GEOLOGICA CARPATHICA, OCTOBER 2008, 59, 5, 395—409
www.geologicacarpathica.sk
New methods for ichnofabric analysis and correlation with
orbital cycles exemplified by the Baden-Sooss section
(Middle Miocene, Vienna Basin)
PETER PERVESLER
1
, ALFRED UCHMAN
2
and JOHANN HOHENEGGER
1
1
Department of Paleontology, University of Vienna, Althanstrasse 14,
1090 Vienna, Austria;
peter.pervesler@univie.ac.at; johann.hohenegger@univie.ac.at
2
Institute of Geological Sciences, Jagiellonian University, Oleandry 2a, 30-063 Kraków, Poland; alfred.uchman@uj.edu.pl
(Manuscript received December 13, 2007; accepted in revised form June 12, 2008)
Abstract: A two step cluster analysis based on log-likelihood measures for categorial variables using ‘Schwarz’s Bayesian
Criterion’ for grouping allows the automatic detection of ichnofabric categories from a large data set. Preferred succes-
sions of these ichnofabrics were tested by ‘Embedded Markov Chains’. This leads to the following ichnofacies interpreta-
tion: Alternating periods of higher/lower accumulation rates with higher/lower inputs of particulate food and higher/
lower oxygen contents in pore waters led to sequential colonization of the substrate. The trace fossils Phycosiphon and
Nereites represent opportunistic colonization of oxygenated sediments rich in particulate organic matter (POM) by de-
posit-feeding animals, quickly after an increased sediment input. A further stage of colonization caused by the decrease of
POM induced by consumption and oxidation forced the animals to search for food on sediment surfaces and from the water
column. The open burrows Thalassinoides, Chondrites, Trichichnus and Zoophycos indicate stable-bottom conditions in
periods of low accumulation rates. Zoophycos, Phycosiphon, Nereites and Teichichnus suggest the Zoophycos ichnofacies
for the lower section of the core; a transition to the distal part of the Cruziana ichnofacies is suggested for the upper section
of the core with the appearance of Thalassinoides. The changes between stable and unstable bottom conditions signifi-
cantly correlate with periods in magnetic susceptibility and calcium carbonate content, both forced by orbital cycles.
Key words: Miocene, Badenian, Vienna Basin, statistical analysis, orbital cycles, trace fossils, ichnofabrics.
Introduction
Trace fossil and ichnofabric analysis is a powerful tool in the
recognition of ecological parameters such as energy level, oxy-
gen content, food supply, salinity or stability of the environ-
ment. Several ichnological researches refer to the Badenian of
the Central Paratethys (e.g. Abel 1928; Kühnelt 1931; Klee-
mann 1982; Hohenegger & Pervesler 1985; Pervesler & Uch-
man 2004; Pervesler & Zuschin 2004), but almost all of them
concern littoral sandy sediments or rocky shores.
By drilling close to the Badenian stratotype (Middle Mi-
ocene) near the western margin of the southern Vienna Ba-
sin, a continuous 102 m section of mostly fine-grained
sublittoral Badenian deposits was obtained. Biostratigraphy,
paleoecology, sedimentology, geochemistry, magneto-
stratigraphy and magnetic climate proxies such as magnetic
susceptibility (Khatun et al. 2006; Hohenegger et al. 2008)
of the core were studied in FWF-Project P13743 – BIO.
The main aim of this paper is to present the results of the ich-
nological analysis.
Geological setting
The drill site is situated near the western margin of the
southern Vienna Basin (Fig. 1). The basin formed during the
Neogene lateral extrusion within the Eastern Alps (Ratsch-
bacher et al. 1991), is situated at the junction of the Eastern
Alps and the Western Carpathians (e.g. Decker 1996; Hamil-
ton et al. 2000). The scientific borehole at Baden-Sooss pene-
trated a succession of Badenian (Langhian, Middle Miocene)
deposits, below the type section of the Badenian stage, the old
Baden-Sooss brickyard near Baden (Papp et al. 1978). The
“Badener Tegel” is placed into the Baden Group which is sub-
divided into the Jakubov Formation and the Lanžhot Forma-
tion (e.g. Kováč et al. 2004). The part of the Badenian within
the Baden-Sooss borehole can be correlated to the Lanžhot
Formation.
Material and methods
Preparation and documentation
After the core was split longitudinally and the cross-section
was smoothed, it was scanned and photographed digitally.
The image series was used for ichnological analyses. Trace
fossils were detected from 8 to 102 meters of the core depth.
Additional cuts were made parallel to bedding. The contrast
obtained by moistening the core sections was further im-
proved by graphic software elaboration of the images.
Statistical methods
The core was divided into 25 cm intervals and ichnofossils
were determined for each interval as present/absent, resulting
396
PERVESLER, UCHMAN and HOHENEGGER
in a data matrix of 376 samples by 12 ichnospecies. Lamina-
tion was included as a further qualitative character. Samples
were clustered using the ‘Two Step Cluster Analysis’ de-
signed to handle very large data sets (SPSS 2006; Zhang et al.
1996). Log-likelihood measures appropriate to categorical
variables functioned as distances between samples.
‘Schwarz’s Bayesian Criterion’ was used for cluster finding
with an automatic determination of cluster numbers (SPSS
2006).
Preferred successions of clusters were tested by ‘Embedded
Markov Chains’ (Davis 2002), thus excluding self-transforma-
tions. General tendencies in ichnofabric composition along the
core were shown in percentages of ichnofabric types calculat-
ed over an interval of 3 meters with intervals moving in 25 cm
steps (‘moving percentages’).
Finally, correlations of each ichnospecies with quantitative
environmental data were tested using the ‘Point-Biserial Cor-
relation’ (Gibbons 1976). Statistical analyses were performed
with EXCEL (for Markov Chains) and the program packages
SPSS 15.0 and PC-ORD (McCune & Mefford 1999) for com-
plex analyses.
Systematic ichnology
Except for several layers with primary lamination, the core
is completely bioturbated.
Trace fossils of the ichnogenera Asterosoma, Chondrites,
Nereites, Ophiomorpha, Palaeophycus, Phycosiphon, Scoli-
cia, Siphonichnus, Teichichnus, Thalassinoides, Trichichnus
and Zoophycos were distinguished in cross-sections and occa-
sionally on surfaces parallel to bedding. Their distribution in
the core is shown in Fig. 2.
Asterosoma von Otto, 1854
Asterosoma isp.
Fig. 3D
D e s c r i p t i o n : In cross-section, clusters of variably ori-
ented ovals, with faint concentric lamination around a central
lumen. The ovals are 7—18 mm wide and 15—45 mm long. Lo-
cally, the lumen is oval, 5 mm in diameter, and in some cases
filled with slightly coarser and darker sediment than in sur-
rounding laminae. In many cases the centre is poorly outlined
and seen as a dark dot.
R e m a r k s : The ovals are cross-sections of vertical to in-
clined elongated bulbs tapering at both ends, with concentric
internal lamination. Clusters of such bulbs form tree-like
structures spread out from a common vertical or inclined
shaft. The morphology of such cross-sections is typical of As-
terosoma (e.g. Bromley & Uchman 2003; Pervesler & Uch-
man 2004). Asterosoma is interpreted as a selective-feeding
burrow of a worm (Pemberton et al. 2001). It occurs in soft
(mostly siliciclastic, rarely carbonate) substrates (e.g. Gibert
1996), typically in various shallow-marine settings, especially
in the upper lower shoreface (Pemberton et al. 2001).
Chondrites von Sternberg, 1833
Chondrites isp.
Fig. 3A
D e s c r i p t i o n : In cross-section, clusters of straight, locally
branched, light bars and dots, 1.5—2 mm wide.
R e m a r k s : The described cross-section morphology is
typical of Chondrites (compare e.g. Werner & Wetzel 1981;
Ekdale & Bromley 1991; Wetzel & Uchman 1998), which in
three dimensions is a branched tunnel system ramifying at
Fig. 1. Location map.
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ICHNOFABRIC ANALYSIS AND CORRELATION WITH ORBITAL CYCLES (MIDDLE MIOCENE, VIENNA BASIN)
Fig. 2. Section of the core, distribution of ichnotaxa and ichnofabrics.
398
PERVESLER, UCHMAN and HOHENEGGER
depth from a master shaft in a dendritic manner. Chondrites is
interpreted as a chemosymbiotic feeding structure (Fu 1991;
Uchman 1999, and references therein).
Nereites MacLeay, 1839
Nereites isp.
Fig. 3C, E—H
D e s c r i p t i o n : Horizontal to subhorizontal, rarely oblique,
winding subcylindrical structures seen rarely on surfaces of
parted rocks parallel to the bedding as winding or loosely me-
andering dark ribbons bounded by lighter side zones. The rib-
bon is 3.5—4.0 mm wide and the bounding zone is 1.0—1.5 mm
wide. Locally, the ribbon displays scalariform menisci. Much
more commonly this trace fossil is seen in cross-section,
where it appears as clusters of elongated dark, oval spots sur-
rounded by a lighter halo (Fig. 3C,E,F,G). Some of the dark
spots are asymmetric and pointed from one side. They are
1.2—2.0 mm high and mostly 3—10 mm wide. Some of them
are more elongated as bars up to 30 mm long, with local me-
niscate structure. The lighter halo is 1.0—2.5 mm wide. The
clusters are commonly as much as 60 mm wide and as much
as 40 mm high.
R e m a r k s : The ribbons, visible in cross-section as dark
spots, are faecal strings having locally meniscate filling. The
bounded lighter zones seen in cross-section as the light halo
represents a reworking zone around the faecal string. These
features are typical of Nereites, which is interpreted as a pasc-
ichnion made just above the redox boundary (Wetzel 2002).
For discussion of this ichnogenus see Chamberlain (1971), Ben-
ton (1982), Uchman (1995, 1999) and Mángano et al. (2002).
The described trace fossil, by its relatively narrow reworking
zone and local scalariform menisci, resembles Nereites missou-
riensis (Weller 1899) (see also Conkin & Conkin 1968).
Ophiomorpha Lundgren, 1891
Ophiomorpha isp.
D e s c r i p t i o n : Vertical or oblique, curved tubes, whose
lumen diameter ranges from 5 to 10 mm, and whose wall
thickness ranges from 3 to 6 mm. The lumen diameter and
thickness of the wall are more or less constant in each speci-
men. The wall is built of a material that is slightly lighter than
the host rock. The outer margin of the wall is poorly expressed
or slightly lobate in cross-section.
R e m a r k s : Size, orientation and thick wall suggest that
this trace fossil belongs to Ophiomorpha. The lobate outer
margin of the wall corresponds to the knobby wall exterior
typical of this ichnogenus. Ophiomorpha nodosa is one of the
most common shallow-marine trace fossils and is produced
mostly by thalassinoid shrimps (Frey et al. 1978; Ekdale
1992). It is most typical of the Skolithos ichnofacies (Frey &
Seilacher 1980; Pemberton et al. 2001), but also occurs in
deeper shelf tempestites (Frey 1990; Frey & Goldring 1992).
Palaeophycus Hall, 1847
Palaeophycus tubularis Hall, 1847
D e s c r i p t i o n : Horizontal to oblique simple tubes, 3—5 mm
in diameter, with a thin, light wall.
R e m a r k s : Palaeophycus tubularis is a facies-crossing
form produced by carnivorous or omnivorous animals, mostly
polychaetes (Pemberton & Frey 1982). For discussion of
Palaeophycus see also Keighley & Pickerill (1995).
Phycosiphon Fischer-Ooster, 1858
Phycosiphon incertum Fischer-Ooster, 1858
Fig. 3A,C,E—H
D e s c r i p t i o n : In cross-section, clusters of curved dark
bars and spots surrounded by lighter halo. The dark spots and
bars are up to 1mm thick and the bars are up to 6 mm long.
Most of the bars and spots are parallel or sub-parallel to the
bedding. The clusters are commonly as much as 35 mm wide
and as much as 25 mm high. This trace fossil commonly oc-
curs in the filling of larger burrows.
R e m a r k s : The described structures are typical of poorly
expressed Phycosiphon incertum Fischer-Ooster (Wetzel &
Bromley 1994). It is seen on bedding planes as horizontal,
curved small repeated lobes encircled by thin marginal tun-
nels. Phycosiphon incertum is a feeding structure (fodinich-
nion) (e.g. Ekdale & Mason 1988).
Scolicia de Quatrefages, 1849
Scolicia isp.
Fig. 3C
D e s c r i p t i o n : Horizontal subcylindrical structures with
a complex meniscate backfill. In cross-section, they are seen
as oval structures with slightly concave top and concave bot-
tom. The bottom concavity is bounded by two oval protru-
sions, which are about 5 mm in diameter. The structures are
20—35 mm high and 35—75 mm wide. In some cross-sections,
the structures are dissected obliquely or along their course and
in such cases the meniscate backfill is highly visible.
R e m a r k s : The oval protrusions at the bottom are cross-
sections of basal strings (see Uchman 1995). Scolicia is a fos-
sil locomotion and feeding structure (pascichnion) produced
by irregular echinoids (e.g. Bromley & Asgaard 1975; Smith
& Crimes 1983).
Siphonichnus Stanistreet, Le Blanc Smith & Cadle, 1980
Siphonichnus isp.
D e s c r i p t i o n : Siphonichnus is a steeply oblique struc-
ture, in the studied section about 9 mm wide and at least
107 mm long. It displays a thin margin, concave-down menis-
ci and a central, straight vertical shaft crossing the menisci.
The shaft 1 mm wide is distinctly darker than the surrounding
sediment.
R e m a r k s : Siphonichnus is interpreted as a bivalve bur-
rowing structure, where menisci related to the burrowing ac-
tion are crosscut by the shaft produced by the siphon
(Stanistreet et al. 1980).
Teichichnus Seilacher, 1955
Teichichnus isp.
Fig. 3B
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ICHNOFABRIC ANALYSIS AND CORRELATION WITH ORBITAL CYCLES (MIDDLE MIOCENE, VIENNA BASIN)
Fig. 3. Trace fossils from the Baden-Sooss core. A – Ichnofabric 4 with Phycosiphon (Ph) and primary lamination (Lam) penetrated by
Chondrites (Ch). Meter 87.14—87.24. B – Ichnofabric 5_4 with Teichichnus (Te). Meter 26.43—26.53. C – Ichnofabric 3 with Phycosi-
phon (Ph), Nereites (Ne), Scolicia (Sc) and Chondrites (Ch). Meter 42.26—42.35. D – Ichnofabric 3 with Asterosoma (As). Meter 86.56—
86.67. E – Ichnofabric 6 with Phycosiphon (Ph) and Nereites (N). Meter 95.47—95.57. F – Ichnofabric 1 with Nereites (Ne) and Thalassi-
noides (Th) filled with Phycosiphon (Ph). Meter 46.20—46.29. G – Ichnofabric 2 with Phycosiphon (Ph), and Nereites (Ne) cut by
Zoophycos (Zo). Meter 96.44—96.54. H – Ichnofabric 2 with Phycosiphon (Ph), Nereites (Ne) and Trichichnus (Tr). Meter 85.83—85.93.
400
PERVESLER, UCHMAN and HOHENEGGER
D e s c r i p t i o n : Oblique or vertical, structure without wall
consisting of dense, parallel or sub-parallel convex down spre-
ite. The structure is about 100 mm high and about 50 mm
wide, however the width can easily be overestimated because
this trace fossil is observed in oblique cross-section. Side mar-
gins of the structure are uneven. The causative burrow at the
top shows concentric lamination (Fig. 3B).
D i s c u s s i o n : Teichichnus is a typical feeding structure.
For discussion of this ichnogenus see e.g. Fillion & Pickerill
(1990) and Schlirf (2000).
Thalassinoides Ehrenberg, 1944
Thalassinoides isp.
Fig. 3F
D e s c r i p t i o n : In cross-section, variably oriented, branch-
ing 6—18 mm wide bars and spots. They are filled with sand
from the overlying bed and are surrounded by mudstone and
siltstone. They represent a system of a boxwork burrow sys-
tem without wall. Density of the burrows increases in proxim-
ity of the overlying sand bed. The lowest part of the system is
located 165 mm below the base of the sand bed.
R e m a r k s: Thalassinoides is characterized as a system of
tunnels and shafts produced by crustaceans, mostly decapods
in many marine environments (Fürsich 1973; Frey et al. 1984;
Ekdale 1992; Bromley 1996; Schlirf 2000).
Trichichnus Frey, 1970
?Trichichnus linearis Frey, 1970
Fig. 3H
D e s c r i p t i o n : Variably oriented, very thin (sub-millimet-
ric), rarely branched cylinders. They are filled with ferrugi-
nous material and commonly surrounded by a yellowish halo.
R e m a r k s : Trichichnus occurs mostly in fine-grained,
shallow-water deposits (e.g. Frey 1970) as well as deep-sea
deposits (e.g. Kennedy 1975; Wetzel 1981). A strong tenden-
cy to pyritization is typical of this form (e.g. Werner & Wetzel
1981). It is a deep-tier trace fossil produced by opportunistic
organisms in poorly oxygenated sediments (McBride & Pi-
card 1991), which may belong to the chemosymbiotic meio-
benthos (Uchman 1999).
Zoophycos Massalongo, 1855
Zoophycos isp.
Fig. 3G
D e s c r i p t i o n : Spreite structures seen in core cross-sec-
tions as horizontal or oblique, rarely steeply inclined stripes,
filled with spreites, 4—9 mm thick, which in cross-section look
like densely packed menisci. In oblique cross-sections, the
spreite laminae can be seen (Fig. 3G). They contain very
small, sub-millimetric pellets. In some specimens the stripes
converge in the axial part, where they are strongly wrapped up
forming inverted V-structures.
R e m a r k s : Zoophycos s.l. is generally regarded as a struc-
ture produced by some as-yet undiscovered deposit-feeder,
which has been referred to sipunculids (Wetzel & Werner
1981), polychaete annelids, arthropods (Ekdale & Lewis
1991), and echiurans (Kotake 1992). The feeding strategy is,
however, controversial (e.g. Bromley 1991; Locklair & Savr-
da 1998; MacEachern & Burton 2000). Bromley & Hanken
(2003) suggested that the upper helical part of a large Pliocene
Zoophycos from Rhodes, Greece, is a deposit-feeding struc-
ture, and lateral lobes developing from its lower part are sul-
phide wells for chemosymbiotic bacteria. The same interpreta-
tion refers to a similar but smaller Zoophycos from the
Miocene of Austria (Grund Formation), which displays very
steep lobes in its lowermost part (Pervesler & Uchman 2004).
Results
‘Ichnofabric’ types
The cluster analysis of ichnofabrics resulted in seven auto-
matically determined groups, which can be interpreted as ‘ich-
nofabric’ types. Six clusters are homogeneous in their
composition possessing one or two dominant species, while
the seventh cluster (Type 5) is heterogeneous, consisting ei-
ther of a single species that is not found or underrepresented in
the other types, or none.
Type 1
All samples are characterized by the concurrence of
Thalassinoides (in all samples of the cluster = 100 %) and Phy-
cosiphon (in all samples of the cluster = 100 %) with an impor-
tant proportion of Nereites (in 61 % of the samples within the
cluster; Table 1).
Type 2
This is the most heterogeneous group, where Phycosiphon
again is represented in all samples. The major concurrent spe-
cies are Trichichnus (72 %) and Nereites (65 %), followed by
minor proportions of Zoophycos (26 %), Thalassinoides
(14 %) and Ophiomorpha (12 %).
Type 3
This is similar to the former with identical proportions of
Phycosiphon (100 %) and Nereites (65 %). Trichichnus is less
important (10 %), while its position as the second important
species is overtaken by Scolicia (92 %). Trichichnus, Astero-
soma (both 10%), Chondrites (8 %) and Thalassinoides (6 %)
are rare; Zoophycos, Siphonichnus and Teichichnus have ex-
tremely low proportions (2 % each). The relatively high per-
centage of lamination within this type (24 %) demonstrates
the close relationship to the following ichnofabric type.
Type 4
This is characterized by the predominance of lamination
(100 %) in combination with a high proportion of Phycosiphon
(92 %). Nereites (18 %) and Thalassinoides (8 %) are of minor
importance and a very few Chondrites, Zoophycos, Siphonich-
nus, Trichichnus and Asterosoma (all 3 %) can be found.
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ICHNOFABRIC ANALYSIS AND CORRELATION WITH ORBITAL CYCLES (MIDDLE MIOCENE, VIENNA BASIN)
Table 1: Ecological interpretation of ichnofabric types.
Type 5
The 5
th
group is heterogeneous according to the low number
or lack of ichnospecies and thus must be divided into several
homogeneous sub-types:
Type 5_1: Although bioturbated, no distinct ichnofossil
could be identified in this ichnofabric type.
Type 5_2: Only Nereites (100 %) is represented in this type.
Type 5_3: Nereites (100 %) and Thalassinoides (100 %) are
the concurrent ichnofossils.
Type 5_4: Teichichnus is the single ichnofossil here.
Type 5_5: Only Thalassinoides can be found.
Type 6
Nereites (100 %) is combined with Phycosiphon (100 %,
Table 1).
Ichnofabric
type
Presence of trace fossils and
lamination in the measured
intervals
Opportunistic
colonization
(number of
ichnotaxa)
Stable colonization
(number of
ichnotaxa)
Remarks
Type 1
Thalassinoides
Phycosiphon
Nereites
100 %
100 %
61 %
2
1
opportunistic colonization, followed
by stabilization phase
Type 2
Phycosiphon
Trichichnus
Nereites
Zoophycos
Thalassinoides
Ophiomorpha
100 %
72 %
65 %
26 %
14 %
12 %
3
3
high organic matter content
Type 3
Phycosiphon
Scolicia
Nereites
Trichichnus
Asterosoma
Thalassinoides
Zoophycos
Siphonichnus
Teichichnus
lamination
100 %
92 %
65 %
10 %
10 %
6 %
2 %
2 %
2 %
24 %
3
5
high sediment input
Type 4
Phycosiphon
Nereites
Thalassinoides
Chondrites
Zoophycos
Siphonichnus
Trichichnus
Asterosoma
lamination
92 %
18 %
8 %
3 %
3 %
3 %
3 %
3 %
100 %
3
6
less food
Type 5_1
bioturbated 100
%
shallow
bioturbation
Type 5_2
Nereites 100
%
1
Type 5_3
Nereites
Thalassinoides
100 %
100 %
1 1
Type 5_4
Teichichnus
lowered
salinity?
Type 5_5
Thalassinoides
discontinuity
surface
Type 6
Phycosiphon
Nereites
100 %
100 %
2
only opportunistic colonization
Type 7
Phycosiphon
Teichichnus
100 %
3 %
1
opportunistic colonization, less food
Type 7
Phycosiphon dominates (100 %), accompanied by very few
Teichichnus (3 %).
Regarding proportions of ichnofabric types in the core,
Type 6 (Nereites and Phycosiphon) is the most abundant
(24.7 %) followed by Type 7 (19.7 %). The remaining types
show similar proportions from 10.4 to 13 %, while the insig-
nificant group 5 with different singular or lacking major ich-
nofossils (8.5 %) is less important (Fig. 4).
Figure 4 demonstrates the importance of ichnofossils to ich-
nofabric types. While Phycosiphon is characteristic for all
types (except the heterogeneous group 5), Nereites and
Thalassinoides follow with decreasing proportions (Fig. 4).
Scolicia is a marker species for Type 3, while Trichichnus is
abundant in Type 2 and less important for Types 3 and 4. All
remaining ichnofossils are of low importance in all ichnofab-
402
PERVESLER, UCHMAN and HOHENEGGER
ric types. Lamination signalizing lack of bioturbation is char-
acteristic for Type 4, being only of minor importance in
Type 3 and rare in Type 2.
Transition between ichnofabric types
Transitions from one into another ichnofabric type could be
random, then characteristic for rather stable environments
with minor alterations. Otherwise, preferred transitions dem-
onstrate significant reactions to monotonously or periodically
changing environments. Proving these transitions by embed-
ded Markov Chains resulted in significant transitions at the
5% error probability level (Table 2). The representation of
Fig. 4. Percentages of ichnofabric types in the core (pie-diagram) and percentages of ichnogenera in these ichnofabric types (bar-diagram).
transitions between ichnofabric types and their intensities are
represented as a directed graph (Fig. 5). Within this figure,
preferred transition from the heterogeneous Type 2 to the ho-
mogeneous Type 6 that combines dominating Phycosiphon
and Nereites is more frequent (46 % of all transitions starting
from Type 2) than the reverse transition (only 9 % of transi-
tions starting from Type 6). Similar preferred transitions can
be found from Type 4 (with predominant laminations) to
Type 7 (23 % of transitions versus 7 % reverse transitions)
and in weaker form to Type 3 (31 % of transitions versus
14 % reverse transitions). Similar types of transitions, where
the one direction is of a two-fold intensity than the opposite
can be found in relations from Type 3 to Type 6 (25 % of tran-
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ICHNOFABRIC ANALYSIS AND CORRELATION WITH ORBITAL CYCLES (MIDDLE MIOCENE, VIENNA BASIN)
sitions versus 12 % reverse transitions)
and from Type 3 to Type 7 (34 % of tran-
sitions versus 17 % reverse transitions).
The closest transitions without preferred
directions are between Types 6 and 7
(24 % versus 26 %).
Ichnofabric Type 5 must be separately
treated and is thus excluded from this
analysis, because it represents an inhomo-
geneous group. Therefore, normal Mark-
ov Chains were calculated, since self-
transformations within the 25 cm intervals
are rare in these subtypes (Table 3). Char-
acteristic transitions are from Subtype 5_1
(no visible macro-burrows) to Type 7
(dominated by Phycosiphon) with 30 %
forward versus 2 % reverse transitions,
and from Subtype 5_3 (Nereites and
Thalassinoides exclusively) and Sub-
type 5_5
(Thalassinoides
solely)
to
Type 1 (Thalassinoides and Phycosi-
phon), both with 33 % versus 5 % reverse
transitions. Both transitions are of similar
intensity between Subtypes 5_2 (only
Nereites) and 5_3 (16.7 % in both direc-
tions), while no visible burrows (Sub-
type 5_1) mainly overlay Subtypes 5_3
(16.7 %) and 5_2 (8 %).
Embedded Markov Chains allow the
detection of preferred transitions starting
from non-bioturbated (laminated) core
segments (Fig. 6). After the period with
laminated (undisturbed) sediments, the
Phycosiphon as a pioneer species reaches
a high proportion, accompanied by a few
Nereites. Preferred transitions from
Type 4 to Type 3 (probability 0.32) that
shows less lamination and the addition of
abundant Scolicia follow this pioneer
stage. Type 7 with dominance of Phycosi-
phon can also be directly derived from
Type 4 (probability 0.24), while transi-
tions to Type 6 (Phycosiphon and Nere-
ites) are less important (probability 0.16).
Type 1 (Thalassinoides additional to
both former species) mainly derives from
Type 4 (probability 0.12) and from
Type 6 (probability 0.17; Figs. 5, 6).
Type 3 as the main derivative from the
laminated type mostly transforms to
Type 7 (probability 0.34) and Type 6
(probability 0.25), with additional re-
verse transitions to the laminated Type 4
(probability 0.14) or to the most hetero-
geneous Type 2 (probability 0.11). It is
important that the latter as the ‘climax’
community type mainly derives from
Type 3, although Scolicia is completely
lacking here.
Fig. 5. Transitions between ichnofabric types represented as directed graphs.
Fig. 6. Transition probabilities between ichnofabric types.
404
PERVESLER, UCHMAN and HOHENEGGER
Type 7 and Type 6 shows the closest connections as men-
tioned before with transition probabilities of 0.26 from
Types 7 to 6 and 0.27 for the reverse. Transitions to Type 3
are more abundant from Type 7 (probability 0.17) than from
Type 6 (probability 0.12) confirming the non-directed transi-
tions between Types 3, 6 and 7.
Types 1 and 2 are not easily derived from the other types.
After verification, Type 1 intensively changes to Type 6
(probability 0.22), Type 3 (probability 0.16) and Type 7
(probability 0.22). The most intensive transitions are from
Type 2 to Type 6 (probability 0.46) meaning a restriction of
the heterogeneous ichnofossils of Type 2 to Phycosiphon and
Nereites in Type 6. Further important transitions are from
Type 5.1 Type 5.2 Type 5.3 Type 5.4 Type 5.5
Type 1
Type 2
Type 4
Type 6
Type 7
sum
Type 5.1
5
1
1
3
10
Type 5.2
1
6
2
1
1
1
12
Type 5.3
1
1
2
2
6
Type 5.4
1
1
2
Type 5.5
1
2
3
Type 1
1
2
2
17
2
3
7
6
40
Type 2
2
23
10
4
40
Type 4
1
3
2
17
4
6
33
Type 6
1
2
11
6
6
42
16
84
Type 7
1
2
1
1
1
6
5
5
19
23
64
sum
11 12 7 3 3 43 38 32 83 62 294
Type 5.1 Type 5.2 Type 5.3 Type 5.4 Type 5.5
Type 1
Type 2
Type 4
Type 6
Type 7
sum
Type 5.1
0.500
0.100
0.100 0.300
1.0
Type 5.2
0.083
0.500
0.167
0.083
0.083
0.083
1.0
Type 5.3
0.167
0.167
0.333
0.333
1.0
Type 5.4
0.500
0.000
0.500
1.0
Type 5.5
0.333
0.667
1.0
Type 1
0.025 0.050
0.050 0.425 0.050 0.075 0.175 0.150
1.0
Type 2
0.025
0.050
0.575
0.000
0.250
0.100
1.0
Type 4
0.030
0.091
0.061
0.515
0.121
0.182
1.0
Type 6
0.012
0.024
0.131
0.071
0.071
0.500
0.190
1.0
Type 7
0.016 0.031 0.016 0.016 0.016 0.094 0.078 0.078 0.297 0.359
1.0
sum
0.833 0.747 0.566 0.616 0.066 1.541 0.835 0.840 1.426 2.532 10.0
Table 3: Transformation matrix between ichnofabric subtypes of group 5 treated as normal Markov Chains and matrix of transition probabilities.
Type 2 to Type 7 (solely Phycosiphon; probability 0.18) and
to Type 3 (probability 0.14).
General tendencies in ichnofabrics along the core could be
shown in percentages calculated over an interval of 2.75 m,
whereby this interval moves in 25 cm steps (Figs. 2, 7). The
results of ‘moving percentages’ demonstrate core segments
preferred by special ichnofabric types and confirms the transi-
tions between types explained above. Climax Type 2 domi-
nates in the deepest core (95 to 100 m), only accompanied by
a few Type 3 and Type 6 ichnofabrics. After vanishing around
90 m, Type 6 dominates, accompanied from 90 m upward by
Types 3 and 7 (Figs. 2, 7). Type 4 with lamination becomes
abundant between 78 and 87 m getting its maximum around
Table 2: Transformation matrix between ichnofabric types (diagonal calculated for embedded Markov Chains) and matrix of transition proba-
bilities.
Type 1
Type 2
Type 3
Type 4
Type 5
Type 6
Type 7
sum
Type 1
3.7
2
5
3
5
7
6
31.7
Type 2
2
1.8
3
0
1
10
4
21.8
Type 3
1
4
4.7
5
0
9
12
35.7
Type 4
3
2
8
2.4
0
4
6
25.4
Type 5
7
0
0
0
1.1
1
8
17.1
Type 6
11
6
8
6
3
16.2
16
66.2
Type 7
6
5
12
5
6
19
19.4
72.4
sum
33.7
20.8
40.7
21.4
16.1
66.2
71.4
270.3
Type 1
Type 2
Type 3
Type 4
Type 5
Type 6
Type 7
sum
Type 1
0.12
0.06
0.16
0.09
0.16
0.22
0.19
1.0
Type 2
0.09
0.08
0.14
0.00
0.05
0.46
0.18
1.0
Type 3
0.03
0.11
0.13
0.14
0.00
0.25
0.34
1.0
Type 4
0.12
0.08
0.32
0.09
0.00
0.16
0.24
1.0
Type 5
0.41
0.00
0.00
0.00
0.06
0.06
0.47
1.0
Type 6
0.17
0.09
0.12
0.09
0.05
0.25
0.24
1.0
Type 7
0.08
0.07
0.17
0.07
0.08
0.26
0.27
1.0
sum
1.0
0.5
1.0
0.5
0.4
1.7
1.9
7.0
χ
2
= 55.1; df = 36; p(H
0
) = 0.022
405
ICHNOFABRIC ANALYSIS AND CORRELATION WITH ORBITAL CYCLES (MIDDLE MIOCENE, VIENNA BASIN)
84 m. After this period with abundant lamination, Type 3
briefly becomes important, but is suddenly replaced by Type 6
interchanging with Type 7 (Figs. 2, 7). These abundance
changes between Types 6 and 7 are characteristic for the core
from 87 m upwards, except the interval between 42 and 67 m,
where Type 1 characterized by Thalassinoides additional to
Phycosiphon and Nereites disturb these alterations. Further
disturbance can be found in the upper core by temporal occur-
rence of laminated core intervals (Type 4) and by the indefi-
nite group 5 with different subtypes or no visible macro-bur-
rows. The latter is mainly restricted to the core interval
between 16 and 34 m with two maxima around 27 and 22 m
(Figs. 2, 7). While the ‘climax’ Ichnofabric Type 2 dominates
the deepest core parts, it is rare between 47 and 82 m and com-
pletely lacking between 18 and 47 m.
Correlations and dependencies from the parameters of lami-
nation and magnetic susceptibility were tested using the com-
plete core, while relations to calcium carbonate and organic
carbon could be tested only for the deeper part (40 m to
102 m). Phycosiphon is significantly positively correlated
with magnetic susceptibility, explaining its presence in almost
all ichnofabric types (Table 4). Nereites, which is an equiva-
Fig. 7. General tendencies in ichnofabrics along the core shown in percentages calculated over an in-
terval of 2.75 m moving in 25 cm steps.
lent partner in Ichnofabric
Type 6 and less prominent –
compared to Phycosiphon – in
Types 3, 2 and 1, is signifi-
cantly positively correlated
with organic carbon and mag-
netic susceptibility, but highly
negatively correlated with lam-
ination (Table 4). This is sur-
prising, because lamination is
positively correlated with mag-
netic susceptibility. Thalassi-
noides, characteristic for Ich-
nofabric
Type 1
and
the
Subtypes 5_3 (together with
Nereites), and for 5_5 where it
represents the only trace fossil,
is negatively correlated with
magnetic susceptibility and
lamination, but highly posi-
tively correlated with calcium
carbonate (Table 4). Scolicia
as a typical component of Ich-
nofabric Type 3 is, contrary to
Thalassinoides, highly posi-
tively correlated with both
lamination and magnetic sus-
ceptibility. Trichichnus is the
only abundant ichnofossil that
is exclusively correlated with
organic carbon, thus similar in
demands to the rare Zoophy-
cos; both are elements of the
climax Type 2. The rare ichno-
fossils Asterosoma and Chon-
drites behave similarly in their
positive relations to lamina-
tion and magnetic susceptibility, which are more intensive in
Asterosoma compared to Chondrites (Table 4).
Discussion
Classification of ichnofabric types and their relations by sta-
tistical methods can be applied in ichnofabric analysis (Erba &
Premoli Silva 1994). However, its limitations must be taken
into account, since co-occurrence of trace fossils in ichnofab-
ric types may result from overlapping of different horizons.
For example for Ophiomorpha and Zoophycos, the coloniza-
tion surface can be above the 25-cm observation interval taken
as the basic unit of observation. Such trace fossils are more
connected with the environment at the colonization surface
than in the horizon at which they are observed. Such situations
influence the source data.
Accumulation rate, trophic changes and bottom stability
The fill of some open burrows, mainly Thalassinoides,
shows coarser grains than the surrounding, totally bioturbated
406
PERVESLER, UCHMAN and HOHENEGGER
A
st
ero
so
m
a C
ho
nd
ri
te
s
Ne
re
ite
s
Op
hi
om
or
ph
a
P
al
ae
op
hy
cu
s
P
hy
co
si
pho
n
Sc
ol
ic
ia
Si
ph
oni
chnus
T
ei
chi
chnus
T
ha
la
ss
inoi
de
s
T
ri
ch
ic
hnu
s Zo
op
hy
co
s
lam
in
at
ion
0.
13
3
0.
08
8
–0.
193
–0
.0
51
–0
.0
36
0.
04
9
0.
13
7
0.
05
1
–0
.0
47
–0
.1
28
–0
.0
55
0.
00
9
0.
01
0
0.
08
8
0.
00
0
0.
32
4
0.
48
7
0.
34
5
0.
00
8
0.
32
7
0.
36
8
0.
01
3
0.
29
0
0.
86
9
37
6
37
6
37
6
37
6
37
6
37
6
37
6
37
6
37
6
37
6
37
6
37
6
0.
15
7
0.
10
7
0.
17
2
–0
.032
–
0.
094
0.
17
6
0.
22
7
–0.
037
–0.
129
–0.
175
–
0.
011
–0.
013
0.
22
7
0.
00
2
0.
03
8
0.
00
1
0.
53
7
0.
06
8
0.
00
1
0.
00
0
0.
48
0
0.
01
2
0.
00
1
0.
83
2
0.
79
6
0.
00
0
37
6
37
6
37
6
37
6
37
6
37
6
37
6
37
6
37
6
37
6
37
6
37
6
37
6
–0
.1
35
–0
.0
48
–0
.0
22
0.
02
8
0.
07
6
–0
.0
82
–0
.1
06
0.
00
7
0.
02
9
0.
18
0
0.
08
7
0.
02
1
–0.
220
0.
03
4
0.
45
1
0.
73
4
0.
66
2
0.
23
2
0.
19
6
0.
09
4
0.
91
6
0.
65
3
0.
00
4
0.
16
9
0.
73
6
0.
00
0
24
9
24
9
24
9
24
9
24
9
24
9
24
9
24
9
24
9
24
9
24
9 24
9
24
9
0.
03
3
–0
.0
24
0.
21
7
0.
11
7
0.
06
7
0.
09
2
–0
.0
45
0.
04
5
–0
.0
04
–0
.0
43
0.
32
0
0.
14
2
–0.
272
0.
60
3
0.
70
5
0.
00
1
0.
06
5
0.
28
9
0.
14
9
0.
48
2
0.
47
9
0.
95
3
0.
50
4
0.
00
0
0.
02
5
0.
00
0
24
9
24
9
24
9
24
9
24
9
24
9
24
9
24
9
24
9
24
9
24
9
24
9
24
9
hig
hl
y s
ig
nif
ica
nt
po
sit
iv
e co
rr
el
atio
n
s
ig
nif
ic
an
t po
si
tiv
e co
rr
el
atio
n
hi
gh
ly
s
ign
if
ic
ant
n
ega
ti
ve c
orrela
ti
on
s
igni
fi
ca
nt
n
ega
ti
ve c
orrela
ti
on
Table 4:
Point-biserial
correlation
matrix
between
ichnogenera
and
the
p
arameters
lamination,
magnetic
susceptibility,
CaCO
3
and
organic
carbon.
rock. This suggests that deposition of coarser and finer sedi-
ments alternated but that the sediments were homogenized by
bioturbation. The sediment of the original grain size avoided
this process only in deep burrows. Thus, it can be supposed
that accumulation rate of the finer and coarser sediments var-
ied in periods of higher and lower accumulation rates. Proba-
bly, coarser sediments of higher accumulation rate derived
from shallower zones and contained higher amounts of partic-
ulate food and better oxygenated pore waters than the finer
sediments, which no doubt influenced ichnofauna.
Phycosiphon dominates the core, occurs in nearly all hori-
zons, and is accompanied in many layers by Nereites (Ichno-
fabric Type 6). These two trace fossils are produced by
deposit-feeding animals that have no permanent connection to
the seafloor and use oxygen from pore waters. They use par-
ticulate organic matter by horizontal reworking. Their abun-
dance suggests an opportunistic colonization. In turbiditic
sediments, they are typical of the initial colonization of turbid-
itic muds, whose pore waters are oxygenated and which con-
tain abundant food; Nereites usually crosscuts Phycosiphon
(Wetzel & Uchman 2001). By analogy, these trace fossils rep-
resent opportunistic colonization of oxygenated sediment rich
in particulate organic matter, probably quickly after increased
sediment input.
In most cases, Nereites crosscuts Phycosiphon and both are
crosscut by Scolicia, which is another horizontally reworking
infaunal deposit-feeding and omnivorous trace fossil. This or-
der of crosscutting relationships resulted rather from sequen-
tial colonization of sediment than from upward migration of
steady tiers in sediment occupied by the trace makers, accord-
Fig. 8. Position of ichnofossils along environmental gradients based
on correlation with magnetic susceptibility (related to accumulation
rate and oxygenation) and organic carbon (related to particulate
food content) (Table 1).
407
ICHNOFABRIC ANALYSIS AND CORRELATION WITH ORBITAL CYCLES (MIDDLE MIOCENE, VIENNA BASIN)
ing to sediment accumulation. Nereites followed Phycosi-
phon; Scolicia was produced later by irregular echinoids. Tak-
ing into account the size of these trace fossils, their trace
makers were larger and probably more effective deposit feed-
ers in their order of crosscutting. In turn, this can be correlated
with decrease of particulate food in the sediment. It is intrigu-
ing that Nereites has a tendency to occur in sediments richer in
TOC than Phycosiphon (Fig. 8). Probably, the Nereites stage
of colonization is missing when the food content is lower in
the colonized sediment. In such situations, Phycosiphon is al-
ways present as an unfailing opportunistic colonizer followed
directly by ichnotaxa for which horizontal sediment rework-
ing is less important.
When the sedimentation rate decreased, oxygen content
gradually decreased in pore waters and the redox boundary
migrated up. Particulate food also became less available due
to consumption by deposit feeders and by oxidation. In such a
situation, open and more stationary burrows were constructed,
which allowed use of oxygen from the water column instead
of pore waters. Intense sediment reworking for particulate
food deeper in the sediment was replaced by searching for
food at the sediment surface and from the water column. Some
of the trace makers applied chemosymbiotic feeding, such as
Chondrites or Trichichnus and partly Zoophycos (chemichnia
sensu Bromley 1996). Trichichnus is present only in sedi-
ments rich in TOC (Fig. 8), mainly in Ichnofabric Type 2,
where abundant particulate organic matter was buried below
the redox boundary.
The open burrows (Thalassinoides, Chondrites, Trichich-
nus, Zoophycos) indicate more specialized feeding related to
lower trophic level above the redox boundary and competition
for food, which are both probably caused by decreasing accu-
mulation rate. Thus, they indicate more stable-bottom condi-
tions. Based on such assumptions it is possible to interpret the
ichnofabric types and to distinguish tendencies in bottom sta-
bility change in the core.
Ichnofabric Types 6 and 7 record opportunistic colonization
(Phycosiphon, Nereites) of well-oxygenated sediments
(Fig. 7), interrupted by stable-periods allowing construction of
open burrows and their maintenance. In the Ichnofabric
Type 2, the stable periods after opportunistic colonization
were probably longer and characterized by some deficiency of
food above the redox boundary, but still with high food con-
tent below the redox boundary. In such conditions, stationary
chemosymbiotic feeding (Trichichnus and probably partly
Zoophycos) was effective. Ophiomorpha can be related to
shallowing or the effects of storms.
Ichnofabric Type 3 records the stabilized phase of coloniza-
tion and work of vagrant, opportunistic, omnivorous bioturba-
tors, namely irregular echinoids producing Scolicia. This was
evidently caused by higher input of detritus and slightly coars-
er sediment.
The lamination in Ichnofabric Type 4 indicates a high rate
of sedimentation. Phycosiphon is abundant and Nereites quite
rare. The sediment was only shallowly reworked and the pri-
mary lamination partly destroyed. In the Ichnofabric
Type 5_1, the sediment was reworked in a very shallow semi-
fluid zone, in which discrete trace fossils cannot be produced.
In Ichnofabric Type 5_2, only opportunistic colonization of
oxygenated, moderately food-rich-sediment took place, with-
out more stable-periods. In Ichnofabric Type 5_3, opportunis-
tic periods of colonization (Nereites) were followed by more
stable periods (Thalassinoides).
Bathymetry, salinity, oxygenation, deposition and consis-
tency of the substrate
The presence of Zoophycos and associated Zoophycos, Phy-
cosiphon, Nereites and Teichichnus suggests the Zoophycos
ichnofacies for the deeper part of the core. Upcore, Thalassi-
noides is more common, suggesting a transition to the distal
Cruziana ichnofacies. Such relations indicate shallowing up
to the upper offshore zone (cf. Pemberton et al. 2001).
The trace fossil Scolicia, produced by stenohaline irregular
echinoids, indicates fully marine conditions (e.g. Bromley &
Asgaard 1975; Smith & Crimes 1983). The salinity-tolerant
crustacean burrow Thalassinoides (Frey et al. 1984) replaces
Scolicia in the higher parts of the core. Teichichnus, which is
especially common in fine-grained brackish sediments (Pem-
berton et al. 2001), occurs infrequently in Ichnofacies
Types 3, 5_4 and 7. Thus, lowering salinity is not excluded in
the upper part of the core, especially in Ichnofacies Type 5_4,
where Teichichnus occurs alone.
The horizons with primary horizontal lamination in fine-
grained sediments can be related to anoxic conditions. The
laminae are partly disturbed by trace fossils connected to sub-
sequent improvement of oxygenation.
Commonly, Thalassinoides is filled with slightly coarser
sediment (fine sandy siltstone) than the host rock (siltstone,
mudstone). This suggests deposition of coarser sediment beds,
maybe distal tempestites or other event deposits, which were
later completely obliterated by bioturbation and the only less
mixed sediment was trapped in open Thalassinoides burrows,
similarly to the so-called tubular tempestites (Wanless et al.
1988; Tedesco & Wanless 1991). This is quite clear for the
boxwork of Thalassinoides filled with sand from the overly-
ing bed (Fig. 3F). Probably, rare Ophiomorpha isp. is connect-
ed with colonization of such event beds.
Small compaction of the Thalassinoides galleries indicates
stiffground substrate sensu Wetzel & Uchman (1998), espe-
cially in Ichnofabric Type 5_5, where Thalassinoides marks a
discontinuity surface (Glossifungites ichnofacies sensu Pem-
berton et al. 2001).
Periodicity
Occurrences of the ichnofabric types show some periodici-
ty. Ichnofabric Types 6 and 7, related to opportunistic coloni-
zation and bottom instability, display maxima every 10—15 m
(Figs. 2, 7). They are intercalated by ichnofabrics that show
generally higher trace fossil diversity and are related to higher
bottom stability. Periods of stability and instability are related
to changes in sedimentation rate and resultant oxygenation
and food availability changes. Changes of these factors, in
turn, can be related to the precession cycles calculated for
11—15 m periods (Hohenegger et al. 2008). The other cycles,
namely obliquity cycles with 20-m periods and the eccentrici-
ty cycles with around 40-m periods, are less distinct but still
408
PERVESLER, UCHMAN and HOHENEGGER
visible (Fig. 2). Thus, analysis of ichnofabric can help the rec-
ognition of orbital cycles in lithologically monotonous sections.
Conclusions
1. Trace fossil and ichnofabric analyses are powerful tradi-
tional tools in the recognition of ecological parameters such as
energy level, oxygen content, food supply, salinity or stability
of ancient environments.
2. Statistical classification analyses help us to distinguish
types of recurrent ichnofabrics.
3. Calculation of transition probabilities between ichnofab-
ric types allows the recognition of general tendencies in ichno-
fabric distribution along a section.
4. Correspondence of these general tendencies with envi-
ronmental gradients like magnetic susceptibility and organic
carbon results in detailed information on tolerance of burrow-
ing organisms against accumulation rate, oxygenation and
particulate food content.
5. Based on these correspondences the distribution of ichno-
fabrics significantly marks the influence of periods in orbital
cycles.
Acknowledgments: The study was supported by the Aus-
trian Science Fund (FWF, Projects P13743-BIO, P 13740,
P 18203), the Austrian Oriental Society Hammer-Purgstall,
and the OAD, Austrian Exchange Service. A. Uchman was
supported additionally by the Jagiellonian University,
Kraków (DS funds).
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