ELSEVIER Palaeogeography, Palaeoclimatology, Palaeoecology 140 (1998) 367–382

Trace fossil analysis of lacustrine facies and basins

Luis A. Buatois Ł , M. Gabriela Mángano
Kansas Geological Survey, 1930 Constant Avenue, Lawrence, Kansas 66047, USA
Accepted 18 January 1997

Abstract

Two ichnofacies are typical of lacustrine depositional systems. The Scoyenia ichnofacies characterizes transitional
terrestrial=nonmarine aquatic substrates, periodically inundated or desiccated, and therefore is commonly present in lake
margin facies. The Mermia ichnofacies is associated with well oxygenated, permanent subaqueous, fine-grained substrates
of hydrologically open, perennial lakes. Bathymetric zonations within the Mermia ichnofacies are complicated by the wide
variability of lacustrine systems. Detected proximal–distal trends are useful within particular lake basins, but commonly
difficult to extrapolate to other lakes. Other potential ichnofacies include the typically marine Skolithos ichnofacies for
high-energy zones of lakes and substrate-controlled, still unnamed ichnofacies, associated to lake margin deposits. Trace
fossils are useful for sedimentologic analysis of event beds. Lacustrine turbidites are characterized by low-diversity suites,
reflecting colonization by opportunistic organisms after the turbidite event. Underflow current beds record animal activity
contemporaneous with nearly continuous sedimentation. Ichnologic studies may also help to distinguish between marine
and lacustrine turbidites. Deep-marine turbidites host the Nereites ichnofacies that consists of high diversity of ornate
grazing traces and graphoglyptids, recording highly specialized feeding strategies developed to solve the problem of the
scarcity of food in the deep sea. Deep lacustrine environments contain the Mermia ichnofacies, which is dominated by
unspecialized grazing and feeding traces probably related to the abundance and accessibility of food in lacustrine systems.
The lower diversity of lacustrine ichnofaunas in comparison with deep-sea assemblages more likely reflects lower species
diversity as a consequence of less stable conditions. Increase of depth and extent of bioturbation through geologic time
produced a clear signature in the ichnofabric record of lacustrine facies. Paleozoic lacustrine ichnofaunas are typically
dominated by surface trails with little associated bioturbation. During the Mesozoic, bioturbation depth was higher in lake
margin facies than in fully lacustrine deposits. While significant degrees of bioturbation were attained in lake margin facies
during the Triassic, major biogenic disruption of primary bedding in subaqueous lacustrine deposits did not occur until the
Cretaceous.  1998 Elsevier Science B.V. All rights reserved.

Keywords: trace fossils; lakes; ichnofacies; ichnofabrics; sedimentology; palaeolimnology

1. Introduction tional systems. However, our understanding of the

Trace fossils represent an important component
of nonmarine sedimentary successions and deposi-
Ł Corresponding author. Fax: C1 913 864 5317; E-mail:
buagano@kgs.ukans.edu
ichnology of nonmarine environments still lags be-
hind our understanding of trace fossils in the marine
stratigraphic record. Eight marine ichnofacies have
been formally defined, namely the Psilonichnus,
Skolithos, Cruziana, Zoophycos, Nereites, Teredo-
lites, Glossifungites and Trypanites ichnofacies, the

0031-0182/98/$19.00 c 1998 Elsevier Science B.V. All rights reserved.

PII S 0 0 3 1 - 0 1 8 2 ( 9 8 ) 0 0 0 2 0 - 0
368 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 140 (1998) 367–382

latter being further subdivided into the Gnathich-
nus and Entobia ichnofacies by Bromley and As-
gaard (1993). In contrast, only one recurrent assem-
blage, the Scoyenia ichnofacies, has been widely
recognized in nonmarine environments (Seilacher,
1967; Frey and Seilacher, 1980; Pemberton et al.,
1992a). However, our knowledge of nonmarine ich-
nology is growing rapidly, and several studies have
documented the trace fossil content of lacustrine
facies and sequences (e.g. Bromley and Asgaard,
1979; Pollard et al., 1982; Walker, 1985; Gier-
lowski-Kordesch, 1991; Pickerill, 1992; Buatois and
Mángano, 1993a; Metz, 1995; Buatois et al., 1996a).
Furthermore, a nonmarine ichnofacies model con-
sisting of three recurrent associations, Termitichnus,
Scoyenia and Mermia, has been proposed (Buatois
and Mángano, 1995a).
This paper reviews recent work on the ichnol-
ogy of lake deposits, emphasizing the significance
of detailed trace fossil analysis in the recognition
and characterization of lacustrine ichnofacies, the
ichnologic signatures of lake event stratigraphy, the
strengths and limitations of ichnofabric analysis, and
the use of ichnology in the delineation of specific
facies and environments.
2. Nonmarine ichnofacies models
2.1. Background
Ichnofacies are trace fossil assemblages that re-
cur through geologic time and are characteristic of
certain environmental conditions (Seilacher, 1967;
Frey and Pemberton, 1984; Pemberton et al., 1992a).
Only a single archetypical association, the Scoye-
nia ichnofacies, is widely recognized in nonma-
rine settings (Seilacher, 1963, 1967). However, the
Scoyenia ichnofacies is only one of the recurrent
trace fossil assemblages of continental environments
(Frey and Pemberton, 1987; Maples and Archer,
1989; Gierlowski-Kordesch, 1991; Pickerill, 1992;
Hasiotis and Bown, 1992; Buatois and Mángano,
1993a; MacNaughton and Pickerill, 1995). Buatois
and Mángano (1995a) reviewed the current status of
nonmarine associations and proposed the existence
of three archetypical assemblages: the Termitichnus,
Scoyenia, and Mermia ichnofacies (Fig. 1). The Ter-
mitichnus ichnofacies was introduced by Smith et al.
(1993) as a subdivision of the Scoyenia ichnofacies.
However, in the model by Buatois and Mángano
(1995a), the three ichnofacies were considered at

Fig. 1. Model of nonmarine ichnofacies. Typical ichnogenera include: (1) Termitichnus, (2) Scaphichnium, (3) Celliforma, (4) Edaphich-
nium, (5) Coprinisphaera, (6) Krausichnus, (7) Vondrichnus, (8) Scoyenia, (9) Rusophycus, (10) Beaconites, (11) Merostomichnites, (12)
Umfolozia, (13) Cruziana, (14) Taenidium, (15) tetrapod tracks, (16) Diplichnites, (17) Fuersichnus, (18) Mermia, (19) Helminthopsis,
(20) Tuberculichnus, (21) Maculichna, (22) Vagorichnus, (23) Helminthoidichnites, (24) Cochlichnus, (25) Undichna, (26) Palaeophycus,
(27) Circulichnis, (28) Gordia, (29) Treptichnus, (30) Planolites (modified from Buatois and Mángano, 1995a).
 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 140 (1998) 367–382
the same hierarchical level. The Termitichnus ich-
nofacies characterizes subaerial, fully terrestrial en-
vironments and will not be discussed in this paper.
Discussion is focused on the remaining two ichno-
facies that are associated with lacustrine deposits:
the Scoyenia ichnofacies, characterizing transitional
fluvio-lacustrine environments, and the Mermia ich-
nofacies, characterizing fully lacustrine settings (Bu-
atois and Mángano, 1995a).
2.2. Scoyenia ichnofacies
Seilacher (1967) proposed the Scoyenia ichno-
facies for “non-marine sands and shales, often red
beds, with a distinctive association of trace fos-
sils” and referred to a previous illustration of this
ichnofauna (Seilacher, 1963; Fig. 7), which in-
cluded meniscate burrows, arthropod trackways, and
bilobed traces, as well as several physical sedimen-
tary structures.
Subsequent workers tended to use the Scoye-
nia ichnofacies as a catchall for all assemblages of
nonmarine trace fossils, and as a consequence the
concept lost its original meaning (Frey et al., 1984).
Trying to solve this problem, Frey et al. (1984)
suggested a three-fold definition of the Scoyenia
ichnofacies based on (1) relatively low diversity,
(2) very few unique ichnogenera, and (3) overall
similarity to some marine trace fossil assemblages.
Additionally, they proposed that “future designations
of this ichnofacies be restricted to trace-fossil as-
semblages in which Scoyenia gracilis, Ancorichnus
coronus, or ethologically and ecologically equivalent
burrows predominate” (Frey et al., 1984, p. 511).
Their review also showed that this ichnofacies char-
acterizes nonmarine deposits periodically exposed to
air or inundated, intermediate between aquatic and
nonaquatic, typical of floodplains and lake margins
(see also Frey and Pemberton, 1984, 1987).
Buatois and Mángano (1995a) suggested that non-
marine trace fossil assemblages dominated by arthro-
pod trackways be included in the Scoyenia ichnofa-
cies, regardless of the presence of meniscate bur-
rows. Arthropod trackways are particularly dominant
in Paleozoic fluvio-lacustrine deposits (e.g., Pollard
et al., 1982; Walter, 1983; Walker, 1985). A narrower
definition was recently proposed by Bromley (1996),
who considered the Scoyenia ichnofacies as a non-
369
marine firmground assemblage. Bromley (1996) also
tentatively proposed a Fuersichnus ichnofacies for
lacustrine settings below the fairweather wavebase.
This ichnofacies is based on examples in which
Fuersichnus occurs in nonmarine settings. However,
this ichnotaxon has been recorded in fluvial deposits
(MacNaughton and Pickerill, 1995) and ephemeral
alluvial plain=sandflat facies (Gierlowski-Kordesch,
1991), and is best regarded as an element of the
fluvio-lacustrine Scoyenia ichnofacies.
Typical components of the Scoyenia ichno-
facies include meniscate burrows (e.g., Scoye-
nia, Beaconites, and Taenidium), bilobate traces
(e.g., Cruziana and Rusophycus, which sometimes
are recorded as Isopodichnus), arthropod track-
ways (e.g., Umfolozia, Merostomichnites, Diplich-
nites, and Acripes), simple horizontal burrows (e.g.,
Planolites and Palaeophycus), simple vertical bur-
rows (e.g., Skolithos), sinuous crawling traces (e.g.,
Cochlichnus), spreiten burrows (e.g., Fuersichnus),
and various vertebrate tracks (Figs. 2 and 3). In
lacustrine complexes, this ichnofacies typically char-
acterizes lake margin deposits, being present in both
open and closed lake basins, and in both ephemeral
and perennial systems.
Buatois and Mángano (1995a, table 3) summa-
rized selected examples of this ichnofacies. Lacus-
trine examples of the Scoyenia ichnofacies were
recorded for the Devonian (Pollard et al., 1982;
Pollard and Walker, 1984; Walker, 1985), Permian
(Aceñolaza, 1978; Aceñolaza and Buatois, 1993;
Van Amerom et al., 1993), Triassic (Seilacher, 1963;
Bromley and Asgaard, 1979, 1991; Pollard, 1981,
1985; Metz, 1995, 1996), Jurassic (Gierlowski-
Kordesch, 1991; Metz, 1992), and Tertiary (Toots,
1967). Because the Scoyenia ichnofacies records the
activity of a benthos adapted to low-energy condi-
tions in either very slightly submerged sediments that
are periodically desiccated or in waterside subaerial
substrates that are periodically submerged (Frey and
Pemberton, 1987), it is particularly useful to de-
lineate marginal lacustrine facies in sedimentologic
studies.
2.3. Mermia ichnofacies
The Mermia ichnofacies, proposed by Buatois and
Mángano (1995a), is characterized by (1) dominance
370 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 140 (1998) 367–382

Fig. 2. Trace fossil association characteristic of the Scoyenia ichnofacies (modified from Buatois and Mángano, 1995a).

of horizontal to subhorizontal grazing and feeding Helminthopsis, Helminthoidichnites, Gordia, Mer-

traces produced by mobile deposit feeders, (2) subor-
dinate occurrence of locomotion traces, (3) generally
high to moderate ichnodiversity and abundance, and
(4) low degree of specialization of grazing patterns.
Typical components of this ichnofacies include a
variety of unspecialized grazing traces (e.g., Mer-
mia, Gordia, Helminthopsis, Helminthoidichnites,
and Cochlichnus), simple feeding structures (e.g.,
Treptichnus, Tuberculichnus, Vagorichnus, and Cir-
culichnis), locomotion traces (e.g., Maculichna), and
fish trails (e.g., Undichna) (Figs. 4 and 5). High
ichnodiversity does not necessarily equate here with
species richness. Different ichnogenera recorded in
this ichnofacies probably result from minor varia-
tions of behavior of a very simple, unspecialized
grazing pattern developed by one tracemaker (e.g.,
mia). However, these ichnogenera have been tradi-
tionally considered as distinct ichnotaxa.
The Mermia ichnofacies is characteristic of fine-
grained sediments that occur in well-oxygenated,
low-energy, permanently subaqueous zones of lacus-
trine systems (Buatois and Mángano, 1995a). Ex-
amples of the Mermia ichnofacies range from Car-
boniferous to Recent. Recordings of this recurrent
assemblage include the Carboniferous of Argentina
(Buatois and Mángano, 1993a) and Canada (Picker-
ill, 1992), Permian of Antarctica (Miller et al., 1991),
Jurassic of China (Wu, 1985; Buatois et al., 1996a),
Cretaceous of Spain (Fregenal-Martinez et al., 1995),
and Pleistocene of Finland (Gibbard, 1977), Canada
(Gibbard and Dreimanis, 1978), and England (Gib-
bard and Stuart, 1974). It is typical of open perennial

Fig. 3. Elements of the Scoyenia ichnofacies. (A) Taenidium isp. Tertiary, Nanyang oil field, China. (B) Merostomichnites aicuñai. Per-
mian, Bordo Atravesado, Argentina. Bar 1 cm. (C) Monomorphichnus isp. Permian, Bordo Atravesado, Argentina. (D) Mirandaichnium
famatinense. Permian, Bordo Atravesado, Argentina. Bar 1 cm. (E) Umfolozia sinuosa. Note associated desiccation crack. Permian,
Bordo Atravesado, Argentina. (F) detailed view of (E) showing association of tracks with desiccation cracks. Bar 1 cm.
L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 140 (1998) 367–382 371
372 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 140 (1998) 367–382
 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 140 (1998) 367–382 373

siliciclastic lacustrine systems of variable depths and
dimensions. However, recent research indicates the
presence of the Mermia ichnofacies in carbonate lake
systems also (Fregenal-Martinez et al., 1995).
This ichnofacies comprises sediments deposited
in fully lacustrine environments, extending from
shallow to deep bathymetric zones. There are no
universally accepted trace fossil associations that
clearly distinguish shallow and deep lacustrine set-
tings, probably because of the wide variability of
lacustrine systems. Accordingly, the Mermia ichno-
facies can be regarded as a nonmarine equivalent
of the Cruziana, Zoophycos, and Nereites ichno-
facies in the classical Seilacherian scheme. How-
ever, bathymetric zonations have been reported in
some localized settings (Walter, 1986; Buatois and
Mángano, 1993b). In general, lacustrine facies mod-
els may be enhanced and refined by careful analysis
of the associated biogenic structures, taking into
account not only ichnotaxonomic composition, but
also ethologic, trophic, and taphonomic features of
the trace fossil fauna. In general, trends seem to
show a decrease in locomotion traces, and an in-
crease in ichnodiversity and grazing and feeding
structures from nearshore lacustrine facies to off-
shore and deep lacustrine environments (Buatois and
Mángano, 1995a). Local subdivisions of the Mermia
ichnofacies into discrete ichnocoenoses may be use-
ful for sedimentologic purposes. Subdivisions may
reflect different subenvironments within the lake
basin, or pre- vs. post-depositional suites in the
case of lakes dominated by event deposition. In
such cases, the pre-turbidite ichnocoenosis records
the activity of the resident benthic fauna, while the
post-turbidite suite represents an impoverished op-
portunistic ichnocoenosis, reflecting colonization of
the newly produced substrate.
2.4. Prospectus for future research
The nonmarine ichnofacies model described here
does not include all potential continental ichnofa-
cies. With respect to lake systems, the Scoyenia
and Mermia ichnofacies both record benthic activ-
ity in low-energy environments, and therefore other
potential ichnofacies may be recognized in high-
energy settings. Additionally, the problem of sub-
strate-controlled ichnofacies in nonmarine environ-
ments should be addressed.
Sediments deposited in high-energy nonmarine
environments, such as lacustrine wave-dominated
shorelines and delta mouth bars, commonly contain
simple vertical burrows (Skolithos), U-shaped verti-
cal burrows (Arenicolites), and escape structures (e.g.,
Bromley and Asgaard, 1979; Mángano et al., 1994).
Such assemblages are hardly distinctive from the ma-
rine Skolithos ichnofacies, so it is best to consider
them as nonmarine occurrences of the Skolithos ich-
nofacies (Buatois and Mángano, 1995a). The Areni-
colites ichnofacies proposed by Bromley and Asgaard
(1991) is here included in the Skolithos ichnofacies
(see discussions in Pemberton et al., 1992b; Frey and
Goldring, 1992; Goldring, 1993).
Recognition of substrate-controlled nonmarine
ichnofacies is a promising field. Firmground trace
fossil suites have been recognized in continental sed-
imentary successions, usually associated with des-
iccated fluvial floodplains (e.g., Bromley and As-
gaard, 1979, 1991; Fürsich and Mayr, 1981; Smith
et al., 1993; Buatois et al., 1996b). Firmground as-
semblages also have been documented in lacustrine
deposits (e.g., Metz, 1993, 1995; Mángano et al.,
1994). Metz (1993, 1995) recorded the presence of
Spongeliomorpha milfordensis in Triassic lacustrine
deposits of New Jersey and Pennsylvania. This form
consists of horizontal burrow systems that display
sharply incised wall scratchings. Such kind of or-
namentation is typical of dewatered, firm substrates.
These structures are associated with marginal lacus-
trine deposits that have experienced significant peri-
ods of dryness. Striated, vertical burrows (Skolithos
serratus) were reported by Mángano et al. (1994)
in Triassic lacustrine deposits of central China. If
the definition of the Scoyenia ichnofacies proposed

Fig. 4. Elements of the Mermia ichnofacies. (A) Undichna insolentia. Carboniferous, Cantera La Laja, Argentina. (B) Mermia carickensis.
Carboniferous, Cantera La Laja, Argentina. (C) Treptichnus pollardi. Carboniferous, Cantera La Laja, Argentina. (D) Vagorichnus anyao.
Jurassic, Jiyuan, China. (E) Helminthoidichnites tenuis. Carboniferous, Cantera La Laja, Argentina. (F) Cochlichnus anguineus. Jurassic,
Jiyuan, China. (G) Circulichnis montanus. Carboniferous, Cantera La Laja, Argentina. In all cases, bar is 1 cm.
374 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 140 (1998) 367–382
Fig. 5. Trace fossil association characteristic of the Mermia ichnofacies (modified from Buatois and Mángano, 1995a).
by Bromley (1996) is adopted, all these occurrences
should be included in such ichnofacies. Inclusion of
both firmground and softground suites in the Scoye-
nia ichnofacies seems to be the best alternative.
A single report of trace fossils in lacustrine hard-
grounds is known at present. Ekdale et al. (1989) de-
scribed bioerosion traces in Pleistocene stromatolites
in lake margin deposits of Kenya. This hardground
suite comprises the ichnogenera Trypanites, recorded
from the first time in a nonmarine environment, and
Sertaterebrites, a new ichnotaxon.
Buatois and Mángano (1995a) suggested some
guidelines for the recognition of additional nonma-
rine ichnofacies, including (1) accurate taxonomy of
nonmarine trace fossils, trying to avoid the use of
open nomenclature, (2) identification of additional
ichnofacies, rather than subdivision of the Scoyenia
ichnofacies, (3) emphasis primarily on environmen-
tal conditions and subsequently on sedimentary envi-
ronments, (4) consideration of the wide variability of
nonmarine ichnofaunas through geologic time, and
(5) integration of invertebrate and vertebrate trace
fossil information.
3. Ichnologic signatures of event stratification
3.1. Density underflows vs. turbidity currents
Trace fossils may be used to distinguish between
specific types of lacustrine sedimentary facies and to
delineate event beds. Density underflow and turbidity
currents occur in open lacustrine systems. However,
distinction between both processes is difficult, and
the two terms sometimes are used interchangeably in
the literature (e.g., Sturm and Matter, 1978; Lambert
and Hsü, 1979). The conceptual differences between
the two were summarized by Pharo and Carmack
(1979), who explained that underflow currents are
relatively continuous currents that represent the un-
interrupted extension of river-borne sediment into the
lake basin, and are influenced by geostrophic effects.
On the other hand, turbidity currents are episodic
currents that involve redeposition of sediment ini-
tially emplaced under unstable conditions, and mate-
rial concentrations dominate the fluid density, driving
the downslope flow. Buatois and Mángano (1995b)
stressed the potential of such distinction in sequence
 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 140 (1998) 367–382
stratigraphic studies because underflow currents tend
to dominate during periods of high discharge and
lake highstand, and turbidity currents are triggered
during lowstands.
Trace fossil analysis may provide valuable keys
to the distinction between density underflows and
turbidity currents. Buatois and Mángano (1995b)
noted that lacustrine turbidites from the Carbonifer-
ous Lake Narváez (northwest Argentina) host trace
fossils at bedding tops, either on Bouma division
E or division C in the case of top-missing tur-
bidites. A similar assemblage was documented by
Buatois and Mángano (1995c) from the Carbonif-
erous Lake Malanzán in western Argentina. These
ichnocoenoses are characterized by a low diver-
sity of traces, dominance of surface grazing trails,
and extremely simple grazing patterns (Fig. 6).
Gordia, Mermia, and Helminthoidichnites are com-
mon elements. This peculiar trace fossil assemblage
records colonization of opportunistic organisms after
episodic emplacement of the turbidite bed (Bua-
tois and Mángano, 1993a,b). Such postdepositional
suites usually are easily distinguished from the more
diverse pre-turbidite ichnocoenoses that record the
activity of the resident fauna.
Underflow current beds from the Carboniferous
Lake Narváez display distinctive suites of ichno-
fossils in each lamina or lamina-set (Buatois and
Fig. 6. Trace fossils from lacustrine turbidites.
375
Mángano, 1993a,b). Each bed consists of normally
graded to stratified units formed of thin lamina-sets
of fine- to very fine-grained sandstones and mud-
stones. Careful splitting of individual laminae in
several beds allows us to detect a recurrent pattern of
trace fossil distribution (Fig. 7). The basal, thicker,
fine-grained sandstone lamina-set lacks trace fossils
and probably records high sedimentation rates dur-
ing catastrophic floods. High episodic influx of sand
may have prevented activity of the benthic fauna
and hence precluded the formation of biogenic struc-
tures. The uppermost fine-grained sandstone lamina-
set contains a low density, but relatively high diver-
sity of ichnofossils, consisting mainly of locomotion
traces (Undichna, Aulichnites, Orchesteropus) and
to a lesser extent resting (Rusophycus) and grazing
traces (Gordia, Mermia). The overlying very fine-
grained lamina-sets host ichnocoenoses dominated
by grazing trails (Mermia, Gordia, Helminthopsis,
Helminthoidichnites), whereas the mudstone lam-
ina-set at the top includes also feeding structures
(Treptichnus). Accordingly, ichnologic evidence in-
dicates animal activity contemporaneous with the
sedimentation instead of colonization after a major
break in deposition. This fact suggests near-contin-
uous deposition from river-fed density underflows
rather than episodic sedimentation from turbidity
currents.
376 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 140 (1998) 367–382
Fig. 7. Trace fossils from underflow current deposits.
3.2. Lacustrine vs. marine turbidites
Salinity is one of the most important factors con-
trolling distribution of benthic faunas (Pemberton
and Wightman, 1992). While physical processes of
sedimentation are commonly independent of salin-
ity, faunal distribution, and hence trace fossil dis-
tribution is not. Accordingly, in certain depositional
environments, detailed ichnologic analysis may pro-
vide higher resolution than sedimentologic studies,
supplying valuable information for facies model con-
struction.
Marine and lacustrine turbidites are virtually
indistinguishable in terms of physical sedimen-
tary structures. However, their biogenic sedimentary
structures are clearly different. Therefore, turbidites
represent a case where trace fossils can distinguish
between marine or nonmarine.
Deep-sea turbidites are characterized by the Nere-
ites ichnofacies (Seilacher, 1967). Traces of the Nere-
ites ichnofacies are typically preserved in positive
relief on the sole of sandstone turbidites and com-
prises a high diversity of ornate grazing traces (Pas-
cichnia) and graphoglyptids (Agrichnia) that record
highly specialized feeding strategies of the deep
marine fauna (Ekdale et al., 1984; Miller, 1991;
Pemberton et al., 1992a,b; Uchman, 1995). This
fauna is adapted to periodic disruption by event
sedimentation and displays complex feeding strate-
gies, possibly including trapping of meiobenthos
and cultivation of bacteria, in order to solve the
problem of the scarcity of food in the deep-sea
biotope (Seilacher, 1977). Typical morphologic pat-
terns include spirals (Spirorhaphe), guided mean-
ders (Helminthorhaphe, Cosmorhaphe), uni- and
biramous meanders (Urohelminthoida, Desmograp-
ton), radial structures (Glockerichnus, Lorenzinia,
Capodistria), irregular networks (Acanthorhaphe,
Megagrapton, Protopaleodictyon), and regular net-
works (Paleodictyon) (Seilacher, 1977; Miller, 1991).
In contrast, deep lacustrine settings are charac-
terized by the Mermia ichnofacies (Buatois and
Mángano, 1995a), which is dominated by nonspe-
cialized grazing and feeding traces. Although the
Mermia ichnofacies displays higher diversity than
the majority of other nonmarine assemblages, trace
fossils are less varied than their marine counterparts
(e.g. Wu, 1985; Buatois and Mángano, 1993a; Bu-
atois et al., 1996a). Nonspecialized feeding patterns
are exemplified by the ichnogenus Mermia, which
displays looping and a high level of self-overcross-
ing, recording the repeated passage of the tracemaker
across the same portion of sediment. Such nonspe-
cialized trophic strategies probably are related to
the abundance and accessibility of food in lacustrine
systems (Buatois and Mángano, 1995a).
Differences between diversity of trace fossils in
deep marine and lacustrine environments can be ex-
plained by the stability–time hypothesis developed
by Sanders (1968). This author suggested that species
 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 140 (1998) 367–382
diversity reflects stability or predictability of the en-
vironment, as well as its geologic history. Deep ma-
rine environments are typically more stable than la-
custrine systems, and host a more diverse benthic
fauna. Lakes are generally shorter-lived than deep ma-
rine basins. Sanders (1968) noted that lakes from re-
cently glaciated regions show limited diversification.
Although ichnodiversity should not be equated with
biodiversity (Bromley, 1990), the former may also
provide some very general trends in species richness.
We suggest that the comparatively lower ichnodiver-
sity of lakes with respect to deep seas also results
from the less stable conditions of the former. Lacus-
trine ichnofaunas, however, are typically more diverse
than ichnofaunas in other nonmarine settings, such as
eolian, fluvial, and alluvial fan environments. This
situation is probably also explained by the fact that
deep lakes are the most stable environments in the
nonmarine realm. Sanders (1968) found high species
diversity in the deep zones of the Lake Baikal, which
has a long geologic history and is the deepest fresh-
water lake of the world. It is therefore not surprising
that some of the most diverse nonmarine trace fos-
sil assemblages have been described from long-lived,
large, and deep lakes of the late Paleozoic Gondwana
(e.g. Buatois and Mángano, 1993a).
4. Lacustrine ichnofabrics: a matter of time
Ichnofabric comprises all aspects of the texture
and internal structure of a sediment that result from
bioturbation at all scales (Ekdale and Bromley, 1983;
Bromley and Ekdale, 1986). The ichnofabric ap-
proach became very popular during the last ten years
and the concept has been applied to the study of
a considerable number of marine sedimentary suc-
cessions (e.g. Ekdale and Bromley, 1991; Goldring
et al., 1991; Pollard et al., 1993). However, little is
known about the nature and genesis of nonmarine
ichnofabrics.
The progressive colonization of nonmarine envi-
ronments resulted in a rather complex history of con-
tinental ichnofaunas (Buatois and Mángano, 1993c).
Evolutionary innovations of the terrestrial and fresh-
water biotas established a strong signature and con-
strained the development of nonmarine ichnofabrics.
A brief review of some key ichnofaunas will be of
377
help to understand the applicability and limitations
of the ichnofabric concept in lacustrine successions.
Devonian and Carboniferous lacustrine ichnofau-
nas are well known from several localities all around
the world, such as in Scotland (Pollard and Walker,
1984; Walker, 1985), Norway (Pollard et al., 1982),
Canada (Pickerill, 1992), and Argentina (Buatois
and Mángano, 1993a). Assemblages are dominated
by surface traces, mostly grazing trails and arthro-
pod trackways (see also Maples and Archer, 1989;
Buatois and Mángano, 1993c). The dominance of
epifaunal traces result in little or no bedding disrup-
tion. Accordingly, such lacustrine deposits tend to be
unbioturbated. Trace fossils are typically restricted
to bedding planes of thinly laminated sediment.
The development of deposit-feeder, back-filled
meniscate burrows during the Permian may have
favored increasing disruption in lake margin de-
posits. However, recording of such structures in Per-
mian lacustrine rocks is rare, with most examples of
meniscate burrows reported from floodplain settings
(Aceñolaza and Buatois, 1993). Consequently, Per-
mian lacustrine facies typically display low degrees
of bioturbation.
The establishment of deep burrows in lake margin
deposits lead to considerable disruption of the sedi-
mentary fabric in the Triassic. Hasiotis et al. (1993),
and Hasiotis and Mitchell (1993) documented the
crayfish burrow Camborygma from Triassic flood-
plain to lake margin facies of Colorado, New Mex-
ico, Arizona, and Utah. According to these authors,
burrows penetrate as deep as 1 m into the sub-
strate. Relatively high degrees of bioturbation were
also detected from the Fuersichnus ichnocoenosis
described by Bromley and Asgaard (1979, 1991)
from Triassic successions of East Greenland, which
are interpreted as having been deposited in playa
lake complexes (Dam and Stemmerik, 1994). Firm-
ground burrows were also common in lake margin
deposits during the Triassic. The ornamented burrow
system Spongeliomorpha has been documented from
Triassic marginal lacustrine desiccated substrates of
New Jersey and Pennsylvania by Metz (1993, 1995).
Accordingly, composite ichnofabrics (including both
softground and firmground suites) associated with
periods of dryness may have been formed in Triassic
transitional fluviolacustrine deposits. Although infor-
mation suggests an increase in bioturbation degree
378 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 140 (1998) 367–382
in lake margin facies (Scoyenia ichnofacies) during
the Triassic, few data are available on ichnofabrics
from permanent subaqueous environments (Mermia
ichnofacies).
An increase in bioturbation depth also from fully
lacustrine deposits is recorded in the Jurassic. Buatois
et al. (1996a) documented a moderately diverse as-
semblage of grazing and feeding traces from Jurassic
lacustrine turbidites of central China. Post-turbidite
ichnofossils (e.g., Vagorichnus) are preserved on the
sole of up to 5 cm thick sandstone beds (Buatois et
al., 1995). The presence of this intermediate-depth
infauna is in contrast with the dominance of surface
trails in Paleozoic assemblages from the Mermia ich-
nofacies. However, as trace fossils are restricted to
the sand=mud interface, bioturbation is sparse and the
primary sedimentary fabric is fully preserved.
This situation seems to have changed during the
Cretaceous. Lacustrine shoreface deposits of the Cre-
taceous Coqueiro Seco Formation in northeast Brazil
Fig. 8. Bioturbated lacustrine shoreface deposits recording the activity of an infaunal benthic community. Coqueiro Seco Formation,
Cretaceous, Brazil.
are intensely bioturbated by the activity of a de-
posit-feeder infauna (Azambuja Filho and Spadini,
1994) (Fig. 8). This suggests a significant increase in
bioturbation extent in fully lacustrine deposits.
Cenozoic lacustrine basins in Southeast Asia pro-
vide some information on ichnofabric trends. Whate-
ley and Jordan (1989, fig. 8) illustrated intensely
bioturbated lacustrine facies from the Paleogene of
Indonesia. High degrees of bioturbation in compa-
rable facies also were documented by Flint et al.
(1989) from the Neogene of Thailand. Furthermore,
cores of Paleogene lake deposits of east China also
show significant bioturbation and disruption of the
sedimentary fabric (Wang Pixian, pers. commun.,
1992). Accordingly, available data seem to indicate
intense activity of a fully lacustrine, infaunal de-
posit-feeding biota during the Cenozoic, and hence
the accentuation of trends previously initiated during
the Cretaceous.
Although our record of lacustrine ichnofaunas
 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 140 (1998) 367–382
throughout the Phanerozoic is still sparse and in-
complete, some general, albeit preliminary, overall
trends in burrowing depth, bioturbation degree, and
ichnofabric traits seem to be apparent. Available
information suggests that bioturbation depth was
comparatively higher in lake margin facies than in
fully lacustrine deposits. Additionally, while signifi-
cant degrees of bioturbation were attained in Triassic
fluviolacustrine transitional deposits, major biogenic
disruption of sedimentary bedding was delayed until
the Cretaceous in permanent subaqueous lacustrine
facies. Accordingly, the ichnofabric approach to the
study of Paleozoic lacustrine facies seems to be
rather limited. On the other hand, the ichnofabric
concept may be useful for the analysis of post-Pale-
ozoic lake ichnofaunas.
5. Conclusions
(1) Two ichnofacies are typically associated with
lacustrine depositional systems: the Soyenia and
Mermia ichnofacies. Because the Scoyenia ichno-
facies characterizes low-energy conditions in either
very slightly submerged sediments that are periodi-
cally desiccated or waterside subaerial substrates that
are periodically submerged, it is useful to delineate
marginal lacustrine facies in sedimentologic stud-
ies. The Mermia ichnofacies typifies fine-grained
sediments at well-oxygenated, low-energy, perma-
nently subaqueous zones of lacustrine systems. Lo-
cal subdivisions of the ichnofacies into discrete ich-
nocoenoses may record different subenvironments
within the lake, or pre- vs. post-depositional suites in
the case of event deposition.
(2) The typically marine Skolithos ichnofacies
also may be present in high-energy zones of lakes
(e.g., deltaic mouth bars, wave-dominated shore-
lines).
(3) Recognition of substrate-controlled ichnofa-
cies may be of help to delineate successive stages of
consolidation typically associated with progressive
dryness in lake margin facies.
(4) Trace fossils may yield valuable insights into
facies analysis of event beds. Lacustrine turbidites
are characterized by low-diversity suites that record
colonization by opportunistic organisms after the
turbidite event. Underflow current deposits reflect
379
animal activity contemporaneous with nearly contin-
uous sedimentation. Although marine and lacustrine
turbidites are virtually identical in terms of physi-
cal sedimentary structures, their biogenic structures
are completely different. Deep-sea turbidites host the
Nereites ichnofacies that consists of a high diversity
of ornate grazing traces (Pascichnia) and grapho-
glyptids (Agrichnia), recording highly specialized
feeding strategies employed by the marine benthos
to solve the problem of the scarcity of food in the
deep sea. Deep lacustrine environments contain the
Mermia ichnofacies, which is dominated by unspe-
cialized grazing and feeding traces probably related
to abundance and accessibility of food in lacustrine
systems. The lower diversity of lacustrine ichnofau-
nas in comparison with deep-sea assemblages more
likely reflects lower species diversity as a conse-
quence of less stable conditions.
(5) Increase of depth and extent of bioturbation
through geologic time in lacustrine deposits pro-
duced a clear signature in the ichnofabric record.
Surface trails, which cause negligible bedding dis-
ruption, were dominant during the Paleozoic. Biotur-
bation depth was comparatively higher in lake mar-
gin facies than in fully lacustrine deposits throughout
the Mesozoic. Furthermore, while significant degrees
of bioturbation were attained in Triassic lake margin
facies, major biogenic disruption of sedimentary fab-
ric in fully lacustrine deposits did not occur until the
Cretaceous. The ichnofabric approach to the study
of lacustrine facies seems to be rather limited for
Paleozoic sequences, but it is useful for the analysis
of post-Paleozoic lake ichnofaunas.
Acknowledgements
We thank the Antorchas Foundation for providing
financial support for our studies on nonmarine ich-
nofaunas. This paper was written under tenure of an
external grant of the Argentinean Research Council
(CONICET) at the Kansas Geological Survey. We
also thank Wang Pixian for sharing his information
on the ichnology of lacustrine basins of east China,
N. Azambuja Filho and A. Spadini for guiding one
of us (LAB) in the Sergipe–Alagoas Basin in Brazil,
journal reviewers for their valuable comments, and
Mark Schoneweis for the drawings.
380 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 140 (1998) 367–382
References
Aceñolaza, F.G., 1978. Traza fósiles de la Formación Parquı́a en
el Bordo Atravesado, Sierra de Famatina, La Rioja. Acta Geol.
Lilloana 15, 19–29.
Aceñolaza, F.G., Buatois, L.A., 1993. Nonmarine perigondwanic
trace fossils from the Late Paleozoic of Argentina. Ichnos 2,
183–201.
Azambuja Filho, N.C., Spadini, A.R., 1994. Field Trip Guide to
Sergipe–Alagoas Basin. 14th Int. Sedimentol. Congr. Recife,
26 pp.
Bromley, R.G., 1990. Trace Fossils. Biology and Taphonomy.
Unwin Hyman, London, 280 pp.
Bromley, R.G., 1996. Trace Fossils. Biology, Taphonomy and
Applications. Chapman and Hall, London, 361 pp.
Bromley, R.G., Asgaard, U., 1979. Triassic freshwater ichno-
coenoses from Carlsberg Fjord, East Greenland. Palaeogeogr.,
Palaeoclimatol., Palaeoecol. 28, 39–80.
Bromley, R.G., Asgaard, U., 1991. Ichnofacies: a mixture of
taphofacies and biofacies. Lethaia 24, 153–163.
Bromley, R.G., Asgaard, U., 1993. Endolithic community re-
placement on Pliocene rocky coast. Ichnos 2, 93–116.
Bromley, R.G., Ekdale, A.A., 1986. Composite ichnofabrics and
tiering of burrows. Geol. Mag. 123, 59–65.
Buatois, L.A., Mángano, M.G., 1993a. Trace fossils from a
Carboniferous turbiditic lake: Implications for the recognition
of additional nonmarine ichnofacies. Ichnos 2, 237–258.
Buatois, L.A., Mángano, M.G., 1993b. The paleoenvironmen-
tal and paleoecological significance of turbiditic lake ich-
nocoenoses from the Late Carboniferous of the Paganzo
Basin. XII Int. Congr. Carboniferous and Permian Geology
and Stratigraphy, Buenos Aires, Compt. Rend. 2, pp. 409–420.
Buatois, L.A., Mángano, M.G., 1993c. Ecospace utilization, pa-
leoenvironmental trends and the evolution of early nonmarine
biotas. Geology 21, 595–598.
Buatois, L.A., Mángano, M.G., 1995a. The paleoenvironmental
and paleoecological significance of the lacustrine Mermia ich-
nofacies: an archetypical subaqueous nonmarine trace fossil
assemblage. Ichnos 4, 151–161.
Buatois, L.A., Mángano, M.G., 1995b. Sedimentary dynamics
and evolutionary history of a Late Carboniferous Gondwanic
lake in northwestern Argentina. Sedimentology 42, 415–436.
Buatois, L.A., Mángano, M.G., 1995c. Post glacial lacustrine
event sedimentation in an ancient mountain setting: Carbonif-
erous Lake Malanzán (Western Argentina). J. Paleolimnol. 12,
1–22.
Buatois, L.A., Mángano, M.G., Wu, X., Zhang, G., 1995.
Vagorichnus, a new ichnogenus for feeding burrow systems
and its occurrence as discrete and compound ichnotaxa in
Jurassic lacustrine turbidites of central China. Ichnos 3, 265–
272.
Buatois, L.A., Mángano, M.G., Wu, X., Zhang, G., 1996a. Trace
fossils from Jurassic lacustrine turbidites of the Anyao Forma-
tion (central China) and their environmental and evolutionary
significance. Ichnos 4, 287–303.
Buatois, L.A., Mángano, M.G., Aceñolaza, F.G., 1996b. Icnofau-
nas paleozoicas en sustratos firmes no marinos: evidencias del
Pérmico de la cuenca Paganzo. Ameghiniana 33, 265–270.
Dam, G., Stemmerik, L., 1994. East Greenland lacustrine com-
plexes. In: Gierlowski-Kordesh, E., Kelts, K. (Eds.), Global
Geological Record of Lake Basins, 1. Cambridge University
Press, Cambridge, pp. 19–27.
Ekdale, A.A., Bromley, R.G., 1983. Trace fossils and ichnofabric
in the Kjolby Gaard Marl, Upper Cretaceous, Denmark. Bull.
Geol. Soc. Denm. 31, 107–119.
Ekdale, A.A., Bromley, R.G., 1991. Analysis of composite ich-
nofabrics in Cretaceous chalks, Denmark. Palaios 6, 232–249.
Ekdale, A.A., Bromley, R.G., Pemberton, S.G., 1984. Ichnology:
trace fossils in sedimentology and stratigraphy. Soc. Econ.
Paleontol. Mineral., Short Course 15, 317 pp.
Ekdale, A.A., Brown, F.H., Feibel, C.S., 1989. Nonmarine mac-
roborings in Early Pleistocene algal biolithites (Stromatolites)
of the Turkana Basin, northern Kenya. Palaios 4, 389–396.
Flint, S., Stewart, D.J., Van Riessen, E.D., 1989. Reservoir geol-
ogy of the Sirikit oilfield, Thailand: lacustrine deltaic sedimen-
tation in a Tertiary intermontane basin. In: Whateley, M.K.G.,
Pickering, K.T. (Eds.), Deltas: Sites and Traps for Fossil Fuels.
Geol. Soc. Spec. Publ. 41, 223–237.
Fregenal-Martinez, M.A., Buatois, L.A., Mángano, M.G., 1995.
Invertebrate trace fossils from Las Hoyas fossil site (Ser-
ranı́a de Cuenca, Spain). Paleoenvironmental interpretations.
2nd Int. Symp. Lithographic Limestones, Lleida-Cuenca, Ext.
Abstr. pp. 67–70.
Frey, R.W., Goldring, R., 1992. Marine event beds and recol-
onization surfaces as revealed by trace fossil analysis. Geol.
Mag. 129, 325–335.
Frey, R.W., Pemberton, S.G., 1984. Trace fossil facies models.
In: Walker, R.G. (Ed.), Facies Models, 2nd ed. Geosci. Can.,
Reprint Ser. 1, 189–207.
Frey, R.W., Pemberton, S.G., 1987. The Psilonichnus ichno-
coenose, and its relationship to adjacent marine and nonmarine
ichnocoenoses along the Georgia coast. Bull. Can. Pet. Geol.
35, 333–357.
Frey, R.W., Seilacher, A., 1980. Uniformity in marine inverte-
brate ichnology. Lethaia 13, 183–207.
Frey, R.W., Pemberton, S.G., Fagerstrom, J.A., 1984. Morpho-
logical, ethological, and environmental significance of the
ichnogenera Scoyenia and Ancorichnus. J. Paleontol. 58, 511–
528.
Fürsich, F.T., Mayr, H., 1981. Non-marine Rhizocorallium (trace
fossil) from the Upper Freshwater Molasse (Upper Miocene)
of southern Germany. Neues Jahrb. Geol. Paläontol., Monatsh.
1981, 321–333.
Gibbard, P.L., 1977. Fossil tracks from varved sediments near
Lammi, south Finland. Bull. Geol. Soc. Finl. 49, 53–57.
Gibbard, P.L., Dreimanis, A., 1978. Trace fossils from late Pleis-
tocene glacial lake sediments in southwestern Ontario, Canada.
Can. J. Earth Sci. 15, 1967–1976.
Gibbard, P.L., Stuart, A.J., 1974. Trace fossils from proglacial
lake sediments. Boreas 3, 69–74.
Gierlowski-Kordesch, E., 1991. Ichnology of an ephemeral
lacustrine=alluvial plain system: Jurassic East Berlin Forma-
tion, Hartford Basin, USA. Ichnos 1, 221–232.
 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 140 (1998) 367–382
Goldring, R., 1993. Ichnofacies and facies interpretation. Palaios
8, 403–405.
Goldring, R., Pollard, J.E., Taylor, A.M., 1991. Anconichnus
horizontalis: a pervasive ichnofabric-forming trace fossil in
Post-Paleozoic offshore siliciclastic facies. Palaios 6, 250–
263.
Hasiotis, S.T., Bown, T.M., 1992. Invertebrate trace fossils: the
backbone of continental ichnology. In: Maples, C.G., West,
R.R. (Eds.), Trace Fossils. Paleontol. Soc. Short Course 5,
64–104.
Hasiotis, S.T., Mitchell, C.E., 1993. A comparison of crayfish
burrow morphologies: Triassic and Holocene fossil, paleo-
and neo-ichnological evidence, and the identification of their
burrowing signatures. Ichnos 2, 291–314.
Hasiotis, S.T., Mitchell, C.E., Dubiel, R.F., 1993. Application
of morphologic burrow interpretations to discern continental
burrow architects: lungfish or crayfish? Ichnos 2, 315–333.
Lambert, A., Hsü, K.J., 1979. Non-annual cycles of varve-like
sedimentation in Walensee, Switzerland. Sedimentology 23,
87–105.
MacNaughton, R.B., Pickerill, R.K., 1995. Invertebrate ichnol-
ogy of the nonmarine Lepreau Formation (Triassic), southern
New Brunswick, Eastern Canada. J. Paleontol. 69, 160–171.
Mángano, M.G., Buatois, L.A., Wu, X., Sun, J., Zhang, G.,
1994. Sedimentary facies, depositional processes and climatic
controls in a Triassic lake, Tanzhuang Formation, western
Henan Province, China. J. Paleolimnol. 11, 41–65.
Maples, C.G., Archer, A.W., 1989. The potential of Paleozoic
nonmarine trace fossils for paleoecological interpretations.
Palaeogeogr., Palaeoclimatol., Palaeoecol. 73, 185–195.
Metz, R., 1992. Trace fossils from the Lower Jurassic nonmarine
Towaco Formation, New Jersey. Northeast. Geol. 14, 29–34.
Metz, R., 1993. A new ichnospecies of Spongeliomorpha from
the Late Triassic of New Jersey. Ichnos 2, 259–262.
Metz, R., 1995. Ichnologic study of the Lockatong Formation
(Late Triassic), Newark Basin, southeastern Pennsylvania. Ich-
nos 4, 43–51.
Metz, R., 1996. Newark Basin ichnology: the Late Triassic
Perkasie Member of the Passaic Formation, Sanatoga, Penn-
sylvania. Northeast. Geol. Environ. Sci. 18, 118–129.
Miller, M.F., Collinson, J.W., Frisch, R.A., 1991. Depositional
setting and history of a Permian post-glacial black shale:
Mackellar Formation, Central Transantarctic Mountains. In:
Ulbrich, H., Rocha Campos, A.C. (Eds.), Gondwana Seven:
Proc. 7th Int. Gondwana Symp., Sao Paulo, pp. 201–215.
Miller III, W., 1991. Paleoecology of graphoglyptids. Ichnos 1,
305–312.
Pemberton, S.G., Wightman, D.M., 1992. Ichnological character-
istics of brackish water deposits. In: Pemberton, S.G. (Ed.),
Applications of Ichnology to Petroleum Exploration, a Core
Workshop. Soc. Econ. Paleontol. Mineral., Core Workshop 17,
141–167.
Pemberton, S.G., Mac Eachern, J.A., Frey, R.W., 1992a. Trace
fossils facies models: environmental and allostratigraphic sig-
nificance. In: Walker, R.G., James, N.P. (Eds.), Facies Models
Response to Sea Level Change. Geological Association of
Canada, pp. 47–72.
381
Pemberton, S.G., Mac Eachern, J.A., Ranger, M.J., 1992b. Ich-
nology and event stratigraphy: the use of trace fossils in recog-
nizing tempestites. In: Pemberton, S.G. (Ed.), Applications of
Ichnology to Petroleum Exploration, a Core Workshop. Soc.
Econ. Paleontol. Mineral., Core Workshop 17, 85–117.
Pharo, C.H., Carmack, E.C., 1979. Sedimentation processes in
a short residence-time intermontane lake, Kamloops Lake,
British Columbia. Sedimentology 26, 523–541.
Pickerill, R.K., 1992. Carboniferous nonmarine invertebrate ich-
nocoenoses from southern New Brunswick, eastern Canada.
Ichnos 2, 21–35.
Pollard, J.E., 1981. A comparison between the Triassic of
Cheshire and south Germany. Palaeontology 24, 555–588.
Pollard, J.E., 1985. Isopodichnus, related arthropod trace fossils
and notostracans from Triassic fluvial sediments. Trans. R.
Soc. Edinburgh 76, 273–285.
Pollard, J.E., Walker, E., 1984. Reassessment of sediments and
trace fossils from Old Red Sandstone (Lower Devonian) of
Dunure, Scotland, described by John Smith (1909). Geobios
17, 567–576.
Pollard, J.E., Steel, R.J., Undersrud, E., 1982. Facies sequences
and trace fossils in lacustrine=fan-delta deposits, Hornelen
Basin (M. Devonian), western Norway. Sediment. Geol. 32,
63–87.
Pollard, J.E., Goldring, R., Buck, S.G., 1993. Ichnofabrics con-
taining Ophiomorpha: significance in shallow-water facies in-
terpretation. J. Geol. Soc. London 150, 149–164.
Sanders, H.L., 1968. Marine benthic diversity: a comparative
study. Am. Nat. 102, 243–282.
Seilacher, A., 1963. Lebensspuren und Salinitatsfazies. Fortschr.
Geol. Rheinland Westfalens 10, 81–94.
Seilacher, A., 1967. Bathymetry of trace fossils. Mar. Geol. 5,
413–428.
Seilacher, A., 1977. Pattern analysis of Paleodictyon and related
trace fossils. In: Crimes, T.P., Harper, J.C. (Eds.), Trace Fossils
2. Geol. J. Spec. Iss. 9, 289–334.
Smith, R.M.H., Mason, T.R., Ward, J.D., 1993. Flash-flood sed-
iments and ichnofacies of the Late Pleistocene Homeb Silts,
Kuiseb River, Namibia. Sediment. Geol. 85, 579–599.
Sturm, M., Matter, A., 1978. Turbidites and varves in Lake
Brienz (Switzerland): deposition of clastic detritus by density
currents. In: Matter, A., Tucker, M.E. (Eds.), Modern and
Ancient Lake Sediments. Spec. Publ. Int. Assoc. Sedimentol.
2, 147–168.
Toots, H., 1967. Invertebrate burrows in the nonmarine Miocene
of Wyoming. Contrib. Geol. 6, 93–96.
Uchman, A., 1995. Taxonomy and paleoecology of flysch trace
fossils: The Marnoso-arenacea Formation and associated facies
(Miocene, Northern Apennines, Italy). Beringeria 15, 1–116.
Van Amerom, H.W.J., Broutin, J., Ferrer, J., Gamez-Vintaned,
J.A., Gisbert, J., Linan, E., 1993. Les flores du Permien basal
et la paléoichnologie de la fosse de Fombuena (province de
Zaragoza, Espagne). Meded. Rijks Geol. Dienst 48, 1–63.
Walker, E., 1985. Arthropod ichnofauna of the Old Red Sand-
stone at Dunure and Montrose, Scotland. Trans. R. Soc. Edin-
burgh, Earth Sci. 76, 287–297.
Walter, H., 1983. Zur Taxonomie, Ökologie und Biostratigraphie
382 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 140 (1998) 367–382

der Ichnia limnisch–terrestrischer Arthropoden des mitteleu-
ropäischen Jungpaläozoikums. Freiberger Forschungsh. C 382,
146–193.
Walter, H., 1986. Arthropodenfahrten in eiszeitlichen Banderto-
nen bei Liebegast, Kreis Hoyerswerda. Natur Landschaft Bez.
Cottbus, NLBC 8, 51–58.
Whateley, M.K.G., Jordan, G.R., 1989. Fan-delta-lacustrine sedi-
mentation and coal development in the Tertiary Omlibin Basin,
W. Sumatra, Indonesia. In: Whateley, M.K.G., Pickering, K.T.
(Eds.), Deltas: Sites and Traps for Fossil Fuels. Geol. Soc.
Spec. Publ. 41, 317–332.
Wu, X., 1985. Trace fossils and their significance in non-marine
turbidite deposits of Mesozoic coal and oil bearing sequences
from Yima–Jiyuan basin, western Henan, China (in Chinese
with English summary). Acta Sedimentol. Sin. 3, 23–31.