Palaeogeography, Palaeoclimatology, Palaeoecology 272 (2009) 127–142

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Palaeogeography, Palaeoclimatology, Palaeoecology

j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / p a l a e o

Applications of ichnology in lacustrine sequence stratigraphy: Potential

and limitations
Luis A. Buatois ⁎, M. Gabriela Mángano
Department of Geological Sciences, University of Saskatchewan, 114 Science Place, Saskatoon, SK, Canada S7N 5E2

a r t i c l e i n f o a b s t r a c t

Article history: In comparison with their marine counterparts, lacustrine ichnofaunas have not been extensively used in
Received 1 March 2008 sequence stratigraphy. Application of trace fossils in continental sequence stratigraphy cannot be based
Received in revised form 18 August 2008 simply on the extrapolation of marine sequence stratigraphy. Marine substrate-controlled ichnofacies, in
Accepted 24 October 2008
particular the firmground Glossifungites ichnofacies, develop in stable and cohesive substrates, reflecting
erosive exhumation of the sediment. In contrast, in lacustrine sequences, substrate-controlled ichnofossils
Keywords:
Lakes
only rarely indicate erosional exhumation because they are commonly related to desiccation of water bodies.
Trace fossils The tripartite division of overfilled, balanced-fill, and underfilled lakes provides a comprehensive framework
Sequences stratigraphy to evaluate the potential and limitations of lacustrine ichnofaunas in sequence stratigraphy. Based on a
Overfilled lakes number of case studies spanning the three lake-basin types, a model of trace-fossil distribution in overfilled,
Balanced-fill lakes balanced-fill, and underfilled basins is proposed.
Underfilled lakes Overfilled lake basins contain well-developed softground trace fossils (Mermia and Skolithos ichnofacies, and
the softground suite of the Scoyenia ichnofacies) that are useful to delineate parasequences and parasequence
sets. Fluvial discharge into overfilled lakes usually generates density currents that oxygenate lake bottoms,
allowing the establishment of epifaunal and infaunal communities. Additionally, these are freshwater lakes
where no stress due to hypersalinity occurs, leading to the development of a relatively diverse benthos.
Land–plant derived organic matter is the prime source of nutrients, favoring the development of a deposit-
feeding benthic fauna in permanently subaqueous, low-energy zones. Firmground suites are rare because
such large lakes usually do not experience desiccation.
Abundant firmground trace-fossil suites of the Scoyenia ichnofacies occur in balanced-fill lakes, but
softground assemblages are usually depauperate. Lowstand deposits contain abundant and widespread
ichnofaunas, commonly meniscate trace fossils and arthropod trackways of the Scoyenia ichnofacies. Lake
hydrology is closed during lowstands and salinity usually increases, imposing a stress factor on the lake biota
and, therefore, softground ichnofaunas are depauperate. Ichnofaunas from turbidite systems in balanced-fill
lakes are less abundant and diverse than those from overfilled lacustrine turbidites. Freshwater conditions
are common during transgression, but dysaerobic conditions may prevail, imparting a stress factor on the
benthic biota. In hydrologically closed lakes, the Scoyenia ichnofacies is widespread, but the Mermia
ichnofacies is commonly absent or impoverished.
The Scoyenia ichnofacies is associated with lowstand desiccated substrates in underfilled lakes. The density
of arthropod trackways may be high, forming tracked omission surfaces. Some of these omission surfaces
may represent sequence boundaries expressed by co-planar surfaces of lowstand and subsequent flooding.
Rapid changes in depositional conditions reflecting desiccation during vertical aggradation can lead to the
formation of composite ichnofabrics that reflect successive bioturbation events.
© 2008 Elsevier B.V. All rights reserved.

1. Introduction increase in the number of studies applying ichnology to refine

During the 1990s the potential of trace fossils in sequence
stratigraphy was realized (e.g., Pemberton et al., 1992; MacEachern
et al., 1992). In little more than a decade, the field experienced a rapid
⁎ Corresponding author.
E-mail addresses: luis.buatois@usask.ca (L.A. Buatois), gabriela.mangano@usask.ca
(M.G. Mángano).
sequence-stratigraphic analyses (e.g., Savrda et al., 1993; Pemberton
and MacEachern, 1995; Ghibaudo et al., 1996; MacEachern et al., 1999;
Buatois et al., 2002; Pemberton et al., 2004; Gibert and Robles, 2005;
Carmona et al., 2006; Rodríguez-Tovar et al., 2007). At present, trace
fossils are regarded as useful tools in sequence-stratigraphic analyses
of cores and outcrops, and ichnologic aspects are covered in recent
sequence-stratigraphic textbooks (e.g. Catuneanu, 2006). However,
the use of trace fossils in continental sequence stratigraphy has

0031-0182/$ – see front matter © 2008 Elsevier B.V. All rights reserved.
doi:10.1016/j.palaeo.2008.10.012
128 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 272 (2009) 127–142
remained essentially underexplored until very recently (Buatois and
Mángano, 2004).
Buatois and Mángano (2004) noted that application of trace fossils
in continental sequence stratigraphy cannot be based simply on the
extrapolation of ichnologic concepts derived from marine sequence
stratigraphy. Substrate-controlled ichnofacies, in particular the firm-
ground Glossifungites ichnofacies, develop in stable and cohesive
substrates, reflecting erosive exhumation of the sediment, which is
commonly (albeit not invariably) linked to relative sea-level changes
(MacEachern et al., 1992). However, in continental successions,
substrate-controlled trace fossils only rarely indicate erosional exhu-
mation because these suites are commonly related to desiccation of
water bodies, which in turn, is usually linked to autogenic processes
(e.g. Buatois et al., 1996). In addition, continental firmgrounds may
develop rapidly under subaerial exposure, without implying a
significant hiatus (Fürsich and Mayr, 1981).
Furthermore, lacustrine sequence stratigraphy historically has
lagged behind its marine counterpart. Bohacs et al. (2000) pioneered
the potential of sequence stratigraphy to address the complexities of
lacustrine basins (see also Carroll and Bohacs, 1999, 2001; Bohacs
et al., 2003). As noted by these authors, lacustrine systems differ from
oceans in several ways, including the smaller volumes of sediment
and water present in lakes, the direct link between lake level and
sediment supply, and the fact that shoreline migration may be due not
only to progradation but also to withdrawal of water. Bohacs et al.
(2000) recognized three different types of lake basins: overfilled,
balanced-fill and underfilled, which correspond to three broadly
defined facies assemblages, namely fluvial-lacustrine, fluctuating
profundal, and evaporative, respectively (see Carroll and Bohacs,
1999). This model provides an elegant way to summarize in a
systematic fashion the variability of lacustrine systems. The same
framework has been used by Gierlowski-Kordesch and Park (2004) to
analyze changes in species diversity in modern and ancient lakes.
Recently, Buatois and Mángano (2004, 2007) integrated ichnologic
data within this classification framework, and provided well-reasoned
explanations for the occurrence of specific ichnofaunas in overfilled,
balanced-fill, and underfilled lakes. However, only the most general
aspects of the model were addressed, and no examples in support of
the suggested pattern were presented. The aim of this paper is to use
case studies to further explore the potential of lacustrine ichnology in
sequence stratigraphy.
2. Conceptual framework
In this section we briefly address the classification framework of
lake-basin types and associated facies assemblages, which is adopted
in this paper (Carroll and Bohacs, 1999; Bohacs et al., 2000). Also, we
review some ichnologic concepts that are of use to understand po-
tential links between ichnofaunal distribution and lake–basin types.
Overfilled lake basins occur when the rate of sediment/water input
exceeds potential accommodation. Overfilled lakes are commonly
hydrologically open and freshwater, contain fluvio-lacustrine silici-
clastic deposits and display parasequences driven mainly by shoreline
progradation and delta-channel avulsion (Carroll and Bohacs, 1999;
Bohacs et al., 2000). Overfilled lake deposits typically consist of clastic
sedimentary rocks, mostly mudstone and sandstone, but also marls,
coquina, coal, and coaly shale.
Balanced-fill lake basins are characterized by rates of sediment/
water supply in balance with potential accommodation. Carbonate
and siliciclastic facies can accumulate in lakes that are alternatively
hydrologically open and closed, fluctuating from freshwater to saline
(Carroll and Bohacs, 1999; Bohacs et al., 2000). Successions record not
only progradational parasequences, but also aggradation of chemical
sediments due to desiccation. Balanced-fill lake deposits reflect an
alternation of siliciclastic and chemical sediments, including different
types of carbonates, marl, mudstone, siltstone, and sandstone.
Underfilled-lake basins occur when rates of accommodation ex-
ceed rate of supply of sediment/water. In hypersaline and hydro-
logically closed lakes, deposition of evaporites dominates, and
parasequences record vertical aggradation (Carroll and Bohacs,
1999; Bohacs et al., 2000). Deposits are dominated by chemical
sediments, including various types of evaporites and carbonates,
although clastic sediments may be present locally. Structures indi-
cative of subaerial exposure (e.g. desiccation cracks, raindrop im-
prints) are pervasive.
A number of ichnologic concepts are useful to address trace-fossil
distribution in the three lake-basin types. A simple checklist
approach is of little use because none of these lake-basin types is
characterized by specific ichnotaxa. In fact, recent studies have
shown significant overlap of ichnotaxa in the three lake–basin types
(Bohacs et al., 2007). In particular, we favor a combined ichnofacies
and ichnofabric approach to the study of overfilled, balanced-fill, and
underfilled lakes. Ichnofacies (trace fossil assemblages that recur
through long intervals of geologic time, and are characteristic of
a given set of environmental conditions) are produced through a
"distillation" process that concentrates the diagnostic features of
various ichnofaunas, and eliminates the local peculiarities or the
"noise" of the particular examples (Pemberton et al., 1992). Different
invertebrate ichnofacies have been defined for continental envi-
ronments, including Scoyenia (transitional subaerial to subaqueous
freshwater), Mermia (permanent subaqueous freshwater), Coprini-
sphaera (terrestrial, paleosols associated with herbaceous commu-
nities), and Octopodichnus–Entradichnus (eolian dunes) (Seilacher,
1963; Frey et al., 1984; Buatois and Mángano, 1995; Genise et al.,
2000; Ekdale et al., 2007; Hunt and Lucas, 2007). More recently,
archetypal vertebrate ichnofacies have also been introduced, namely
the Chelichnus, Grallator, Brontopodus, Batrachichnus, and Characichich-
nos ichnofacies (Hunt and Lucas, 2007).
The study of ichnofabrics (any aspect of the texture and internal
structure of a substrate resulting from bioturbation and bioerosion at
any scale) emphasizes taphonomic aspects (Bromley and Ekdale,
1986). The notion of taphonomic pathways is particularly useful to
evaluate the interplay between depositional and taphonomic pro-
cesses in specific sedimentary environments, bridging potential gaps
between the ichnofabric approach and the ichnofacies model (Buatois
and Mángano, 2004, 2007). In fact, Bromley and Asgaard (1991) noted
that some ichnofacies are closely related to biofacies, while others are
more akin to taphofacies. In particular, analysis of trace-fossil pre-
servational styles in lacustrine deposits suggests that ichnofaunas
result from various taphonomic pathways that reflect depositional
conditions (subaqueous vs. subaerial), and time spans between
flooding events (Zhang et al., 1998; Buatois and Mángano, 2002,
2004; Minter et al., 2007; Buatois and Mángano, 2007). In addition,
taphonomic pathways help to understand the role of substrate and
rapid environmental fluctuations as major controlling factors in
ichnofacies development and replacement (Buatois and Mángano,
2002, 2004, 2007). This combined approach allows characterization of
vertical changes of archetypal ichnofacies as a response to secular
changes in depositional conditions (e.g. progressive shallowing), and
delineation of palimpsest surfaces resulting from desiccation along
lake margins.
3. Case studies
In this section we review different case studies of lacustrine
ichnofaunas that may serve to detect patterns of trace-fossil distri-
bution in different types of lacustrine basins. The examples discussed
fulfill the following requirements: (1) appropriate systematic treat-
ment of the ichnotaxa present; (2) careful description and interpreta-
tion of the associated sedimentary facies; (3) adequate understanding
of strata stacking patterns and basin-fill history, and (4) documenta-
tion of vertical ichnofaunal changes.
 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 272 (2009) 127–142 129

3.1. The Patquía Formation (Permian, western Argentina)
The Patquía Formation records part of the infill of the Carboniferous–
Permian Paganzo Basin of western Argentina. This basin is regarded as a
foreland basin related to subduction of the Pacific plate beneath the
western continental margin of Gondwana, which may have evolved to a
rift system during the Permian (Ramos, 1988). In the area of Bordo
Atravesado, this unit consists of fluvial conglomerate, sandstone and
mudstone, which are replaced upwards by playa-lake mudstone and
sandstone, and eolian sandstone (Caselli, 1996; Zhang et al., 1998). The
sedimentary facies, stratal stacking pattern, and trace-fossil content of
the ephemeral lacustrine deposits have been analyzed in detail by Zhang
et al. (1998). Lacustrine sedimentation occurs in an underfilled basin
characterized by arid conditions (López Gamundí et al., 1992).
The Bordo Atravesado ichnofauna includes Cruziana problematica,
Diplocraterion isp., cf. Diplopodichnus biformis, Kouphichnium? isp.,
Merostomichnites aicuñai, Mirandaichnium famatinense, Monomor-
phichnus lineatus, Palaeophycus tubularis, Umfolozia sinuosa, and Um-
folozia cf. longula (Fig. 1). Trace fossils are mostly preserved as
hypichnial ridges on the soles of climbing and wave-ripple cross-
laminated fine-grained sandstone deposited by sheet floods under
wave influence in the margins of the playa-lake complex. Desiccation
cracks and raindrop imprints are common throughout the succession
and, in some cases, trace fossils are preserved on top of cracked
surfaces. Trace-fossil preservation was linked to rapid influx of sand
via sheet floods entering into the lake (Zhang et al., 1998). During
pluvial periods, the lake experienced expansion, and flash floods
reached the lake basin, leading to deposition of sandy inundites.
Gypsum layers occur locally, suggesting evaporation and the incipient
formation of saline crusts. Zhang et al. (1998) recognized four
taphonomic variants essentially controlled by substrate consistency
and time averaging. Type 1 consists of low-density assemblages of

Fig. 1. Selected trace fossils from the Permian Patquía Formation of western Argentina. All are sole views, with the exception of C and E, which are upper bedding-plane views. A, B, C.
High density of arthropod trackways, showing numerous superimposed specimens. Note desiccation cracks in A and C, and Diplocraterion isp. overprinted to the background
ichnofauna in A. D. Monomorphichnus lineatus. E. Mirandaichnium famatinense (right) and Umfolozia sinuosa (left). F. Merostomichnites aicuñai. G. Close-up of surface densely covered
by arthropod trackways. H. Cruziana problematica. All scale bars are 1 cm and coins are 1.2 cm.
130 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 272 (2009) 127–142
poorly defined trackways, suggesting that arthropods crawled in soft,
probably slightly subaqueous substrates. Type 2 consists of low- to
moderate-density suites of sharply defined trackways. The style of
preservation and the association with mud cracks indicate that the
arthropods inhabited a firm, desiccated substrate. Type 3 records the
overprint of terrestrial ichnocoenoses over previously formed soft-
ground suites, representing palimpsestic bedding surfaces, Type 4 is
similar to type 2, but displays a high density of trace fossils, showing
intense overlap, and representing omission surfaces formed during
major time gaps of subaerial exposure before sheet-flood events.
Virtually no trace fossils occur in the sediments of the lake-center
environment due to hypersaline conditions. The playa-lake ichno-
fauna from the Patquía Formation illustrates the role of taphonomic
factors and time gaps in controlling the expression of lake-margin
surfaces delineated by the high density of arthropod trackways.
3.2. The Agua de la Peña Group (Middle to Upper Triassic, western
Argentina)
The Middle to Upper Triassic Agua de la Peña Group contains
several lacustrine units (e.g. Chañares, Ischichuca and Los Rastros
formations) of the Ischigualasto–Villa Unión Basin of western
Argentina. This basin is one of the various rift basins developed on
the western margin of Gondwana during the breakup of Pangea
(Uliana and Biddle, 1988; Milana and Alcober, 1994). The sedimentary
facies and stratal architecture of these lacustrine deposits have been
analyzed in detail by Milana (1998) and Melchor (2007), while their
ichnologic content has been documented by Melchor (2001, 2004)
and Melchor et al. (2003). The Agua de la Peña Group has been
subdivided into four depositional sequences (SS 1–4) recording
overfilled, balanced-fill, and underfilled lake–basin types (Melchor,
2007). The combination of variable depositional regimes and careful
sedimentologic, ichnologic and stratigraphic studies (Melchor et al.,
2003; Melchor, 2004, 2007) makes the Agua de la Peña Group an ideal
unit to address trace-fossil distribution in the different lake-basin
types.
The Chañares Formation and the lowermost part of the Ischichuca
Formation represent deposition in underfilled lakes that fluctuated
from mildly saline to playa lakes surrounded by mud flats (Melchor,
2007). Tetrapod trackways occur in the mud-flat deposits associated
with desiccation cracks. These units illustrate the stressful conditions
prevalent in underfilled lakes, but also the high potential for trackway
preservation in surrounding areas.
The bulk of the Ischichuca Formation records deposition in a
balanced-fill lake characterized by alternating shallow- and deep-
water conditions, represented by the fluctuating profundal facies
assemblage (Melchor, 2007). Deposits form distinctive coarsening-
upward parasequences reflecting progradation of wave- and river-
dominated deltas, and are stacked forming two depositional sequences
(SS 2–3 of Melchor, 2007). The boundary between both sequences is
represented by a forced-regressive surface. Offshore underflow-
current deposits do not contain trace fossils, probably reflecting anoxic
conditions in the hypolimnion of a freshwater stratified lake (SS 2).
However, delta-plain channel deposits present towards the top of SS 2
contain escape trace fossils, recording rapid sedimentation in a river-
dominated context (Melchor, 2004). Lower delta-plain deposits
present at the top of SS 3 contain a moderately diverse ichnofauna
dominated by locomotion trace fossils (Cruziana problematica, Un-
dichna britannica, Diplichnites isp., Stiaria isp.), together with resting
(Rusophycus stromnessi), and grazing (Cochlichnus anguineus) trace
fossils (Melchor, 2004). The Ischichuca Formation illustrates the
complex distribution of stress factors (e.g. anoxic conditions) in a
balanced-fill lake affected by deltaic progradation.
By far, the richest ichnologic record is present in the Los Rastros
Formation, which records deposition in a shallow overfilled lake (SS 4
of Melchor, 2007). Deposits are commonly arranged in coarsening-
upward parasequences reflecting deltaic progradation. Upper delta-
plain deposits contain simple dwelling trace fossils, some of which
contain striations (e.g. Palaeophycus striatus) and vertebrate trackways
(Rhynchosauroides isp.). Upper delta-front to lower delta-plain
deposits contain a few ichnotaxa, mostly dwelling trace fossils of
suspension feeders (Palaeophycus tubularis, Skolithos isp.), although
Cochlichnus anguineus is also present. Some of the delta-front trace
fossils (Skolithos isp., Palaeophycus tubularis) occur in hummocky cross-
stratified sandstone, recording opportunistic colonization after storm
events. Middle delta-front deposits also contain a relatively diverse
ichnofauna dominated by simple grazing trails (Helminthoidichnites
tenuis, Helminthopsis abeli, Gordia indianaensis, Archaeonassa fossulata,
C. anguineus). Fish locomotion trails (Undichna britannica) and simple
facies-crossing dwelling trace fossils (P. tubularis) are also present
(Melchor et al., 2003). Underflow-current deposits of the distal delta
front are the most ichnologically diverse, containing a wide variety of
simple grazing trails (H. tenuis, Gordia marina, A. fossulata, C. anguineus)
and fish trails (U. britannica, U. bina, U. cf. insolentia). Feeding structures
(Treptichnus pollardi), horizontal dwelling structures (Palaeophycus
tubularis), and arthropod trails (Cruziana problematica, Diplopodichnus
biformis, Didymaulichnus lyelli), resting traces (Rusophycus stromnessi,
Avolatichnium isp.) and trackways (Bifurculapes isp., Diplichnites isp.,
Protichnites isp.) are also present (Melchor, 2001). The Los Rastros
Formation illustrates vertical patterns of trace-fossil distribution in an
overfilled lake affected by wave-dominated deltaic progradation.
3.3. The Lockatong Formation (Upper Triassic, Pennsylvania, United States)
The Upper Triassic–Lower Jurassic Newark Supergroup of north-
east United States represents the infill of the Newark Basin, a rift basin
formed during the initial breakup of Pangaea (Olsen, 1989; Schlische,
2003). Lacustrine deposits occur in a number of stratigraphic units
within this basin. In particular, the Upper Triassic Lockatong Formation
of southeastern Pennsylvania represents deposition under balanced-
fill conditions characterized by recurrent lake-level fluctuations,
exhibiting a series of vertically stacked transgressive–regressive cycles
(Olsen, 1980). The sedimentary facies of this unit have been addressed
by Olsen (1980, 1989), while the invertebrate trace fossils were
described by Metz (1995) and the vertebrate trackways by Olsen and
Flynn (1989).
The invertebrate ichnofauna from the Lockatong Formation
consists of Cochlichnus anguineus, Lockeia siliquaria, Planolites mon-
tanus, Scoyenia gracilis, Spongeliomorpha milfordensis, Treptichnus
pollardi, and an undetermined arthropod trackway (Metz, 1995). The
reptile trackway Gwyneddichnium is present also (Olsen and Flynn,
1989). Biogenic structures are essentially restricted to lake-margin
deposits. Some of these ichnotaxa may have been emplaced in wet
substrates (e.g. T. pollardi and L. siliquaria), but the distinct orna-
mentation in other forms (e.g. S. gracilis and S. milfordensis) clearly
indicates firm substrates due to desiccation. Metz (1995) noted that
the greatest diversity and abundance of trace fossils occur at the top of
regressive packages. In contrast, the onset of the transgression and the
lowstand are characterized by sparse bioturbation. Under extremely
dry conditions, Spongeliomorpha and Scoyenia are the only ichnotaxa
present. Lake-center deposits only contain sparse indistinct trace
fossils, as a result of oxygen-depleted conditions. The Lockatong
Formation ichnofauna illustrates the importance of oxygen-depleted
conditions in lake-center environments and the development of
substrate-controlled suites due to progressive desiccation at the lake
margins of balanced-fill basins.
3.4. The Anyao Formation (Lower Jurassic, central China)
The Lower Jurassic Anyao Formation is approximately 100 m thick,
and represents part of the infill of the Late Triassic–Middle Jurassic
Jiyuan–Yima Basin. It records sedimentation in a deep overfilled lake
 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 272 (2009) 127–142 131

system developed in a pull-apart basin (Buatois et al., 2000a).
Sedimentary facies have been discussed by Buatois et al. (2000a),
and the ichnology was documented by Wu (1985) and Buatois et al.
(1996). Turbidite systems were fed from fan deltas advancing from the
south. Vertical aggradation of turbidite lobes fed from multiple
sources is revealed by the stacking of thin-bedded turbidites, classical
turbidites, and massive turbidites forming coarsening-upward succes-
sions. Final infill of the lake basin is indicated by delta progradation
recorded in the overlying Middle Jurassic Yangshuzhuang Formation.
The thin-bedded turbidites record sedimentation in lobe fringe
areas, and contain relatively diverse ichnofaunas, representing both
pre- and post-event suites (Buatois et al., 1996). The pre-event suite
consists of Helminthopsis abeli, Helminthoidichnites tenuis, discrete
specimens of Tuberculichnus vagans, Monomorphichnus lineatus, Par-
acanthorhaphe togwunia, and thin irregular trails (Fig. 2). The post-
event suite consists of Vagorichnus anyao, Cochlichnus anguineus and
Helminthopsis hieroglyphica; specimens of Gordia marina, and Tuber-
culichnus vagans intergrading with V. anyao are also part of the post-
event suite. The ichnofauna is dominated by feeding and grazing trace
fossils produced by deposit feeders. The Anyao ichnofauna illustrates
environmental conditions in a deep, freshwater overfilled lake, in
which turbidity currents provide oxygenation to the lake bottom,
allowing the establishment of a relatively diverse benthos. High
sedimentation rate, high energy, continuous erosion, and intense soft-
sediment deformation preclude preservation of biogenic structures in
more proximal areas of the turbidite system.
3.5. La Huérguina Limestone Formation (Lower Cretaceous, Las Hoyas
fossil site, central Spain)
Las Hoyas sub-basin of the Iberian Basin was an asymmetrical and
subsiding block tilted towards the SE and filled with alluvial and
lacustrine sediments included in the La Huérguina Limestones
Formation. The sedimentary facies and depositional dynamics of the
lacustrine system have been addressed by Fregenal-Martínez (1998)
and Fregenal-Martínez and Meléndez (2000). The vertebrate ichno-
fauna has been documented by Moratalla et al. (1995), Lockley et al.
(1995) and Gibert et al. (1999), while the invertebrate ichnofossils
were analyzed by Buatois et al. (2000b). These deposits are organized
in four sedimentary sequences separated by local unconformities
(Fregenal-Martínez, 1998; Fregenal-Martínez and Meléndez, 2000).
The Las Hoyas Fossil Site was formed during the deposition of the
second depositional sequence (La Rambla de las Cruces II). The lacustrine
deposits preserved in this sequence contain four facies belts, one

Fig. 2. Selected trace fossils from the Lower Jurassic Anyao Formation of central China. All trace fossils are preserved at the base of sandstone turbidites. A. Vagorichnus anyao.
B. Paracanthorhaphe togwunia. C. Tuberculichnus vagans. D. Intergrading Vagorichnus anyao and Tuberculichnus vagans. E. Helminthopsis hieroglyphica. F. Helminthopsis abeli.
G. Helminthoidichnites tenuis. All scale bars are 1 cm.
132 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 272 (2009) 127–142
palustrine belt, and three belts of lacustrine facies (supralittoral and
eulittoral realm, intralittoral and sublittoral realm, and basinal realm)
(Fregenal-Martínez, 1998; Fregenal-Martínez and Meléndez, 2000). The
Las Hoyas paleolake was a shallow and permanent, balanced-fill lake,
dominated by carbonate deposition. The fossil site occurs in the
rhythmically laminated facies of the basinal belt, consisting of pale
grey to black, varve-scale laminated carbonate mudstone deposited
from low-density turbidity currents and underflow currents (Fregenal-
Martínez, 1998; Sanz et al., 2001). The majority of the trace fossils
occur in rhythmically laminated limestone at the central part of the fossil
site and are concentrated along a few bedding surfaces. Root traces occur
in the associated palustrine carbonates (Gierlowski-Kordesch et al.,
1991).
The vertebrate ichnofauna consists of crocodilian and turtle
trackways (Moratalla et al., 1995), two isolated tracks, probably pro-
duced by a pterosaurian and comparable to Pteraichnus (Lockley et al.,
1995), and the fish trail Undichna unisulca (Gibert et al., 1999). The
invertebrate ichnofauna includes Cruziana problematica, Helminthoi-
dichnites tenuis, Lockeia isp., Palaeophycus tubularis, and Treptichnus
pollardi (Buatois et al., 2000b) (Fig. 3). Invertebrate trace fossils are
rare, and represented only by tiny epifaunal or very shallow trace
fossils restricted to bedding planes. Dominance of epifaunal structures
and paucity of infaunal traces may indicate short periods of oxy-
genated bottom waters due to the passage of underflow and turbidity
currents, but permanently anoxic interstitial waters existed within the
sediment. The Las Hoyas ichnofauna underscores the role of oxygen
depletion as a controlling factor in carbonates from a balanced-fill
lake.
3.6. The Yenimahalle Formation (Lower Miocene, southern Turkey)
The Neogene Ermenek Basin of southern Turkey contains up to
300 m of thick carbonate and siliciclastic lacustrine deposits that
accumulated in an intramontane graben affected by strike–slip
movements (Andrew and Robertson, 2001; Ilgar and Nemec, 2005).
Clastic-dominated lacustrine deposits, mostly recording balanced-fill
conditions, are included in the Lower Miocene Yenimahalle Forma-
tion. The sedimentary facies and sequence architecture of these
deposits have been documented by Ilgar and Nemec (2005), while the
trace fossils were analyzed by Uchman et al. (2007). The lacustrine
basin-fill has been subdivided into four depositional sequences
Fig. 3. Selected trace fossils from the Lower Cretaceous La Huérguina Limestones Formation of central Spain. All trace fossils are preserved at the base of rhythmically laminated
limestones. A. Helminthoidichnites tenuis. B. Palaeophycus tubularis. C. Cruziana problematica. D. Undichna unisulca. All scale bars are 1 cm.
bounded by forced-regression surfaces (Ilgar and Nemec, 2005). In
proximal settings, a thick lowstand systems tract of alluvial fan
deposits is overlain by variably developed transgressive and highstand
systems tracts represented by fan-delta deposits. In distal settings, a
thin lowstand systems tract of shoreface deposits is overlain by
thick transgressive deposits formed by sandstone tempestites and
delta-generated turbidites with poor or no preservation of highstand
deposits.
Lacustrine offshore-transition tempestites contain an assemblage
dominated by Vagorichnus cf. anyao together with some indistinct
forms (undulating ridges and small discontinuous ridges) (Uchman
et al., 2007). The pre-event suite consists of V cf. anyao and probably
some of the small discontinuous ridges, while the post-event suite is
represented by the undulating ridges and most of the small
discontinuous ridges. The relative shallowness of the lake (maximum
depth of 10 m, and storm wave base around 5 m have been estimated)
may have provided a considerable wave fetch for fair-weather and
storm-wave generation. Modest waves occur in the lake embayment,
punctuating background sedimentation characterized by deposition
of marls and micritic limestone. The pre-event suite in this setting is
represented by oblique cylindrical trace fossils, while the post-event
suite is dominated by large tubular structures and planar wall struc-
tures. No trace fossils were found in the sandy turbidites, although
burrow mottlings occur in the associated muddy deposits. Lacustrine
bottom waters were most likely oxygen depleted, hard, acidic and rich
in CO2, imposing stressful conditions on the benthic fauna, and being
conducive to a reduction in ichnodiversity. The Yenimahalle ichno-
fauna illustrates how lake chemistry and oxygen conditions strongly
control ichnodiversity levels in a dominantly siliciclastic succession of
a balanced-fill lake.
3.7. The “Gypsum and grey shales” unit (Middle to Upper Miocene, northeast
Spain)
The Neogene Calatayud–Teruel Basin of northeast Spain is a half-
graben structure that contains thick (up to 500 m) and laterally
extensive lacustrine deposits dominated by carbonates and evapora-
tive facies (e.g. Alonso-Zarza and Calvo, 2000; Aziz et al., 2003; Ortí
et al., 2003). In particular, the Calatayud sub-basin hosts proximal to
medial alluvial-fan conglomerate, breccia and sandstone near the sub-
basin margin that interfinger with mudstone, marl, carbonate and
 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 272 (2009) 127–142 133

evaporite towards the sub-basin center (Uchman and Alvaro, 2000;
Aziz et al., 2003). At the Terrer locality, a unit referred to as the
“Gypsum and grey shales” by del Olmo et al. (1983) represents an
intermediate facies belt. This unit consists of reddish shale, green
sandy marlstone and gypsum, the latter increasing in abundance
towards the top (Uchman and Alvaro, 2000). These deposits record
sedimentation in evaporatic underfilled lakes.
The trace-fossil content of the “Gypsum and grey shales” has been
analyzed in detail by Uchman and Alvaro (2000), who recognized a
complex array of trace-fossil suites mostly concentrated in a single
sandy marlstone bed and the underlying red mudstone. The sandy
marlstone bed contains a number of overprinted suites. Celliforma isp.,
Celliforma? ispp., Roselichnus cf. arabicus, and Labyrintichnus terrer-
ensis are part of a suite dominated by bee nests and were emplaced in
a dry, subaerially exposed substrate reflecting incipient paleosol
development. Cohesive, desiccated firmgrounds were colonized by
crustaceans that constructed striated burrow systems attributed to
Spongeliomorpha isp. Taenidium barreti, a meniscate trace fossil
without ornamentation, was most likely emplaced in moist substrates.
Polykladichnus aragonensis and Arenicolites isp. are also present in the
sandy marlstone bed, but their substrate affinities are unclear. The red
mudstone contains Beaconites filiformis, a lined meniscate trace fossil
probably produced by salinity-tolerant chironomid larvae. This
ichnotaxon seems to record a wide range of environmental conditions
because it not only occurs in association with desiccation cracks, but
also in deposits that are thought to have been formed under sub-
aqueous conditions (Uchman and Alvaro, 2000). The “Gypsum and
grey shales” ichnofauna provides an excellent example of the impact
of stress factors (i.e. hypersalinity) in the subaqueous portion of
evaporite lakes, and of the complex cross-cutting relationships of
suites in the margins of underfilled lakes.
4. Summary of observations and discussion of the model
The case studies discussed provide useful examples to illustrate
trace-fossil distribution in different lake-basin types. These examples,
together with some additional ones discussed below, constitute a
valuable database to evaluate the potential of ichnology in lacustrine
sequence stratigraphy. Table 1 summarizes the most important
sedimentologic and ichnologic characteristics of overfilled, balanced-
fill, and underfilled lakes. Available information suggests that trace-
fossil distribution in these lacustrine systems essentially reflects the
specific combination of stress factors, sediment type, and the position
of the water table. Stress factors are of paramount importance in
controlling ichnodiversity, and sediment type is a key limiting factor in
benthic colonization (Buatois and Mángano, 1998, 2004; Hasiotis,
2004; Bromley et al., 2007; Bohacs et al., 2007). The importance of the
water table has been emphasized in a number of ichnologic studies
(e.g. Gierlowski-Kordesch, 1991; Buatois and Mángano, 2004; Gibert
and Sáez, 2009-this issue). Because stress factors, sediment type, and
water table position vary in a systematic way in the different lake-basin
types, an integrated sedimentologic-ichnologic model within the
classification framework of overfilled, balanced-fill, and underfilled
lakes can be proposed (Buatois and Mángano (2004, 2007). Cohen
(2003) listed a number of abiotic and biotic factors that control animal
distribution in lakes. Abiotic factors include energy, light, oxygen,
temperature, salinity, substrate and nutrients, while biotic factors,
such as competition, grazing, predation and symbiosis, have complex
feedback loops and are difficult to interpret (see also Miller and White,
2007). Species diversity results from a complex interplay of these
different factors. Taphonomic factors also play a major role in
controlling ichnodiversity, and are essential to understand the
ichnologic expression of key stratal surfaces.

Table 1
Characteristics of lake-basin types and associated ichnofaunas

Lake–basin Lacustrine Stratal stacking Lake Water Environmental controls Ichnofacies distribution Ichnofabric Surface expression
type facies pattern hydrology table types
association
Overfilled Fluvial– Dominantly Open High Freshwater Well oxygenated
Mermia in subaqueous Dominantly Poorly developed
lacustrine progradational High rates of clastic portions Scoyenia simple Flooding surfaces and
Parasequences sedimentation High energy
(softground suite) in low- sequence boundaries
related to deltaic and intense erosion inenergy lake margins revealed by vertical
progradation proximal areas MinimumSkolithos in high-energy changes in softground
desiccation lake margins Progressive suites
replacement of softground
ichnofacies in
parasequences
Balanced-fill Fluctuating Mixed Open during Fluctuating Freshwater during TST-HST Depauperate Mermia in Dominantly Well developed
profundal progradational TST-HST and and hypersaline during LST subaqueous portions (albeit composite (palimpsestic) Flooding
and aggradational closed during Poorly oxygenated during rare) Scoyenia (dominantly surfaces and sequence
Parasequences LST TST-HST and well oxygenated firmground suite boundaries revealed
related to delta during LST High rates of overprinted to the by vertical changes in
progradation and clastic sedimentation during softground suite) in low- softground suites and
vertical aggradation HST, but low during TST energy lake margins by suite overprinting
due to desiccation High rates of chemical Skolithos in high-energy
sedimentation during LST lake margins Limited
High energy and intense replacement of softground
erosion in proximal areas ichnofacies in
during HST Minimum parasequences
desiccation during TST–LST,
but maximum during LST
Underfilled Evaporitive Dominantly Closed Low Hypersaline Variable oxygen Scoyenia (dominantly Dominantly Well developed
aggradational content High rates of firmground suite composite (palimpsestic) Co-
Parasequences chemical sedimentation Low overprinted to the planar surfaces
related to vertical energy and little erosion softground suite) in lake revealed by tracked
aggradation due Maximum desiccation margins Octopodichnus– omission surfaces and
to desiccation Entradichnus in associated suite overprinting
eolian dunes No
replacement of softground
ichnofacies in
parasequences
134 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 272 (2009) 127–142
4.1. Overfilled lake basins (Figs. 4 and 5)
Overfilled lakes are the least stressful of the three types of lake
basins. These are freshwater lakes where no stress due to hypersa-
linity occurs, favoring the establishment of a relatively diverse
benthos. In addition, fluvial discharge into overfilled lakes usually
generates density currents that oxygenate lake bottoms, allowing the
establishment of epifaunal and infaunal communities (Buatois and
Mángano, 1998; Melchor, 2004). In the case of deep overfilled lakes,
relatively large basin-floor turbidite systems can develop. Land-plant
derived organic matter is the prime source of nutrients, favoring the
development of a deposit-feeding benthic fauna in permanently
subaqueous, low-energy zones (Figs. 4 and 5).
The Mermia ichnofacies is typical of the middle to distal regions of
turbidite-lobe successions, comprising both pre- and post-deposi-
tional suites in thin-bedded turbidite sandstone (e.g., Buatois et al.,
1996; Buatois and Mángano, 1998) (see Section 3.4). Frequent erosion
and high rates of sedimentation in the proximal portion of turbidite
systems prevent development or preservation of biogenic structures.
Integration of ichnologic and sedimentologic data may help to
delineate environmental zonations in aggradational and prograda-
tional turbidite lobes. In shallow overfilled lakes, distal facies
commonly consist of delta-fed underflow-current and background-
fallout deposits hosting the Mermia ichnofacies (see Section 3.2).
Intermediate facies may contain wave-dominated delta-front and
nearshore deposits, including storm-emplaced hummocky cross-
stratified sandstone and fair-weather wave- and combined-flow ripple
cross-laminated sandstone. Grazing trails of the Mermia ichnofacies
may form colonization suites at the top of storm beds in such settings.
However, assemblages are usually impoverished with respect to those
of the more distal facies (Buatois and Mángano, 1998). Under con-
ditions of moderate to high energy due to continuous wave agitation,
the Skolithos ichnofacies may be present. Proximal facies include
distributary-channel, trough and tabular cross-bedded sandstones
that are commonly unbioturbated. Locally, these deposits may contain
escape traces and vertical domiciles of suspension feeders, represent-
ing the Skolithos ichnofacies (e.g. Melchor et al., 2003).
Fig. 4. Trace-fossil assemblages, environmental controls, and lacustrine sequence stratigraphy in overfilled lakes. Modified from Buatois and Mángano (2004) with stratal patterns
illustrated after Bohacs et al. (2000).
Overfilled lake basins contain well-developed softground trace
fossils that are useful to delineate parasequences and parasequence
sets (e.g. Melchor et al., 2003; Melchor, 2004). Upward shallowing
successions due to delta and shoreline progradation are typical.
Firmground suites are rare because such large lakes usually do not
experience desiccation. As a result, low-energy lake margins com-
monly contain the pre-desiccation suite of the Scoyenia ichnofacies. In
these settings, Taenidium and Beaconites replace Scoyenia as the
dominant meniscate trace fossils.
4.2. Balanced-fill lake basins (Figs. 6 and 7)
Balanced-fill lakes are extremely complex ichnologically because
they are characterized by fluctuating conditions, which commonly
result in a combination of stress factors that impact negatively on the
benthic fauna. Abundant firmground trace-fossil suites occur in
balanced-fill lakes, but softground assemblages are usually depaupe-
rate, particularly with respect to fully subaqueous suites. During
lowstands, shallow balanced-fill lakes are characterized by relatively
thin aggradational parasequences due to desiccation (Bohacs et al.,
2000). Lowstand deposits contain abundant and widespread ichno-
faunas of the Scoyenia ichnofacies. In particular, the firmground suite
of this ichnofacies, containing striated trace fossils, such as Scoyenia
and Spongeliomorpha, shows widespread development due to lake
desiccation, commonly cross-cutting the softground pre-desiccation
suite (e.g. Bromley and Asgaard, 1979; Metz, 1995; Clemmensen et al.,
1998). Biogenic structures are usually preserved during subsequent
flooding by rapid influx of sand (Figs. 6 and 7).
Relatively thick aggradational parasequence sets form in lake-floor
turbidite systems in deep balanced-fill lakes during lowstands (Bohacs
et al., 2000). Under these conditions, firmground trace fossils are
absent. Lake hydrology is closed during lowstands and salinity usually
increases (Bohacs et al., 2000), imposing a stress factor on the lake
biota and, therefore, softground ichnofaunas are depauperate. During
transgression, parasequences are relatively thick, displaying retro-
gradational stacking patterns, while highstand parasequences are
variable in thickness, and are either aggradational or progradational
 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 272 (2009) 127–142 135

Fig. 5. Trace-fossil distribution in sedimentary sequences of overfilled lake basins. Note the dominantly progradational stacking pattern related to deltaic progradation, the
overwhelming dominance of softground suites, and the progressive replacement of ichnofacies due to shallowing.

(Bohacs et al., 2000). Freshwater conditions are common during As a result, ichnofaunas from turbidite systems in balanced-
transgression, but dysaerobic conditions may prevail, imparting a fill lakes are less abundant and diverse than those from overfilled
stress factor on the benthic biota. lake turbidites. For example, Buatois and Mángano (2007) noted
136 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 272 (2009) 127–142
Fig. 6. Trace-fossil assemblages, environmental controls, and lacustrine sequence stratigraphy in balanced-fill lakes. Modified from Buatois and Mángano (2004) with stratal patterns
illustrated after Bohacs et al. (2000).
that trace fossils are notably absent in the thin-bedded turbidites
of the balanced-fill Cretaceous Candeias Formation (Recôncavo
Basin of Brazil), but abundant and diverse in identical facies of the
overfilled Jurassic Anyao Formation (see Section 3.2). Also, Uchman
et al. (2007) noted an absence of trace fossils in thin-bedded
sandstone turbidites of the Lower Miocene Yenimahalle Formation
(see Section 3.6).
Trace fossils may occur locally in transgressive and highstand
carbonates. However, ichnodiversity is low and trace fossils are pro-
duced by epifaunal organisms. The presence of very shallow structures
and a paucity of infaunal traces are associated with brief periods of
oxygenated bottom waters, but the interstitial waters permanently are
anoxic. This is illustrated by the Cretaceous Las Hoyas ichnofauna,
which records the presence of a depauperate Mermia ichnofacies (e.g.
Buatois et al., 2000b) (see Section 3.5). In some cases, presence of
arthropod trackways in anoxic lacustrine deposits reveals the activity
of doomed pioneers (Gibert et al., 2000). Additionally, the pre-
servation potential of trace fossils in carbonates is low due to
diagenetic alteration. Lake-margin palustrine carbonates commonly
contain a wide variety of root structures (Alonso-Zarza, 2003).
Elements of the Skolithos ichnofacies may occur in delta mouth bars
during highstand progradation of deltaic systems (Bromley and
Asgaard, 1979; Mángano et al., 1994, 2000).
The paleoecological and paleoenvironmental significance of
lacustrine ichnofabrics dominated by vertical burrows is unclear.
Suites containing Arenicolites, Skolithos, and Polykladichnus have been
commonly recorded in lacustrine deposits (e.g. Bromley and Asgaard,
1979, 1991; Uchman and Alvaro, 2000; Magyar et al., 2006; Gingras
et al., 2007). In some cases, they are associated with sandstone
tempestites and have been regarded as produced by opportunistic
colonization (e.g. Bromley and Asgaard, 1979, 1991; Magyar et al.,
2006). Some of these burrows occur in sandstone layers that are
interbedded with poorly oxygenated mudstone and, therefore, it has
been suggested that they represent temporary oxygenation of the lake
bottom due to storms (Magyar et al., 2006). This situation may
represent an example of the Skolithos ichnofacies (or Arenicolites
ichnofacies of Bromley and Asgaard, 1991) in a lacustrine context (see
Buatois and Mángano, 1998). However, it has been shown that similar
structures are formed in modern lakes by chironomid larvae to harvest
oxygen from the uppermost photosynthetic layer in otherwise
oxygen-deployed sediments (Gingras et al., 2007). In addition, vertical
burrows may be preserved in sandstone emplaced by relatively high-
energy events, but reflect colonization but subsequent withdrawal of
surface water and groundwater (e.g. Bohacs et al., 2007; Netto, 2007).
Evaluation of the significance of ichnofabrics dominated by vertical
burrows in continental deposit should be carefully assessed on a case-
by-case basis.
Species diversity in balanced-fill lakes is also affected by changes
from hydrologically open to closed systems (Gierlowski-Kordesch and
Park, 2004). In particular, the freshwater fish fauna shows a decrease
in diversity under closed hydrology due to a salinity stress, retreating
via streams to freshwater sanctuaries (e.g. Joyce et al., 2005). Sub-
sequently, with the return to freshwater conditions during times of
open hydrology an increase in species diversity increase takes place.
As a result, Gierlowski-Kordesch and Park (2004) noted that overall
species diversity in balanced-fill lakes is even higher than in overfilled
lakes.
4.3. Underfilled-lake basins (Figs. 8 and 9)
Hypersalinity is undoubtedly the most stressful factor in under-
filled lakes and, accordingly, species diversity is extremely low. Locally
oxygen-depleted conditions may occur in some deep underfilled lakes
as a result of water stratification during an oligohaline phase (e.g. Ortí
 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 272 (2009) 127–142 137

Fig. 7. Trace-fossil distribution in sedimentary sequences of balanced-fill lake basins. Note the mixed progradational and aggradational stacking pattern, paucity of subaqueous suites,
and the common superimposition of softground and firmground suites in lake-margin deposits.

et al., 2003), imposing an additional stress factor. Lowstand deposition lack biogenic structures. In the remaining zones, sediments that
is characterized by evaporite accumulation in remnant pools devel- accumulated during the previous highstand experience extreme
oped in the zones of maximum subsidence (Bohacs et al., 2000). desiccation during lowstand (Bohacs et al., 2000). A species-diversity
Evaporite pools are very stressful environments, and almost invariably increase is most evident at the lake margins due to lake-level rise and
138 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 272 (2009) 127–142
Fig. 8. Trace-fossil assemblages, environmental controls, and lacustrine sequence stratigraphy in underfilled lakes. Modified from Buatois and Mángano (2004) with stratal patterns
illustrated after Bohacs et al. (2000).
the associated decrease in salinity, while low-diversity levels are
maintained in the central part of the lake due to continuous accu-
mulation of saline groundwater and chemical stratification (mero-
mixis) (Cohen, 2003) (Figs. 8 and 9).
The Scoyenia ichnofacies is widespread in underfilled lake basins,
but the Mermia ichnofacies is commonly absent. The Scoyenia
ichnofacies is associated with lowstand desiccated substrates in
underfilled lakes (e.g. Gierlowski-Kordesch, 1991; Metz, 1996, 2000).
Large meniscate trace fossils filled with gypsum and micrite, locally
containing pellets and forming moderate- to high-density assem-
blages, have been observed in saline lake deposits (Rodriguez-Aranda
and Calvo, 1998; Ortí et al., 2003). The density of arthropod trackways
and insect trace fossils may be high, forming tracked omission
surfaces, commonly at the top of sheet-flood deposits (e.g. Zhang et al.,
1998; Minter et al., 2007; Scott et al., 2007; Bohacs et al., 2007) (see
Section 3.1). Some of these omission surfaces may represent sequence
boundaries expressed by co-planar surfaces of lowstand and sub-
sequent flooding (e.g. Scott et al., 2009-this issue). These surfaces may
contain several trace-fossil suites reflecting specific conditions of
water availability and substrate consistency. Lake-level fluctuations,
particularly in gently dipping lacustrine coastal plains, allow the
establishment of complex cross-cutting relationships due to suite
overprinting. This is particularly evident where these surfaces involve
more than one transgressive–regressive cycle (Scott et al., 2009-this
issue). Lake-margin omission surfaces commonly contain extensive
surfaces with high density of tetrapod trackways or tracksites (e.g.
Farlow and Galton, 2003; Szajna and Hartline, 2003). In terms of
vertebrate ichnofacies, lake-margin trackway suites commonly belong
to the Grallator ichnofacies (Hunt and Lucas, 2007). Bird footprints
(e.g. Charadriipeda, Gruipeda) are also common (e.g. Bohacs et al.,
2007; Gibert and Sáez, 2009-this issue), illustrating the shorebird
ichnofacies of Hunt and Lucas (2007).
Rodriguez-Aranda and Calvo (1998) showed that the relative
proportion of grass roots versus bush roots may be used as indicator of
saline conditions in mudflats. Bush root traces of halophilic plants
occur in gypsum deposits that accumulated in mud-flat areas,
indicating that vegetation colonized calcium sulfate-rich substrates
after desiccation. Root traces of halophilic grasses are present in
carbonate ponds formed in mud flats, revealing germination and rapid
growth after episodic floodings.
Hypersalinity usually prevents the establishment of a subaqueous
Mermia ichnofacies during transgression and highstand. However,
elements of the Mermia ichnofacies may occur, albeit in reduced
numbers, in very shallow-water thin deposits immediately above
flooding surfaces at the base of parasequences. This assemblage is
abruptly replaced upward by the Scoyenia ichnofacies reflecting lake
regression (e.g. Metz 1996, 2000). Elements of the Mermia ichnofacies
are commonly cross-cut by elements of the Scoyenia ichnofacies,
forming palimpsest surfaces in lake-margin zones (e.g. Metz, 1995;
Uchman and Alvaro, 2000) (see Section 3.7). Additionally, dwelling
traces possibly produced by aquatic chironomid larvae may be present
(Rodriguez-Aranda and Calvo, 1998; Uchman and Alvaro, 2000).
Although high salinity essentially excludes bioturbating organisms
(Gierlowski-Kordesch and Rust, 1994), moderately high salinity allows
establishment of a salinity-tolerant ichnofauna (Rodriguez-Aranda
and Calvo, 1998). In fact, an intensely bioturbated gypsum facies is
common in moderately high salinity lakes of Neogene Basins in Spain,
probably as a result of the work of chironomids (Rodriguez-Aranda
and Calvo, 1998; Ortí et al., 2003). Marl deposition commonly
indicates brine dilution and more benign conditions, which result in
 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 272 (2009) 127–142 139

Fig. 9. Trace-fossil distribution in sedimentary sequences of underfilled lake basins. Note the dominantly aggradational stacking pattern as a result of vertical aggradation due to
desiccation, and typical absence of bioturbation in most of the subaqueous deposits as a result of hypersalinity. Instead of progressive replacement of ichnofacies throughout the
stratigraphic column, a complex overlap of suites characterizes the lake-margin interval reflecting omission surfaces formed in response to desiccation.

the establishment of other tracemakers, most likely coleopterans
(Rodriguez-Aranda and Calvo, 1998).
Transgressive systems tracts recorded by thin backsteping deposits
usually reflect drastic ichnofaunal changes, from terrestrial assem-
blages (Coprinisphaera ichnofacies) to transitional terrestrial-subaqu-
eous assemblages (Scoyenia ichnofacies), and salinity-tolerant
subaqueous monospecific assemblages of Beaconites filiformis attrib-
uted to chironomids (Uchman and Alvaro, 2000). Rapid changes in
depositional conditions reflecting desiccation during vertical aggra-
dation led to the formation of composite ichnofabrics reflecting
successive bioturbation events. Eolian dunes may flank overfilled
lakes in arid settings. The Octopodichnus–Entradichnus ichnofacies,
140 L.A. Buatois, M.G. Mángano / Palaeogeography, Palaeoclimatology, Palaeoecology 272 (2009) 127–142
dominated by arthropod trackways and burrows, may be present in
eolian dune sandstones (Ekdale et al., 2007; Hunt and Lucas, 2007).
5. Conclusions
(1) Application of ichnofossils in lacustrine sequence stratigraphy
cannot be based simply on the extrapolation of concepts
derived from marine sequence stratigraphy. Marine substrate-
controlled ichnofacies, in particular the firmground Glossifun-
gites ichnofacies, develop in stable and cohesive substrates,
reflecting erosive exhumation of the sediment, commonly due
to allogenic processes. In contrast, in lacustrine sequences,
substrate-controlled ichnofossils only rarely indicate erosional
exhumation because they are commonly related to desiccation
of water bodies.
(2) Trace-fossil distribution in lacustrine systems essentially
reflects combinations of stress factors, sediment type, and the
position of the water table. Because these generally vary in a
systematic way in the different lake–basin types, an integrated
sedimentologic–ichnologic model within the classification
framework of overfilled, balanced-fill, and underfilled lakes
can be proposed. A combined ichnofacies–ichnofabric ap-
proach allows characterizing vertical changes of archetypal
ichnofacies as a response to secular changes in depositional
conditions (e.g. progressive shallowing), and delineation of
palimpsest surfaces resulting from desiccation along lake
margins (composite ichnofabrics).
(3) Overfilled lake basins contain well-developed softground trace
fossils (Mermia and Skolithos ichnofacies, and softground suite
of the Scoyenia ichnofacies) that are useful to delineate
parasequences and parasequence sets. Fluvial discharge into
overfilled lakes usually generates density currents that oxyge-
nate lake bottoms, allowing the establishment of epifaunal and
infaunal communities. Additionally, these are freshwater lakes
where no stress due to hypersalinity occurs, leading to the
development of a relatively diverse benthos. Firmground suites
are rare because such large lakes usually do not experience
desiccation.
(4) Balanced-fill lakes are extremely complex ichnologically
because they are characterized by fluctuating conditions,
which commonly result in a combination of stress factors
that impact negatively on the benthic fauna. Abundant firm-
ground trace-fossil suites occur in balanced-fill lakes, but
softground assemblages are usually depauperate due to
oxygen-depleted conditions during transgression, and hyper-
salinity during lowstands. Palimpsest surfaces and composite
ichnofabrics are common in lake-margin deposits, commonly
revealing cross-cutting relationships between the softground
and firmground suites of the Scoyenia ichnofacies. Ichnofaunas
from turbidite systems in balanced-fill lakes are less abundant
and diverse than those from overfilled lake turbidites. The
Mermia ichnofacies is absent or depauperate.
(5) Hypersalinity is undoubtedly the most stressful factor in
underfilled lakes and, as a result, species diversity is extremely
low. The Scoyenia ichnofacies is associated with lowstand de-
siccated substrates in underfilled lakes. The density of arthro-
pod and tetrapod trackways may be high, forming tracked
omission surfaces. Some of these omission surfaces may
represent sequence boundaries expressed by co-planar sur-
faces of lowstand and subsequent flooding. Rapid changes in
depositional conditions reflecting desiccation during vertical
aggradation can lead to the formation of composite ichnofab-
rics that reflect successive bioturbation events. Paleosol
(Coprinisphaera ichnofacies) and eolian dune (Octopodichnus–
Entradichnus ichnofacies) ichnofaunas may be present around
the lake.
Acknowledgments
We are grateful to Ricardo Melchor and Alfred Uchman for sharing
their experience on the Ischigualasto–Villa Unión and Ermenek basins,
respectively, Robin Renaut for critically reading the manuscript, and
Palaeo-3 reviewers Murray Gingras and Kevin Bohacs for their
comments. Financial support for this study was provided by the
Natural Sciences and Engineering Research Council (NSERC) Discovery
Grant 311726-05 and 311727-05 awarded to Buatois and Mángano,
respectively.
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