RESEARCH REPORTS
Deep-Sea Benthic Food Content Recorded by
Ichnofabrics: A Conceptual Model
Based on Observations from Paleogene Flysch,
Carpathians, Poland
ANDREAS WETZEL
Geologisch-Paläontologisches Institut der Universität, Bernoullistrasse 32, CH-4056 Basel, Switzerland
ALFRED UCHMAN
Institute of Geological Sciences, Jagiellonian University, Oleandry 2a; PL-30-063 Kraków, Poland
PALAIOS, 1998, V. 13, p. 533–546
Fluctuations in the input of organic matter into a deep-sea
environment can be deciphered from ichnofabrics if the con-
tinuous processes of hemipelagic sedimentation and biotur-
bation are episodically interrupted by turbidite deposition.
Below turbidites, the bioturbated zone may be preserved al-
most intact; the benthic food content in such frozen tiers can
be interpreted from the ethology, size, penetration depth,
and density of trace fossils. Especially high benthic food
content is characterized by (1) dark sediment color, (2) com-
plete bioturbation, (3) high density of burrows produced
near-surface in several levels, (4) rarity or absence of gra-
phoglyptids, and (5) deep tiers completely bioturbated by
feeding burrows having an open connection to the surface.
Oxygen-deficiency can be eliminated as a reason for the
black color of the sediment; the size, diversity, and penetra-
tion depth of the trace fossils are similar to that of modern
and fossil counterparts formed under well-oxygenated set-
tings that are not restricted in benthic food. An increase in
sedimentation rate enhances the burial of benthic food and
is indicated by the downward extension of the bioturbated
zone and by increasing burrow density. Our analysis sug-
gests that several other black shale occurrences may result
from increased organic matter input rather than from wa-
ter-column anoxia due to sluggish circulation. Our model is
supported by similarities with other deposits that contain
many frozen tiers.
INTRODUCTION
Ichnology has become an important tool for environ-
mental analysis (see review by Savrda, 1995). The most
significant advantage of trace fossils is that they are most-
ly autochthonous indicators of ecologic conditions. Envi-
ronmental changes recorded by trace fossils are detected
via analysis of ichnofabric (Bromley and Ekdale, 1986).
Ichnofabrics result from bioturbation processes that vary
in response to changes in sediment accumulation and oth-
er environmental factors (Taylor and Goldring, 1993;
Goldring, 1995). Ecologic conditions change with depth be-
low the sediment-water interface—sediment strength in-
creases, porosity and permeability decrease, organic mat-
ter becomes decomposed, and oxygen content of the pore
Copyright Q 1998, SEPM (Society for Sedimentary Geology)
533
water decreases (e.g., Bromley, 1996). As a result, the bio-
turbated zone is subdivided into a vertical sequence of
habitats occupied by different organisms. The resultant
traces define tiers. Tiering structure can be inferred based
on cross-cutting relationships of traces; in an accreting
sediment column, co-occurring traces cross-cut each other,
and trace fossils emplaced at deeper levels cross-cut pre-
existing ones emplaced at levels closer to the sediment-
water interface.
Ichnofabric analysis based on the tiering concept has
been used, for instance, to reveal the oxygenation history
of sedimentary units (e.g., Bromley and Ekdale, 1984; Sa-
vrda and Bottjer, 1986). In this way, trace fossils can pro-
vide information about the formation of black shales and
other organic-rich sediments (e.g., Savrda and Bottjer,
1991). In addition, sediment color can be a useful param-
eter; for instance, Leszczyński (1993) found a good corre-
lation between sediment color and ichnofauna. Organic-
rich, dark sediments can preserve their original signature
if organic matter is rapidly buried and organic matter de-
position on the seafloor is high (e.g., Jung et al., 1997).
Sediment color, however, is not only related to Corg con-
tent, but also to the amount of carbonate, iron sulfides,
and paleo-oxygenation (e.g., Arthur et al., 1984). Dark sed-
iments in which unbioturbated intervals alternate with
bioturbated layers are usually interpreted to indicate low-
ered oxygenation levels during the former (e.g., Savrda
and Bottjer, 1986, 1991). It is still controversial whether
input of organic matter or retarded circulation controlled
the formation of specific black shales. For instance, Ped-
ersen and Calvert (1990) proposed that the first-order con-
trol on black shale accumulation was organic matter input
rather than water-column anoxia (e.g., Demaison and
Moore, 1980).
In contrast to oxygenation, ichnofabric responses to
‘‘productivity fluctuations are virtually unknown and war-
rant further study’’ (Savrda, 1995, p. 571). This lack of
knowledge results from the fact that the primary produc-
tion ‘‘signal’’ is heavily attenuated before it reaches the
fossil record (Fig. 1). Organic matter accumulation on the
seafloor depends on primary production and the depth
and oxygen content of the water column (e.g., Stein, 1991).
Only 1–10% of the export production (i.e., the part of pri-
mary production leaving the photic zone) reaches the
deep-sea sediment surface (e.g., Suess, 1980; Betzer et al.,
0883-1351/98/0013-0533/$3.00
534
FIGURE 1—Organic matter delivery to the seafloor and its effects on
benthic activity on and within the sediment in slowly accumulating
(,10–12 cm/ka), oxygenated deep-sea settings. If export production
is set at 100%, only 1–10% of this reaches the deep-sea floor under
3–4 km of water; there, more than the half is oxidized. Hence, 0.5–
5% of the export production is buried. During diagenesis, an unknown
proportion is degraded. The processes and effects involved are de-
scribed by 1Suess (1980), Betzer et al. (1984); 2Moore and Dymond
(1988), Pfannekuche (1993), Brasier (1995a, b), Trauth (1995); 3Müller
and Suess (1979), Emerson et al. (1985), Stein (1991); and 4Wetzel
(1991).
1984). There, the proportion of organic matter buried is
mainly controlled by sedimentation rate (Müller and
Suess, 1979), benthic activity, and oxygenation (e.g., Bra-
sier, 1995a, b). Normally 30–100% of the organic matter
deposited on the deep-sea floor is decomposed (e.g., Sibuet
et al., 1984; Emerson et al., 1985; Stein, 1991).
In oxygenated environments, the abundance of benthic
organisms on or near the sediment surface is directly re-
lated to the organic matter flux to the seafloor (Moore and
Dymond, 1988; Pfannekuche, 1993; Trauth, 1995). Fur-
thermore, the higher the benthic food content, the larger
are the diameters of certain burrows (e.g., Wetzel, 1981,
1991). The thickness of the bioturbated zone, however, de-
pends on the burial of organic matter rather than input of
organic matter. Trace fossils emplaced in deeper tiers are
WETZEL & UCHMAN
preferentially preserved in the fossil record (Werner and
Wetzel, 1982). Therefore, the response of burrowing or-
ganisms in the shallow tiers to high organic matter input
generally is not clearly preserved.
The purpose of this paper is to illustrate ichnofabrics
from dark-colored, deep-sea sediments, and to demon-
strate that they can reflect high benthic food contents
probably related to organic matter input, rather than vari-
ations in other environmental factors such as oxygenation
and sedimentation rate. Turbidite sequences are suitable
for such investigations because rapid depositional events
can preserve the bioturbated zone as a ‘‘frozen profile’’
(e.g., Bromley, 1996). In such profiles, the upper tiers are
unmodified and, therefore, traces of organisms that lived
on or near the sediment surface are present and their re-
sponse to organic matter input can be interpreted (e.g.,
Gage and Tyler, 1991).
GEOLOGICAL SETTING
The Late Paleocene to Early Eocene Szczawnica For-
mation comprises about 1200 m of thin- to medium-bed-
ded flysch deposits. It occurs in the southern part of the
Magura Nappe (Krynica zone) in the Polish Carpathians
(Fig. 2). The stratigraphy and lithology of this formation
were summarized by Birkenmajer and Oszczypko (1989).
In the northern part of its outcrop belt, the Szczawnica
Formation comprises packages of conglomerates and
coarse-grained sandstones. These packages are tens of
meters thick, and are interpreted as deep-sea fan-channel
facies (Oszczypko and Pore˛bski, 1986). The sections stud-
ied are located in the southern outcrop belt of the
Szczawnica Formation. They are interpreted to have been
deposited on a lobe in an unchannelized part of a deep-sea
fan. The sand:mud ratio within the studied sections com-
monly ranges from 1:1 to 2:1. In terms of deep-sea clastic
facies, the sections studied represent facies C2.1 and C2.2
of Pickering et al. (1989), which is consistent with the in-
ferred depositional setting.
To date, the paleobathymetry of the Szczawnica For-
mation has not been estimated. Winkler (1993) calculated
a water depth of about 4 km for an Alpine flysch basin of
similar age and tectonic setting. In analogy, we assume a
water depth of 3 to 4 km for the Szczawnica Formation
(see also Winkler and Slaczka, 1994).
MATERIAL AND METHODS
The thin- to medium-bedded flysch deposits of the
Szczawnica Formation were analyzed on a bed-by-bed
scale for ichnofabrics. The fine-grained parts of turbidites
and background sediment were polished in horizontal sec-
tions millimeter by millimeter using abrasive paper in the
field. Ichnofabrics were studied on wet surfaces. The illus-
trated specimens are housed in the Institute of Geological
Sciences of the Jagiellonian University in Kraków.
Organic carbon contents were determined using a
LECO carbon analyzer. Crushed (,100 mm) material was
gradually heated in oxygen and the resultant CO2 was
measured (see Appendix).
BENTHIC FOOD AND ICHNOFABRICS 535

FIGURE 3—Schematic drawing of section A (shown in Fig. 2) studied

on a bed-by-bed scale; muddy intervals were examined at the mm-
scale in horizontal sections prepared using sandpaper. Note dark up-
per parts of several beds (Td/e black shales).

upwards into greenish to grayish shales. In turn, the

shales typically darken upward, sometimes becoming
black at the top (Figs. 3, 4). The Corg content increases from
,0.1% in gray-green parts to 0.3–0.5% in the black layers
FIGURE 2—Location of the studied sections (A and B). Map simplified
(Appendix). The upper fine-grained, dark parts represent
after Birkenmajer (1992).
hemipelagic background sediments partially mixed with

STUDIED SECTIONS
The best outcrops (sections A and B; Fig. 2) of the
Szczawnica Formation at Ła˛kcica, north of Krościenko,
were selected for ichnofabric analysis. These outcrops and
their microfauna were described by Birkenmajer et al.
(1979), Birkenmajer and Dudziak (1981), and Birkenma-
jer (1992). The general ichnology was described by Ksi-
a˛żkiewicz (1977) and Uchman (1991a, b, 1992), but a more
detailed ichnofabric analysis is presented here.
In the studied sections, the Szczawnica Formation com-
prises 5-to-35-cm-thick turbidites, each usually separated
FIGURE 4—Examples of turbidite bedding in the studied sequences
by approximately 1 cm of dark, hemipelagic background (section B, see Fig. 2). (A) General view of thin- to medium-bedded
sediments. The turbidites consist of fine- to medium- flysch deposits. Scale is 10 cm long. (B) More detailed view of some
grained calcareous sandstones. Most beds have a Bouma turbidites. Note the dark muddy layer (arrow) below the sandstones.
Tb or Tc division at the base (Fig. 3). The sandstones grade Scale is 4 cm long.
536
TABLE 1—Organic matter content of the Szczawnica Formation:
present state, estimate of the original value, and comparison with
modern environments (organic matter content is given in % Corg dry
weight).
Present value
black sediments gray-green sediments
0.3–0.6 0.1
Diagenetic loss (50–60% of the original value1)
Original value (calculated)
black sediments gray-green sediment
0.6–1.2 0.2–0.3
Comparisons with modern deep-sea sediments
South China Sea Off NW Africa
monsoonal trade wind
upwelling upwelling
water depth Corg2
Corg3
2000 m 0.6 1.2
3000 m 0.5 0.8
4000 m 0.4 0.6
1
based on compilation by Stein (1991).
2
data from Kuehl et al. (1993).
3
data from Müller (1975).
turbiditic muds from below, resulting in the grading of col-
or and organic matter content.
It is assumed that a considerable proportion of the or-
ganic content of the mudstones was decomposed at or near
the sediment surface, and later during diagenesis (Table
1). The late diagenetic loss of organic matter cannot be es-
timated. However, decomposition of organic matter within
the bioturbated zone (and somewhat below) has been stud-
ied in some detail (see, for instance, Stein, 1991). At sedi-
mentation rates of 5–10 cm/ka, 35–70% of the deposited
organic matter is decomposed. Emerson et al. (1985) found
decomposition rates of 0.1–0.001% per day at residence
times of 15 to 150 years. Consequently, a loss of 50% Corg
since deposition seems to be a reasonable estimate. Al-
though the Corg content of the Szczawnica Formation
seems low, it is comparable to modern deep-sea sediments
in upwelling areas of high productivity (Table 1). Maps
displaying reconstructed paleo-wind directions and oce-
anic currents (e.g., Parrish and Curtis, 1982) show upwell-
ing-related high primary production in the area where the
Szczawnica Formation accumulated. A high benthic food
supply is also indicated by the presence of the large fora-
minifera Aschemocella grandis which commonly occurs in
the Szczawnica Formation (W. Kuhnt, pers. comm., 1996).
To understand the relative importance of event deposi-
tion versus hemipelagic background sedimentation, the
recurrence time of turbidites was estimated. Sedimenta-
tion rate of hemipelagic material and recurrence time of
turbidites cannot be directly determined because bio-
stratigraphic resolution is too low. To get results as relia-
ble as possible, two independent estimates were made; an
estimate of turbidite frequency in relation to modern set-
tings and an estimate of burial of organic matter.
Biostratigraphic data (Birkenmajer and Dudziak, 1981)
for the formation’s base (nannoplankton zone 7) and top
(nannoplankton zone 11) equate to absolute ages of 57 and
44.5 Ma, respectively, on the time scale of Haq et al.
(1987). Based on an average of 5 turbidites per meter, the
WETZEL & UCHMAN
Szczawnica Formation contains about 6000 turbidites. If
the turbidite deposition was regular, the turbidites had a
recurrence time of about 2000 years. However, in tectoni-
cally active areas, mass movements mainly occur during
periods of falling or low sea level or enhanced tectonic ac-
tivity (e.g., Klein, 1985; Betzler et al., 1991; Winkler, 1993)
and the recurrence time of turbidites is shorter than av-
erage. For the tectonically active areas of the Sulu Sea and
Celebes Sea, recurrence time is about 500 years during
phases of turbidite deposition (Exon et al., 1981; Betzler et
al., 1991; Betzler, pers. comm., 1998) and 5 to 8 thousand
years during times of reduced turbidite activity. Accepting
this setting as a modern analog to the studied sections of
the Szczawnica Formation and taking into account the
lithologic similarities, a turbidite recurrence time of 500
years for the Szczwanica Formation seems to be a reason-
able value. It would follow then that the hemipelagic back-
ground deposits accumulated at decompacted sedimenta-
tion rates of 8–10 cm per 1000 years.
Based on the relation between Corg burial and sedimen-
tation rate as given by Müller and Suess (1979, fig. 3), and
taking a diagenetic loss of 50% of the organic matter into
account, we would infer a sedimentation rate of 2–4 cm/
1000 years (maximum 8 cm/1000 years).
Both ways of estimating the sedimentation rate of the
hemipelagic background sediments are subject to uncer-
tainties. Nonetheless, a range of 4–8 cm/1000 years seems
to be very likely.
ICHNOLOGY
Organisms living on or near the sediment surface in-
tensely mix the surface layer and mainly produce biode-
formational structures (sensu Schäfer, 1956). These or-
ganisms homogenize the record of a seasonally fluctuating
input of particles (e.g., Deuser et al., 1983). Because of the
biogenic use of organic matter, the availability of benthic
food decreases downwards. Animals living within sedi-
ments are adapted to specific environments. Some organ-
isms can change their nutrition strategies, for example,
from deposit feeding to surface grazing (e.g., Kotake, 1989;
Bromley, 1991). The oxygen content within the pore water
and the permeability of muds decline with depth in the
sediment column. Deeper-burrowing animals, therefore,
normally maintain an open connection to the seafloor.
Open, ventilated tubes extend the oxygenated zone down-
ward (Ziebis et al., 1996), as does the bioturbational mix-
ing process itself (Reimers et al., 1986). With depth, the
strength of fine-grained sediments increases as a result of
decreasing water content. In response to these factors, the
number and vertical extent of the tiers depend on environ-
mental conditions, as do the size and the penetration
depth of individual trace fossils.
Ichnofauna
The ichnofauna of the Szczawnica Formation is highly
diverse; about 35 ichnogenera typical of the Nereites ich-
nofacies have been recognized and described (Uchman,
1991a, 1992). In the studied outcrops, however, the diver-
sity of trace fossils is lower. The observed pre-depositional
and post-depositional burrows (sensu Seilacher, 1962) are
listed in Table 2 and the ichnofabric-forming trace fossils
BENTHIC FOOD AND ICHNOFABRICS
TABLE 2—Trace fossils observed in the studied section of the
Szczawnica Formation and their abundance: common [*] (.1 speci-
men per 1 m2), rare [1] (,1 specimen per 5 m2).
Pre-depositional
Desmograpton ichnosp. [1]
Helicolithus tortuosus (Ksia˛żkiewicz) [1]
Helminthopsis ichnosp. [1]
Helminthorhaphe flexuosa Uchman [1]
Megagrapton ichnosp. [1]
Paleodictyon strozzii Meneghini, P. majus Meneghini [1]
Scolicia strozzii (Savi and Meneghini) [1]
Spirophycus bicornis (Heer) [1]
Sublorenzinia pussila Ksia˛żkiewicz [1]
Urohelminthoida dertonensis (Sacco) [1]
Post-depositional
Chondrites intricatus (Brongniart) [*]
Chondrites targionii (Brongniart) [*]
Keckia annulata Glocker sensu Ksia˛żkiewicz (1977)
[5Walcottia ichnosp., Miller and Dyer, 1877] [*]
Nereites irregularis (Schafhäutl) [see Uchman, 1995] [*]
Ophiomorpha annulata (Ksia˛żkiewicz) [*]
Phycosiphon incertum Fischer-Ooster [*]
Planolites ichnosp. [*]
Rhizocorallium ichnosp. [1]
‘‘Rotundusichnium’’ zumayensis (Llarenea) [taxonomy unclear]
[*]
Scolicia prisca de Quatrefages, S. granulata (Ksia˛żkiewicz) [*]
Spirophyton ichnosp. [1]
Strobilorhaphe clavata Ksia˛żkiewicz [1]
Thalassinoides ichnosp. [1]
Tuberculichnus vagans Ksia˛żkiewicz [1]
‘‘Tubulichnium incertum’’ Ksia˛żkiewicz [*]
Zoophycos ichnosp. [*]
are shown in Figure 5. The latter traces are Chondrites,
Phycosiphon incertum, Nereites irregularis, Ophiomorpha
annulata, and, locally, ‘‘Rotundusichnium’’ zumayensis,
and more rarely Scolicia, ‘‘Tubulichnium incertum’’, and
Zoophycos. The ichnofabric-forming taxa mainly occur in
the hemipelagites, in turbiditic muds, and in the upper
sandy part of turbidites. The thickness of the totally bio-
turbated interval is usually between 3 and 5 cm, but can
be up to 10 cm, in those beds containing a dark layer.
Tiering Pattern
The vertical distribution of the trace fossils within the
beds is exemplified in Figure 6 (see also Uchman, 1991a, b,
1992). The dark, muddy top of the beds contains biodefor-
mational structures plus frequent Phycosiphon, Nereites,
and Planolites. Nereites occurs at several levels that are
millimeters apart vertically; it cross-cuts Phycosiphon. In
some beds ‘‘Rotundusichnium’’ occurs (Fig. 7A) at several
levels. All of these traces are cross-cut by Chondrites (Fig.
7B). On the tops of sandstones, Spirophyton and Zoophy-
cos are common while ‘‘Tubulichnium incertum’’ may be
abundant locally. The Tc division below is bioturbated by
Nereites (Fig. 7C) and Planolites to a varying degree. Some
Chondrites and Nereites irregularis penetrate downwards
into Tb. The lower and middle parts of the sandstones are
bioturbated to a very low degree. Post-depositional
Ophiomorpha annulata is common and Scolicia and Plan-
olites isp. occur rarely on the lower surfaces of turbidites
(Fig. 7D). Ophiomorpha obliquely penetrates the sandy
537
parts of the turbidites and continues into the muddy parts
of the underlying turbidites cross-cutting all other traces.
Unspecified sand-filled tubes that are significantly small-
er in diameter than Ophiomorpha also are observed local-
ly.
Ichnofabric and its Interpretation
The following characteristics of the sediments are the
key to interpreting environmental conditions:
(1) Lamination in fine-grained sediments is absent due to
complete bioturbation.
(2) Bioturbation is intense, and normally penetrates
down to at least the Tc interval; even the muddy upper
parts of previous beds were exploited. The decompact-
ed thickness of the completely bioturbated layer con-
sisting of hemipelagic and turbiditic sediments ex-
ceeds 10 cm.
(3) The ichnofauna of the shallow tiers consists of traces
such as Nereites, Phycosiphon, and ‘‘Rotundusich-
nium.’’ These occur in high densities at several levels.
They were produced near the surface and are current-
ly interpreted as feeding traces. Their producers effi-
ciently exploited the sediment and, hence, a relatively
high benthic food content (see Table 1) is inferred (see
Fu, 1991). The producers of these trace fossils did not
maintain an open connection to the water column. The
relatively thick upper tier, therefore, indicates that
the sediments were highly oxygenated at depth. A pos-
itive feedback between oxygenation of the sediment
and bioturbation is possible. Bioturbation by shallow-
ly burrowing organisms enhances the flux of oxygen
into the sediment (Reimers et al., 1986) and extends
the oxygenated zone downward. Hence, this increases
the habitat within which animals that lack a connec-
tion to the seafloor can thrive.
(4) Graphoglyptids are absent from the basal surface of
overlying turbidite beds that are green to black. These
traces are suggested to indicate low benthic food con-
tents (e.g., Seilacher, 1977). Observations in modern
environments seem to support this idea (Ekdale, 1980;
Gaillard, 1991). However, the environmental implica-
tions of the presence of graphoglyptids is not fully un-
derstood (Miller, 1991).
(5) The ichnofauna of the deep tiers comprises burrows
that had an open connection to the seafloor; e.g., Chon-
drites, Spirophyton, ‘‘Tubulichnium’’, and Zoophycos.
Their occurrence indicates that nutritious organic
matter was present deeper in the sediment. Further-
more, some organisms specifically exploited the mud-
dy parts of one, or several, underlying beds; these are
called deep-layer or multi-layer colonizers (Uchman,
1995; Wetzel and Uchman, 1997). They produced Scol-
icia and some types of Chondrites. Individual, sand-
filled Ophiomorpha annulata cross-cut several beds
and all other traces and are probably the most deeply
penetrating burrows. They are common even on soles
of 30-cm-thick turbidites.
(6) Burrow size is similar to that of the ichnofaunas de-
scribed from modern and fossil counterparts that oc-
cur in well-oxygenated environments not restricted in
benthic food (e.g., Wetzel, 1981; Leszczyński, 1992;
538
FIGURE 5—Schematic drawings of the trace fossils often found in the
studied sections. Scale bars represent about 1 cm. Chondrites tunnel
systems consist of a connection to the surface and numerous regularly
ramifying tunnels that form a dendritic network. Nereites is a winding
to regularly meandering, often horizontal trace, consisting of a medial
back-filled tunnel enveloped by an even to lobate zone of reworked
sediment; Helminthoida is a junior synonym of Nereites (Uchman,
1995). Ophiomorpha is composed of cyclindrical, pellet-lined tubes
that branch to form a three-dimensional network connected to the sur-
face by (sub)vertical shafts. Phycosiphon consists of a spreite ar-
ranged in an antler-like system, parallel or obliquely oriented to bed-
ding. Planolites are straight or gently curved, inclined or horizontally
oriented, cylindrical, actively filled tunnels, with diameters of 0.5–1 cm;
Palaeophycus is similar to Planolites in terms of size and geometry,
but has a lining and is passively filled. ‘‘Rotundusichnium’’ is a hori-
zontally oriented, spiral structure composed of inclined, overlapping,
ribbon-like lamellae that are a few mm wide; the whole structure is up
to 20 cm in diameter. Scolicia is a large, bilaterally symmetrical, sub-
cylindrical burrow backfill structure of meniscate lamellae that is pro-
duced by Spatangus-like sea urchins. Thalassinoides is composed of
cyclindrical tubes that branch to form a three-dimensional network
connected to the surface by (sub)vertical shafts; fill structure is often
eccentric, resulting from active fill or tube collapse. ‘‘Tubulichnium’’ is
a mostly obliquely oriented, simple tube up to 10 mm wide and filled
with muddy pellets. Zoophycos consists of backfilled spreiten around
a central axis. Open (collapsed) or actively filled marginal tubes indi-
cate a lower or higher oxygen content, respectively, in the respiration
water.
Gaillard, 1991; Olivero, 1993; Uchman, 1995). There is
no tendency for burrow size (diameter) to decrease as
the sediment becomes darker.
(7) Sediment color of the fine-grained deposits changes
from greenish-gray in the turbiditic muds to black in
the hemipelagites. This is accompanied by an increase
in the organic carbon content by a factor .3 (see Ap-
WETZEL & UCHMAN
FIGURE 6—Ichnofabrics representative of darkening-upward muddy
intervals (example from bed 56 of section A; see Fig. 3). Note high
density of near-surface feeding or grazing traces in upper, dark, mud-
dy part. Tb, Tc, Td/e refer to Bouma turbidite divisions (including hem-
ipelagic background sediment Te). Ichnofossil labels on the right refer
to examples shown in Figure 7. Color grading is due to biogenic mix-
ing of turbiditic and hemipelagic, organic-rich material.
pendix). It did not result from de-oxygenation on the
seafloor, because no ichnological trends diagnostic of
decreasing oxygen content (see e.g., Marintsch and
Finks, 1978; Savrda and Bottjer, 1986; Wetzel, 1991)
are observed. That is, there is no decrease in the ex-
tent of bioturbation, burrow size, or number of tiers,
and there is no evidence for successive disappearance
of burrows.
Given the lack of evidence for variations in benthic oxy-
genation, the upward darkening and increase in the or-
ganic matter content must have resulted from an increase
in either sedimentation rate or the input of organic mat-
ter. The role of these two factors needs to be assessed.
Role of Sedimentation Rate
Increasing sedimentation rate leads to an increased
burial of organic matter because the residence time of or-
ganics on the seafloor is lowered (e.g., Müller and Suess,
1979). In the studied deposits, two cases need to be distin-
guished; turbidite deposition and increased input of hem-
ipelagic sediment.
Turbidite deposition leads to an immediate upward
shift of the redox boundary within the sediment (Rutgers
van der Loeff, 1990) and, hence, to increased preservation
of organic matter below the new turbidite. Some of the in-
terturbidite muds darken-upward while others do not
(Fig. 3); the observed darkening-upward pattern, there-
fore, cannot be explained by turbidite deposition. Alterna-
tively to turbidite deposition, the green or black color of
the fine-grained parts of beds may reflect variation in the
degree of erosion of the top layer by turbidity currents.
The amount of erosion was estimated from the composi-
tion of the preturbidite ichnofossil assemblages preserved
BENTHIC FOOD AND ICHNOFABRICS 539

FIGURE 7—Examples of ichnofabrics in horizontal section occurring in darkening-upward, muddy upper parts of turbidites (examples from bed
56 of section A; schematic occurrence of trace fossils within such a bed is shown in Fig. 6). (A) ‘‘Rotundusichnium’’ (R), Nereites irregularis
(N), and Phycosiphon (Ph); primary sedimentary structures are absent. (B) Chondrites (C), Planolites (P), and Skolithos-like shafts; primary
sedimentary structures are absent. (C) Sand-filled tube (t), Nereites irregularis (N), and Chondrites (C); primary sedimentary structures are
absent. (D) Lower surface of the sandy part of the turbidite with Scolicia (S). Scales are 31 in A–C and 30.5 in D.

on the lower surface of the turbidites using the method of
Wetzel and Aigner (1986). In this method, erosion is esti-
mated on the basis of the missing tiers. Generally, beds
underlain by either green or black muds display traces in-
dicating limited erosion; specifically, the upper tier is pre-
served, as it is typical for distal deep-sea fan settings
(Wetzel and Aigner, 1986; Weaver and Thomson, 1993).
An increase in hemipelagic sedimentation rate can re-
sult in an increased rate of organic matter burial, provided
that sediment input does not result in a dilution effect; the
latter occurs at sedimentation rates .10–12 cm/ka (e.g.,
Einsele, 1992). The hemipelagites studied very likely ac-
cumulated at rates ,8–10 cm/ka (see above) and, thus,
were not likely subject to such dilution effects.
Role of Organic Matter Input
These deductions suggest that increases in organic car-
bon content in the studied section could have resulted
from increases in the input of organic matter or sedimen-
tation rate. These two factors are interrelated (Müller and
Suess, 1979), and determining which is responsible from
analysis of ichnofabrics seems to be impossible (Savrda,
1995). However, if turbidite deposition interrupts the con-
tinuous bioturbation process, all tiers are preserved.
Hence, a conceptual model can be produced to describe
how the tiering structure reflects organic matter input
and sedimentation rate.
Theoretically, the interplay of low and high organic
matter input and low and high sedimentation rate in hem-
ipelagic background deposits creates four end-member
scenarios for well-oxygenated deep-sea environments
(Fig. 8). This model is based on our own observations,
analyses of modern deep-sea sediments (e.g., Wetzel,
1981, 1991) and fossil deposits (e.g., Uchman, 1995), and
the findings of benthic ecologists (e.g., Jumars and Wheat-
croft, 1989).
High organic matter input leads to intense feeding ac-
tivity in a particular zone of the sediment. In conjunction
with low hemipelagic sedimentation rates, burial of organ-
ic matter is limited and feeding activity is restricted to a
near-surface zone documented by a high proportion of
grazing traces and other near-surface feeding burrows.
Because only a little organic matter becomes buried, deep-
ly penetrating burrows are small and sparse (Fig. 8A). In
contrast, high hemipelagic sedimentation rates, in con-
junction with high organic matter input, lead to high ben-
thic food availability deeper in the sediment; the result is
a vertically extended bioturbated zone in which deeply
penetrating, large burrows are common (Fig. 8C).
Low benthic food content results in sparse surface and
near-surface grazing and feeding burrows. However, gra-
phoglyptids are common because their producers are be-
lieved to be adapted to environments restricted in benthic
food (e.g., Seilacher, 1977). At low hemipelagic sedimen-
540 WETZEL & UCHMAN

FIGURE 8—Effects of variation of hemipelagic sedimentation rate and organic matter input on ichnofabrics in environments not restricted with
regard to bottom-water oxygenation. (A) Low organic matter input, low hemipelagic sedimentation rate: brown sediment color, vertical extent
of the bioturbated zone small, small-sized burrows, graphoglyptids common. (B) High organic matter input, low hemipelagic sedimentation rate:
brown to green sediment color, near-surface feeding burrows in high densities, deeply penetrating burrows usually small-sized, bioturbated
zone vertically restricted. (C) Low organic matter input, high hemipelagic sedimentation rate: dark gray to green sediment color, graphoglyptids
common, bioturbated zone vertically extended, burrows not reduced in size, reworking of pre-existing burrows common. (D) High organic matter
input, high hemipelagic sedimentation rate: dark sediment color, bioturbated zone vertically extended, burrows not reduced in size, deeply
penetrating burrows having an open connection to the seafloor, near-surface feeding burrows in high densities, reworking of pre-existing burrows
common.

tation rates (Fig. 8B), little organic matter is buried and
its abundance within the sediment is very low. Therefore,
the bioturbated zone is narrow, the tiering structure is
condensed and deep-tier burrows are generally small
(Wetzel, 1991). At high hemipelagic sedimentation rates
(Fig. 8D), organic matter is buried and burrowing of larger
animals is supported (e.g., Jumars and Wheatcroft, 1989).
A fifth case, extremely high benthic food content, is
characterized by very high proportion of biodeformational
structures. However, this type of ichnofabric is seldom ob-
served in the fossil record (Wetzel, 1981, 1991) and was
not found in the Szczawnica Formation.
On the basis of this model, the darkening-upward inter-
turbidite sediments examined in the current study can be
inferred to reflect high benthic food content and high hem-
ipelagic sedimentation rate. The surface layer is intensely
burrowed in several levels by grazing or sediment-feeding
organisms. Deeper sediment intervals were intensely bio-
turbated by larger animals, indicating that sedimentation
rate was sufficient to bury nutritious organic matter. The
common occurrence of multi- and deep-layer colonizers
also point to a high benthic food content because their pro-
ducers penetrate down to and bioturbate one or several
previous muddy intervals.
As noted above, paleogeographic reconstructions place
the depositional area of the Szczawnica Formation in an
upwelling region (Parrish and Curtis, 1982), and remains
of large benthic foraminifera are indicative of a high ben-
thic food content.
DISCUSSION
In order to evaluate our deductions and assess the valid-
ity of our model, we have examined other trace fossil as-
semblages that probably reflect high benthic food content.
The Paleozoic flysch of Menorca was studied by Orr
(1994). He observed a surface layer that is intensely bio-
turbated by Nereites and comprises ⅓ of the bioturbated
zone, which is 8–12 cm thick in its compacted state (Orr,
pers. comm., 1996). The tier below is only partially biotur-
bated in contrast to the complete bioturbation in the deep
tier of the Szczawnica Formation. The Paleozoic flysch of
Menorca is gray and does not show obvious color grading
(Orr, pers. comm., 1996). The flysch of Menorca can be
grouped within the category of high benthic food content
and intermediate sedimentation rate.
Wetzel and Uchman (1997) reported fluctuations in
benthic food content in Eocene flysch at Ganei in the Alps.
BENTHIC FOOD AND ICHNOFABRICS
Low benthic food contents are suggested by the presence
of graphoglyptids in some parts of the section. In other
parts of the section, higher benthic food contents are re-
flected in the dominance of feeding burrows produced on
or close to the sediment surface, such as Scolicia and Ner-
eites. However, Nereites does not occur at several levels, so
the benthic food content was probably lower than in the
Szczawnica Formation, and hemipelagic sedimentation
rate likely was high.
Ekdale and Mason (1988) interpreted changes in the
ethology of trace-fossil assemblages in late Paleozoic
Oquirrh Basin to be related to oxygenation. With increas-
ing oxygenation, an ichnoassemblage dominated by sub-
surface deposit-feeding structures was replaced by one
dominated by grazing traces, and then one dominated by
dwelling structures. Certainly oxygenation is a controlling
factor for these ichnofossil assemblages. However, some
black hemipelagic intervals of the Oquirrh Formation
studied by Ekdale and Mason (1988) contain large-diame-
ter traces that were produced near the surface, such as
Scalarituba (5Nereites; Uchman, 1995), Phycosiphon, and
Spirophycus. Their presence can be explained more con-
vincingly by organic matter input rather than by de-oxy-
genation.
Leszczyński (1993) investigated relationships between
sediment color and ichnofauna and explained them in
terms of varying degrees of oxygenation. However, the col-
or of modern deep-sea sediments is also related to the con-
tent and burial of organic carbon (Thomson et al., 1987);
high organic matter content leads to gray, green, and dark
colors, whereas sediments low in organic matter are red to
brown. In slowly accumulating, oxygenated, deep-sea sed-
iments, the burial of organic matter depends on sedimen-
tation rate (Müller and Suess, 1979; Stein, 1991). Hence,
the color changes described by Leszczyński (1993) can be
interpreted in terms of fluctuating sedimentation rate and
organic matter input rather than oxygenation. This sug-
gestion is supported by the ichnofossil assemblages he de-
scribed; graphoglyptids preferentially occur in red-brown
to gray sediments, whereas feeding traces of echinoids
dominate in green to black deposits. Generally, it can be
inferred that the benthic food content was lower than in
the Szczawnica Formation, based on the common occur-
rence of graphoglyptids and relative rarity of pascichnia.
However, episodic occurrence of feeding traces produced
by echinoids points to fluctuating input of organic matter.
CONCLUSIONS
(1) In slowly accumulating (,10 cm/ka), oxygenated,
deep-sea settings, the effects of sediment accumula-
tion and organic matter input on the burial of organic
matter normally cannot be differentiated when study-
ing ichnocoenoses because deeply penetrating bur-
rows generally dominate the ichnofabric. However, in-
formation about benthic food content can be interpret-
ed from ichnofabrics if the near-surface layer is pre-
served. This layer stores information on the organisms
that first respond to changes in organic matter flux.
The tiering structure of the bioturbated zone is pre-
served if the continuous process of bioturbation is in-
terrupted by event deposits such as turbidites. How-
ever, the surface layer is preserved only where the tur-
541
bidity currents caused little or no erosion of pre-exist-
ing sediments.
(2) Dark to black color of sediments may be indicative of
oxygen deficiency of bottom waters. However, de-oxy-
genation events can be recognized by ichnofabric anal-
ysis. There is an ordered disappearance of ichnotaxa,
accompanied by a decrease in the extent of bioturba-
tion, the size of burrows, the number of tiers, and the
penetration depth of endobenthic organisms. These
criteria are not evident in the dark-colored Szczawnica
Formation deposits.
(3) Alternatively, we hypothesize that dark sediment col-
or may be caused by high Corg content and is unrelated
to benthic oxygen levels. Criteria indicative of high
benthic food content include a high degree of biotur-
bation, a large number of tiers, deep penetration of
burrows, the presence of relatively large trace fossils,
and the presence or the absence of specific groups of
trace fossils corresponding to specific nutrition strate-
gies. Presence of surface grazing traces and absence of
graphoplyptids reflects a high organic supply.
(4) On the basis of these criteria, complemented by anal-
ysis of other modern and fossil deposits, a conceptual
model is developed for deep-sea sediments. This model
is applicable to hemipelagic deposits that accumulated
at rates ,10 cm/ka in settings where oxygen was not a
limiting factor. Four end-member ichnofabrics are
typical of combinations of low and high benthic food
content and low and high hemipelagic sedimentation
rates. High benthic food content at the sediment sur-
face supports an abundance of grazing and other near-
surface feeding burrows. If hemipelagic sedimentation
rate is low, little organic matter becomes buried for
use deeper in the sediment and, hence, deeply pene-
trating burrows are sparse. In contrast, high hemipe-
lagic sedimentation rates result in availability of ben-
thic food deeper in the sediment; the product is a ver-
tically extended bioturbated zone in which deeply pen-
etrating traces occur. At low benthic food content,
surficial and near-surface grazing and feeding bur-
rows are sparse, whereas graphoglyptids are common.
At low hemipelagic sedimentation rates, deep tiers are
characterized by small-sized burrows, the bioturbated
zone is narrow, and the tiering structure is condensed.
At high hemipelagic sedimentation rates, large-sized
traces occur in deep tiers.
(5) Low sedimentation rates lead to red to brown sedi-
ments whereas high sedimentation rates give rise to
green or gray to black sediments. In the studied case,
intervals completely bioturbated by pascichnia show a
darkening-upward pattern and a concomitant in-
crease in the Corg content. Furthermore, the deep tiers
also display strong bioturbation. The rarity or absence
of graphoglyptids, and the presence of ‘‘Tubulichnium
incertum’’ and ‘‘Rotundusichnium’’ zumayensis, are
probably typical for sediments with high benthic food
content in Upper Cretaceous-Paleocene flysch depos-
its.
(6) Comparison with other ichnofossil assemblages from
flysch environments support our model. Future appli-
cations of this model may aid in the interpretation of
dark, organic rich sediments. For example, it is possi-
ble that several presumed ‘‘anoxic events’’ may, in-
542
stead, be related to an increased input of organic mat-
ter to the deep-sea floor, rather than water-column an-
oxia.
ACKNOWLEDGMENTS
R.G. Bromley (Copenhagen, Denmark), P. Orr (Bristol,
U.K.), and two anonymous journal reviewers critically
read the manuscript and made helpful suggestions. P. Orr
additionally provided information about the flysch of Me-
norca. W. Kuhnt (Kiel, FRG) explained some new ideas in
the ecology of the encountered microfauna. C. Betzler
(Frankfurt, FRG) provided additional information about
the turbidite deposition in the Sulu Sea and Celebes Sea.
A.A. Ekdale (Salt Lake City, Utah) demonstrated Spanish
Fork Canyon (Oquirrh Formation) outcrops during the
Second International Ichnofabric Workshop (1993). W.
Stern (Basel) provided chemical analyses. This study was
made possible by support from the Schweizerische Natio-
nalfond zur Förderung der Wissenschaftlichen Forschung
(Swiss National Science Foundation; grant No. 7 PLPJ041
522) and travel funds provided by the universities of Kra-
ków and Basel in terms of their cooperation contract. All
these contributions are gratefully acknowledged.
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ACCEPTED MARCH 31, 1998
544 WETZEL & UCHMAN

APPENDIX
Sediment color, organic carbon contents, and ichnology of selected beds from Section A. The trace fossils are listed in alphabetical
order. The columns refer from left to right to the lower, middle, and upper parts of the bioturbated zone. Please note that the organic
carbon content of the top layer (right column) controlled the ichnofaunal composition.

Bed 40a
Color gray dark gray green
Corg (%) 0.08 0.13 0.24
Bioturbation
degree ca. 80% 100% 100%
homogeneous ca. 40% ca. 40% ca. 50%
traces Chondrites Chondrites Chondrites
Planolites Planolites Phycosiphon
Planolites
meniscate burrow
Remarks deep-colonizing Chondrites, vertical shafts, small sand-filled tubes, ‘‘Tubulichnium’’, Scolicia at the
base of the overlying bed, Aschemocella present
Bed 40b
Color gray green black
Corg (%) 0.1 0.18 0.38
Bioturbation
degree ca. 80% 100% 100%
homogeneous ca. 40% ca. 50% ca. 60%
traces Chondrites Chondrites Chondrites
Planolites Nereites Nereites
Phycosiphon Phycosiphon
Planolites Planolites
Remarks deep-colonizing Chondrites, small sand-filled tubes, Thalassinoides, vertical shafts, Aschemocella
present
Bed 55a
Color gray green black
Corg (%) 0.1 0.22 0.41
Bioturbation
degree ca. 80% 100% 100%
homogeneous ca. 30% ca. 30% ca. 40%
traces Chondrites Chondrites Chondrites
Nereites Hydrancylus Nereites
Palaeophycus Nereites Phycosiphon
Planolites Planolites Planolites
‘‘Tubulichnium’’ ‘‘Tubulichnium’’
Remarks deep-colonizing Chondrites, sand-filled tubes, meniscate burrows, Rhizocorallium, Scolicia at the
base of overlying bed, Aschemocella present
Bed 55b
Color gray green dark-gray to black
Corg (%) 0.1 0.19 0.34
Bioturbation
degree ca. 80% 100% 100%
homogeneous ca. 40% ca. 30% ca. 40%
traces Chondrites Chondrites Chondrites
Phycosiphon Nereites Nereites
Planolites Phycosiphon Phycosiphon
Thalassinoides Planolites Planolites
Remarks deep-colonizing Chondrites, sand-filled tubes, ‘‘Tubulichnium’’, small vertical shafts, Aschemocella
present
Bed 55c
Color gray green dark-gray to black
Corg (%) ,0.1 0.18 0.32
Bioturbation
degree ca. 80% 100% 100%
homogeneous ca. 30% ca. 50% ca. 60%
traces Chondrites Chondrites Chondrites
Nereites Phycosiphon Hydrancylus
Phycosiphon Nereites Phycosiphon
Planolites Planolites Nereites
stuffed tubes
Remarks deep-colonizing Chondrites, vertical shafts, sand-filled tubes, ‘‘Tubulichnium’’, Aschemocella present
BENTHIC FOOD AND ICHNOFABRICS 545

APPENDIX
Continued.

Bed 55d
Color gray green black
Corg (%) 0.09 0.20 0.29
Bioturbation
degree ca. 70% 100% 100%
homogeneous ca. 30% ca. 50% ca. 60%
traces Chondrites Chondrites Chondrites
Nereites Nereites Nereites
Phycosiphon Phycosiphon Planolites
Planolites Planolites ‘‘Rotundusichnium’’
‘‘Rotundusichnium’’
Remarks deep-colonizing Chondrites, sand-filled tubes, ‘‘Tubulichnium’’; Ophiomorpha at the base of sandy
part; Aschemocella present
Bed 56a
Color gray green black
Corg (%) 0.1 0.18 0.54
Bioturbation
degree ca. 80% 100% 100%
homogeneous ca. 30% ca. 50% ca. 60%
traces Chondrites Chondrites Chondrites
Nereites Nereites Nereites
Palaeophycus Phycosiphon Phycosiphon
Planolites Planolites
‘‘Rotundusichnium’’ ‘‘Rotunsudichnium’’
Remarks deep-colonizing Chondrites, Thalassinoides, sand-filled tubes, ‘‘Tubulichnium’’, Scolicia at the base
of succeeding bed, Ophiomorpha at the base of sandy part, Aschemocella present
Bed 56b
Color gray green black
Corg (%) 0.1 0.14 0.38
Bioturbation
degree ca. 80% 100% 100%
homogeneous ca. 30% ca. 50% ca. 60%
traces Chondrites Chondrites Chondrites
Nereites Nereites Nereites
Planolites Phycosiphon Phycosiphon
Planolites Planolites
‘‘Tubulichnium’’
Remarks deep-colonizing Chondrites, vertical shafts
Bed 56c
Color gray green black
Corg (%) 0.08 0.25 0.39
Bioturbation
degree ca. 80% 100% 100%
homogeneous ca. 30% ca. 50% ca. 50%
traces Chondrites Chondrites Chondrites
Nereites Nereites Nereites
Planolites Phycosiphon Phycosiphon
Planolites ‘‘Rotundusichnium’’
‘‘Rotundusichnium’’
Remarks deep-colonizing Chondrites, vertical shafts
Bed 57a
Color gray green black
Corg (%) 0.1 0.23 0.41
Bioturbation
degree ca. 80% 100% 100%
homogeneous ca. 30% ca. 50% ca. 60%
traces Chondrites Chondrites Chondrites
Nereites Nereites (small) Nereites
Planolites Planolites ‘‘Rotundusichnium’’
Thalassinoides ‘‘Rotundusichnium’’
Remarks deep-colonizing Chondrites, ‘‘Tubulichnium’’
546 WETZEL & UCHMAN

APPENDIX
Continued.

Bed 57b
Color gray gray-green green
Corg (%) 0.1 0.18 0.2 8
Bioturbation
degree ca. 60% 100% 100%
homogeneous ca. 30% ca. 50% ca. 70%
traces Chondrites Chondrites Chondrites
Planolites Phycosiphon Phycosiphon
Planolites
Remarks deep-colonizing Chondrites, vertical shafts, meniscate burrows, ‘‘Tubulichnium’’
Bed 57c
Color gray gray-green black
Corg (%) 0.1 0.15 0.24
Bioturbation
degree ca. 70% 100% 100%
homogeneous ca. 30% ca. 50% ca. 60%
traces Chondrites Chondrites Chondrites
Nereites Nereites Nereites
Phycosiphon Phycosiphon Phycosiphon
Planolites Planolites Planolites
‘‘Rotundusichnium’’ ‘‘Rotundusichnium’’
Remarks deep-colonizing Chondrites, vertical shafts, sand-filled tubes, Aschemocella present
Bed 58a
Color gray gray-green green
Corg (%) 0.1 0.18 0.29
Bioturbation
degree ca. 80% 100% 100%
homogeneous ca. 30% ca. 50% ca. 60%
traces Chondrites Chondrites Chondrites
Planolites Nereites Nereites
Planolites Phycosiphon
Planolites
Remarks deep-colonizing Chondrites, sand-filled tubes, Scolicia at the base of succeeding bed, Ophiomorpha
at the base of sandy part
Bed 58b
Color gray green black
Corg (%) 0.1 0.23 0.37
Bioturbation
degree ca. 80% 100% 100%
homogeneous ca. 30% ca. 50% ca. 60%
traces Chondrites Chondrites Chondrites
Nereites Nereites Nereites
Palaeophycus Planolites Phycosiphon
Planolites ‘‘Rotundusichnium’’ Planolites
‘‘Rotundusichnium’’
Remarks deep-colonizing Chondrites, sand-filled tubes, ‘‘Tubulichnium’’, Scolicia and Ophiomorpha at the
base of sandy part, Aschemocella present
Bed 58c
Color gray gray-green green
Corg (%) 0.1 0.18 0.29
Bioturbation
degree ca. 80% 100% 100%
homogeneous ca. 30% ca. 50% ca. 60%
traces Chondrites Chondrites Chondrites
Palaeophycus Nereites Nereites
Planolites Planolites Planolites
Remarks deep-colonizing Chondrites, sand-filled tubes, ‘‘Tubulichnium’’