Genise 2 PDF

Geological Society, London, Special Publications
doi: 10.1144/GSL.SP.2004.228.01.16
p355-382.
2004, v.228; Geological Society, London, Special Publications
Jorge F. Genise, E. S. Bellosi and M. G. Gonzalez
ichnofabrics in palaeosols
An approach to the description and interpretation of
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An approach to the description and interpretation of
ichnofabrics in palaeosols
J ORGE F. GENI S E l, E. S. BELLOS I 2 & M. G. GONZALEZ 2
1CONICET, Museo Paleontol6gico Egidio Feruglio, Av. Fontana 140, 9100 (Trelew),
Chubut, Argentina (e-mail: j geni se@mef org.ar)
2 CONICET, Laboratorio de Icnologia, Museo Argentino de Ciencias Naturales,
Av. Angel Gallardo 470, (1405) Buenos Aires, Argentina
Abstract: Studies on ichnofabrics have focused mainly on marine environments. Attempts to
apply the ichnofabric methodology and theoretical framework to continental deposits bearing
palaeosols are few and poorly developed. Methodologies analysed in this contribution include
the applicability of current ichnofabric indexes and diagrams, the assessment of the destruction
of the original bedding by ichnofabrics and by other soil characters separately, and the relation-
ships between different stages of palaeosol and ichnofabric development. Many soil features
may be formed without the intervention of bioturbation, or may be the result of interactions
of physical, chemical and biological processes, in which traces of organisms may have only a
subsidiary role. Ichnofabrics can be well developed in palaeosols devoid of other soil characters
and, conversely, palaeosols showing a well-developed soil structure can bear almost no trace
fossils. This fact adds a third component to classical methods that normally consider only
original bedding and ichnofabrics. Theoretical analysis includes the possibility of recording
composite ichnofabrics in palaeosols, and the value of individual ichnotaxa as possible
indicators of subaerial conditions and environmental changes. The ecological preferences
and requirements of trace-makers provide the key to understanding composite ichnofabrics;
however, only complex traces can be certainly attributed to particular modern taxa. Insect
nests, pupal chambers and earthworm burrows are the most reliable indicators of subaerial
exposure and, in many cases, very particular environmental conditions.
Ichnofabri c analysis has become an increasingly
i mpor t ant t ool in i chnol ogi cal analysis, provid-
ing a met hodol ogy for document i ng and compar-
ing the extent of bi ot ur bat i on and relative
chr onol ogy of i nfaunal tiering (Ausich & Bottjer
1982; Ekdal e & Broml ey 1983, 1991; Broml ey &
Ekdal e 1986; Droser & Bottjer 1986; Bottjer &
Droser 1991, 1994; Tayl or & Gol dr i ng 1993;
Tayl or et al. 2003). Most studies of i chnofabri cs
have been focused on mari ne envi ronment s
(Tayl or & Gol dr i ng 1993; Bottjer & Droser
1994). Few, poor l y developed, at t empt s have
been made to appl y i chnofabri c met hodol ogy
and its concepts to cont i nent al deposits beari ng
palaeosols. The few references involve the utiliza-
t i on of the i chnofabri c index (Droser & Bottjer
1986) as a comparat i ve scale to evaluate the
degree of devel opment of soil structure (Retal-
lack 1990), tiering in palaeosols (Hasiotis et al.
1993; Hasiotis & Dubi el 1994; Gonzal ez et al.
1998), the pr oposal of i nformal names for
i chnofabri cs produced by part i cul ar groups of
ar t hr opods (Hasiotis 1997), and the use of the
t erm terrestrial ichnofabries to describe trace
fossil assemblages in palaeosols (Miller &
Mason 2000).
The aim of this cont r i but i on is to present
examples of pal aeosol i chnofabri cs in order to
analyse the utility of different aspects of ichno-
fabric in cont i nent al deposits bearing palaeosols.
In order to appl y the i chnofabri c met hod to
pal aeosol s the effects of bi ot ur bat i on must first
be distinguished from physical and chemical
pedogenic processes. Ot her aspects required for
compl et e pal aeosol descri pt i on are the interrela-
t i onshi ps between different stages of pal aeosol
and i chnofabri c development. Anot her particu-
lar poi nt to be considered herein in detail is the
value of the i ndi vi dual component s of pal aeosol
i chnofabri cs as possible i ndi cat ors of subaerial
condi t i ons and changes in envi ronment , and the
possibility of recording composi t e i chnofabri cs
(sensu Broml ey & Ekdal e 1986) in palaeosols.
Methodol ogy
Review of ex&ting methodology &
ichnofabric analysis
There are different met hods for measuri ng the
extent of bi ot ur bat i on and describing i chno-
fabrics, which are reviewed elsewhere (Tayl or &
Gol dr i ng 1993; Bottjer & Droser 1994; Miller &
Smail 1997; Net t o 2000). Two basic phi l osophi es
are recognized: those t hat use a semi -quant i t at i ve
From: MClLROY, D. (ed.) 2004. The Application of lehnology to Palaeoenvironmental and Stratigraphic Analysis.
Geological Society, London, Special Publications, 228, 355-382. 0305-8719/04/$15.00 9 The Geological Society
of London.
356 J. F. GENISE ET AL.
approach (e.g. Reineck 1963; Droser & Bottjer
1986; Miller & Smail 1997) and those that are
mostly descriptive (e.g. Ekdale & Bromley
1991; Wetzel & Uchman 1998).
Semi-quantitative methods recognize different
categories of ichnofabric (ichnofabric indices or
bioturbation indices) based on the degree of
disruption of the original bedding by biotur-
bation, which are represented in schematic dia-
grams developed for different sedimentary
settings (Droser & Bottjer 1986; Bottjer &
Droser 1994; Miller & Smail 1997). Most of
them have been developed for studying ichno-
fabrics in vertical sections, whereas that of
Miller & Smail (1997) was elaborated for
bedding planes. In turn, descriptive diagrams
(tiering diagrams, TD) were proposed to illus-
trate relationships of ichnofabrics with palaeoen-
vironmental factors, rate of sedimentation, and
biological activity (e.g. Ekdale & Bromley 1991;
Pollard et al. 1993; Wetzel & Uchman 1998 and
references therein) at particular environments
or localities. A comprehensive approach to com-
bine in a single analysis the evaluation of the
extent of bioturbation (bioturbation index, BI),
as in semi-quantitative methods, with a visual
representation of ichnofabrics (ichnofabric con-
stituent diagram, ICD) was proposed by Taylor
& Goldring (1993). The ICD considers the ichno-
taxa present, ichnodiversity, density and order of
emplacement.
However, no single method has been widely
accepted. Critical analysis of the applicability
of different methods has been made elsewhere
(e.g. Frey & Wheatcroft 1989; Ekdale & Brom-
Icy 1991; Pemberton et al. 1992; Miller & Smail
1997). Some authors stated that ichnofabric
indices may be useful for describing simple
ichnofabrics, whereas descriptive methods are
more suitable for complex ones (e.g. Ekdale
& Bromley 1991). Others concluded that
semi-quantitative methods are more appropri-
ate for large-scale studies, involving hundreds
of box cores or thousands of metres of strati-
graphic section, whereas the descriptive
method has been used with core material or
outcrops of limited size (e.g. Miller & Smail
1997). In addition, Taylor & Goldring (1993)
claimed that descriptive methods provide the
possibility of assessing and comparing a more
complete set of data, including also the
degree of bioturbation contemplated in the
semi-quantitative method. In practice, however,
the methodology is so time consuming that its
use is restricted, in most cases, to describing
key stratigraphic surfaces or summarizing
recurring ichnofabrics within a stratigraphic
unit (e.g. McIlroy 2004b).
Me t h o d o l o g y f o r pal ae os ol i chnof abri c
anal ys i s
The case studies presented herein involve com-
plex ichnofabrics, small-scale, high-resolution
observations and detailed palaeoenvironmental
analysis. The inherent complexity of soil fabrics
is such that conventional methodologies were
deemed inadequate, and some modifications to
previous descriptive methods are introduced for
their description and as an aid to their analysis.
Additionally, the bioturbation indices proposed
by Taylor & Goldring (1993) have been used to
complete the descriptions, as well as a semi-
quantitative approach to quantification of bio-
turbation of each tier used for composite
ichnofabrics (as suggested by Ekdale & Bromley
1991). Seven grades of BI, based on Reineck
(1963), were recognized by Taylor & Goldring
(1993), which were adapted herein for descrip-
tion of palaeosols according to burrow density
and the amount of burrow overlap. These BI
are: 0, no bioturbation; 1, sparse bioturbation,
few discrete traces; 2, low bioturbation, low
trace density; 3, moderate bioturbation, traces
discrete, overlap rare; 4, high bioturbation,
high trace density with overlap common; 5,
intense bioturbation, later burrows discrete;
and 6, complete bioturbation, repeated over-
printing. The previous authors also utilized the
sharpness of the primary sedimentary fabric to
define the grades (Taylor & Goldring 1993).
However, in palaeosols the original bedding
can also be heavily disrupted by other soil
processes that are non-biological, or only par-
tially related to bioturbation: thus this disruption
of the sedimentary fabric is not exclusively
related to bioturbation as in the classical
marine examples.
In both the semi-quantitative and descriptive
methods only two variables are considered: the
original bedding and the ichnofabric. In palaeo-
sols, soil structures other than ichnofabrics
should be also considered as destructive agents
of the original bedding. Different soil features
that can be recognized in palaeosols, such as
horizons, peds, cutans, glaebules, crystals, and
root and ani mal traces, modify the original
sedimentary fabric of the deposits in which soils
develop (Teruggi 1971; Yaal on 1971; Andreis
1981; Bown & Kraus 1987; Retallack 1990;
Nettleton et al. 2000). Root and animal traces
are the direct products of the activity of organ-
isms, and as such they are considered trace
fossils and deserve an ichnotaxonomical treat-
ment, whereas other soil characters are the
result of interactions of purely physical-chemical
processes, or mixed physical-chemical-biological
ICHNOFABRIC ANALYSIS OF PALAEOSOLS 357
processes, in which traces of organisms may only
be involved to different extent (e.g. Brewer 1976;
Fitzpatrick 1984; Retallack 1990). Regarding
Ekdale & Bromley' s (1983) definition of ichno-
fabrics, it would be possible to consider the
whole soil structure as an ichnofabric because it
may be a direct or indirect result of bioturbation.
However, soil features such as peds, cutans,
glaebules and crystals may be formed as well,
without the necessary intervention of biotur-
bation (e.g. Buol e t al . 1990; Retallack 1990).
For instance, Blokhuis e t al . (1990) stated that
Vertisols are typical cases of ped formation by
physical processes of shear failure, along inclined
surfaces, and cracking. Repeated wetting and
drying of a dispersed cracking clay soil results
in the fragments parting into fine wedge-shaped
aggregates, because of the stress-strain regime
of the swelling soils (Blokhuis e t al . 1990).
Even when the traces of organisms would have
been involved in the origin of soil characters (e.g.
granular and crumb peds), they might be
unpreserved or so distorted as to become recog-
nizable, thereby precluding an ichnotaxonomical
approach. Apart from its formal definition, the
common usage of the ichnofabric concept
involves fabrics completely produced by traces
of organisms, more commonly t han fabrics in
which traces are only partially involved. Finally,
the development of an ichnofabric may be inde-
pendent of other soil characters. Ichnofabrics
can be well developed in immature palaeosols,
obliterating the original bedding almost com-
pletely (e.g. Genise & Bown 1994b). Conversely,
palaeosols showing a well-developed ped struc-
ture may preserve almost no trace fossils
(Melchor e t al . 2001). In conclusion, the fabric
directly and completely produced by root and
animal traces is considered herein to be the i c hno-
f a b r i c , whereas the fabric produced by other soil
characters, resulting from the interactions of
physical, chemical and biological processes, is
termed herein the p e d o f a b r i c . It should be noted
that pedofabric is distinct from the terms 'soil
fabric' and 'soil microfabric' (e.g. Brewer 1976;
Fitzpatrick 1984; Bullock e t al . 1985; Retallack
1990; Buol e t al . 1990), which are applied
mostly to the microstructure of the fine-grained
part of the soils. The stages of palaeosol develop-
ment may be used to evaluate the degree of
pedofabric development. There are two scales
for estimating stages of palaeosol development,
which were proposed by Bown & Kraus (1987)
and Retallack (1988) respectively. Both scales
are broadly based on similar criteria. The first
three stages are characterized by (1) no horizon
formation; (2) A horizon and incipient B hori-
zon, and (3) A and B horizons well defined,
whereas the last two stages (4, 5) are defined
mostly by the increasing thickness of A and B
horizons. In terms of BI, a highly developed
palaeosol will comprise volumetrically high
percentages of pedofabric and, accordingly, low
to moderate percentages of primary sedimentary
fabric and ichnofabrics. Hence the single BI
value does not necessarily indicate the percentage
of disruption of the original bedding (by animals
and plants), and conversely the single percentage
of disruption is not a true reflection of the degree
ofbi ot urbat i on. In palaeosols, the proportions of
all three processes must be properly described in
order to analyse meaningfully the fabric of the
deposits in which they occur. A comparison of
five study cases is presented herein utilizing a
pedofabric/ichnofabric ternary diagram (PITD).
The PITD (Fig. l) includes the bioturbation
index (BI), and percentages of original bedding
(OB), ichnofabrics (IF) and pedofabrics (PF).
BI and percentage of IF were measured following
Taylor & Goldring' s (1993) adapted table based
on burrow density and overlap. The percentage
of the original bedding (OB) was measured
according to the preservation of primary sedi-
mentary structures. The pedofabric (PF) was
calculated considering the development of soil
features other than the recognizable trace fossils.
Two types of diagram have been proposed for
representing descriptive methodologies: tiering
diagrams (TD) (e.g. Ekdale & Bromley 1991)
and ichnofabric constituent diagrams (ICD) pro-
posed by Taylor and Goldring (1993). The ICD
represents events consecutively, starting with
the primary fabric and then ichnotaxa in order
of emplacement. This order reflects the infaunal
succession produced by changes in the environ-
mental conditions (e.g. Bromley & Ekdale 1986;
Buatois e t al . 1997). In turn, the original tiering
- the vertical partitioning of the habitat in
response to environmental gradients (e.g.
Ausich & Bottjer 1982; Bromley & Ekdale
1986) - may be not represented i f it is not the
result of the order of emplacement. In marine
environments, where colonization of substrates
may progress from the surface downwards, the
deepest tier is also the last emplaced (Bromley
& Ekdale 1986; Ekdale & Bromley 1991; Brom-
ley 1994). In these cases ICDs may represent tier-
ing and order of emplacement simultaneously.
However, in palaeosols the first colonizers of
the deposit may belong to the deepest tier, as
shown herein. Both original tiering (Hasiotis &
Dubiel 1994; Gonzalez e t al . 1998) and infaunal
succession (Hasiotis e t al . 1993) have been
recorded from palaeosols, and so descriptive
diagrams such as TDs, which can illustrate
both features of ichnofabrics simultaneously,
ORI GI NAL
BEDDI NG
I C H N O F A B R I C 5 ( ) % . w - 9 w -
5 1 P E D O F A B R I C
Fig. 1. Proposed diagram for assessing palaeosol ichnofabric, pedofabric, original bedding and grades of
bioturbation index, BI. l, Ischigualasto Formation, well-developed Vertisols; 2, Ischigualasto Formation,
poorly developed Vertisols; 3, Laguna Palacios Formation, Entisols; 4, Laguna Palacios Formation, Alfisols;
5, Asencio Formation, Ultisols; 6, Jebel Qatrani Formation, Inceptisols; 7, Sarmiento Formation, Andisols;
8, Sarmiento Formation, Alfisols.
are of fundamental importance. In addition, per-
centages of bioturbation, which can sometimes
be time consuming to calculate, are instrumental
for the construction of the ICDs (Taylor &
Goldring 1993). In TDs, the thickness of the
study section, ichnotaxa, tiering and cross-
cutting relationships are readily understandable
(e.g. Bromley & Ekdale 1986).
In conclusion, the study cases presented herein
are represented in TDs in which the BI of
different tiers is indicated, the pedofabric is
depicted along with and independently from the
ichnofabric, and are completed with the com-
positional diagram (Fig. 1) showing total
percentages of bioturbation, pedofabric and
original bedding.
Theoretical background
One of the most important results of the study of
ichnofabrics is the recognition of the tiered pat-
tern of organisms in substrates and subsequent
modifications of it through time in relation to
environmental change (Ausich & Bottjer 1982;
Bromley & Ekdale 1986). The vertical stacking
of ichnofossils, or tiering, is the response of
trace-makers to biological, physical and chemical
gradients within a substrate (Bromley & Ekdale
1986). In the marine realm, the vertical accretion
of the seafloor or major changes in palaeo-
environmental conditions produce either the
upward migration of the communities or a shift
from one to another. This is reflected in the
juxtaposition of tiers producing composite ich-
nofabrics (Bromley & Ekdale 1986; McIlroy
2004a). In modern soils the tiering of organisms
is also a well-known phenomenon (e.g. Hasiotis
& Bown 1992), but scarcely described from
palaeosols. A few examples of infaunal changes
and tiering in palaeosols have been documented
until now, which show the importance of soil
moisture and palaeo-water table as controls on
organism distribution (Hasiotis et al. 1993;
Hasiotis & Dubiel 1994).
There are different sedimentological, palaeo-
pedological, geochemical and ichnological indi-
cators that can contribute to the understanding
of fluctuations of ancient water tables (e.g.
Bown & Kraus 1983; Retallack 1990; Hasiotis
et al. 1993). The development of independent
ichnological criteria, which allow comparisons
with data gathered from other sources, may be
based on two different approaches. One method-
ology, proposed by Bown & Kraus (1983), is
based on the detailed record and comparison of
the occurrence of trace fossils in particular
palaeosol types and horizons. This approach
is particularly important when dealing with
trace fossils whose producers are unknown or
uncertain. Regrettably, papers that analyse trace
fossils and palaeosols with such a degree of detail
are too few to provide a broad reference database
(e.g. Bown & Kraus 1983; Hasiotis e t al . 1993;
Hasiotis & Dubiel 1994; RetaUack 2001b;
Genise e t al . 2002). A second evaluation of
palaeoenvironmental changes involves the
recognition of the producers of ichnotaxa
recorded from palaeosols. If the trace-maker
can be identified, the ecological preferences and
requirements of that taxon can then provide the
key to understanding composite ichnofabrics.
A word of caution on the attribution of trace
fossils to modern groups should be introduced
before an analysis. Trace fossils range from
very simple to very complex morphologies.
Along this spectrum, the more complex the
trace, the more reliable will be its attribution to
a particular modern taxon. The principle of
special quality of behavioural homology
(Wenzel 1992) states that the more complicated
the behaviour, the stronger will be the postulate
of homology. Behavioural homologies are
defined as those behaviours that are similar
because of their origin in a common ancestor,
whereas analogies are those similarities that are
shown by groups that are not phylogenetically
related (Wenzel 1992). To extrapolate the ecolo-
gical preferences and requirements of modern
organisms to the producers of trace fossils, it is
necessary to postulate a close phylogenetic rela-
tionship for both. Only behavioural homologies,
confirmed by macro-and micromorphological
studies and/ or the high complexity of traces,
are useful for postulating this phylogenetic
relationship between a modern and a fossil
trace-maker. In subaerial settings, insect nests
and pupation chambers, and earthworm, milli-
pede and crayfish burrows are complex enough
and/or have been well studied macro- and micro-
morphologically to satisfy this special quality
criterion (e.g. Bown & Kraus 1983; Hasiotis &
Mitchell 1993; Genise e t al . 2000; Retallack
2001b). Conversely, other trace fossils in palaeo-
sols remain to be studied in more detail because
their morphologies are very simple and can be
attributed indistinctly to different groups of
organisms (e.g. Ratcliffe & Fagerstrom 1980).
In modern groups, other criteria that cannot be
used with trace fossils are also employed to
establish behavioural homologies (Wenzel
1992). In turn, a certain degree of congruence
with the body fossil record can be proposed as
an additional ichnological criterion to postulate
homologies. For instance, the proposal that
Triassic bee cells (Hasiotis e t al . 1995) largely
predate the appearance of angiosperms and the
oldest fossil bees, known from the Cretaceous,
has received criticism and general lack of accep-
tance (e.g. Engel 2001; Genise 2004).
The compiled information on recorded trace
fossil assemblages from palaeosols shows that
they are composed of insect nests and pupation
chambers, earthworm, millipede and crayfish
burrows, root traces and other ichnofossils
whose affinities are obscure (Genise e t al . 2000,
table 2; Retallack 2001b). Insect nests as a
whole are the most common trace fossils in
palaeosols, comprising a distinct ethological
category termed calichnia (Genise & Bown
1994a). These authors explained their abundance,
arguing that nests are constructed structures and
not merely excavated ones, and consequently
have a high preservational potential. The nesting
activities involve the provision of nests with
different kinds of organic matter to feed larvae
by adults. Both provisions and larvae are very
sensitive to soil temperature and moisture,
requiring very specific environmental conditions
(e.g. Michener 1979; Grass6 1984; Cane 1991;
Hanski & Cambefort 1991; Genise 1999).
Adults achieve these conditions basically in two
ways: (1) by the morphology and constructional
materials of nests and breeding chambers, and
(2) by selecting specific nesting sites and soil
depths to locate them. Constructed walls and lin-
ings, along with the general morphology of nests,
provide the necessary isolation from soil to main-
tain a suitable temperature and humidity inside
breeding chambers (e.g. Halffter & Edmonds
1982; Grass~ 1984). Accordingly, these morpho-
logical devices increase the complexity of the
whole structure, providing the diagnostic charac-
ters for recognition of behavioural homologies
that permit the attribution to particular insect
taxa (e.g. Genise 1999). In contrast, other struc-
tures excavated by different groups of insects
for shelter, feeding or dwelling are in most
cases more simple, and these analogous struc-
tures cannot be attributed to any particular
invertebrate taxa (e.g. Ratcliffe & Fagerstrom
1980).
Hence, for the purpose of palaeoenvironmen-
tal analysis, trace fossils that can be certainly
attributed to particular invertebrate taxa and
those whose attribution is doubtful should be
analysed separately. Some broad-scale relation-
ships between emplacement of trace fossils and
position of an ancient water table can be easily
addressed knowing the physiological require-
ments of producers. For instance, insects that
are air breathers will inhabit the soil above the
water table, whereas crayfishes and other
crustaceans, which need free water to breathe,
live in burrows under the water table or con-
nected to watercourses (Retallack 1990; Hasiotis
360 J. F. GENISE E T AL.
& Mitchell 1993). However, a more accurate
picture of soil moisture preferences and tiering
can be achieved by analysing particular groups
of trace-makers involved within the context of
the resultant ichnofabric.
Sel ect ed study cases
The few examples presented herein were selected
particularly (1) to test the application of present
methodology, (2) to show the independent devel-
opment of ichnofabrics and pedofabrics, (3) to
demonstrate tiering in ancient soils, and (4) to
illustrate the presence of composite ichnofabrics
in palaeosols. It is not the purpose of this con-
tribution to review the palaeoenvironmental
conditions or any other aspects of the sedimen-
tary sequences included in the study cases. In
the case of the Jebel Qatrani Formation from
Egypt, only one of us (JFG) conducted ichno-
logical research at that locality: hence the
sedimentological and palaeoenvironmental set-
ting is taken exclusively from the literature
(Bown 1982; Bown & Kraus 1988). However,
the importance of this case study for the pre-
viously mentioned purposes of this contribution
necessitates its inclusion herein.
P a l a e o s o l s f r o m t h e I s c h i g u a l a s t o F o r m a t i o n
( A r g e n t i n a )
The Ischigualasto Format i on (lower Late Tri-
assic) is a continental unit of the Ischigualasto-
Villa Uni 6n Basin, which crops out in San Juan
Province, western Argentina (Stipanicic 2001).
It is composed mainly of grey claystones, green-
ish tuffaceous siltstones and bentonites, carbon-
aceous mudstones and few thin coal beds
intercalated with fine- to coarse-grained, cross-
bedded sandstones (Stipanicic & Bonaparte
1979; Bellosi et al. 2001a). At the surveyed
localities in the Ischigualasto Provincial Park,
the fluvial facies are organized in fining-upward
cycles formed by channelized sandstone bodies
with trough cross-bedding and downstream
and lateral accretion surfaces, grading to thick
floodplain deposits (Fig. 2). General alluvial
architecture and the anatomy of sandstone
bodies indicate an avulsion-controlled fluvial
system, having perennial, moderate to high
sinuosity, channels, mixed load and frequent
splays (Bellosi et al. 2001a; Spalletti 2001).
Floodplains were predominantly poorly drained,
favouring the development of palustrine condi-
tions locally (Milana & Alcober 1994). Land-
scape and basin configuration varied slightly
during Ischigualasto times. Trunk and more
steady braided rivers occupied the central area
of the Ischigualasto Park, whereas in northern
and southern areas floodplains were better
developed (Milana et al. 1998).
Previous workers (Milana & Alcober 1994)
recognized the presence of ' r oot traces, bioturba-
tion and burrows' at many horizons. In addition,
Melchor et al. (2001) stated that Ischigualasto
palaeosols exhibit vertic features, tabular root
systems and drab colours. Palaeosols of the
lower section are better drained, and present
calcic and mottled horizons, whereas those of
the upper section are more clayey and rooted
(Martinez et al. 1998). In general, these alluvial
palaeosols are simple and show moderate to
poor maturity, in accordance with rapid sedi-
mentation deduced from the presence of fresh
feldspars (Milana & Alcober 1994). Most
mudrocks exhibit greyish to greenish hues,
whereas some beds are reddish, indicating
better-drained conditions. Fossil amphibians,
primitive dinosaurs and therapsids are abundant,
and occur together with diverse plant remains
belonging to the Di c r oi di um Fl ora (Kokogian
et al. 1999; Zamuner et al. 2001). Vegetation
along the margins of fluvial channels was
dominated by subtropical (seasonal) evergreen
and sclerophyllous forests, with low diversity,
whereas on floodplains a herbaceous-arbustive
communi t y developed (Artabe et al. 2001).
The two studied palaeosols are intercalated
in a small-scale (10 m thick in average) sequence
in the upper section of the formation (Fig 2).
Several of these sequences are intercalated
between mai n channel and typical floodplain
deposits, and are associated with carbonaceous
mudstones deposited in lacustrine/backswamp
environments and fine- to medium-grained
sandstones that accumulated in levees and as
splay lobes, or in subordinated channels. Such
sequences are comparable to avulsion belt
deposits (Kraus & Gwi nn 1997; Farrell 2001),
which generally support high sedimentation
rates (Kraus 1996; Kraus & Aslan 1998).
One example is a 60 cm thick, olive grey (5Y5/
1), smectitic clay-rich palaeosol (Figs 2, 4a) in
which no relict of the original bedding is recog-
nized. Pedofabric is uniform and characterized
by small, angular blocky peds (wedge-shaped)
and closely spaced slickensides. The palaeosol
shows no horizonation, and it is eroded at top
by a medium-grained sandstone with convolute
bedding accumulated in a mi nor avulsion
channel. Micromorphologically it has a well-
defined blocky microstructure ( sensu Bullock
et al. 1985) and abundant connected streaks
of bright clay, which are aligned around ped
I CHNOF ABRI C ANALYSI S OF PALAEOSOLS 361
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E q u i s e t a l e s ~ S k o f i t h o s
" . . , " " , , L a r g e w e d g e - s h a p e d
" " , " " p e d s
s. /
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l a m i n a t i o n l a mi n a t i o n
Fig. 2. (a) Locat i on of stacked vertisols (asterisks) in a coarseni ng-upward avul si on sequence ( CU Seq) of the
upper section of the Ischi gual ast o For mat i on. Arrows show posi t i on of Vertisols wi t h in situ Equisetales stems.
The CU sequence is formed by suspension clay deposits (SC), splay sand lobes (SL) wi t h ripple l ami nat i on and
mi nor sandst one channel s (Ch) wi t h deformed t rough cross-bedding. Tiering di agrams from well-developed
Vertisols (b) and poor l y developed ones (c) in this For mat i on.
surfaces. Truncated stems of equisetales in life-
position filled by green sandstone represent the
only bioturbation (BI1) (Genise et al. 2001;
Fig. 4a). This palaeosol probably underwent
advanced pedogenic homogenization. Thus it is
interpreted as a very well-developed Vertisol, as
the original bedding is totally obliterated by the
pedofabric.
Another example is a less-developed Vertisol,
where the original bedding is clearly recognized
(Figs 2, 4b). The upper 20 cm are composed of
a very coarse siltstone, light olive grey (5Y 6/1),
having relict ripple lamination and less-
developed short slickensides and wedge-shaped
peds. In the lower 20cm wedge-shaped clay
peds and slickensides are also present, but a
fine lamination is still preserved. Bioturbation
is higher than in the case described previously
(BI2). Ichnofabric is composed of equisetales
stems and Skol i t hos linearis in the upper siltstone
(Genise et al. 2001; Fig. 4b). The preservation of
parallel and ripple cross-lamination suggests a
poorly developed Vertisol.
Pa l a e o s o l s f r o m t he L a g u n a Pa l a c i o s
Fo r ma t i o n ( A r g e n t &a )
The Laguna Palacios Format i on is an orange to
reddish brown Upper Cretaceous (Sant oni an-
Maastrichtian) tuffaceous succession, which
constitutes the upper part of the Chubut Group
(Feruglio 1949; Sciutto 1981). It records terres-
trial sedimentation at the northwest periphery
of the San Jorge Basin and north Deseado
Massif, in central Patagonia, Argentina (Bellosi
& Sciutto 2002). Palaeogeographic reconstruc-
tion and thickness variation indicate that it
accumulated on basin margin topographic
highs (Sciutto 1981). The Laguna Palacios For-
mation is divided into lower, middle and upper
members (Sciutto 1981). The only recorded
body fossils from this formation are a few
stems and trunks (Sciutto 1981). Two main
facies type are recognized where the unit reaches
its maxi mum thickness (Bellosi & Sciutto 2002).
Pyroclastic facies predominate, including mas-
sive sheet strata of vitric-crystal tufts, very fine
tufts with accretionary lapilli, and bioturbated
tuffaceous mudstones and claystones (Sciutto
1981; Genise et al. 2002). The remaining facies
are fining-upward cross-bedded pyroclastic sand-
stones and scarce intra-formational conglomer-
ates, which are subordinate, albeit significant,
because they are mostly restricted to incised
channels. Several of these channel bodies, in the
middle and upper members, comprise a thin
basal siliciclastic sandstone with (fluvial) current
structures abruptly covered by a primary white
tuff, suggesting non-permanent streams. Most
of the pyroclastic deposits (60% in thickness)
are pedogenized, commonly occurring as stacked
non-calcic pataeosols (Sciutto 1981; Bellosi et al.
2002a). Detailed sedimentary, ichnologic and
pedogenic features were recently published by
Genise (2000) and Genise et al. (2002). Well-
developed and more frequent palaeosols occur
in the middle and upper members (Sciutto
1981). The Laguna Palacios Format i on is
considered to be a volcaniclastic-influenced,
loess-palaeosol sequence, formed in a temperate
subhumid/-seasonal climate and relatively dry
conditions (Bellosi & Sciutto 2002; Bellosi et al.
2002a; Genise et al. 2002). Fluctuations of the
water table, suggested by scarce hydromorphic
features such as Fe- Mn nodules (Genise et al.
2002), are also consistent with a fossil stump in
growth position of the Middle Member, which
would have required rapid burial in a water-
logged habitat (Driese et al. 1997; Retallack
2001a).
The depositional setting of the Laguna
Palacios Format i on can be compared to the
Pleistocene Pampean loess plains, with few
rivers and ponds, and savannah or prairie soils
with herbaceous or low vegetation (Teruggi
1957; Iriondo 1999; Sayago et al. 2001). Most
sediments were supplied as distal volcanic ash
falls and subordinately by fluvial streams
(Genise et al. 2002). The development of
this loess-palaeosol sequence was regulated by
the balance supply versus pedogenesis, which
ultimately could have been governed by
fluctuations in the climate regime (Bellosi et al.
2002a).
The first case of palaeosol ichnofabrics shown
herein occurs in a vitric Entisol from the Upper
Member (Genise et al. 2002, Fig. 3B, 4B; Figs
3, 4c). The parent material is very fine (clay size
grade) vitric volcanic ash. Glass shards are
entire and fresh or slightly weathered. This iso-
lated bed, 60 cm thick, moderately bioturbated,
and very pale orange (10YR8/2), is a distinct
light-coloured pyroclastic deposit interbedded
in a suite of brownish palaeosols. The original
bedding of this air-fall ash is massive or crudely
stratified, and pedofabric is wholly absent.
Micromorphologically no soil feature is present,
and no crystalline mineral was identified in XRD
analysis. The ichnofabric shows two tiers (Fig.
3). The upper tier is dominated by Taeni di um
barret t i and a few root traces, resulting in a
moderate bioturbation (BI3). The lower tier
includes fossil bee nests, Cellicalichnus chubuten-
sis, and also a few root traces, resulting in low
intensities of bioturbation (BI 2) (Genise et al.
|
I I
; , <; , > :, > >
<>< < <
>< > > >< >
< < <
> > > > > >
:r
r
r r ~ r r r , , i
, x. , ,, . , - ,, . J
> > > > > >
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(fluvial channel deposit)
- ~1 Tuff (volcanic ash fall)
~ Well- developed paleosol
~ Bentonitic mudstone
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BI 2
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Taenidium
Rhizoliths
Cellicalichnus
9
BI 4
r-~-/:.t-.:, i . t i T' l . - 9 - : . , . . : - - r - - i :
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i Rhizoliths I I Skolithos
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Taenidium
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. . . . . . . . b - - " 1 I11 I
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" : - : " " oe'd's" : "~' : peds ---- p , - - -.,.
Fig. 3. (a) Section of the upper member of the Laguna Palacios Formation containing two different palaeosol
types. These facies are included in a 300 m thick pyroclastic loess-palaeosol succession, with intercalated fluvial
channel deposits. (b) Entisol (star) developed in very fine vitric tuff (volcanic dust). (c) Intensely bioturbated
Alfisol (asterisk) developed in tuffaceous sandstones.
Fig. 4. (a) Truncated stem of equisetales in growth position (left of hammer), in a well-developed Vertisol.
Ischigualasto Formation. (b) Abundant Skolithos in a poorly developed Vertisol, which preserves ripple
lamination. Ischigualasto Formation. Scale bar: 1 cm. (c) Eroded surface of a vitric Entisol showing fossil bee
nests (Cellicalichnus chubutensis) tunnels in relief. Laguna Palacios Formation. Scale bar: 10 cm. (d) Detail of
Cellicalichnus in plan view in the same Entisol. Laguna Palacios Formation. Coin 2 cm. (e) Structured and
moderately bioturbated Alfisol; the pedofabric is composed mainly of blocky peds. Laguna Palacios
Formation. (f) Coleopteran pupal chambers (RebufJoichnus casamiquelai) from the palaeosol horizon shown in
(e). Laguna Palacios Formation. Lens cap: 5 cm.
2002). The nests, which originally would have
reached the surface of the soil, could not be
traced upwards into the upper tier, suggesting
that the later activity of the shallow Taenidium
trace-makers and roots of plants destroyed this
part of the nests.
The second case is also included in the Upper
Member, separated by 3 m from the previous
one (Fig. 3). The ichnofabric is present in a well-
developed Alfisol, included in a sequence of
composite tuffaceous palaeosols (Genise et al.
2002, figs 3C, F, 4B). It is the most common
palaeosol type in the Laguna Palacios Forma-
tion. The original bedding is only very scarcely
preserved in the upper horizon. The palaeosol
shows a yellowish to light brown colour (5 YR
6/4), a well-developed, upper, elluvial, horizon,
having platy peds, and a lower illuvial one,
having angular to subangular blocky peds (Fig.
4e). The original deposit is a light-coloured,
horizontal-bedded tuffite (vitric medium tuffac-
eous sandstone), preserved as a C horizon in
some cases. Bioturbation is high in the upper
horizons (BI 4) and low to absent in the C hori-
zon. Recognizable trace fossils are coleopteran
pupation chambers, attributable to Rebuffoich-
nus casamiquelai (Fig. 4f), Taenidium barretti,
Skolithos linearis, Beaconites coronus (sensu
Keighley & Pickerill 1994) and thin root traces.
Rebuffoichnus casamiquelai is not cross-cut by
the other traces, which suggests that it was the
last emplaced trace. Its occurrence is patchy, in
contrast to the other ichnogenera that display a
more homogeneous pattern of lateral distribu-
tion in the deposit.
Palaeosols f r om the Asencio Formation
(Uruguay)
The Asencio Format i on is a late Cretaceous-
Palaeogene red-bed unit cropping out in western
Uruguay (Bossi 1966) and partly in eastern
Argentina (Genise & Zelich 2001). It is thin, 5-
15m in average thickness, and composed of
three to six stacked palaeosols, developed on
sandstones of probable fluvial origin (Genise &
Bown 1996; Figs 5a, 7a). Natural outcrops are
small, but the formation is well exposed in
many stone quarries. No body fossils were
recorded from this formation despite its contain-
ing one of the most diverse, well-preserved and
abundant assemblages of bee and coleopteran
nests in palaeosols (Genise & Bown 1996). Origi-
nal bedding is nearly absent with the exception of
a few channelized bodies at the base of the unit
(Pazos et al. 1998; Goso Aquilar & Gu6r6quiz
2001) and poorly preserved in trough cross-
beds in Dumestre quarry. The outstanding pedo-
genized character of this unit was first recognized
by Ford (1988a, 1988 b), who also proposed that
fossil insect nests were restricted to ' conglomer-
ates' [sic] sourced from coeval Oxisols developed
in humid tropical conditions. Regarding the ich-
nological content and sedimentological charac-
teristics of the Asencio Formation, Genise &
Bown (1996) interpreted the clasts of Ford' s con-
glomerates as palaeosol peds. Finally, Gonzalez
et al. (1998) proposed the existence of two differ-
ent horizons characterized by different types of
ped (prismatic and nodular) and ichnological
content. The abundant ichnological assemblage,
composed of 12 ichnogenera, was described by
Roselli (1938, 1987) and redescribed by several
subsequent authors (Genise & Bown 1996;
Genise & Hazeldine 1998a, 1998b; Genise &
Laza 1998; Genise & Verde 2000).
This kind of red ferruginous palaeosot, as
exemplified by the Asencio Formation, is usually
produced by a complex of iterative destruction-
reconstruction stages related to hydromorpho-
logical processes (Tardy 1992). The original Bt
horizons of mature Ultisols (Gonzfilez 1999) sub-
sequently supported lateritization and indura-
tion (crusts) (Caorsi & Gofii 1958; Ford 1988c),
becoming ferricretes or cuirasses (sensu Nahon
1976; Fig. 7a, b). Lateral changes in the structure
from nodular to vacuolar and massive is normal
in cuirasses because of iron transfer, leaching or
dissolution of kaolinite and quartz grains, forma-
tion of voids, and ferruginization (Nahon 1986).
The beds with nodular structure are interpreted
herein as being produced by dismantling of
such iron duricusts (Tardy 1992; Tardy &
Roquin 1992; Temgoua 2001; Figs 7a, c). Most
of them are residual deposits showing no
evidence of transport.
Intense and widespread bioturbation (e.g.
termite activity) is considered to be responsible
for surficial hematite duricrust dismantling
(Eschenbrenner 1986; Grass~ 1986). However,
no undoubtedly termite-made nest structure was
identified. Irregular geometries and transitional
boundaries between the ferricrete-ferricrete lag
are the response of gradual dismantling processes,
which may occur at the surface, inside, or at the
bottom of ferricretes (Temgoua 2001). Most
ferricretes develop under seasonally tropical
climates: mean annual temperature 30~ and
rainfall 1300-1700mm per year, with several
dry months (Tardy & Roqui n 1992). As they
usually develop along large periods of time (105
to 106 years) it is probable that the thin Asencio
sequence represents several million years.
Changes in pedoenvironment produce small-
scale variations in mineralogy and structure.
For instance, red ferralitic soils (with haematite
and kaolinite-haematite micronodules) are
generally located in well-drained upl and areas
(Schwertmann 1988).
In the selected example, two interfingered
horizons are identified, one nodular and poorly
consolidated (dismantling horizon) and the
other well indurated, showing columnar
|
o ~ ~ ~
Paleosurface
Erosional surface
D
Dismantling
horizon
Ferricrete with
columnar/blocky
structure
9
Sandstone
liens
"k
D
Gypsiferous
sandstone
Paleosol
| @
Q o 9
BI 3 BI 4
R
,, ,-~-~-,-:-.::.. E ~
m 31ii~-" i~i:: il [~4.:,,i: i 7: : ~; _i il"
Palmiraichnus Teisseirei
Uruguay
Coprinisphaera Monesichnus I
- 4"- Rhizoliths 11~ - i II
i
Blocky
structure Columnar C)O
structure Dismantling
horizon
Fleaglellius t Rhizoliths
Trough
cross-bedding
Fig. 5. (a) Asencio Formation. The originally structured Ultisols contain abundant insect nests, supported
laterization, ferricrete development and later dismantling (intraformational conglomerates). (b) Tiering diagram
shows the different fabrics in the dismantling horizon and ferruginized crust. (c) Stratigraphic interval in the
middle section of Jebel Qatrani Formation showing high-sinuosity fluvial facies and the position of the studied
Inceptisol (star) (taken from Bown 1982). (d) The Inceptisol developed in fine-grained sandstones that preserve
the original bedding originated in the upper part of a channel point-bar, with abundant rhizoliths and fossil
termite nests ( Fleaglellius pagodus).
structure (duricrust) in which the original bed-
ding is completely disrupted (Figs 5a, 7a). The
dismantling horizons are massive, having
variable quantities of clay matrix (Ford 1988a).
Their individual thickness is 1-3 m, and their
geometry is lenticular. The base is, with rare
exceptions, clearly non-erosive. Lateral bound-
aries of the columnar horizons are always
transitional and irregular (Gonzfilez 1999). In
some exposures, dismantling horizons surround
remains of duricrusts. Nodules are subrounded
and reddish, and their mean size varies from 3
to 10 cm. The mineral composition and micro-
morphol ogy of the nodules are similar to those
of the duricrust, where hematite-kaolinite
aggregates prevail. The smallest nodules are the
more rounded ones. Clay matrix proport i on
also varies from 10% to 60%. Matrix percentage
and roundness increase as nodule size decreases.
The predominance of Monesichnus, Uruguay and
Copr&isphaera in the nodul ar horizons, either in
their original positions or rotated, was pointed
out by Gonz~.lez et al. (1998; Fig. 7c). The bio-
turbation is moderate (BI 3).
The duricrusts are 0.5-2.0 m thick, 200-300 m
long, undulatory, and wedge shaped (Ford
1988b). Internal structure is defined by coarse
columnar to angular blocky peds (Gonzfilez
et al. 1998), with reddish to orange clay cutans,
pedotubules and glaebules (Ford 1988a). Col-
umns are often inclined at 10-30 ~ . Laterally
and subordinately the structure becomes massive
or vacuolar, with empty channels (3~5 mm wide)
and macrovoids (alveoles). Some horizons in the
lower section of the unit include abundant iron
nodules or brecciated carbonate patches, and
others show mottled and bleached (yellowish)
levels (Ford 1988b). Micromorphologically
these horizons are characterized by a clayey-
haematitic ground mass and thick laminated
ferroargillans coating voids and grains (Gonzfi-
lez 1999). Mineral composition is dominated by
weathered and fractured quartz and hematite
cement. Kaolinite is abundant and decreases
upwards in the profile, whereas montmorillonite
increases upwards (Ford 1988a). These horizons
are dominated by Palmiraichnus and Teisseirei
located in situ and rizoliths up to 30cm long
(Gonzalez et al. 1998; Fig. 7b). Bioturbation is
also moderate (BI 3).
Palaeosols f r om the Jebel Qat rani Formation
( Egypt )
The Eocene-Oligocene Jebel Qatrani Format i on
of northern Egypt is a sequence of fine-grained to
gravelly sandstones, mudstones, scarce carbo-
naceous shales and freshwater limestones, most
of them showing evidence of damp pedogenesis
(Bown 1982; Bown & Kraus 1988). This unit
records the sedimentation of high-sinuosity
rivers and overbank floods on a Palaeogene
lowland coastal plain (Bown & Kraus 1988).
Fluvial sediment accumulation was controlled
by sea-level fluctuations in a stable tectonic
setting. Variegation of the rocks in tabular
bands is the most striking feature of palaeosol
formation.
Root and insect traces, cutans, clay-lined vugs
and red duricrusts are additional evidence of
widespread pedogenesis. The rhizolith assem-
blage contains roots and stems of fossil bushy
and reedy plants and trees. Curiously, vegetal
and animal bioturbation are not present in asso-
ciated argillic horizons (Bown & Kraus 1988).
The recognized alluvial palaeosol types include
wet Entisols, Inceptisols, mature gley Spodosols
and very mature Ultisols, some of them with
fragipan horizons (Bown 1982; Bown & Kraus
1988). Fossil leaves, fruits and woods indicate a
wet tropical to subtropical (probably monsoo-
nal) climate. Mangrove swamps prevailed in
coastline areas, whereas in the forested interior
a large and varied vertebrate fauna proliferated,
especially diverse mammal s including Old
World primates (Bown 1982; Bown et al. 1982).
Apart from body fossils, ichnofossils of mam-
mals (rodents), invertebrates (insects, annelids,
crabs and crayfishes) and plants (rhizoliths) are
also very common in the Jebel Qatrani Forma-
tion. Most traces are exceptionally preserved in
pedogenized fluvial-channel sandstones, because
particular geochemical conditions prevailed
during early diagenesis (Bown 1982). In some
localities these trace fossils formed intricate,
dense masses of chambers and tubes (Bown &
Kraus 1988).
The selected example comes from the middle
part of the Jebel Qatrani Formation, where a
yellowish brown, poorly developed Inceptisol
occurs in a fine sandstone deposit, corresponding
to the upper part of a point bar of a meandering
river (Genise & Bown 1994b; Fig. 5b). Relict
trough cross-bedding is the only remaining pri-
mary physical sedimentary structure. Inceptisols
are the most common palaeosol group in the
Jebel Qatrani Formation, especially in the
coarser channel deposits (Bown & Kraus 1988).
Their poor development reflects iterative deposi-
tional-erosional conditions in the active meander
belt and mi nor pauses in the sedimentation
(Bown & Kraus 1988). No horizons.or soil struc-
ture were recognized. The ichnofabric is uniform
along the palaeosol. Bioturbation intensity is
high (BI4), with frequent trace fossil overlap.
Rhizoliths and termite nests are particularly
abundant. Oblate chambers and narrow galleries
compose the fossil termitaria (Fleaglellius pago-
dus), which occur in the upper part of densely
rooted palaeosols, where it is difficult to distin-
guish between galleries and small rhizoliths
(Genise & Bown 1994b; Fig. 7d). The nest
system, comprising polychambered structures
forming high towers interconnected by galleries,
occupies a large volume of the palaeosol.
Palaeosols f r om the Sarmi ent o Formation
(Argent i na)
The Sarmiento Format i on is an Eocene-lower
Miocene pyroclastic succession known mostly
because of its abundant and varied fossil verte-
brates. The mammal assemblages (marsupials,
xenarthrans, astrapotherians, notoungulates,
primates and rodents) are considered the strati-
graphic standards for the South America land
368 J . F. GENISE ET AL.
mammal ages (Ameghino 1906; Simpson 1940;
Cifelli 1985). It is broadly exposed in central
and north Patagonia (Argentina), covering
more than 200 000 km 2 and showing a relatively
uniform lithology, characterized by chonites
(mud and clay-size tufts), fine tufts, bentonites,
and intraformational conglomerates (Mazzoni
1979, 1985). Likewise, the presence of palaeosols
bearing trace fossils is another significant and
well-known feature (Frenguelli 1938; Andreis
1972; Andreis et al. 1975; Spalleti & Mazzoni
1977, 1979). At the type locality, Gran Barranca,
south of Chubut province, where the unit is
divided in three members, there are numerous
well-exposed palaeosols in the middle and
upper members. The domi nant parent material
for palaeosols is fine volcanic ash composed of
rhyolitic-dacitic glass shards and subordinate
plagioclase (andesine). Opal phytoliths are also
frequent or abundant in some levels, reaching
up to 7% of the sample (Mazzoni 1979). Pyro-
clastic deposits resulted from distal ash falls,
sourced from plinian-type eruptions more than
500km to the NNW (Mazzoni 1985). Most of
the Sarmiento Format i on tuffaceous beds are
interpreted as an aeolian loessite (Spalletti &
Mazzoni 1979).
At Gran Barranca, the middle section (Puesto
Al mendra Member) is composed of primary
vitric tufts, bentonites, cross-bedded volcanic
sandstones, intraformational conglomerates
(generally palaeosol fragments) and a few lenti-
cular basaltic flows (Spalletti & Mazzoni 1979;
Fig. 6). A late Eocene-early Miocene age has
been established for this member (Kay et al.
1999). Pyroclastic deposits were partly recycled
by fluvial and aeolian processes, and supported
pedogenesis. Most of the tuffaceous palaeosols
are bioturbated and possess a significant argillic
horizon, Fe and Mn nodules, a moderate to
well-developed b-fabric and significant argilans
and ferromangans (Bellosi et al. 2001b). Domi-
nant trace fossils are dung-beetle nests (Coprini-
sphaera) (Spalletti & Mazzoni 1979; Laza 1986;
Escribano & Delgado 1996; Genise et al. 2000;
Bellosi et al. 2001b), but there are also frequent
root traces, other coleopteran traces (Teisseirei),
bee cells (Celliforma), Beaconi t es coronus, and
large horizontal burrows (Bellosi et al. 2001b).
Scarce calcareous palaeosols (calcretes) contain
almost exclusively Cel l i f orma in association
with charopid land gastropods (Bellosi et al.
2002c).
Two palaeosols of Puesto Al mendra Member
were selected to show tiering and pedofabric/ich-
nofabric relationships (Figs 6, 7e, f). The first
example is in the middle section of the west end
profile of the Gran Barranca (stake MZ-10
from Kay et al. 1999; fig. 1). The palaeosol
occurs in a fine (coarse silt grain size) massive
tuff, 2.4 m thick, highly porous, and light grey
(N 8). Bed lithology and thickness are laterally
very uniform on a kilometre scale (mantle bed-
ding). The bases of beds are sharp, horizontal
erosional surfaces, overlain by a thin upward-
fining intra-formational breccia. Two horizons
may be recognized in this palaeosol (Figs 6,
7f). The upper one is 0. 40m thick, indurated,
very light grey (5YR7/1), and shows intense bio-
turbation and scattered specimens of Coprini-
sphaera (BI5). The intense bioturbation is
composed of a boxwork of sinuous intercon-
nected burrows, 1-2 mm wide and 20 mm long,
having smooth walls and irregular chambers.
Micromorphologically this horizon is character-
ized by abundant glass shards and a vesicular
structure, with an undifferentiated b-fabric,
abundant voids and channels, some of them
coated by thin argilans, organic nodules and a
few braided strands. Some features of this
boxwork resemble modern termite nests. Copri-
nisphaera balls are densely perforated inside
this boxwork, but in the remaining palaeosol
they are undamaged. The lighter-coloured
lower horizon is 2.0 m thick with a transitional
upper contact, showing coarse columnar struc-
ture. Preservation of the original bedding is
difficult to assess because of the massive charac-
ter (ash fall) of the primary sediment. Bioturba-
tion is sparse (BI 1), and composed of very thin,
long root traces. Owing to a lack of clay (which
precluded the formation of an argillic horizon),
the abundance and good preservation of vitric
shards (volcanic ash) and the thick profile, this
palaeosol is classified as a weak to moderately
developed Andisol. This example is comparable
to the Luquem palaeosol of Retallack et al.
(2000), a type of inmature Andisol, which pre-
serves opportunistic vegetation of low specific
diversity that grew on volcanic ash substrates.
The second case is included in a series of
stacked, reddish-orange indurated palaeosols
(stake MZ-17 in Kay et al. 1999; fig. 1) associated
to the upper erosive unconformity of this
member (Kay et al. 2001; Bellosi et al. 2002b;
Figs 6, 7e). Palaeosols differ in horizon thickness
and boundary, textural features, ped type and
abundance of iron nodules (pedofabric), and
especially in the trace fossil content (ichno-
fabric). In the selected palaeosol the original
deposit preserved in the lower 80 cm, is massive,
greyish orange (10YR7/4), with an intraforma-
tional conglomerate. A rough stratification
describes the poor preservation of the original
bedding. The surface horizon is 30-40 cm thick
and argillic. The pedofabric is composed of
< < ( .c .c r ,c ,c .c ,c ~
:. > > :. :. :, :. :. :, :.
:,:,:+1
mm
' "~"r ~2 ~-?,~' I MZ10
l t,' ,' ,t:I' 4' ,l ,' ,l ,1 *
: (>': 1,': > ( :.(:. :.
,I ~, ,{ . . . . E
Bioturbated horizon
Columnar horizon
Intraformational
conglomerate
Tuff (volcanic ash fall)
Volcanic sandstone
Bentonitic mudstone
Basalt flow
Well-developed
stacked paleosols
@ 9
BI 5, BI 2 - ~ !
~ q
~_ ~:_-,-.,,-'r ~ _ ~ :-~ ._~_-',,
BI 3 ~ - ~':::",l
I" " ~ . ~ _ ~ - ~ ', ~- - ~ I
y3- =-' ---~~~--' ,_-_~ .',-. :,_-_'~'~'?-=='~ E ~
BI 1 ~ ~ . ~ : ~ ; [
r - - - ~- a - ~=~' _- _, ' r - L- ~- "- *.'r-,-~_~-'- **rr-~_~_-~- I '
,Large horizontal ~ Ce/litorma
burrows
k../
O coprinisphaera
( ~ Teisseirei ~ traces'Pan'shaped
I T.hin .... ~-. Chambered ( ~ :z.-.z:
rnl zol Kns boxwork Coprinisphaera _' ]. J- Subangular
' blocky structure . l l . . ~
(~o Granular peds '~' Verv coarse
'~t' coldmnar structure
| t |
Fig. 6. (a) Stratigraphic position of the selected palaeosols from the Puesto Almendra Member of Sarmiento
Formation. Facies association records a discontinuous sedimentation (erosional surfaces and abundant
palaeosols), with distal volcanic ash falls, local basalt flows and fluvial reworking by a low-sinuosity system.
(b) The lower example (asterisk) is an Andisol preserving the C horizon. (c) The upper example (star) is an
Alfisol located near the top of a series of stacked palaeosols related to an unconformity.
subangular blocky peds (subordinate crumb
peds), with ferruginized crusts at the top (Fig.
7e). Ped size varies from coarse blocky (2-5 cm)
to medium crumb (2-5 mm) (Soil Survey Staff
1975), and the shape is slightly equidimensional.
Peds are commonly not interlocked, being
separated by clay materials. The crusts are dark
orange (10 YR 5/6), 2- 3cm thick and folded.
Mi cromorphol ogy is characterized by abundant
fresh glass shards, a well-developed spongy
structure, laminated clay, and Fe-cutans, along
with Fe nodules. Some interconnected pores
generate a blocky microstructure. The contact
between both horizons is undul at ory and rela-
tively sharp. This palaeosol is interpreted as a
moderately developed Alfisol because of the
370 J . F. GENISE ET AL.
Fig. 7. (a) Laterized profile of Ultisols from the Asencio Formation. Massive to coarse columnar duricrust
(arrow) covered by a nodular dismantling horizon. (b) Duricrust showing blocky peds, rhizoliths and
Palmiraichnus castellanosi (arrow). Asencio Formation. Scale in cm. (c) Dismantling horizon showing
nodular structure and the fossil bee nest Uruguay auroranormae (arrow). Scale in cm. (d) Inceptisol of the
Jebel Qatrani Formation showing tiered chambers of the fossil termite nests (Fleaglellius pagodus) and
abundant vertical rhizoliths. Lens cap: 5 cm. (e) Alfisol showing subangular blocky peds and folded
ferruginized crusts showing a dung-beetle brood mass (Coprinisphaera) (arrow). Sarmiento Formation.
(f) Andisol showing an upper horizon with granular peds and intense bioturbation possibly produced by
termites and Coprinisphaera (arrows). The lower horizon show a coarse columnar structure and low
bioturbation. Sarmiento Formation.
coarse well-defined peds, thick argillans and
undifferentiated b-fabric in the ground mass.
The ichnofabric is arranged in two tiers (Fig. 6).
In the upper one, Celliforma is the only discrete
trace, resulting in a low bioturbation (BI2).
The lower tier includes Coprinisphaera, Teisseirei
barattinia, pan-shaped traces, and large hori-
zontal burrows 1.8cm in diameter (Fig. 7e).
Bioturbation is moderate (BI 3), discrete traces
are frequent, and cross-cutting is rare.
Discussion
I c h n o f a b r i c s a n d p e d o f a b r i c s
The different cases described herein show how
pedofabrics and ichnofabrics may independently
exhibit variable degrees of development, how
bioturbation and other soil processes may dis-
rupt the original bedding by themselves or in
combination, and how soil processes (e.g. homo-
genization, lateritization) may also disrupt
ichnofabrics. In palaeosols of the Ischigualasto
Format i on ichnofabric is almost absent - j u s t a
few stems of equisetales and Skol i t hos linearis
in the less-developed palaeosols. Even so, the
degree of ped development in these Vertisols is
such that the original bedding is visible only in
one of the study cases. The absence of a well-
developed ichnofabric in those palaeosols may
be attributed to different causes:
9 the absence of constructed, more preservable,
structures, such as those made by bees, dung-
beetles and termites before the Cretaceous
(Genise & Bown 1994a);
9 the frequent waterlogging and reducing condi-
tions found in these soils (Retallack 1990);
9 ichnofabric destruction by intense pedogenic
homogeneization in Vertisols (Melchor et al.
2001).
The last hypothesis suggests not only that non-
biological soil processes may disrupt the original
bedding by itself, but also that they may destroy
the ichnofabric of the deposits, thus emphasizing
the importance of studying both pedofabric
and ichnofabric in palaeosols. According to
independent sedimentological data, these alluvial
Vertisols are immature, as maturity was defined
as a function of time by Bown & Kraus (1987).
In addition, the scales of palaeosol development
are based mostly on particular types of horizon
(Bown & Kraus 1987; Retallack 1988) that are
not present in palaeovertisols (Nettleton et al.
2000). Consequently, and in the absence of
important bioturbation, the stage of develop-
ment of these palaeosols is evaluated herein in
terms of destruction of the original bedding by
the ped-forming processes.
A still clearer case of ichnofabric destruction
by other soil processes occurs in the Ultisols of
the Asencio Formation. These palaeosols show
well-developed pedofabrics and ichnofabrics,
both responsible for the obliteration of the
original bedding, which is absent in almost all
outcrops. In addition, the lateritization process
dismantled the duricrusts, partially destroying
the original ichnofabric. The only preserved
trace fossils in these dismantling horizons are
insect nests with thick constructed walls. Many
nests are disturbed relative to their original
positions, as evidenced by random orientation
of their nest entrances, as most bees and dung-
beetles are very constant in the orientation of
their cells and brood masses (e.g. Halffter &
Matthews 1966; Stephen et al. 1969). The ferri-
cretization process also modified the original
pedofabric of Ultisols in which insects nested
and bioturbation developed. At the other
extreme, the study case from the Jebel Qatrani
Format i on shows Inceptisols almost deprived
of pedofabric, where intense bioturbation results
in a complex intense ichnofabric of BI 4.
Entisols from the Laguna Palacios Format i on
and palaeosols from the Sarmiento Format i on
show intermediate cases in which pedofabric is
poorly or undeveloped, whereas ichnofabric
shows different degrees of development. Alfisols
of the Laguna Palacios Format i on show well-
developed ichnofabrics and pedofabrics, whose
relative degree of development varies laterally
along the exposed palaeosol on a 100m scale.
Most of the Laguna Palacios and Sarmiento
palaeosols differ from the remaining alluvial
examples in that they developed in pyroclastic
aeolian systems. The contrasting mechanisms of
sediment supply and hiatuses controlled the
soil-forming processes and soil stratigraphy in a
variety of different ways.
The independent evaluation of ichnofabrics
and pedofabrics in these cases favours a more
complete analysis of the evolution of these
deposits. Different characters of the pedofabric
can be used to estimate the relative duration of
formation of soils (Retallack 1994). For instance,
the clayey Bt horizon of the Alfisol from Laguna
Palacios Format i on suggests 104 to 103 years of
formation because of the well-developed clay
skins, ground mass, and homogeneous sepic
microfabric (Retallack 1994). During this
period of subaerial exposure water tables fluctu-
ated seasonally, as revealed by hydromorphic
features and illuviation of clays (Genise et al.
2002). The evidence of subaerial exposure pro-
vided by coleopteran pupation chambers in
these palaeosols falls in a very different timescale.
Insect activity involved in the construction of
Rebu.ffoichnus probably indicates the subaerial
exposure of the deposits during only one or two
seasons. Pedofabrics of the Ischigualasto Forma-
tion appear to have been developed in less time
t han t hat of Laguna Palacios because of the
rapid sedimentation rate in the first formation,
as discussed previously.
The nesting activity of termites t hat produced
the ichnofabric dominated by Fleaglellius pago-
dus in the Jebel Qat rani Format i on suggests
several years of colony growth by apposition of
chambers and simultaneous foundation of new
colonies (Genise & Bown 1994b). This fact,
along with the abundance of rhizoliths, indicates
a longer period of subaerial exposure and lower
water tables t han that suggested by the nesting
of solitary insects in the Laguna Palacios,
Asencio and Sarmiento Formations. Nests of
solitary insects may be constructed, provisioned
and closed in only one day, whereas those of
social insects may be maintained for years, and
their micromorphology can even reveal their
age (e.g. Jonkman 1980). Regardless of this,
high concentrations of nests of solitary insects,
along with well-developed pedofabrics, such as
from the Asencio Formation, may indicate a
very long period of subaerial exposure and low
water tables, interrupted only by heavy seasonal
rains, attested to by the presence of abundant
dismantled horizons. The Egyptian example
demonstrates the independence of ichnofabric
and pedofabric development, as the intense sub-
aerial bioturbation produced by termites and
roots is not accompanied by the development
of other soil characters.
Be e a n d c o l e o p t e r a n t r ac e f o s s i l s
The ichnofabrics of Laguna Palacios, Asencio
and Sarmiento show fossil bee cells and/ or
dung-beetle brood masses as their dominant
components. Remains of nests or cells attribu-
table to bees are included in the ichnofamily
Celliformidae, comprising the ichnogenera
Pal mi rai chnus, Celliforma, Cori mbat i chnus,
Uruguay, Ellipsoideichnus, Rosellichnus and
Cellicalichnus (Genise 2000). In soil bee nests,
excessive moisture causes liquefaction or decay
of provisions by fungal attack, which may also
destroy the larvae (Michener 1979; Rozen in.
litt. in Michener 1979). In addition, in water-
logged soils, larvae are exposed to poor oxygen
diffusion (Visscher et al. 1994). There are a
few exceptional records of bees nesting in
periodically or sporadically submerged sites
(e.g. Michener 1979; Roubik & Michener 1980;
Cane 1991; Visscher et al. 1994). The water-
proofed breeding cells of an aggregation of Epi-
charis zonat a (Anthophorinae) remain beneath
the water table during the wet season (Roubik
& Michener 1980). An aggregation of Calliopsis
pugi oni s (Andreninae) that nested in a site t hat
was flooded after a heavy, unusual winter rain
was recorded by Visscher et al. (1994). Flooding
was not fatal, although bees emerged later in the
season, affecting the pollen collection and repro-
ductive success. Accordingly, most bee nests are
located in well-drained soils (Linsley 1958;
Bat ra 1984), and the cell wall is lined with
water-repellent lipids to maintain the moisture
conditions (Cane 1991). The ichnogenera Copri-
nisphaera, Font anai and Monesi chnus, present in
the Asencio and Sarmiento ichnofabrics, are
considered to be brood-masses of dung-beetles
(Sauer 1955; Roselli 1987; Genise 1993; Genise
& Laza 1998). Poorly drained and occasionally
flooded soils are unfavourable for many digging
species of dung-beetle, whereas the period and
duration of flooding determine the success or
failure of reproduction (Lumaret 1983). One spe-
cies nesting in soils that were flooded during
winter showed a high mortality of eggs and
larvae (Kirk 1983). In addition, Hanski and
Cambefort (1991) stated that waterlogged soils
are generally poor for all dung-beetles owing to
alteration of the droppings t hat these insects
utilize to provision their nests. In conclusion, in
nests of bees and dung-beetles, excessive water
content in the soil produces problems related to
larval mortality, decay of provisions, oxygen dif-
fusion and alteration of droppings. Thus the ich-
not axa mentioned previously may be regarded as
indicators of well drained to sporadically flooded
palaeoenvironments and low water tables at the
time of their emplacement in the soils.
Apar t from calichnia, other recognizable
coleopteran structures, fossil pupation cells are
common in palaeosols (Roselli 1938, 1987; Retal-
lack 1984; Johnston et al. 1996; Genise 1999;
Genise et al. 2002). They are represented in the
present study by the ichnogenera Teisseirei in
the Asencio and Sarmiento ichnofabrics and by
Rebuf f oi chnus in the Laguna Palacios locality.
Such structures are chambers constructed by
larvae of different groups of coleopterans to
contain and protect pupae before emergence as
adult (Retallack 1984; Johnston et al. 1996). It
is difficult to attribute these chambers to par-
ticular coleopteran taxa with known ecological
preferences (Genise et al. 2002), but at least as
the trace-makers are air breathers it is possible
to ascertain that they were constructed under
subaerial conditions above the water table.
An t and t ermi t e t race f ossi l s
The ichnogenus Fleaglellius, which is domi nant
in the ichnofabric of the study case from the
Jebel Qatrani Formation, is considered to be a
termite nest (Genise & Bown 1994b). Anot her
possible termite nest presented herein is that
recorded from the second case study of the
Sarmiento Formation. Fossil termite nests are
one of the most common trace fossils in palaeo-
sols (Bown & Laza 1990; Genise & Bown
1994b; Genise 1997), sometimes comprising
complex termitic ichnofabrics (Genise & Bown
1994b). Termites as a whole prefer high atmos-
pheric and soil moisture (e.g. Collins 1969;
Grass6 1986), and consequently their nests may
be found in periodically waterlogged soils (e.g.
Lee & Wood 1971; Grass~ 1984). However, as
with bee or dung-beetle nests, waterlogging
produces similar problems of gas exchange (e.g.
Schmidt 1960; Roy-Noel 1972; San Jos6 et al.
1989) or fungal or microbial infections (e.g.
Grass6 1984). Thus species that inhabit soils
t hat are seasonally waterlogged show particular
behaviours or nest features, such as those that
retreat into epigeous mounds during the wet
season (Lee & Wood 1971; Matthews 1977).
Other species provide their nests with special
chimneys (Roy-Noel 1972), perforation systems,
and air or sand envelopes (Schmidt 1960) to
maintain the microenvironmental conditions
inside nests, which are very specific in terms of
moisture and concentrations of 02 and CO2
(Grass6 1984). A similar case is that of fossil
ant nests, also common in ichnofabrics of
Tertiary palaeosols in Nort h and South America
(Laza 1982; Bown et al. 1997). In contrast to
termites, which are restricted mostly to the
stable microenvironment of their nests, most
ants construct less elaborate structures but have
the ability to move their eggs and larvae from
place to place in response to environmental
changes (e.g. Wheeler 1910; H611dobler &
Wilson 1990). Colonies move frequently - flood-
ing is one of the factors that can trigger these
movements - although there are records of ants
surviving several hours or even days submerged
by floodwater (H611dobler & Wilson 1990).
There are few records of ants t hat nest in low
lands and also construct mounds to which they
retreat during the wet season to avoid high
water tables (e.g. Bruch 1916; Bonetto et al.
1961), like termites. In summary, ants and -
particularly - termites nest in well-drained to
seasonally flooded soils. In the latter case nests
are provided with particular devices, such as
epigeous mounds, chimneys, or special walls. In
terms of soil moisture, ichnofabrics dominated
by fossil termite nests such as those of the Jebel
Qatrani Format i on would indicate higher moist-
ure conditions and more frequently flooded soils
than ichnofabrics dominated by bee and dung-
beetle traces. Fossil plants and mammal s of the
Jebel Qat rani Format i on also indicate a wet
climate (Bown 1982; Bown et al. 1982; Bown &
Kraus 1988).
In conclusion, ichnofabrics dominated by
calichnia and pupation chambers indicate sub-
aerial conditions. The construction of nests and
other underground activities take place during
periods of subaerial exposure of deposits. It is
i mport ant to note that insect nests can be trans-
ported, and that they have been recorded as
clasts in conglomerates (e.g. Andreis 1981;
Bown & Ratcliffe 1988). Therefore not only the
identification of nests, but also their position in
situ, are needed before the subaerial exposure
of deposits can be determined.
Ear t hwor m t race f ossi l s
Fossil earthworm burrows indicate a moist soil
environment. Edaphichnium lumbricatum Bown
and Kraus 1983 is considered to be an earth-
worm burrow because of its morphology, the
presence of faecal pellets, and the concentration
of calcium carbonate in the burrows and pellets.
One of the most i mport ant requirements of
earthworms is adequate soil moisture, because
respiration depends on the diffusion of gases
through the moistened body wall (Lee 1985).
Consequently, earthworms inhabit soils in
which water is confined to films on the surface
of soil aggregates or is held in pore spaces by
capillary forces, and the relative humidity of
air-filled spaces is 100% or slightly less (Lee
1985). In waterlogged soils or those where free
water is no longer present, earthworms cannot
survive (Lee 1985). In seasonal tropical climates
earthworms construct spherical aestivation
chambers to spend the dry season at deeper
layers in the soil profile (Jim6nez et al. 2000).
This type of trace, of great environmental
value, has recently been found in palaeosols
(Verde et al. 2002). In accordance with the
moist environments preferred by earthworms,
E. lumbricatum was found in gleyed palaeosols
of the Willwood Format i on (Bown & Kraus
1983). It should be noted that a complete ichno-
taxonomical review of pellet-filled burrows from
different environments (e.g. Richter & Richter
1939; Pickerill 1989 and references therein) is
still lacking, and would be critical to correctly
distinguish eart hworm from other similar
burrows.
Tr a c e f o s s i l s o f unc e r t ai n af f i ni t i es
In contrast, another group of palaeosol ichno-
fossils is that composed of those ichnotaxa that
(1) cannot be attributed unequivocally to a parti-
cular group of producers, and (2) are recorded
from different continental and marine deposits.
As Retallack (1990) pointed out, these tubular
trace fossils, which are common in modern
soils, are not diagnostic of them because they
are also found in lacustrine and marine environ-
ments (e.g. Ratcliffe & Fagerstrom 1980). Ichno-
fabrics of Ischigualasto, Laguna Palacios, and
Sarmiento show this type of trace fossil. In
contrast, they are absent from those showing
the most developed subaerial assemblages,
Asencio and Jebel Qatrani. It is possible that,
in the future, micromorphological studies may
aid in the attribution of these trace fossils to
modern taxa, but currently they are of uncertain
affinities. The identification of ichnotaxa belong-
ing to air breathers is critical for assessing the
subaerial exposure of the deposits, particularly
for those showing poorly developed palaeosols.
Palaeosols and/or subaerial trace fossils may
occur in fluvial (e.g. Bown & Kraus 1983),
lacustrine (e.g. Edwards et al. 1998) and marine
(e.g. Curran 1994) deposits during periods of
subaerial exposure. In theory, i f palaeosol devel-
opment is very weak, a trace fossil suite related to
the original subaqueous conditions of these
deposits could be preserved along with trace
fossils produced under subaerial conditions
(e.g. Curran 1994; Buatois et al. 1998; Retallack
2001b). It has been stressed that the presence
of root traces is a post-depositional process,
independent from the origin of deposits, that
indicates subaerial exposure and change of
environment (Bockelie 1994; Curran 1994).
Similarly, a subaqueous assemblage could be
developed i f a subaerially exposed deposit were
later to be submerged (e.g. Driese & Foreman
1991; Curran 1994), and subaerial and sub-
aqueous trace fossils may occur together in
deposits subsequently submerged and exposed
to air (Frey et al. 1984 and references therein).
Recently, Retallack (2001b) analysed the possi-
bility of recording fossil burrows representing
aquatic environments predating soil formation
or resulting from later inundations by lake or
lagoonal waters. Different criteria, such as
density of burrows in relation to other soil
characters, cross-cutting relationships with
carbonate nodules, burrow collapse and fillings
and associated ichnofauna, were used to demon-
strate that Scoyeni a beerboweri was produced
during the period of soil formation (Retallack
1985, 2001b).
Ichnotaxa such as Skol i t hos, Scoyeni a, Taeni-
di um, Beaconites, Macanopsi s, Cyl &dri cum and
other named and unnamed trace fossils have
been cited from palaeosols (e.g. Genise et al.
2000; Hasiotis 2000; Retallack 2001b) as well as
from marine and lacustrine environments (e.g.
Alpert 1974; H~intzschel 1975; Bown & Kraus
1983; Frey et al. 1984; Keighley & Pickerill
1994; Buatois & M~ingano 1995). With the excep-
tion of Scoyeni a beerboweri (Retallack 2001b), it
is impossible to distinguish between specimens in
subaerial environments from those occurring in
subaqueous ones. The attribution of most of
these trace fossils to particular groups of insects,
or even invertebrates, is thus, in such cases, at
best speculative. Furthermore, to extract
palaeoecological and other inferences from
them (e.g. Hasiotis & Dubiel 1994; Hasiotis
2000) without a previous accurate analysis of
their affinities is of dubious utility. The Triassic
meniscate burrows attributed to soil bugs by
Hasiotis & Dubiel (1994) are similar to those
recorded from subaqueous environments
(Keighley & Pickerill 1994). In addition, in the
original contribution by Willis & Roth (1962)
cited as source by Hasiotis & Dubiel (1994),
it is stated that no tunnels or channels are pre-
sent, a fact already mentioned by Ratcliffe and
Fagerstrom (1980). Soil bugs have their mouth-
parts adapted for sucking fluids from roots or
other plant materials (e.g. Carver et al. 1991).
Observations of the feeding habits of these
insects are particularly important in the inter-
pretation of Triassic meniscate burrows. Their
menisci are ungraded, and stained with alter-
nating zones of oxidized and unoxidized iron
compounds (Hasiotis & Dubiel 1994). This
pattern is commonly interpreted as the result of
the original alternation of organic-rich (faecal)
material with organic-poor (sediment) material
(D' Alessandro & Bromley 1987; Hasiotis et al.
1993; Keighley & Pickerill 1994). The presence
of such alternating menisci containing faecal
material in burrows made by insects, which do
not ingest sediment, is unlikely (e.g. Frey et al.
1984), and even more in soil bugs that feed on
root fluids. This type of meniscate burrow in
Willwood palaeosols was attributed to earth-
worms by Bown & Kraus (1983).
The only possibility of gathering some
environmental data from this group of trace fos-
sils of uncertain affinities is when its presence is
analysed in combination with other palaeosol
and sedimentological data and compared with
those from other sequences (Bown & Kraus
1983; Retallack 1985, 2001b). The ' adhesive
meniscate burrows' from palaeosols of the
Willwood Format i on described by Bown &
Kraus (1983) were later interpreted as indicators
of increased soil moisture and fluctuating water
table, owing to their presence in the lower half
of the palaeosol profile (Hasiotis et al. 1993). In
addition, Hasiotis et al. (1993) recorded the pre-
sence of Scaphichnium hamatum, a dung-beetle
nest, in the upper half of the same profile in a
better-drained and drier environment. Similarly,
in the Gleysols from the Triassic Chinle Forma-
tion the adhesive meniscate burrows are abundant
in the deepest tier just above the water table,
characterized as having high, stable moisture con-
tent (Hasiotis & Dubiel 1994). In these two cases
Taenidium is recorded from the deepest tiers of
the vadose zone of gleyed palaeosols. The possible
producers of meniscate burrows were analysed in
detail by Frey et al. (1984), who pointed out the
difficulties in establishing their affinities and
palaeoenvironmental significance. In conclusion,
these authors stated that some meniscate burrows
(e.g. Scoyenia gracilis and Beaconites coronus)
occur preferentially in moist to wet substrates in
shallow aquatic deposits periodically exposed to
air or in subaerial deposits periodically submerged
(Frey et al. 1984). These scarce dat a suggest a
clear distinction between the drier and well-
drained conditions preferred by insect to nest or
pupate and the moister and less-drained condi-
tions selected by the producers of these meniscate
trace fossils. In contrast, Scoyenia beerboweri
was described from well-drained palaeosols in a
tropical, seasonally dry, semi-arid palaeoclimate
(Retallack 2001b).
Apart from vertical tiering, soil organisms
show patchy lateral distributions controlled by
soil texture, soil carbon content, vegetation,
and population dynamics (Ettema & Wardle
2002). Accordingly, insect fossil nests and
pupation chambers commonly show a laterally
heterogeneous distribution in palaeosols, as in
the case of Rebuffoichnus described herein from
one example of the Laguna Palacios Formation.
In contrast, meniscate trace fossils in the same
ichnofabric, such as Taenidium and Beaconites,
have an extended and homogeneous lateral dis-
tribution, suggesting t hat these trace fossils
were not controlled by the lateral heterogeneity
of soils. Horizontal patterns of variability of
ichnofabrics have been less documented than
vertical ones, but more homogeneous lateral
distributions are known from subaqueous trace
fossils, so much t hat concentration of burrows
(e.g. Skolithos, Ophiomorpha, Zoophycos) along
bedding planes has been used as marker beds in
stratigraphy (e.g. Ekdale et al. 1984).
In conclusion, this group of ichnogenera of
simple morphology is by now not clearly
indicative of a particular set of palaeoenviron-
mental conditions. The few possible clues are
that:
9 they are mostly recorded from subaqueous
environments;
9 in some cases their lateral distribution is
homogeneous, suggesting t hat it is not con-
trolled by soil environmental factors; and
9 several studies suggest t hat some of them occur
in moister conditions t han insect nests.
Considering that soils may be sporadically,
seasonally or permanently waterlogged (e.g.
Retallack 1990), a possibility is t hat these ichno-
genera, in contrast to those attributed to
insect nests and earthworms, may be recording
periods of high moisture or even waterlogging
of soils.
Composi t e ichnofabrics in the st udy cases
In the Ischigualasto Format i on ichnofabrics,
Skolithos are found at the top of poorly devel-
oped palaeosols in association with remains of
the original bedding (e.g. ripple cross-lamina-
tion), suggesting t hat these trace fossils were
probabl y produced by subaquatic organisms. In
contrast, in Entisols of the Laguna Palacios
Formation, the upper tier of Taenidium can be
demonstrated to obliterate the entrance of Celli-
caliehnus, suggesting t hat bees opportunistically
colonized the deepest layers of the soil first, and
that Taenidium is a later component of this
ichnofabric. This composite ichnofabric would
thus reflect a shift of palaeoenvironmental
conditions from drier conditions (early deepest-
tier Cellicalichnus ichnocoenosis) to moister
conditions (later shallow-tier Taenidium ichno-
coenosis), probably because of a raised water
table. Curiously, Savrda et al. (2000) found a
similar, but extreme, form of composite ichno-
fabric in the Cretaceous Tuscaloosa Format i on,
having recent Cellicaliehnus-like nests cross-
cutting Taenidium-dominated fabrics.
In Alfisols of the Laguna Palacios Formation,
ichnofabrics are dominated by Taenidium,
Beaconites and Skolithos, which show a homoge-
neous horizontal and vertical distribution in the
palaeosol, whereas, Rebuffoichnus - a clear
indicator of subaerial conditions - is laterally
restricted in the same palaeosol, in which they
cross-cut the previously mentioned group of
traces. This composite ichnofabric thus indicates
changes in the soil conditions from moist or
even waterlogged palaeonvironments (Taenidium,
Beaconites, Skolithos) to subaerial exposure
(Rebuffoichnus). Accordingly, illuviation of clays
and hygromorphic features of this palaeosol
i ndi cat e seasonal i t y and a mobi l e wat er t abl e
(Geni se et al. 2002).
On t he ot her hand, tiers composed of different
subaeri al i chnot axa (1) shar i ng c o mmo n envi ron-
ment al preferences, such as bee cells a nd dung-
beetle br ood masses, a nd (2) showi ng little, at
r a ndom, or no cross-cut t i ng ar e i nt er pr et ed
herei n as t he ori gi nal tiering of soils. I n t he case
st udy of t he Asenci o For ma t i on, i chnof abr i c
reveals a tiered st ruct ure, albeit s omewhat bl ur r ed
by t he l at er i zat i on processes. I n t he di smant l i ng
hor i zon t he upper tier is compos ed most l y of
Uruguay, Monesi chnus a nd Coprinisphaera,
wher eas in t he crust s Teisseirei and Palmiraichnus
prevail. Similarly, in t he Sar mi ent o For ma t i on
Celliforma const i t ut es t he upper tier, wher eas in
t he l ower one Coprinisphaera, Teisseirei and t wo
ot her undet er mi ned t r ace fossils ar e domi nant .
Conclusions
Ma n y soil f eat ur es t ha t di s r upt t he or i gi nal bed-
di ng of t he deposi t s ma y be f or me d wi t hout t he
i nt er vent i on of bi ot ur bat i on, or ma y be t he
result of its i nt er act i ons wi t h physi cal a nd chemi -
cal processes. F o r i chnof abr i c anal ysi s t hese soil
f eat ur es ar e consi der ed t o const i t ut e t he pedof ab-
ric o f t he deposi t , whi ch is di st i ngui shed f r om t he
i chnof abr i c (t he f abr i c di rect l y a nd compl et el y
pr oduced by pl ant a nd ani mal t races). The use-
ful ness of t hi s di st i nct i on is s uppor t e d by t he
anal ysi s of s t udy cases t ha t show t ha t i chno-
f abr i cs can be i nt ense in pal aeosol s devoi d of
ot her soil char act er s and, conver sel y, pal aeosol s
showi ng a wel l -devel oped pedof abr i c can bear
al mos t no t r ace fossils. I n addi t i on, bi ot ur ba t i on
a nd ot her soil processes ma y di sr upt t he or i gi nal
beddi ng by t hemsel ves or in combi nat i on, a nd
soil processes (e.g. homogeni zat i on, l at er i t i za-
t i on) ma y al so di s r upt i chnof abr i cs. The pedo-
f abr i c is an addi t i onal c ompone nt t o cl assi cal
i chnof abr i c anal ysi s, whi ch nor mal l y consi der s
onl y or i gi nal beddi ng a nd i chnof abr i cs. Thi s
compl exi t y of pal aeosol f abr i cs r equi r es some
modi f i cat i ons t o pr evi ous met hodol ogi es t o
bet t er descri be a nd i nt er pr et pal aeosol i chnof ab-
rics. The anal ysi s compr i ses:
9 t i eri ng di agr ams , in whi ch t he bi ot ur ba t i on
i ndex of di fferent tiers is i ndi cat ed;
9 t he pedof abr i c, depi ct ed al ong wi t h a nd
i ndependent l y f r o m t he i chnof abr i c; a nd
9 a pedof abr i c/ i chnof abr i c t e r na r y di a gr a m
showi ng per cent ages of bi ot ur ba t i on, pedo-
f abr i c a nd or i gi nal beddi ng.
The r ecor d of pa l a e oe nvi r onme nt a l changes
i n compos i t e i chnof abr i cs of pal aeosol s ma y be
bas ed on t wo di fferent appr oaches . One
a p p r o a c h compr i ses t he det ai l ed r ecor d a nd
c ompa r i s on o f t he occur r ence of t r ace fossils in
par t i cul ar pal aeosol t ypes a nd hor i zons, whi ch
is par t i cul ar l y i mp o r t a n t when deal i ng wi t h t he
mor e si mpl e t r ace fossils, whose pr oducer s ar e
u n k n o wn or uncer t ai n. A second a ppr oa c h, par -
t i cul ar l y i mp o r t a n t when deal i ng wi t h compl ex
t r ace fossils such as insect nests, is bas ed on t he
i dent i f i cat i on of t r a c e - ma ke r s a nd t he eval uat i on
o f ecol ogi cal pr ef er ences a nd r equi r ement s of
t hei r recent r epr esent at i ves.
Di fferent gr oups of t race fossils in t he st udy
cases show a spect r um f r om t he dri er condi t i ons
pr ef er r ed by bees a nd dung-beet l es t o t he moi st er
ones of ant s, t ermi t es a nd t r ace fossils of uncer t ai n
affinities. The subaer i al envi r onment is cl earl y
i ndi cat ed onl y by t r ace fossils cert ai nl y at t r i but a-
ble t o ai r br eat her s (insect nests a nd pupa t i on
cells, e a r t hwor m a nd millipede bur r ows) . I n con-
t rast , t r ace fossils of uncer t ai n affinities ma y be
subaer i al or s ubaqueous in origin. A possibility
is t hat t hey ma y be r ecor di ng per i ods of hi gh
moi st ur e, wat er l oggi ng of soils, or even a sub-
aqueous envi r onment bef or e or af t er t he soil-
f or mi ng process.
The dur a t i on of subaer i al exposur e will be
di rect l y r el at ed t o i chnof abr i c devel opment by
t he subaer i al sui t e and, as such, it will be a key
f act or , al ong wi t h t he devel opment of t he
pedof abr i c, in unde r s t a ndi ng t he dynami cs of
pos t - depos i t i onal soil processes.
The authors thank D. McIlroy, G. Retallack and J. de
Gibert for the critical review of the manuscript. This
research was partially supported by grants from the
National Scientific Research Council of Argentina
(CONICET-PIP 717/98), the National Agency of
Scientific and Technical Promotion of Argentina
(FONCYT-PICT 6156/99), and the National Science
Foundation of the USA (EAR 00-87636).
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Geological Society, London, Special Publications

Jorge F. Genise, E. S. Bellosi and M. G. Gonzalez

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