ICHNOFOSSILS FROM THE TYPE AREA OF THE GRUND FORMATION 103
GEOLOGICA CARPATHICA, 55, 2, BRATISLAVA, APRIL 2004
ICHNOFOSSILS FROM THE TYPE AREA OF THE GRUND
FORMATION (MIOCENE, LOWER BADENIAN) IN NORTHERN
LOWER AUSTRIA (MOLASSE BASIN)
and ALFRED UCHMAN
Department of Paleontology, University of Vienna, Althanstrasse 14, A-1090 Wien, Austria; email@example.com
Institute of Geological Sciences, Jagiellonian University, ul. Oleandry 2a, 30-063 Kraków, Poland; firstname.lastname@example.org
(Manuscript received June 5, 2003; accepted in revised form December 16, 2003)
Abstract: The trace fossils Arenicolites, Asterosoma, Diplocraterion, Zoophycos, Ophiomorpha, Saronichnus, Scolicia
and Thalassinoides have been recognized in the siliciclastics of the Grund Formation. Their occurrence and distribution
is related to storm deposition. In proximal storm deposits, only monospecific Asterosoma occurs. It is typical of the
transition between the Skolithos and the proximal Cruziana ichnofacies. A more diverse trace fossil association of the
proximal and archetypical Cruziana ichnofacies occurs in more distal storm deposits. The vertical structures (Arenicolites,
Diplocraterion, Ophiomorpha) are related to opportunistic colonization of the storm beds (post-event community). The
horizontal forms (Scolicia, Thalassinoides) represent fair weather conditions. The chemosymbiotic structures (Saronichnus,
Zoophycos) are a record of trophic competition that pressures trace makers to deeper and more complex feeding than
simple deposit feeding. The horizontal and chemosymbiotic trace fossils represent the resident community. The develop-
ment from the Skolithos via the proximal Cruziana ichnofacies to the proximalarchetypical Cruziana ichnofacies indi-
cates a deepening from the middle shoreface to upper offshore environments.
Key words: Miocene, Austria, chemosymbiotic structures, molasse, storm deposits, trace fossils.
Trace fossils from the North Alpine Molasse Basin are poten-
tially a good source of information about benthic infaunal life,
paleoenvironmental parameters, and sequence stratigraphy.
Unfortunately, they are underrepresented in the literature (but
see e.g. Ehrenberg 1938, 1944; Hohenegger & Pervesler 1985;
Uchman & Krenmayr 1995, for the Austrian molasse). An in-
teresting trace fossil assemblage has been discovered in the
Fig. 1. Location map. Sections A, B1, B2, C, D, E were excavated in 1998, and sections F, G and H in 1999.
lower Middle Miocene (Lower Badenian) Grund Formation
(Pervesler & Zuschin 2002; Pervesler & Roetzel 2002). It was
studied during two excavation campaigns in 1998 and 1999 at
the type locality of this formation (former Grunder Schicht-
en). Several deep trenches were excavated in the farmland
between the villages of Grund and Guntersdorf, north of Hol-
labrunn in Northern Lower Austria (compare Roetzel & Per-
vesler 2003). Trace fossils were found in the sections B2, C,
D, E, F, G and H (Fig. 1). Their description and interpretation
is the main aim of this paper.
104 PERVESLER and UCHMAN
The sediment containing the trace fossils was carefully re-
moved using a jet of compressed air (Fig. 2). A pistol-shaped
valve helped to control the direction and volume of the airflow
(Fig. 2). This method, mainly developed for uncemented sands,
allowed even very delicate structures to be excavated and ob-
served in three dimensions. For further studies in the laborato-
ry, the trace fossils were collected together with the host sedi-
ment using boxes made of plastic or wood, whereby the
samples were held in position fixed by polyurethane foam.
Fig. 2. Sediment was carefully removed with an air jet. The pistol-
shaped valve helped to control the direction and volume of the air-
Scolicia De Quatrefages, 1849
Material: 2 field photographs.
Location: Grund, excavation 1999, section F, 247.6 m
above sea level.
Host sediment: Silt.
Description: Subcylindrical, slightly winding horizon-
tal structures with meniscate backfilling, 6070 mm wide.
Horizontal sections of this trace fossil visible on erosion sur-
faces, about 30 mm wide, resemble Laminites Ghent et Hend-
erson, which was included in Scolicia (Uchman 1995; Uch-
man & Krenmayr 1995). Scolicia is a fossil burrow produced
by irregular echinoids (e.g. Bromley & Asgaard 1975; Smith
& Crimes 1983).
Arenicolites Salter, 1857
Material: 8 field photographs.
Location: Grund, excavation 1999, section G, 248.1 m
above sea level.
Host sediment: Fine- to medium-grained sand.
Description: Simple U-shaped cylindrical structures
without spreiten. Diameters of the cylinders attain about
3.5 mm (Fig. 3.2), but decrease towards the upper termina-
tions. Limbs of the U-structure are about 30 mm apart. Some
limbs are inclined outward (Fig. 3.2) and give the impression
of a J-shaped structure. Large specimens attain diameters of
14 mm with their limbs about 80 mm apart (Fig. 3.3). Arenico-
lites is interpreted as a dwelling and feeding burrow of suspen-
sion-feeding annelids (e.g. Hakes 1976) or small crustaceans
(Goldring 1962). It occurs in different environments, but is
typical of shallow-marine settings (Crimes 1977). For discus-
sion of this ichnogenus see Fillion & Pickerill (1990) and
Ekdale & Lewis (1991).
Diplocraterion Torell, 1870
Material: 5 field photographs, 3 thin sections (micro-
Locations: Grund, excavation 1998, section E, 245.3 m
above sea level.
Host sediment: Fine- to medium-grained sand.
Description: U-shaped retrusive spreiten-structures, 50
130 mm long, with diverging outwardly inclined limbs. The
tops of the limbs are 70 to 100 mm apart. Up to 13 spreiten
laminae per structure were counted. Some limbs display fun-
nel-shaped entrances (Fig. 3.4). The shallowly inclined limbs
are features of Diplocraterion parallelum var. arcum Ekdale et
Lewis, 1991 (compare Fürsich 1974; Corner & Fjalstad 1993).
Diplocraterion is a typical component of the Skolithos ich-
nofacies, and occurs commonly in very shallow subtidal to in-
tertidal facies (e.g. Fürsich 1974, 1981). In the Jurassic at least,
this trace fossil also characterizes transgressive surfaces (e.g.
Mason & Christie 1986; Dam 1990; Taylor & Gawthorpe
1993; Goldring et al. 1998). Horizons with abundant D. paral-
lelum in the Upper Jurassic of Spain have been used by Olóriz
& Rodríguez-Tovar (2000) to recognize more energetic and
physically unstable environments (a transition from the Cruzi-
ana ichnofacies to the mixed CruzianaSkolithos ichnofacies)
in relation to sea level fall, and used to delineate sequence
boundaries. Diplocraterion is also common in event beds,
where it documents the opportunistic post-event colonization
(Frey & Goldring 1992).
Ophiomorpha Lundgren, 1891
Ophiomorpha nodosa Lundgren, 1891
Material: 16 field photographs, 12 tunnel-fragments.
Location: Grund, excavation 1999, section G, 248.1 m
above sea level.
Host sediment: Medium-grained sand.
Description: Boxwork of long, slightly inclining tunnels
and short, steeper shafts. Branching shows a 120° pattern.
ICHNOFOSSILS FROM THE TYPE AREA OF THE GRUND FORMATION 105
Fig. 3. Trace fossils from the type area of the Grund Formation. 1 Oblique view of Scolicia isp. at the top of a pelitic layer. Excavation
1999, section F, 247.6 m above sea level. 2 Arenicolites isp. in fine- to medium-grained sands. Excavation 1999, section G. 3
Arenicolites isp. and irregularly winding tubes (arrow) that enter the coarser layers below. Excavation 1999, section G, 248.1 m above sea
level. 4 Saronichnus abeli (left side) and Diplocraterion isp. (right side; only a half of the trace fossil is visible). Diplocraterion isp.
shows funnel-shaped burrow openings with a horizontal shift of the limbs. Excavation 1998, section E. 5 Thin section (small magnitude
picture) of Diplocraterion isp. from the deepest part of the U-shaped burrow. Excavation 1998, section E. 6 Oblique view of Asterosoma
radiciforme. Excavation 1998, section C, 239.8 m above sea level. The bulb-shaped trace fossils show tiny shaft-like connections to the
pelitic layer representing the former sea floor. 7 Lateral view of Asterosoma radiciforme in a fining upward sequence, from a fine-
grained sand at the base to a silt at the top. Intervening tunnels (arrow) connect different generations of Asterosoma. Excavation 1999, sec-
tion G, 247.5 m above sea level. 8 Boxwork of the Ophiomorpha nodosa in medium-grained sand with shell fragments. Excavation 1999,
section G, 248.1 m above sea level. 9 Two specimens of Saronichnus abeli (S) associated with their producer, the chemosymbiotic bi-
valve Thyasira michelottii (T).
106 PERVESLER and UCHMAN
Slight swellings occur at the branching points. The exterior of
the tunnel walls is generally lined with sandy granules. The
interior is smooth; the bottom surface of some horizontal and
inclined tunnels is unlined. The wall is 3 to 9.4 mm thick.
Cross-sections of the compressed horizontal tunnels are up to
37 mm wide and up to 31 mm high. The vertical extension of
this trace fossil measures up to 40 cm.
Ophiomorpha nodosa is one of the most common shallow-
marine trace fossils and is produced mostly by callianassid
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).
Thalassinoides Ehrenberg, 1944
Thalassinoides suevicus (Rieth, 1932)
Material: 7 field photographs.
Location: Grund, excavation 1999, section F, 247.6 m
above sea level.
Host sediment: Silt.
Description: System of shallowly inclined tunnels and
steep shafts, which are about 20 mm in diameter. There are
chamber-like swellings around the branching points, up to
35 mm wide, resembling turning chambers of recent crusta-
ceans (see Bromley 1996 for review). This trace fossil was
constructed in a semi-consolidated mud and later truncated by
erosion. The burrow wall is smooth. The vertical extension
measures 15 cm. Thalassinoides was produced by crusta-
ceans, mostly decapods (Frey et al. 1984). For further discus-
sion of this ichnogenus and its ichnotaxonomic problems see
Fürsich (1973), Ekdale (1992) and Schlirf (2000).
Fig. 4. Erosion surface truncates a Thalassinoides burrow system
formed in a silty mud. Grund, excavation 1999, section F, 247.6 m
above sea level.
Asterosoma von Otto, 1854
Asterosoma radiciforme von Otto, 1854
Material: One box sample, 36
26 cm wide and 25 cm
high, with numerous specimens in four horizons, 11 field pho-
Location: Grund, excavation 1998, sections C, D; exca-
vation 1999, sections F, G, H, 239.8249.2 m above sea level.
Host sediment: Fine- to medium-grained sand capped
Description: Vertical to inclined elongated bulbs, up to
14 mm wide and up to 25 mm long, tapering at both ends,
with concentric internal lamination. Clusters of such bulbs
form tree-like structures, up to 10 cm wide, spreading out
from a common vertical or inclined shaft that is 34 mm in
diameter. The main portions of the clusters are located in the
sands directly below the overlying pelitic layers. Steep
shafts connect the clusters from subsequent sedimentary
sand-pelite cycles. 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.
Saronichnus Pervesler et Zuschin, 2004
Saronichnus abeli Pervesler et Zuschin, 2004
Fig. 3.4, 9
Material: 14 field photographs, type specimens: Holo-
type IPUW No. 2004-0001-0001, paratype IPUW No. 2004-
Location: Grund, excavation 1999, section E, G, H, 244
249.5 m above sea level.
Host sediment: Medium- to fine-grained sand covered
Description: Bundles of vertical to steeply inclined,
simple, blade- or club-shaped probes, up to 100 mm long and
about 3 mm wide. They spread downwards below the life po-
sition of the chemosymbiotic bivalve Thyasira michelottii (R.
Hörnes, 1875) (compare Pervesler & Zuschin 2004). The bun-
dles resemble a downward-oriented fan or a broom.
The probes, produced by the bivalve foot, are thought to be
wells for sulphides for the use of chemosymbiotic bacteria
(Zuschin et al. 2001; Pervesler & Zuschin 2004).
The trace fossil is similar to Pragichnus fascis Chlupáè
(1987) known from the Ordovician of Bohemia, which how-
ever, displays rather only club-shaped probes, which are ac-
tively back-filled and branched, especially in distal (lower)
part where dichotomous branches occur. This trace fossil was
also interpreted as sulphide wells produced by an animal us-
ing chemosymbiotic bacteria (Mikulá 1997).
Zoophycos Massalongo, 1856
Material: 19 field and 8 laboratory photographs, one box
50 centimeters wide and 70 centimeters high,
with two specimens (section E), fragments from a further
specimen (section G).
ICHNOFOSSILS FROM THE TYPE AREA OF THE GRUND FORMATION 107
Fig. 5. Zoophycos isp. from the type area of the Grund Formation. Photographs 1, 2, 5, 6 and 7 from excavation 1998, section E. Photographs
3 and 4 from excavation 1999, section G. 1 Two interfingering specimens of Zoophycos isp. Arrows indicate their upper terminations.
2 Transitional area between the planar structure and the Rhizocorallium-like steep lobes. Detail of 1. 3 Single Zoophycos isp. consist-
ing of the helical part with 3 whorls passing into the planar spreite structures and lobes. Most of the structures were destroyed during excava-
tion. 4 Fragment of a single lobe illustrated in 3. X-ray image. 5 Cross-section of a single lobe showing the passively filled marginal
tube (at the top) and actively backfilled tube (at the bottom). 6 A specimen with shallowly inclined lobes. 7 Steep, twisted lobes in the
lower part of Zoophycos isp.
Location: Grund, excavation 1998, section E, 245.5 m
above sea level (Fig. 5.1,2,57); excavation 1999, section G,
247.4 m above sea level (Fig. 5.34).
Host sediment: Fine- to medium-grained sand.
Description: Planar helical spreite structures transition-
ing in the lower part into numerous Rhizocorallium-like long
lobes (Fig. 5.1,3). The whole trace fossil is up to 350 mm
high. The upper helical part forms three to five whorls, which
are subcircular in outline (Fig. 5.3), showing clockwise or
anti-clockwise development; they are inclined downward at
the angle of 10 to 30° and penetrate 60 to 100 mm into the
sediment. The diameter of the whorls increases downward.
108 PERVESLER and UCHMAN
The uppermost whorls are about 20 mm, and the deepest at
least 100 mm in diameter. The helical part develops into larger,
planar, one- or two- spreite structures, which are at least
250 mm wide and are inclined up to 30°. In the distal part they
can be horizontal or even slightly ascending. Locally, the
Rhizocorallium-like lobes (Fig. 5.4), surrounded partly by a
marginal tube, start from the margin of the deepest planar
structures. Exceptionally, lobes can develop directly from the
axial part of the helical portion. They show more or less equal
inclination within the same lobe, or slight steepening from
about 30° in the proximal part to about 40° in the distal part.
The lobes developing from the planar structures overlap in
their proximally. The marginal tubes of the older lobes give
the impression of rays with pinnulae on one side only as de-
scribed by Girotti (1970). Proximally, the lobes are inclined
about 30°. Distally, they steepen rapidly steepening to a verti-
cal Diplocraterion-like position (Fig. 5.13). The lobes are
twisted according to the general development of the burrow
system. They are up to 250 mm long and about 20 mm wide.
The steep part is up to 200 mm long. Their marginal tube al-
ways displays a passive filling from one side and active back-
filling from the other side of the lobe (Fig. 5.5). This indicates
a J-shaped causative burrow, whose shifting resulted in for-
mation of the lobe structures. Interestingly some of the pas-
sively filled marginal tubes display sandy or carbonate linings,
in which the sand is brownish and darker than the sur-
rounding sediment (Fig. 5), similar to oxygenated shafts of re-
cent burrows. Occasionally, the lobes interpenetrate (Fig. 5.2).
In one case, the vertical lobes terminate at a silty layer (Fig.
5.1). Their lowest part deviates from the vertical position and
develops along the layer over a distance of several millimeters.
The U-shaped lobes can be considered to be a product of J-
shaped causative burrows. They correspond to the U-burrow
model proposed for Zoophycos sensu lato by Ekdale et Lewis
(1991) and discussed later by Uchman et Demírcan (1999).
The described trace fossil displays general similarity to Zoo-
phycos rhodensis described by Bromley et Hanken (2003),
but the latter is much larger and shows phobotaxis of the
lobes. The upper, helical part of Z. rhodensis Bromley et Han-
ken (2003) (skirt-like zone) was interpreted as a deposit-
feeding structure, and the lobes as sulphide wells for chemo-
symbiotic bacteria. This interpretation can be generally
accepted for Zoophycos isp. from the Grund Formation. The
steep and deep lobes were probably produced in anoxic sedi-
ment. The spreite laminae in the lobes can be produced when
the trace maker exploited the sediment to obtain the bacteria.
The Zoophycos trace maker is thought to have migrated to
deeper environments during the Jurassic (Bottjer et al. 1988;
Olivero 2003). In Upper Quaternary sediments, Zoophycos
occurs at depths below 1000 m (Löwemark & Schäfer 2003).
Thus, the described Zoophycos is one of the shallowest (upper
offshorelower shoreface; see discussion) occurrences of this
ichnogenus after the Jurassic. Pemberton et al. (2001) men-
tioned the occurrence of this ichnogenus at similar depths in
the Cretaceous. These depths probably delineate the upper
bathymetric range of this trace fossil. The small size of the de-
scribed Zoophycos may be related to stress close to the border
of its environmental range.
The sections at the type locality of the Grund Formation
show sediments from a shallow marine, highly erosive envi-
ronment with small channels (compare Roetzel & Pervesler
2004). The channels, maximally 78 m wide and 0.51 m
deep, always have a sharp erosive base. Their fill with densely
packed bioclast-supported shell layers at the base, fining up-
ward cycles of coarse- to fine-grained sands, and thin pelitic
layers on the top, indicate periodically high-energy events
with rapidly decreasing energy level.
In the lower part of the sequence (sections A, B), sandy
beds, 60 to 120 cm thick, contain up to 40 cm thick layers of
pelitic clasts, showing strong physical reworking. Moreover,
they contain a mixed allochthonous fauna of marine molluscs,
terrestrial gastropods and bones of different vertebrates (tur-
tles, whales, rhinoceroses, small carnivores and micro-mam-
mals). These deposits are distinguished as lithofacies A (Roet-
zel & Pervesler 2004).
Towards the top of the sequence, the thickness and grain
size of these beds decrease. In the middle part of the sequence,
in the sections C and D, 20 to 45 cm thick medium- to fine-
grained sand beds show even lamination. Current ripples and
plant debris at the top of some beds, together with the fining
of grain size, indicate reduced current velocity. Pelitic layers
several centimeters thick commonly cover the laminated
sands. Intense but monospecific bioturbation (Asterosoma)
starts from these layers but only reaches down maximally
3 cm into the sandy layers. The sand bodies display mostly
tabular, locally slightly wedge-shaped geometry. They are cut
by sparse, small and narrow runnels, 60 to 80 cm wide and 10
to 25 cm deep, with erosive base and filled with pelitic clasts.
These deposits are distinguished as lithofacies B (Roetzel &
The uppermost part of the studied section (sections E, F, G,
H) also contains evenly laminated, medium- to fine-grained
sands with graded bedding, but the thickness of the pelitic
layers increases towards the top of the sequence. These pelitic
layers 10 to 20 cm thick contain a diverse trace fossil assem-
blage starting from the pelitic horizons and penetrating down
into the coarser fine- to medium-grained sands. The erosive
deposition of coarse material stopped the work of the burrow-
ers. Arenicolites, Diplocraterion, Zoophycos and Ophiomor-
pha are common in these sediments. Thicker pelitic layers
contain rare Scolicia and Thalassinoides. The chemosymbiot-
ic bivalve Thyasira michelottii (R. Hörnes, 1875) occurs as an
exclusively autochthonous inhabitant of these layers. Some of
the Thyasira-shells are associated with Saronichnus. These
deposits are distinguished as lithofacies C (Roetzel & Per-
The trace fossil assemblage and its distribution can be inter-
preted by means of the model related to the hydrodynamic
level that was elaborated on the basis of long-term studies of
the North American Cretaceous Seaway and summarized by
Pemberton et al. (2001).
The basal sections of the sequence (lithofacies B) show the
highest but mostly monospecific biological disturbance. Only
opportunistic organisms that are able to colonize mobile sedi-
ICHNOFOSSILS FROM THE TYPE AREA OF THE GRUND FORMATION 109
ment (trace makers of Asterosoma) can settle during short pe-
riods of quiet conditions. This is a typical situation for the
middle shoreface settings in the transitional zone between the
Skolithos and the proximal Cruziana ichnofacies (Pemberton
et al. 2001: p. 121).
During deposition of the lithofacies C, longer periods of
benthic recovery after physical disturbances lead to greater
burrowing depth and higher trace fossil diversity. Deposit
feeding and chemosymbiotic strategies are characteristic fea-
tures of the uppermost parts of the excavated sections of the
Grund Formation. The lateral and vertical change of the hy-
drodynamic energy level is the main factor governing the de-
velopment and distribution of different trace fossil assemblag-
es. There are vertical trace fossils typical of higher energy
(Ophiomorpha, Diplocraterion, Arenicolites); these are char-
acteristic for the Skolithos ichnofacies. There are also typical,
mostly horizontal components of the Cruziana ichnofacies
(Scolicia, Thalassinoides), and trace fossils typical of the Zoo-
phycos ichnofacies (Zoophycos, Saronichnus as an equivalent
of Chondrites). Ichnofacies clearly cannot be interpreted on
the basis of single ichnotaxa, but rather on the whole ichnoas-
semblage (e.g. Frey & Seilacher 1980). Such a mixture of
trace fossils of different ethology is characteristic of the upper
offshorelower shoreface settings, where the proximal and ar-
chetypical Cruziana ichnofacies typically occurs (Pemberton
et al. 2001). The vertical structures are related to opportunistic
colonization of the storm beds (post-event community). Storm
currents can transport the trace makers to the deeper environ-
ments (Frey & Goldring 1992). The horizontal structures are
related to fair weather conditions. The chemosymbiotic struc-
tures (Saronichnus and probably Zoophycos) are a record of
trophic competition which forces trace makers to shift to
deeper and more complex feeding than simple deposit feed-
ing. The horizontal and chemosymbiotic trace fossils repre-
sent the resident community.
The studied trace fossil assemblage shows a transition from
the Skolithos to the proximal Cruziana ichnofacies to the
proximalarchetypical Cruziana ichnofacies up the section.
This indicates a deepening from the middle shoreface to upper
offshore environments dominated by storms.
Acknowledgments: Project 13743-BIO (Temporal and spa-
tial changes of microfossil associations and ichnofacies in the
Austrian marine Miocene) and Project P 13745-BIO (Evolu-
tion Versus Migration: Changes in Austrian Marine Miocene
Molluscan Paleocommunities) of the Austrian Science Fund,
and the Department of Paleontology at the University of Vien-
na, supported this study. We extend our special thanks to Rein-
hard Roetzel (Vienna) for his support in sedimentology and to
all helpers during fieldwork. Special thanks are due to Rich-
ard Bromley (Copenhagen) and Radek Mikulá (Prague), who
were the official reviewers, and due to Reinhard Roetzel (Vien-
na) and Michael Stachowitsch (Vienna) for critical reading.
Bottjer D.J., Droser M.L. & Jablonski D. 1988: Paleoenvironmental
trends in the history of trace fossils. Nature 333, 252255.
Bromley R.G. 1996: Trace fossils: Biology, taphonomy and applica-
Edition. Chapman & Hall, London, 1361.
Bromley R.G. & Asgaard U. 1975: Sediment structures produced by
a spatangoid echinoid: a problem of preservation. Bull. Geol.
Soc., Denmark 24, 261281.
Bromley R.G. & Hanken N.-M. 2003: Structure and function of
large, lobed Zoophycos, Pliocene of Rhodes, Greece. Palaeo-
geogr. Palaeoclimatol. Palaeoecol. 192, 79100.
Chlupáè I. 1987: Ordovician ichnofossils in the metamorphic man-
tle of the Central Bohemian Pluton. Èas. Mineral. Geol. 32,
Corner G.D. & Fjalstad A. 1993: Spreite trace fossils (Teichichnus)
in a raised Holocene fjord-delta, Breidvikeidet, Norway. Ich-
nos 2, 155164.
Crimes T.P. 1977: Trace fossils of an Eocene deep-sea sand fan,
northern Spain. In: Crimes T.P. & Harper J.C. (Eds.): Trace
fossils 2. Geol. J. Spec. Iss. 9, 7190.
Dam G. 1990: Paleoenvironmental significance of trace fossils from
the shallow marine Lower Jurassic Neill Klinter Formation,
East Greenland. Palaeogeogr. Palaeoclimatol. Palaeoecol. 79,
Ehrenberg K. 1938: Bauten von Decapoden (Callianassa sp.) aus
dem Miozän (Burdigal) von Burgschleinitz bei Eggenburg im
Gau Nieder-donau (Niederösterreich). Paläont. Z. 20, 263284.
Ehrenberg K. 1944: Ergänzende Bemerkungen zu den seinerzeit aus
dem Miozän von Burgschleinitz beschriebenen Gangkernen
und Bauten dekapoder Krebse. Paläont. Z. 23, 354359.
Ekdale A.A. 1992: Muckraking and mudslinging; the joys of depos-
it-feeding. In: Maples C.G. & West R.R. (Eds.): Trace fossils.
Short Courses in Paleontology 5. Univ. Tennessee, Knoxville,
Ekdale A.A. & Lewis D.W. 1991: Trace fossils and paleoenviron-
mental control of ichnofacies in a late Quaternary gravel and
loess fan delta complex, New Zealand. Palaeogeogr. Palaeo-
climatol. Palaeoecol. 81, 253279.
Fillion D. & Pickerill R.K. 1990: Ichnology of the Lower Ordovi-
cian Bell Island and Waban Group of Eastern Newfoundland.
Palaeontographica Canadiana 7, 1119.
Frey R.W. 1990: Trace fossils and hummocky cross-stratification,
Upper Cretaceous of Utah. Palaios 5, 203218.
Frey R.W. & Goldring R. 1992: Marine event beds and recoloniza-
tion surfaces as revealed by trace fossil analysis. Geol. Mag.
Frey R.W., Curran A.H. & Pemberton G.S. 1984: Tracemaking ac-
tivities of crabs and their environmental significance: the ich-
nogenus Psilonichnus. J. Paleontology 58, 511528.
Frey R.W., Howard J.D. & Pryor W.A. 1978: Ophiomorpha: its
morphologic, taxonomic and environmental significance.
Palaeogeogr. Palaeoclimatol. Palaeoecol. 23, 199229.
Frey R.W. & Seilacher A. 1980: Uniformity in marine invertebrate
ichnology. Lethaia 23, 183207.
Fürsich F.T. 1973: A revision of the trace fossils Spongeliomorpha,
Ophiomorpha and Thalassinoides. Neu. Jb. Geol. Paläont.,
Monatshefte 1972, 719735.
Fürsich F.T. 1974: On Diplocraterion Torell 1870 and the signifi-
cance of morphological features in vertical, spreiten-bearing,
U-shaped trace fossils. J. Paleontology 48, 952962.
Fürsich F.T. 1981: Invertebrate trace fossils from the Upper Jurassic
of Portugal. Comun. Serv. Geol. Port. 67, 11531168.
Ghent H.D. & Henderson R.A. 1966: Petrology, sedimentation, and
paleontology of the Middle Miocene graded sandstones and
mudstone, Kaiti Beach, Gisborne. Trans. Roy. Soc. N. Z., Part
Geol. 4, 147169.
Gibert J.M. de 1996: Diopatrichnus odlingi n. isp. (annelid tube)
and associated ichnofabrics in the White Limestone (M. Juras-
sic) of Oxfordshire: sedimentological and palaeoecological
110 PERVESLER and UCHMAN
significance. Proc. Geologists Assoc. 107, 189198.
Girotti O. 1970: Echinospira pauciradiata g. n., sp. n., ichnofossil
from the Seravallian-Tortonian of Ascoli Piceno (central Italy).
Geologica Romana 9, 5962.
Goldring R. 1962: The trace fossils of the Baggy beds (Upper Devo-
nian) of north Devon, England. Paläont. Z. 36, 34, 232251.
Goldring R., Layer M.G., Magyari A., Palotas K. & Dexter J. 1998:
Facies variation in the Corallian Group (U. Jurassic) of the Far-
ingdon-Shellingford area (Oxfordshire) and the rockground
base to the Faringdon Sponge Gravels (L. Cretaceous). Proc.
Geologists Assoc. 109, 115125.
Hakes W.G. 1976: Trace fossils and depositional environment of
four clastic units, Upper Pennsylvanian megacyclothems,
northeast Kansas. Paleont. Contr. Univ. Kan. 63, 146.
Hohenegger J. & Pervesler P. 1985: Orientation of crustacean bur-
rows. Lethaia 18, 323339.
Hörnes R. 1875: Die Fauna des Schliers von Ottnang. Jb. K.-Kön.
Geol. Reichsanst. 25, 4, 333431.
Löwemark L. & Schäfer P. 2003: Ethological implications from a
detailed X-ray radiograph and
C study of the modern deep-
sea Zoophycos. Palaeogeogr. Palaeoclimatol. Palaeoecol. 192,
Lundgren S.A.B. 1891: Studier öfver fossilförande lösa block. Geol.
Fören. Förh., Stockholm 13, 111121.
Mason T.R. & Christie A.D. 1986: Palaeoenvironmental signifi-
cance of ichnogenus Diplocraterion (Torell) from the Permian
Vryheid Formation of the Karoo Supergroup, South Africa.
Palaeogeogr. Palaeoclimatol. Palaeoecol. 52, 249265.
Massalongo A. 1855: Zoophycos, novum genus plantorum fossili-
um. Antonelli, Verona, 152.
Mikulá R. 1997: Ethological interpretation of the ichnogenus
Pragichnus Chlupáè 1987 (Ordovician, Czech Republic). Neu.
Jb. Geol. Paläont., Mh. 1997, 2, 93108.
Olivero D. 2003: Early Jurassic to Late Cretaceous evolution of Zoo-
phycos in the French Subalpine Basin (southeastern France).
Palaeogeogr. Palaeoclimatol. Palaeoecol. 192, 5978.
Olóriz F. & Rodríguez-Tovar F.J. 2000: Diplocraterion: a useful
marker for sequence stratigraphy and correlation in the Kim-
meridgian, Jurassic (Prebetic Zone, Betic Cordillera, southern
Spain). Palaios 15, 546552.
Otto E. von 1854: Additamente zur Flora des Quadergebirges in
Sachsen. Part 2. G. Mayer, Leipzig, 153.
Pemberton S.G., Spila M., Pulham A.J., Saunders T., MacEachern
J.A., Robbins D. & Sinclair I.K. 2001: Ichnology and sedimen-
tology of shallow to marginal marine systems. Geological As-
sociation of Canada, Short Course Notes 15, 1343.
Pervesler P. & Roetzel R. 2002: Environmental significance of bio-
turbations in the Grund Formation (Miocene, Lower Badenian)
in northern Lower Austria. PANGEO AUSTRIA, Salzburg, June
Pervesler P. & Zuschin M. 2002: Chemosymbiosis, fossil lucinoid
bioturbations and the Chondrites enigma. ESSEWECA Interna-
tional Conference, Bratislava, June 57 2002.
Pervesler P. & Zuschin M. 2004: A lucinoid bivalve trace fossil Sa-
ronichnus abeli igen. et isp. nov. from the Miocene molasse
deposits of Lower Austria, and its environmental significance.
Geol. Carpathica 55, 2, 111115.
Quatrefages M.A. de 1849: Note sur la Scolicia prisca (A. de Q.)
annélide fossile de la Craie. Ann. Sci. Natur., 3 série, Zoologie
Rieth A. 1932: Neue Funde spongeliomorpher Fucoiden aus dem Jura
Schwabens. Geol. Paläont. Abh., Neue Folge 19, 257294.
Roetzel R. & Pervesler P. 2004: Storm-induced event deposits in the
type area of the Grund Formation (Middle Miocene, Lower
Badenian) in the Molasse Zone of Lower Austria. Geol. Car-
pathica 55, 2, 87102.
Plaziat J.C. & Mahmoudi M. 1988: Trace fossils attributed to bur-
rowing echinoids: A revision including new ichnogenus and
ichnospecies. Geobios 21, 209233.
Salter J.W. 1857: On annelide-burrows and surface-markings from
the Cambrian rocks of the Longmynd. Quart. J. Geol. Soc.,
London 13, 199-206.
Schlirf M. 2000: Upper Jurassic trace fossils from the Boulonnais
(northern France). Geologica et Palaeont. 34, 145213.
Smith A.B. & Crimes T.P. 1983: Trace fossils formed by heart urchins
a study of Scolicia and related traces. Lethaia 16, 7992.
Torell O.M. 1870: Petrifacta Suecana Formationis Cambricae.
Lunds Univ. Årsskr. 6, 2, 8, 114.
Taylor A.M. & Gawthorpe R.L. 1993: Application of sequence
stratigraphy and trace fossil analysis to reservoir description
examples from the Jurassic of the North Sea. In: Parker J.R.
(Ed.): Petroleum geology of Northwest Europe. Proceedings of
the 4th Conference. Geol. Soc., London 317335.
Uchman A. 1995: Taxonomy and palaeoecology of flysch trace fos-
sils: The Marnoso-arenacea Formation and associated facies
(Miocene, Northern Apennines, Italy). Beringeria 15, 3115.
Uchman A. & Krenmayr H.G. 1995: Trace fossils from Lower Mi-
ocene (Ottnangian) molasse deposits of Upper Austria.
Paläont. Z. 69, 503524.
Uchman A. & Demírcan H. 1999: A Zoophycos group trace fossil
from Miocene Flysch in Southern Turkey: Evidence for a U-
shaped causative burrow. Ichnos 6, 4, 251259.
Zuschin M., Mandic O., Harzhauser M. & Pervesler P. 2001: Fossil
evidence for chemoautotrophic bacterial symbiosis in the thya-
sirid bivalve Thyasira michelottii from the Middle Miocene
(Badenium) of Austria. Historical Biology 15, 223234.