer Western
Carpathians;  biostratigraphy,  sedimentology,  geochemistry”.
The authors thank the reviewers of the paper: Alfred Uchman
(Cracow), Priska Schäfer (Kiel) and Jozef Michalík (Bratisla-
va)  contributed  by  valuable  comments;  however,  the  authors
are fully responsible to the final version of the paper. The au-
thors are grateful to Jiří Adamovič (Prague) for his assistance
with language presentation and to Martin Mazuch (Prague) for
his  technical  assistance  during  the  preparation  of  the  manu-
script. It is a contribution to the IGCP 463 Project.

250

MIKULÁŠ, SKUPIEN, BUBÍK and VAŠÍČEK

References

Bąk K. 1995: Trace fossils and ichnofabrics in the Upper Cretaceous

red deep-water marly deposits of the Pieniny Klippen Belt, Pol-
ish Carpathians. Ann. Soc. Geol. Polon. 64, 1—4, 81—97.

Bromley R.G. & Hanken N.M. 2003: Structure and function of large,

lobed Zoophycos, Pliocene of Rhodes, Greece. Palaeogeogr.
Palaeoclimatol. Palaeoecol. 192, 79—100.

Droser M.L. & Bottjer D.J. 1986: A semiquantitative field classifica-

tion of ichnofabric. J. Sed. Petrology 56, 558—559.

Ekdale A.A. 1992: Muckraking and mudslinging: the joys of deposit

feeding. In: Maples C.G. & West R.R. (Eds.): Trace Fossils. Pa-
leont. Soc. Short Course 5, 145—171.

Ekdale A.A., Bromley R.G., Bockelie J.F., Droser M.L. & Bottjer

D.J. 1991: “Ichnofabric” it is! Palaios 6, 1, 100—101.

Eliáš M. 1979: Facies and paleogeography of the Silesian unit in the

western part of the Czechoslovak flysch Carpathians. Věst. Ústř.
Úst. Geol. 54, 6, 327—339.

Fillion D. & Pickerill R.K. 1990: Ichnology of the Upper Cambrian?

to Lower Ordovician Bell Island and Wabana groups of Eastern
Newfoundland, Canada. Palaeontographica Canad. 7, 1—83.

Fu S. 1991: Funktion, Verhalten und Einteilung fucoider und lo-

phocteniider Lebensspuren. Cour. Forsch.-Inst. Senckenberg
135, 1—79.

Glocker F.E. 1841: Über die kalkführende Sandsteinformation auf

beiden Seiten der mittleren March, in der Gegend zwischen Kwas-
sitz und Kremsier. Academia Caesarea Leopoldino”Carolina
Germanica Naturae Curiosorum 19 (Suppl. 2), 309—334.

Häntzschel W. 1975: Treatise on Invertebrate Paleontology. Part W:

Miscelanea Supplement 1. Trace fossils and problematica. The
Geological Society of America, 1—269.

Hu X., Jansa L., Wang C., Sarti M., Bąk K., Wagreich M., Michalík

J. & Soták J. 2005: Upper Cretaceous oceanic red beds (CORB)
in the Tethys: occurrences, lithofacies, age and environments.
Cretaceous Research 26, 3—20.

Keighley D.G. & Pickerill R.K. 1995: The ichnotaxa Palaeophycus

and Planolites: historical perspectives and recommendations.
Ichnos 3, 301—309.

Książkiewicz M. 1977: Trace fossils in the flysch of the Polish Car-

pathians. Palaeont. Pol. 36, 1—228.

Leszczyński S. 1993: Ichnocoenosis versus sediment colour in Upper

Albian to lower Eocene turbidites, Guipúzcoa province, northern
Spain. Palaeogeogr. Palaeoclimatol. Palaeoecol. 100, 251—265.

Leszczyński S. & Uchman A. 1991: To the origin of variegated

shales. Geol. Carpathica 42, 5, 279—289.

Leszczyński S. & Uchman A. 1993: Biogenic structures of organic-

poor sediments: examples from the Paleogene variegated shales,
Polish Outer Carpathians. Ichnos 2, 267—275.

Menčík E., Adamová M., Dvořák J., Dudek A., Jetel J., Jurková A.,

Hanzlíková E., Houša V., Peslová H., Rybářová L., Šmíd B.,
Šebesta J., Tyráček J. & Vašíček Z. 1983: Geology of Morav-
skoslezské Beskydy Mts and Podbeskydská pahorkatina Up-
lands. Ústř. Úst. Geol., 1—304 (in Czech).

Michalík J. 1994: Notes on the paleogeography and paleotectonics of

the Western Carpathian area during the Mesozoic. Mitt. Österr.
Geol. Ges. 86, 101—110.

Mišík M. 1992: Pieniny Klippen Belt in relationship with Mesozoic

and Tertiary volcanism. Západ. Karpaty, Geol. 16, 47—64.

Pemberton S.G. & Frey R.W. 1982: Trace fossil nomenclature and

the Planolites—Palaeophychus dilemma. J. Paleontology 56,
843—881.

Potter P.E., Maynard B.J. & Pryor W.A. 1980: Sedimentology of

Shale. Study guide and reference source. Springer, New York,
1—310.

Skupien P. & Vašíček Z. 2003: Lithostratigraphical and biostrati-

graphical knowledge of the Bystrý potok section by Frenštát pod
Radhoštěm (Upper Cretaceous, Silesian Unit of the Outer West-
ern Carpathians). Trans. VŠB – Techn. Univ. Ostrava, Mining
and Geol. Ser. 8, 64—94 (in Czech).

Skupien P., Bubík M., Švábenická L., Mikuláš R., Vašíček Z. &

Matýsek D. 2009: Cretaceous Oceanic Red Beds in the Outer
Western Carpathians of Czech Republic. Final Volume IGCP
463, SEPM. (in print).

Uchman A. 1995: Taxonomy and palaeoecology of flysch trace fos-

sils: The Marnoso-arenacea Formation and associated facies
(Miocene, Northern Apennines, Italy). Beringeria 15, 3—115.

Uchman A. 1998: Taxonomy and ethology of fysch trace fossils: re-

vision of the Marian Ksiazskiewicz collection and studies of
complementary material. Ann. Soc. Geol. Pol. 68, 105—218.

Uchman A. 1999: Ichnology of the Rhenodanubian Flysch (Lower

Cretaceous-Eocene) in Austria and Germany. Beringeria 25,
67—173.

Wetzel A. & Bromley R.G. 1994: Phycosiphon incertum revisited:

Anconichnus horizontalis is its junior subjective synonym. J.
Paleontology 68, 1396—1402.

Wetzel A. & Uchman A. 1998a: Biogenic sedimentary structures in

mudstones – an overview. In: Schieber J., Zimmerle W. & Set-
hi P. (Eds.): Shales and mudstones. I. E. Schweizerbart’sche
Verlag (Nägele u. Obermiller), Stuttgart, 351—369.

Wetzel A. & Uchman A. 1998b: Deep-sea benthic food content re-

corded by ichnofabrics: a conceptual model based on observa-
tions from Paleogene flysch, Carpathians, Poland. Palaios 13,
533—546.

Wetzel A. & Uchman A. 2001: Sequential colonization of muddy tur-

bidites: examples from Eocene Beloveža Formation, Car-
pathians, Poland. Palaeogeogr. Palaeoclimatol. Palaeoecol.
168, 1—2, 171—186.