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RESEARCH REPORTS 533

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 water decreases (e.g., Bromley, 1996). As a result, the bio-
turbated zone is subdivided into a vertical sequence of
Fluctuations in the input of organic matter into a deep-sea habitats occupied by different organisms. The resultant
environment can be deciphered from ichnofabrics if the con- traces define tiers. Tiering structure can be inferred based
tinuous processes of hemipelagic sedimentation and biotur- on cross-cutting relationships of traces; in an accreting
bation are episodically interrupted by turbidite deposition. sediment column, co-occurring traces cross-cut each other,
Below turbidites, the bioturbated zone may be preserved al- and trace fossils emplaced at deeper levels cross-cut pre-
most intact; the benthic food content in such frozen tiers can existing ones emplaced at levels closer to the sediment-
be interpreted from the ethology, size, penetration depth, water interface.
and density of trace fossils. Especially high benthic food Ichnofabric analysis based on the tiering concept has
content is characterized by (1) dark sediment color, (2) com- been used, for instance, to reveal the oxygenation history
plete bioturbation, (3) high density of burrows produced of sedimentary units (e.g., Bromley and Ekdale, 1984; Sa-
near-surface in several levels, (4) rarity or absence of gra- vrda and Bottjer, 1986). In this way, trace fossils can pro-
phoglyptids, and (5) deep tiers completely bioturbated by vide information about the formation of black shales and
feeding burrows having an open connection to the surface. other organic-rich sediments (e.g., Savrda and Bottjer,
Oxygen-deficiency can be eliminated as a reason for the 1991). In addition, sediment color can be a useful param-
black color of the sediment; the size, diversity, and penetra- eter; for instance, Leszczyński (1993) found a good corre-
tion depth of the trace fossils are similar to that of modern lation between sediment color and ichnofauna. Organic-
and fossil counterparts formed under well-oxygenated set- rich, dark sediments can preserve their original signature
tings that are not restricted in benthic food. An increase in if organic matter is rapidly buried and organic matter de-
sedimentation rate enhances the burial of benthic food and position on the seafloor is high (e.g., Jung et al., 1997).
is indicated by the downward extension of the bioturbated Sediment color, however, is not only related to Corg con-
zone and by increasing burrow density. Our analysis sug- tent, but also to the amount of carbonate, iron sulfides,
gests that several other black shale occurrences may result and paleo-oxygenation (e.g., Arthur et al., 1984). Dark sed-
from increased organic matter input rather than from wa- iments in which unbioturbated intervals alternate with
ter-column anoxia due to sluggish circulation. Our model is bioturbated layers are usually interpreted to indicate low-
supported by similarities with other deposits that contain ered oxygenation levels during the former (e.g., Savrda
many frozen tiers. 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-
INTRODUCTION
trol on black shale accumulation was organic matter input
Ichnology has become an important tool for environ- rather than water-column anoxia (e.g., Demaison and
mental analysis (see review by Savrda, 1995). The most Moore, 1980).
significant advantage of trace fossils is that they are most- In contrast to oxygenation, ichnofabric responses to
ly autochthonous indicators of ecologic conditions. Envi- ‘‘productivity fluctuations are virtually unknown and war-
ronmental changes recorded by trace fossils are detected rant further study’’ (Savrda, 1995, p. 571). This lack of
via analysis of ichnofabric (Bromley and Ekdale, 1986). knowledge results from the fact that the primary produc-
Ichnofabrics result from bioturbation processes that vary tion ‘‘signal’’ is heavily attenuated before it reaches the
in response to changes in sediment accumulation and oth- fossil record (Fig. 1). Organic matter accumulation on the
er environmental factors (Taylor and Goldring, 1993; seafloor depends on primary production and the depth
Goldring, 1995). Ecologic conditions change with depth be- and oxygen content of the water column (e.g., Stein, 1991).
low the sediment-water interface—sediment strength in- Only 1–10% of the export production (i.e., the part of pri-
creases, porosity and permeability decrease, organic mat- mary production leaving the photic zone) reaches the
ter becomes decomposed, and oxygen content of the pore deep-sea sediment surface (e.g., Suess, 1980; Betzer et al.,
Copyright Q 1998, SEPM (Society for Sedimentary Geology) 0883-1351/98/0013-0533/$3.00
534 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-
FIGURE 1—Organic matter delivery to the seafloor and its effects on monly ranges from 1:1 to 2:1. In terms of deep-sea clastic
benthic activity on and within the sediment in slowly accumulating
(,10–12 cm/ka), oxygenated deep-sea settings. If export production facies, the sections studied represent facies C2.1 and C2.2
is set at 100%, only 1–10% of this reaches the deep-sea floor under of Pickering et al. (1989), which is consistent with the in-
3–4 km of water; there, more than the half is oxidized. Hence, 0.5– ferred depositional setting.
5% of the export production is buried. During diagenesis, an unknown To date, the paleobathymetry of the Szczawnica For-
proportion is degraded. The processes and effects involved are de- mation has not been estimated. Winkler (1993) calculated
scribed by 1Suess (1980), Betzer et al. (1984); 2Moore and Dymond
(1988), Pfannekuche (1993), Brasier (1995a, b), Trauth (1995); 3Müller
a water depth of about 4 km for an Alpine flysch basin of
and Suess (1979), Emerson et al. (1985), Stein (1991); and 4Wetzel similar age and tectonic setting. In analogy, we assume a
(1991). water depth of 3 to 4 km for the Szczawnica Formation
(see also Winkler and Slaczka, 1994).

1984). There, the proportion of organic matter buried is


mainly controlled by sedimentation rate (Müller and MATERIAL AND METHODS
Suess, 1979), benthic activity, and oxygenation (e.g., Bra-
sier, 1995a, b). Normally 30–100% of the organic matter The thin- to medium-bedded flysch deposits of the
deposited on the deep-sea floor is decomposed (e.g., Sibuet Szczawnica Formation were analyzed on a bed-by-bed
et al., 1984; Emerson et al., 1985; Stein, 1991). scale for ichnofabrics. The fine-grained parts of turbidites
In oxygenated environments, the abundance of benthic and background sediment were polished in horizontal sec-
organisms on or near the sediment surface is directly re- tions millimeter by millimeter using abrasive paper in the
lated to the organic matter flux to the seafloor (Moore and field. Ichnofabrics were studied on wet surfaces. The illus-
Dymond, 1988; Pfannekuche, 1993; Trauth, 1995). Fur- trated specimens are housed in the Institute of Geological
thermore, the higher the benthic food content, the larger Sciences of the Jagiellonian University in Kraków.
are the diameters of certain burrows (e.g., Wetzel, 1981, Organic carbon contents were determined using a
1991). The thickness of the bioturbated zone, however, de- LECO carbon analyzer. Crushed (,100 mm) material was
pends on the burial of organic matter rather than input of gradually heated in oxygen and the resultant CO2 was
organic matter. Trace fossils emplaced in deeper tiers are 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 WETZEL & UCHMAN

TABLE 1—Organic matter content of the Szczawnica Formation: Szczawnica Formation contains about 6000 turbidites. If
present state, estimate of the original value, and comparison with the turbidite deposition was regular, the turbidites had a
modern environments (organic matter content is given in % Corg dry
weight).
recurrence time of about 2000 years. However, in tectoni-
cally active areas, mass movements mainly occur during
Present value
periods of falling or low sea level or enhanced tectonic ac-
tivity (e.g., Klein, 1985; Betzler et al., 1991; Winkler, 1993)
black sediments gray-green sediments
0.3–0.6 0.1 and the recurrence time of turbidites is shorter than av-
erage. For the tectonically active areas of the Sulu Sea and
Diagenetic loss (50–60% of the original value1)
Celebes Sea, recurrence time is about 500 years during
Original value (calculated)
phases of turbidite deposition (Exon et al., 1981; Betzler et
black sediments gray-green sediment al., 1991; Betzler, pers. comm., 1998) and 5 to 8 thousand
0.6–1.2 0.2–0.3
years during times of reduced turbidite activity. Accepting
Comparisons with modern deep-sea sediments this setting as a modern analog to the studied sections of
South China Sea Off NW Africa the Szczawnica Formation and taking into account the
monsoonal trade wind lithologic similarities, a turbidite recurrence time of 500
upwelling upwelling
water depth Corg2
Corg3 years for the Szczwanica Formation seems to be a reason-
able value. It would follow then that the hemipelagic back-
2000 m 0.6 1.2
ground deposits accumulated at decompacted sedimenta-
3000 m 0.5 0.8
4000 m 0.4 0.6 tion rates of 8–10 cm per 1000 years.
Based on the relation between Corg burial and sedimen-
1
based on compilation by Stein (1991). tation rate as given by Müller and Suess (1979, fig. 3), and
2
data from Kuehl et al. (1993). taking a diagenetic loss of 50% of the organic matter into
3
data from Müller (1975). 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
turbiditic muds from below, resulting in the grading of col- hemipelagic background sediments are subject to uncer-
or and organic matter content. tainties. Nonetheless, a range of 4–8 cm/1000 years seems
It is assumed that a considerable proportion of the or- to be very likely.
ganic content of the mudstones was decomposed at or near
the sediment surface, and later during diagenesis (Table ICHNOLOGY
1). The late diagenetic loss of organic matter cannot be es-
timated. However, decomposition of organic matter within Organisms living on or near the sediment surface in-
the bioturbated zone (and somewhat below) has been stud- tensely mix the surface layer and mainly produce biode-
ied in some detail (see, for instance, Stein, 1991). At sedi- formational structures (sensu Schäfer, 1956). These or-
mentation rates of 5–10 cm/ka, 35–70% of the deposited ganisms homogenize the record of a seasonally fluctuating
organic matter is decomposed. Emerson et al. (1985) found input of particles (e.g., Deuser et al., 1983). Because of the
decomposition rates of 0.1–0.001% per day at residence biogenic use of organic matter, the availability of benthic
times of 15 to 150 years. Consequently, a loss of 50% Corg food decreases downwards. Animals living within sedi-
since deposition seems to be a reasonable estimate. Al- ments are adapted to specific environments. Some organ-
though the Corg content of the Szczawnica Formation isms can change their nutrition strategies, for example,
seems low, it is comparable to modern deep-sea sediments from deposit feeding to surface grazing (e.g., Kotake, 1989;
in upwelling areas of high productivity (Table 1). Maps Bromley, 1991). The oxygen content within the pore water
displaying reconstructed paleo-wind directions and oce- and the permeability of muds decline with depth in the
anic currents (e.g., Parrish and Curtis, 1982) show upwell- sediment column. Deeper-burrowing animals, therefore,
ing-related high primary production in the area where the normally maintain an open connection to the seafloor.
Szczawnica Formation accumulated. A high benthic food Open, ventilated tubes extend the oxygenated zone down-
supply is also indicated by the presence of the large fora- ward (Ziebis et al., 1996), as does the bioturbational mix-
minifera Aschemocella grandis which commonly occurs in ing process itself (Reimers et al., 1986). With depth, the
the Szczawnica Formation (W. Kuhnt, pers. comm., 1996). strength of fine-grained sediments increases as a result of
To understand the relative importance of event deposi- decreasing water content. In response to these factors, the
tion versus hemipelagic background sedimentation, the number and vertical extent of the tiers depend on environ-
recurrence time of turbidites was estimated. Sedimenta- mental conditions, as do the size and the penetration
tion rate of hemipelagic material and recurrence time of depth of individual trace fossils.
turbidites cannot be directly determined because bio-
stratigraphic resolution is too low. To get results as relia- Ichnofauna
ble as possible, two independent estimates were made; an
estimate of turbidite frequency in relation to modern set- The ichnofauna of the Szczawnica Formation is highly
tings and an estimate of burial of organic matter. diverse; about 35 ichnogenera typical of the Nereites ich-
Biostratigraphic data (Birkenmajer and Dudziak, 1981) nofacies have been recognized and described (Uchman,
for the formation’s base (nannoplankton zone 7) and top 1991a, 1992). In the studied outcrops, however, the diver-
(nannoplankton zone 11) equate to absolute ages of 57 and sity of trace fossils is lower. The observed pre-depositional
44.5 Ma, respectively, on the time scale of Haq et al. and post-depositional burrows (sensu Seilacher, 1962) are
(1987). Based on an average of 5 turbidites per meter, the listed in Table 2 and the ichnofabric-forming trace fossils
BENTHIC FOOD AND ICHNOFABRICS 537

TABLE 2—Trace fossils observed in the studied section of the parts of the turbidites and continues into the muddy parts
Szczawnica Formation and their abundance: common [*] (.1 speci- of the underlying turbidites cross-cutting all other traces.
men per 1 m2), rare [1] (,1 specimen per 5 m2).
Unspecified sand-filled tubes that are significantly small-
er in diameter than Ophiomorpha also are observed local-
Pre-depositional
ly.
Desmograpton ichnosp. [1]
Helicolithus tortuosus (Ksia˛żkiewicz) [1]
Helminthopsis ichnosp. [1] Ichnofabric and its Interpretation
Helminthorhaphe flexuosa Uchman [1]
Megagrapton ichnosp. [1] The following characteristics of the sediments are the
Paleodictyon strozzii Meneghini, P. majus Meneghini [1]
Scolicia strozzii (Savi and Meneghini) [1]
key to interpreting environmental conditions:
Spirophycus bicornis (Heer) [1]
(1) Lamination in fine-grained sediments is absent due to
Sublorenzinia pussila Ksia˛żkiewicz [1]
Urohelminthoida dertonensis (Sacco) [1] complete bioturbation.
(2) Bioturbation is intense, and normally penetrates
Post-depositional
down to at least the Tc interval; even the muddy upper
Chondrites intricatus (Brongniart) [*]
parts of previous beds were exploited. The decompact-
Chondrites targionii (Brongniart) [*]
Keckia annulata Glocker sensu Ksia˛żkiewicz (1977) ed thickness of the completely bioturbated layer con-
[5Walcottia ichnosp., Miller and Dyer, 1877] [*] sisting of hemipelagic and turbiditic sediments ex-
Nereites irregularis (Schafhäutl) [see Uchman, 1995] [*] ceeds 10 cm.
Ophiomorpha annulata (Ksia˛żkiewicz) [*] (3) The ichnofauna of the shallow tiers consists of traces
Phycosiphon incertum Fischer-Ooster [*] such as Nereites, Phycosiphon, and ‘‘Rotundusich-
Planolites ichnosp. [*]
nium.’’ These occur in high densities at several levels.
Rhizocorallium ichnosp. [1]
‘‘Rotundusichnium’’ zumayensis (Llarenea) [taxonomy unclear] They were produced near the surface and are current-
[*] ly interpreted as feeding traces. Their producers effi-
Scolicia prisca de Quatrefages, S. granulata (Ksia˛żkiewicz) [*] ciently exploited the sediment and, hence, a relatively
Spirophyton ichnosp. [1] high benthic food content (see Table 1) is inferred (see
Strobilorhaphe clavata Ksia˛żkiewicz [1] Fu, 1991). The producers of these trace fossils did not
Thalassinoides ichnosp. [1]
maintain an open connection to the water column. The
Tuberculichnus vagans Ksia˛żkiewicz [1]
‘‘Tubulichnium incertum’’ Ksia˛żkiewicz [*] relatively thick upper tier, therefore, indicates that
Zoophycos ichnosp. [*] 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
are shown in Figure 5. The latter traces are Chondrites, into the sediment (Reimers et al., 1986) and extends
Phycosiphon incertum, Nereites irregularis, Ophiomorpha the oxygenated zone downward. Hence, this increases
annulata, and, locally, ‘‘Rotundusichnium’’ zumayensis, the habitat within which animals that lack a connec-
and more rarely Scolicia, ‘‘Tubulichnium incertum’’, and tion to the seafloor can thrive.
Zoophycos. The ichnofabric-forming taxa mainly occur in (4) Graphoglyptids are absent from the basal surface of
the hemipelagites, in turbiditic muds, and in the upper overlying turbidite beds that are green to black. These
sandy part of turbidites. The thickness of the totally bio- traces are suggested to indicate low benthic food con-
turbated interval is usually between 3 and 5 cm, but can tents (e.g., Seilacher, 1977). Observations in modern
be up to 10 cm, in those beds containing a dark layer. environments seem to support this idea (Ekdale, 1980;
Gaillard, 1991). However, the environmental implica-
Tiering Pattern tions of the presence of graphoglyptids is not fully un-
derstood (Miller, 1991).
The vertical distribution of the trace fossils within the (5) The ichnofauna of the deep tiers comprises burrows
beds is exemplified in Figure 6 (see also Uchman, 1991a, b, that had an open connection to the seafloor; e.g., Chon-
1992). The dark, muddy top of the beds contains biodefor- drites, Spirophyton, ‘‘Tubulichnium’’, and Zoophycos.
mational structures plus frequent Phycosiphon, Nereites, Their occurrence indicates that nutritious organic
and Planolites. Nereites occurs at several levels that are matter was present deeper in the sediment. Further-
millimeters apart vertically; it cross-cuts Phycosiphon. In more, some organisms specifically exploited the mud-
some beds ‘‘Rotundusichnium’’ occurs (Fig. 7A) at several dy parts of one, or several, underlying beds; these are
levels. All of these traces are cross-cut by Chondrites (Fig. called deep-layer or multi-layer colonizers (Uchman,
7B). On the tops of sandstones, Spirophyton and Zoophy- 1995; Wetzel and Uchman, 1997). They produced Scol-
cos are common while ‘‘Tubulichnium incertum’’ may be icia and some types of Chondrites. Individual, sand-
abundant locally. The Tc division below is bioturbated by filled Ophiomorpha annulata cross-cut several beds
Nereites (Fig. 7C) and Planolites to a varying degree. Some and all other traces and are probably the most deeply
Chondrites and Nereites irregularis penetrate downwards penetrating burrows. They are common even on soles
into Tb. The lower and middle parts of the sandstones are of 30-cm-thick turbidites.
bioturbated to a very low degree. Post-depositional (6) Burrow size is similar to that of the ichnofaunas de-
Ophiomorpha annulata is common and Scolicia and Plan- scribed from modern and fossil counterparts that oc-
olites isp. occur rarely on the lower surfaces of turbidites cur in well-oxygenated environments not restricted in
(Fig. 7D). Ophiomorpha obliquely penetrates the sandy benthic food (e.g., Wetzel, 1981; Leszczyński, 1992;
538 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


FIGURE 5—Schematic drawings of the trace fossils often found in the
seafloor, because no ichnological trends diagnostic of
studied sections. Scale bars represent about 1 cm. Chondrites tunnel decreasing oxygen content (see e.g., Marintsch and
systems consist of a connection to the surface and numerous regularly Finks, 1978; Savrda and Bottjer, 1986; Wetzel, 1991)
ramifying tunnels that form a dendritic network. Nereites is a winding are observed. That is, there is no decrease in the ex-
to regularly meandering, often horizontal trace, consisting of a medial tent of bioturbation, burrow size, or number of tiers,
back-filled tunnel enveloped by an even to lobate zone of reworked and there is no evidence for successive disappearance
sediment; Helminthoida is a junior synonym of Nereites (Uchman,
1995). Ophiomorpha is composed of cyclindrical, pellet-lined tubes
of burrows.
that branch to form a three-dimensional network connected to the sur- Given the lack of evidence for variations in benthic oxy-
face by (sub)vertical shafts. Phycosiphon consists of a spreite ar- genation, the upward darkening and increase in the or-
ranged in an antler-like system, parallel or obliquely oriented to bed-
ding. Planolites are straight or gently curved, inclined or horizontally ganic matter content must have resulted from an increase
oriented, cylindrical, actively filled tunnels, with diameters of 0.5–1 cm; in either sedimentation rate or the input of organic mat-
Palaeophycus is similar to Planolites in terms of size and geometry, ter. The role of these two factors needs to be assessed.
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 Role of Sedimentation Rate
to 20 cm in diameter. Scolicia is a large, bilaterally symmetrical, sub-
cylindrical burrow backfill structure of meniscate lamellae that is pro- Increasing sedimentation rate leads to an increased
duced by Spatangus-like sea urchins. Thalassinoides is composed of burial of organic matter because the residence time of or-
cyclindrical tubes that branch to form a three-dimensional network ganics on the seafloor is lowered (e.g., Müller and Suess,
connected to the surface by (sub)vertical shafts; fill structure is often 1979). In the studied deposits, two cases need to be distin-
eccentric, resulting from active fill or tube collapse. ‘‘Tubulichnium’’ is guished; turbidite deposition and increased input of hem-
a mostly obliquely oriented, simple tube up to 10 mm wide and filled ipelagic sediment.
with muddy pellets. Zoophycos consists of backfilled spreiten around
a central axis. Open (collapsed) or actively filled marginal tubes indi- Turbidite deposition leads to an immediate upward
cate a lower or higher oxygen content, respectively, in the respiration shift of the redox boundary within the sediment (Rutgers
water. 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
Gaillard, 1991; Olivero, 1993; Uchman, 1995). There is (Fig. 3); the observed darkening-upward pattern, there-
no tendency for burrow size (diameter) to decrease as fore, cannot be explained by turbidite deposition. Alterna-
the sediment becomes darker. tively to turbidite deposition, the green or black color of
(7) Sediment color of the fine-grained deposits changes the fine-grained parts of beds may reflect variation in the
from greenish-gray in the turbiditic muds to black in degree of erosion of the top layer by turbidity currents.
the hemipelagites. This is accompanied by an increase The amount of erosion was estimated from the composi-
in the organic carbon content by a factor .3 (see Ap- 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 Theoretically, the interplay of low and high organic
Wetzel and Aigner (1986). In this method, erosion is esti- matter input and low and high sedimentation rate in hem-
mated on the basis of the missing tiers. Generally, beds ipelagic background deposits creates four end-member
underlain by either green or black muds display traces in- scenarios for well-oxygenated deep-sea environments
dicating limited erosion; specifically, the upper tier is pre- (Fig. 8). This model is based on our own observations,
served, as it is typical for distal deep-sea fan settings analyses of modern deep-sea sediments (e.g., Wetzel,
(Wetzel and Aigner, 1986; Weaver and Thomson, 1993). 1981, 1991) and fossil deposits (e.g., Uchman, 1995), and
An increase in hemipelagic sedimentation rate can re- the findings of benthic ecologists (e.g., Jumars and Wheat-
sult in an increased rate of organic matter burial, provided croft, 1989).
that sediment input does not result in a dilution effect; the High organic matter input leads to intense feeding ac-
latter occurs at sedimentation rates .10–12 cm/ka (e.g., tivity in a particular zone of the sediment. In conjunction
Einsele, 1992). The hemipelagites studied very likely ac- with low hemipelagic sedimentation rates, burial of organ-
cumulated at rates ,8–10 cm/ka (see above) and, thus, ic matter is limited and feeding activity is restricted to a
were not likely subject to such dilution effects. near-surface zone documented by a high proportion of
grazing traces and other near-surface feeding burrows.
Role of Organic Matter Input Because only a little organic matter becomes buried, deep-
These deductions suggest that increases in organic car- ly penetrating burrows are small and sparse (Fig. 8A). In
bon content in the studied section could have resulted contrast, high hemipelagic sedimentation rates, in con-
from increases in the input of organic matter or sedimen- junction with high organic matter input, lead to high ben-
tation rate. These two factors are interrelated (Müller and thic food availability deeper in the sediment; the result is
Suess, 1979), and determining which is responsible from a vertically extended bioturbated zone in which deeply
analysis of ichnofabrics seems to be impossible (Savrda, penetrating, large burrows are common (Fig. 8C).
1995). However, if turbidite deposition interrupts the con- Low benthic food content results in sparse surface and
tinuous bioturbation process, all tiers are preserved. near-surface grazing and feeding burrows. However, gra-
Hence, a conceptual model can be produced to describe phoglyptids are common because their producers are be-
how the tiering structure reflects organic matter input lieved to be adapted to environments restricted in benthic
and sedimentation rate. 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 As noted above, paleogeographic reconstructions place
its abundance within the sediment is very low. Therefore, the depositional area of the Szczawnica Formation in an
the bioturbated zone is narrow, the tiering structure is upwelling region (Parrish and Curtis, 1982), and remains
condensed and deep-tier burrows are generally small of large benthic foraminifera are indicative of a high ben-
(Wetzel, 1991). At high hemipelagic sedimentation rates thic food content.
(Fig. 8D), organic matter is buried and burrowing of larger
animals is supported (e.g., Jumars and Wheatcroft, 1989). DISCUSSION
A fifth case, extremely high benthic food content, is
characterized by very high proportion of biodeformational In order to evaluate our deductions and assess the valid-
structures. However, this type of ichnofabric is seldom ob- ity of our model, we have examined other trace fossil as-
served in the fossil record (Wetzel, 1981, 1991) and was semblages that probably reflect high benthic food content.
not found in the Szczawnica Formation. The Paleozoic flysch of Menorca was studied by Orr
On the basis of this model, the darkening-upward inter- (1994). He observed a surface layer that is intensely bio-
turbidite sediments examined in the current study can be turbated by Nereites and comprises ⅓ of the bioturbated
inferred to reflect high benthic food content and high hem- zone, which is 8–12 cm thick in its compacted state (Orr,
ipelagic sedimentation rate. The surface layer is intensely pers. comm., 1996). The tier below is only partially biotur-
burrowed in several levels by grazing or sediment-feeding bated in contrast to the complete bioturbation in the deep
organisms. Deeper sediment intervals were intensely bio- tier of the Szczawnica Formation. The Paleozoic flysch of
turbated by larger animals, indicating that sedimentation Menorca is gray and does not show obvious color grading
rate was sufficient to bury nutritious organic matter. The (Orr, pers. comm., 1996). The flysch of Menorca can be
common occurrence of multi- and deep-layer colonizers grouped within the category of high benthic food content
also point to a high benthic food content because their pro- and intermediate sedimentation rate.
ducers penetrate down to and bioturbate one or several Wetzel and Uchman (1997) reported fluctuations in
previous muddy intervals. benthic food content in Eocene flysch at Ganei in the Alps.
BENTHIC FOOD AND ICHNOFABRICS 541

Low benthic food contents are suggested by the presence bidity currents caused little or no erosion of pre-exist-
of graphoglyptids in some parts of the section. In other ing sediments.
parts of the section, higher benthic food contents are re- (2) Dark to black color of sediments may be indicative of
flected in the dominance of feeding burrows produced on oxygen deficiency of bottom waters. However, de-oxy-
or close to the sediment surface, such as Scolicia and Ner- genation events can be recognized by ichnofabric anal-
eites. However, Nereites does not occur at several levels, so ysis. There is an ordered disappearance of ichnotaxa,
the benthic food content was probably lower than in the accompanied by a decrease in the extent of bioturba-
Szczawnica Formation, and hemipelagic sedimentation tion, the size of burrows, the number of tiers, and the
rate likely was high. penetration depth of endobenthic organisms. These
Ekdale and Mason (1988) interpreted changes in the criteria are not evident in the dark-colored Szczawnica
ethology of trace-fossil assemblages in late Paleozoic Formation deposits.
Oquirrh Basin to be related to oxygenation. With increas- (3) Alternatively, we hypothesize that dark sediment col-
ing oxygenation, an ichnoassemblage dominated by sub- or may be caused by high Corg content and is unrelated
surface deposit-feeding structures was replaced by one to benthic oxygen levels. Criteria indicative of high
dominated by grazing traces, and then one dominated by benthic food content include a high degree of biotur-
dwelling structures. Certainly oxygenation is a controlling bation, a large number of tiers, deep penetration of
factor for these ichnofossil assemblages. However, some burrows, the presence of relatively large trace fossils,
black hemipelagic intervals of the Oquirrh Formation and the presence or the absence of specific groups of
studied by Ekdale and Mason (1988) contain large-diame- trace fossils corresponding to specific nutrition strate-
ter traces that were produced near the surface, such as gies. Presence of surface grazing traces and absence of
Scalarituba (5Nereites; Uchman, 1995), Phycosiphon, and graphoplyptids reflects a high organic supply.
Spirophycus. Their presence can be explained more con- (4) On the basis of these criteria, complemented by anal-
vincingly by organic matter input rather than by de-oxy- ysis of other modern and fossil deposits, a conceptual
genation. model is developed for deep-sea sediments. This model
Leszczyński (1993) investigated relationships between is applicable to hemipelagic deposits that accumulated
sediment color and ichnofauna and explained them in at rates ,10 cm/ka in settings where oxygen was not a
terms of varying degrees of oxygenation. However, the col- limiting factor. Four end-member ichnofabrics are
or of modern deep-sea sediments is also related to the con- typical of combinations of low and high benthic food
tent and burial of organic carbon (Thomson et al., 1987); content and low and high hemipelagic sedimentation
high organic matter content leads to gray, green, and dark rates. High benthic food content at the sediment sur-
colors, whereas sediments low in organic matter are red to face supports an abundance of grazing and other near-
brown. In slowly accumulating, oxygenated, deep-sea sed- surface feeding burrows. If hemipelagic sedimentation
iments, the burial of organic matter depends on sedimen- rate is low, little organic matter becomes buried for
tation rate (Müller and Suess, 1979; Stein, 1991). Hence, use deeper in the sediment and, hence, deeply pene-
the color changes described by Leszczyński (1993) can be trating burrows are sparse. In contrast, high hemipe-
interpreted in terms of fluctuating sedimentation rate and lagic sedimentation rates result in availability of ben-
organic matter input rather than oxygenation. This sug- thic food deeper in the sediment; the product is a ver-
gestion is supported by the ichnofossil assemblages he de- tically extended bioturbated zone in which deeply pen-
scribed; graphoglyptids preferentially occur in red-brown etrating traces occur. At low benthic food content,
to gray sediments, whereas feeding traces of echinoids surficial and near-surface grazing and feeding bur-
dominate in green to black deposits. Generally, it can be rows are sparse, whereas graphoglyptids are common.
inferred that the benthic food content was lower than in At low hemipelagic sedimentation rates, deep tiers are
the Szczawnica Formation, based on the common occur- characterized by small-sized burrows, the bioturbated
rence of graphoglyptids and relative rarity of pascichnia. zone is narrow, and the tiering structure is condensed.
However, episodic occurrence of feeding traces produced At high hemipelagic sedimentation rates, large-sized
by echinoids points to fluctuating input of organic matter. traces occur in deep tiers.
(5) Low sedimentation rates lead to red to brown sedi-
CONCLUSIONS ments whereas high sedimentation rates give rise to
green or gray to black sediments. In the studied case,
(1) In slowly accumulating (,10 cm/ka), oxygenated, intervals completely bioturbated by pascichnia show a
deep-sea settings, the effects of sediment accumula- darkening-upward pattern and a concomitant in-
tion and organic matter input on the burial of organic crease in the Corg content. Furthermore, the deep tiers
matter normally cannot be differentiated when study- also display strong bioturbation. The rarity or absence
ing ichnocoenoses because deeply penetrating bur- of graphoglyptids, and the presence of ‘‘Tubulichnium
rows generally dominate the ichnofabric. However, in- incertum’’ and ‘‘Rotundusichnium’’ zumayensis, are
formation about benthic food content can be interpret- probably typical for sediments with high benthic food
ed from ichnofabrics if the near-surface layer is pre- content in Upper Cretaceous-Paleocene flysch depos-
served. This layer stores information on the organisms its.
that first respond to changes in organic matter flux. (6) Comparison with other ichnofossil assemblages from
The tiering structure of the bioturbated zone is pre- flysch environments support our model. Future appli-
served if the continuous process of bioturbation is in- cations of this model may aid in the interpretation of
terrupted by event deposits such as turbidites. How- dark, organic rich sediments. For example, it is possi-
ever, the surface layer is preserved only where the tur- ble that several presumed ‘‘anoxic events’’ may, in-
542 WETZEL & UCHMAN

stead, be related to an increased input of organic mat- BROMLEY, R.G., 1996, Trace Fossils—Biology, Taphonomy and Appli-
ter to the deep-sea floor, rather than water-column an- cations (2nd edition): Chapman and Hall, London, 361 p.
BROMLEY, R.G., and EKDALE, A.A., 1984, Chondrites: A trace fossil in-
oxia. dicator of anoxia in sediments: Science, v. 224, p. 872–874.
BROMLEY, R.G., and EKDALE, A.A., 1986, Composite ichnofabrics and
ACKNOWLEDGMENTS tiering of burrows: Geological Magazine, v. 123, p. 59–65.
DEMAISON, G.J., and MOORE, G.T., 1980, Anoxic environments and oil
source bed genesis: American Association of Petroleum Geologists
R.G. Bromley (Copenhagen, Denmark), P. Orr (Bristol, Bulletin, v. 64, p. 1179–1209.
U.K.), and two anonymous journal reviewers critically DEUSER, W.G., BREWER, P.G., JICKELLS, T.D., and COMMEAU, R.F.,
read the manuscript and made helpful suggestions. P. Orr 1983, Biological control of the removal of abiogenic particles from
additionally provided information about the flysch of Me- the surface ocean: Science, v. 219, p. 388–391.
norca. W. Kuhnt (Kiel, FRG) explained some new ideas in EINSELE, G., 1992, Sedimentary Basins: Springer-Verlag, Berlin,
the ecology of the encountered microfauna. C. Betzler 628 p.
EKDALE, A.A., 1980, Graphoglyptid burrows in modern deep-sea sed-
(Frankfurt, FRG) provided additional information about iments: Science, v. 207, p. 304–306.
the turbidite deposition in the Sulu Sea and Celebes Sea. EKDALE, A.A., and MASON, T.R., 1988, Characteristic trace-fossil as-
A.A. Ekdale (Salt Lake City, Utah) demonstrated Spanish sociations in oxygen-poor sedimentary environments: Geology, v.
Fork Canyon (Oquirrh Formation) outcrops during the 16, p. 720–723.
Second International Ichnofabric Workshop (1993). W. EMERSON, S., FISCHER, K., REIMERS, C., and HEGGIE, D., 1985, Organ-
Stern (Basel) provided chemical analyses. This study was ic carbon dynamics and preservation in deep-sea sediments: Deep-
Sea Research, v. 32, p. 1–21.
made possible by support from the Schweizerische Natio- EXON, N.F., HAAKE, F.-W., HARTMANN, M., KÖGLER, F.-C., MÜLLER,
nalfond zur Förderung der Wissenschaftlichen Forschung P.J., and WHITICAR, M.J., 1981, Morphology, water characteristics
(Swiss National Science Foundation; grant No. 7 PLPJ041 and sedimentation in the silled Sulu Sea, southeast Asia: Marine
522) and travel funds provided by the universities of Kra- Geology, v. 39, p. 165–195.
ków and Basel in terms of their cooperation contract. All FU, S., 1991, Funktion, Verhalten, und Einteilung fucoider und lo-
these contributions are gratefully acknowledged. phocteniider Lebensspuren: Courier Forschungs-Institut Senck-
enberg, v. 135, 79 p.
GAGE, J.D., and TYLER, P.A., 1991, Deep-Sea Biology: Cambridge
REFERENCES University Press, Cambridge, 504 p.
GAILLARD, C., 1991, Recent organism traces and ichnofacies on the
deep-sea floor off New Caledonia, southwestern Pacific: PA-
ARTHUR, M.A., DEAN, W.E., and STOW, D.A.V., 1984, Models for the
deposition of Mesozoic-Cenozoic fine-grained organic-carbon-rich LAIOS, v. 6, p. 302–315.
sediment in the deep-sea: in Stow, D.A.V., and Piper, D.J.W., eds., GOLDRING, R., 1995, Organisms and the substrate: Response and ef-
Fine-Grained Sediments: Deep-Water Processes and Facies: Geo- fect: in Bosence, D.J.W., and Allison, P.A., eds., Marine Palaeoen-
logical Society of London Special Publication No. 15, p. 527–560. vironmental Analysis from Fossils: Geological Society of London
BETZER, P.R., SHOWERS, W.J., LAWS, E.A., WINN, C.D., DITULLIO, Special Publication No. 83, p. 151–180.
G.R., and KROOPNICK, P.M., 1984, Primary production and parti- HAQ, B.U., HARDENBOL, J., and VAIL, P.R., 1987, Chronology of fluc-
cle fluxes on a transect of the equator at 1538W in the Pacific tuating sea levels since the Triassic: Science, v. 235, p. 1156–1167.
Ocean: Deep-Sea Research, v. 31, p. 1–12. JUMARS, P.A., and WHEATCROFT, R.A., 1989, Responses of benthos to
BETZLER, C., NIEDERBRAGT, A.J., and NICHOLS, G.J., 1991, Signifi- changing food quality and quantity, with a focus on deposit feed-
cance of turbidites at Site 767 (Celebes Sea) and Site 768 (Sulu ing and bioturbation: in Berger, W.H., Smetacek, V.S., and Wefer,
Sea): in Silver, E.A., Rangin, C. et al., eds., Proceedings of the G., eds., Productivity of the Ocean: Present and Past: J. Wiley,
Ocean Drilling Project, Scientific Results, v. 124: Ocean Drilling New York, p. 235–253.
Program, College Station, p. 431–446. JUNG, M., ILMBERGER, J., MANGINI, A., and EMEIS, K.-C., 1997, Why
BIRKENMAJER, K., 1992, Stop A.1.5. La˛kcica. Poludniowe obrzeżenie some Mediterranean sapropels survived burn-down (and others
plaszczowiny magurskiej: in Przewodnik 63 Zjazdu Polskiego To- did not): Marine Geology, v. 141, p. 51–60.
warzystwa Geologicznego (63rd Annual Meeting of the Polish KLEIN, G. DEVRIES, 1985, The frequency and periodicity of preserved
Geological Society), p. 50–54. turbidites in submarine fans as a quantitative record of tectonic
BIRKENMAJER, K., and DUDZIAK, J., 1981, Wiek fliszu magurskiego uplift in collision zones: Tectonophysics, v. 119, p. 181–193.
(Paleogen) pólnocnego obrzeżenia pienińskiego pasa skalkowego KOTAKE, N., 1989, Paleoecology of the Zoophycos producers: Lethaia,
w Polsce na podstawie nannoplanktonu: Studia Geologica Poloni- v. 22, p. 327–341.
ca, v. 70, p. 7–34. KUEHL, S.A., FUGLSETH, T.J., and THUNELL, R.C., 1993, Sediment
BIRKENMAJER, K., and OSZCZYPKO, N., 1989, Cretaceous and Paleo- mixing and accumulation rates in the Sulu Sea and South China
gene lithostratigraphic units of the Magura Nappe, Krynica Sub- Seas: Implications for organic carbon preservation in deep-sea en-
unit, Carpathians: Annales Societatis Geologorum Poloniae, v. 59, vironments: Marine Geology, v. 111, p. 15–35.
p. 145–181. KSIA˛ŻKIEWICZ, M., 1977, Trace fossils in the flysch of the Polish Car-
BIRKENMAJER, K., DUDZIAK, J., and JEDNOROWSKA, A., 1979, Wgle˛b- pathians: Palaeontologia Polonica, v. 36, p. 1–208.
na budowa geologiczna pólocnej strefy dyslokacyjnej pienińskiego LESZCZYŃSKI, S., 1992, Controls on trace fossil distribution in flysch
pasa skalkowego w Szczawnicy: Studia Geologica Polonica, v. 61, deposits: Uniwersytet Jagiellońskii Rozprawy Habilitacyjne No.
p. 9–36. 236: Nakladem Uniwersytetu Jagiellońskiego, Kraków, 88 p.
BRASIER, M.D., 1995a, Fossil indicators of nutrient levels. 1: Eutro- LESZCZYŃSKI, S., 1993, Ichnocoenosis versus sediment colour in Up-
phication and climate change: in Bosence, D.W., and Allison, P.A., per Albian to Lower Eocene turbidites, Guipúzcoa province, north-
eds., Marine Palaeoenvironmental Analysis from Fossils: Geolog- ern Spain: Palaeogeography, Palaeoeclimatology, Palaeoecology,
ical Society of London Special Publication No. 83, p. 113–132. v. 100, p. 251–265.
BRASIER, M.D., 1995b, Fossil indicators of nurient levels. 2: Evolution MARINTSCH, E.J., and FINKS, R.M., 1978, Zoophycos size may indicate
and extinction in relation to oligotrophy: in Bosence, D.W., and Al- environmental gradients: Lethaia, v. 11, p. 273–279.
lison, P.A., eds., Marine Palaeoenvironmental Analysis from Fos- MILLER, S.A., and DYER, C.B., 1878, Contributions to paleontology.
sils: Geological Society of London Special Publication No. 83, p. 1.2.: Journal of the Society of Natural History of Cincinnati, v. 1, p.
133–150. 24–39.
BROMLEY, R.G., 1991, Zoophycos: Strip mine, refuse dump, cache or MILLER, W., III, 1991, Paleoecology of graphoglyptids: Ichnos, v. 1, p.
sewage farm?: Lethaia, v. 24, p. 460–462. 305–312.
BENTHIC FOOD AND ICHNOFABRICS 543

MOORE, W.S., and DYMOND, J., 1988, Correlation of 210-Pb removal SUESS, E., 1980, Particulate organic carbon flux in the oceans—Sur-
with organic carbon fluxes in the Pacific Ocean: Nature, v. 331, p. face productivity and oxygen utilization: Nature, v. 288, p. 260–
339–341. 263.
MÜLLER, P.J., 1975, Diagenese stickstoffhaltiger organischer Sub- TAYLOR, A.M., and GOLDRING, R., 1993, Description and analysis of
stanzen in oxischen und anoxischen marinen Sedimenten. ‘‘Mete- bioturbation and ichnofabric: Journal of the Geological Society
or’’-Forschungsergebnisse, Reihe C, v. 22: Schweizerbart-Verlag, London, v. 150, p. 141–148.
Stuttgart, p. 1–60. THOMSON, J., COLLEY, S., HIGGS, N.C., HYDES, D.J., WILSON, T.R.S.,
MÜLLER, P.J., and SUESS, E., 1979, Productivity, sedimentation rate and SOERENSEN, J., 1987, Geochemical oxidation fronts in NE At-
and sedimentary organic matter in the oceans. I. Organic carbon lantic distal turbidites and their effects in the sedimentary record:
preservation: Deep-Sea Research, v. 26, p. 1347–1362. in Weaver, P.P.E., and Thomson, J., eds., Geology and Geochem-
OLIVERO, D., 1993, L’ichnofacies a Zoophycos et sa signification pa- istry of Abyssal Plains: Geological Society of London Special Pub-
leoenvironnementale. Mise au point d’un outil bio-sedimentaire: lication No. 31, p. 167–177.
Unpublished PhD Thesis, L’Université Claude Bernard—Lyon I, TRAUTH, M.H., 1995, Bioturbate Signalverzerrung hochauflösender
270 p. paläoozeanographischer Zeitreihen: Berichte—Reports, Geolo-
ORR, P.J., 1994, Trace fossil tiering within event beds and preserva- gisch-Paläontologisches Institut der Universität Kiel No. 74,
tion of frozen profiles: An example from the Lower Carboniferous 167 p.
of Menorca: PALAIOS, v. 9, p. 202–210. UCHMAN, A., 1991a, Skamienialości śladowe z warst inoceramowych
OSZCZYPKO, N., and PORE˛BSKI, S., 1986, Wycieczka B.15. Potok Ży- oraz utworów formacji szczawnickiej (fm) w strefie krynickiej i
czanowski: in Birkenmajer, K., and Poprawa, D., eds., Przewodnik bystrzyckiej plaszczowiny magurskiej (Trace fossils of the Inocer-
57 Zjazdu Polskiego Towarzystwa Geologicznego (57th Annual amian beds and the Szczawnica Formation in the Krynica and
Meeting of the Polish Geological Society), p. 120–122. Bystrzyca Zones of the Magura Nappe): Przegla˛d Geologiczny, v.
PARRISH, J.T., and CURTIS, R.L., 1982, Atmospheric circulation, up- 4, p.207–212.
welling, and organic-rich rocks in the Mesozoic and Cenozoic: Pa- UCHMAN, A., 1991b, Diversified tiering patterns in Paleogene flysch
laeogeography, Palaeoclimatology, Palaeoecology, v. 40, p. 31–66. trace fossils of the Magura nappe, Carpathians Mountains, Po-
PEDERSEN, T.F., and CALVERT, S.E., 1990, Anoxia vs. productivity: land: Ichnos, v. 1, p. 287–292.
What controls the formation of organic-carbon-rich sediments and UCHMAN, A., 1992, Stop A.1.5. Ła˛kcica. Ichnofauna formacji szczawn-
sedimentary rocks: American Association of Petroleum Geologists ickiej w Ła˛kcicy: in Przewodnik 63 Zjazdu Polskiego Towarzystwa
Bulletin, v. 74, p. 454–466. Geologicznego (63rd Annual Meeting of the Polish Geological So-
PFANNEKUCHE, O., 1993, Benthic response to the sedimentation of ciety), p. 55.
particulate organic matter at the BIOTRANS station 478N 208W: UCHMAN, A., 1995, Taxonomy and palaeoecology of flysch trace fos-
Deep-Sea Research, v. 40, p. 135–149. sils: The Marnoso-arenacea Formation and associated facies (Mio-
PICKERING, K.T., HISCOTT, R.N., and HEIN, F.J., 1989, Deep Marine cene, Northern Apennines, Italy): Beringeria, v. 15, p. 3–115.
Environments: Clastic Sedimentation and Tectonics: Unwin Hy- WEAVER, P.P.E., and THOMSON, J., 1993, Calculating erosion by deep-
man, London, 416 p. sea turbidity currents during initiation and flow: Nature, v. 364, p.
REIMERS, C.E., FISCHER, K.M., MEREWETHER, R., SMITH, K.L., JR., 136–138.
and JAHNKE, R.A., 1986, Oxygen microprofiles measured in situ in WERNER, F., and WETZEL, A., 1982, Interpretation of biogenic struc-
deep ocean sediments: Nature, v. 320, p. 741–744. tures in oceanic sediments: Bulletin de’l Institut Géologie du Bas-
RUTGERS VAN DER LOEFF, M.M., 1990, Oxygen in pore waters of deep- sin d’Aquitaine, v. 31, p. 275–288.
sea sediments: Philosophical Transactions of the Royal Society WETZEL, A., 1981, Ökologische und stratigraphische Bedeutung bio-
London, Series A, v. 331, p.69–84. gener Gefüge in quartären Sedimenten am NW-afrikanischen
SAVRDA, C.E., 1995, Ichnologic applications in paleoceanographic, pa- Kontinentalrand: ‘‘Meteor’’ Forschungs-Ergebnisse, Reihe C, v.
leoclimatic, and sea-level studies: PALAIOS, v. 10, p. 565–577. 34: Schweizerbart-Verlag, Stuttgart, p. 1–47.
SAVRDA, C.E., and BOTTJER, D.J., 1986, Trace-fossil model for recon- WETZEL, A., 1991, Ecologic interpretation of deep-sea trace fossil com-
munities: Palaeogeography, Palaeoclimatology, Palaeoecology, v.
struction of paleo-oxygenation in bottom waters: Geology, v. 4, p.
85, p. 47–69.
3–6.
WETZEL, A., and AIGNER, T., 1986, Stratigraphic completeness:
SAVRDA, C.E., and BOTTJER, D.J. 1991, Redox-related benthic events:
Tiered trace fossils provide a measuring stick: Geology, v. 14, p.
in Einsele, G., Seilacher, A., and Ricken, W., eds., Cycles and
234–237.
Events in Stratigraphy: Springer-Verlag, Berlin, p. 525–541.
WETZEL, A., and UCHMAN, A., 1997, Ichnology of deep-sea fan over-
SCHÄFER, W., 1956, Wirkungen der Benthos-Organismen auf den
bank deposits of the Ganei Slates (Eocene, Switzerland)—A clas-
jungen Schichtverband: Senckenbergiana lethaea, v. 37, p. 183– sical flysch trace fossil locality first studied by Oswald Heer: Ich-
263. nos, v. 5, 139–162.
SEILACHER, A., 1962, Paleontological studies on turbidite sedimenta- WINKLER, W., 1993, Control factors on turbidite sedimentation in a
tion and erosion: Journal of Geology, v. 70, p. 227–234. deep-sea trench setting—The example of the Schlieren Flysch
SEILACHER, A., 1977, Pattern analysis of Paleodictyon and related (Upper Maastrichtian-Lower Eocene, Central Switzerland): Geo-
trace fossils: in Crimes, T.P., and Harper, J.C., eds., Trace Fossils dinamica Acta, v. 6, p. 81–102.
2: Geological Journal Special Issue No. 9, p. 289–334. WINKLER, W., and SLACZKA, A., 1994, A Late Cretaceous to Paleogene
SIBUET, M., MONNIOT, C. DESBRUYÈRES, D., DINET, A., KHRIPOUNOFF, geodynamic model for the western Carpathians in Poland: Geolo-
A., ROWE, G., and SEGONZAC, M., 1984, Peuplements benthiques gica Carpathica, v. 45, p. 71–82.
et caractéristiques trophiques du milieu dans la plaine abyssale de ZIEBIS, W., FORSTER, S., HUETTEL, M., and JOERGENSEN, B.B., 1996,
Demerara dans l’Océan Atlantique: Oceanologica Acta, v. 7, p. Complex burrows of the mud shrimp Callianassa truncata and
345–358. their geochemical impact in the sea bed: Nature, v. 382, p. 619–
STEIN, R., 1991, Accumulation of Organic Carbon in Marine Sedi- 622.
ments: Lecture Notes in Earth Sciences No. 34: Springer-Verlag,
Berlin, 217 p. 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’’