Sedimentology (2020) doi: 10.1111/sed.

12736

An integrative physical, mineralogical and ichnological

approach to characterize underfilled lake-basins
ADRIANA C. MANCUSO* , VERONICA KRAPOVICKAS†,
C E C I L I A A . B E N A V E N T E * and C L A U D I A A . M A R S I C A N O †
*IANIGLA, CCT-CONICET, Adri
an Ruiz Leal s/n, Parque Gral. San Mart
ın, CC330, 5500 Mendoza,
Argentina (E-mail: amancu@mendoza-conicet.gov.ar)
†Departamento de Ciencias Geol
ogicas, Facultad de Ciencias Exactas y Naturales, Universidad de
Buenos Aires, IDEAN-CONICET, Ciudad Universitaria Pabell
on II, 1428, Buenos Aires, Argentina

Associate Editor – Gonzalo Veiga

ABSTRACT
Lakes are particularly sensitive to environmental fluctuations, which are
recorded in their facies and stratigraphy. Ephemeral lakes reveal their sensi-
tivity to palaeoenvironmental changes in the overprinting of the sedimentary
features in every single bed. Tetrapod-track taphonomic-modes and ichno-
logical taphonomic-pathways can be used as sensitive indicators of environ-
mental conditions of the track-bearing beds during deposition and
imprinting. The Middle Triassic Cerro de las Cabras succession (Cuyana
Basin, Argentina) provides an excellent opportunity to these environmental
indicators in an underfilled palaeolake. A model of ichnological preservation
for underfilled lake systems is proposed and the role of the ichnology record
in the sequence stratigraphy analysis is evaluated, based on the integration
of tetrapod-track modes, taphonomic-pathways of playa-lake ichnofauna,
mineralogy and physical data. Soft-ground suites include those dominated
by invertebrate grazing traces and arthropod locomotion traces (Suite 1), and
those overprinted by horizontal-vertical dwelling burrows with tetrapod
tracks preserved in taphonomic modes B and C (Suite 2). The firm-ground
suite (Suite 3) comprises tetrapod-tracks with the best preservation styles
(modes A and B) along less abundant invertebrate dwelling and feeding
traces as found in Suite 2. Clay mineralogy (dominated by illite with subor-
dinate smectite) suggests low plasticity of the layers, in agreement with low-
relief deformation structures observed in tetrapod-track taphonomic-modes.
The well-preserved track tetrapod features documented in the Cerro de las
Cabras succession, together with the absence of pedogenic disturbance, tram-
pling obliterating the footprints, and/or evidence of strong disturbance by
wind, desiccation and/or precipitation, supports short periods of exposure of
the imprinted surface particular to this succession. An integrated multiproxy
approach is proposed to evaluate the evolutionary interpretation and identi-
fication of autogenic versus allogenic controls in underfilled lake-basin histo-
ries. The observed aggradational-trend suggests an equilibrium between rates
of accommodation change and sediment supply, and that the basin-centre
did not experience prolonged sediment-starved conditions.
Keywords Cuyana Basin, footprint taphonomy, ichnology, mineralogy, sed-
imentology, sequence stratigraphy.

© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists 1

2 A. C. Mancuso et al.
INTRODUCTION
The sedimentary and fossil records of lake
basins developed within extensional tectonic
contexts have a high preservation potential, with
distinctive features throughout both space and
time indicated in their deposits (Gierlowski-Kor-
desch & Kelts, 1994, 2000). The relation between
accommodation, sediment supply, and water
supply and flow paths determines what kind of
depositional system predominates (Carroll &
Bohacs, 1999; Bohacs et al., 2000, 2003). Three
main lake-basin types result from those rela-
tions: underfilled, balanced-fill and overfilled
(Bohacs et al., 2000). Overfilled lakes are hydro-
logically open, characterized by a progradational
stratal stacking pattern and a high groundwater
table. Balanced-fill lakes are intermittently open
lakes characterized by a mixture of siliciclastic
and biogenic-chemical deposits, dominated by
an alternation of progradational and aggrada-
tional stacking of facies associations related to a
fluctuating lake level and groundwater table.
Underfilled lakes are usually linked to closed
hydrography (Bohacs et al., 2000, and references
therein) although, in most cases, they have open
groundwater systems (Benavente et al., 2017a,b;
Benavente et al., 2019). This lake-basin type is
characterized by aggradational stratal stacking
patterns (with potential contribution of evapor-
itic deposits) and the development of extensive
dry, desiccated mudflats related to a frequently
low groundwater table.
Lakes, defined as permanent or ephemeral
water bodies, are particularly sensitive to envi-
ronmental fluctuations reflected in their extre-
mely variable facies and depositional histories
(Gierlowski-Kordesch & Kelts, 1994, 2000). In
ephemeral lakes, the overprinting of the sedi-
mentary features in every single bed reveals
their sensitivity to palaeoenvironmental change,
with the stacking pattern recording repeated
extreme fluctuations of base-level (Carroll &
Bohacs, 1999).
Lake-basin types, sequence stratigraphy and
continental ichnofacies models have been suc-
cessfully integrated (Buatois & M
angano, 2004,
2009; Scott & Smith, 2015). The distribution of
trace fossils in lacustrine systems essentially
reflects a combination of such factors as sedi-
mentation (for example, energy level and mixing
regime), the position of the water table, hydro-
chemical conditions (for example, salinity, oxy-
genation and pH) and the ecology of trace-
producing organisms. In addition, vertical
© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology
changes in depositional conditions can be deter-
mined through combined ichnofacies–ichnofab-
ric approaches.
In overfilled lakes, the Mermia and Skolithos
ichnofacies are usually well-represented, as well
as the soft-ground suite of the Scoyenia ichnofa-
cies. Fluvial discharge generates density cur-
rents that oxygenate lake bottoms, where
epifaunal and infaunal benthic communities are
established. In balanced-fill lakes, abundant
firm-ground suites of the Scoyenia ichnofacies
occur, and soft-ground suites are depauperate.
In underfilled-lake basins and lowstand system
tracts of balanced-fill lake-basins the Scoyenia
ichnofacies is widespread, but the Mermia ich-
nofacies is usually absent or impoverished (Bua-
tois & M
angano, 2004, 2009).
In particular, tetrapod tracks and trackways
are usually part of the Scoyenia ichnofacies and
are found in all three lake-basin types (Bohacs
et al., 2007; Marsicano et al., 2010; Scott et al.,
2010). Tetrapod tracks provide additional evi-
dence about the sedimentological conditions of
the substrate at the time of their imprinting,
which controlled their final morphology, preser-
vation, and their spatial and stratigraphic distri-
butions (Avanzini et al., 1997; Laporte &
Behrensmeyer, 1980; Hay & Leakey, 1982; Phil-
lips et al., 2007; Marsicano et al., 2010). There-
fore, detailed studies about the different
tetrapod track taphonomic modes integrated
with ichnofacies observed in a succession can
be used as sensitive indicators of the environ-
mental conditions of the track-bearing beds in
lacustrine systems during deposition and
imprinting.
As a case study, the Middle Triassic Cerro de
las Cabras Formation is analyzed. This unit is
represented by extensive outcrops of a thick
playa-lake succession characterized by a rich
ichnological record, mainly tetrapod footprints
and trackways. With such rich datasets, an inte-
gration of the invertebrate and vertebrate ichno-
logical content with their taphonomy in a
detailed sedimentological context was proposed.
This integration is combined with sequence-stra-
tigraphic, sedimentological and ichnological
datasets of an underfilled lake basin to refine its
evolutionary interpretation and discriminate
autogenic versus allogenic controls in this lake
basin history. The processes on this multiproxy
approach have been summarized, and it has a
high potential for application in other under-
filled lake basins that present the preservation of
ichnological datasets.
 Underfilled lake characterized by multi-approach 3

GEOLOGICAL SETTING The continental Triassic strata of the Cuyana

Triassic continental sedimentation in Argentina
occurred in extensional basins associated with
the breakup of Pangea during the latest Per-
mian–earliest Triassic (Uliana & Biddle, 1988).
The Cuyana Basin (Fig. 1) trends NNW–SSE,
and it spans several asymmetrical half-grabens
due to the alternating location of the master
fault between the western and eastern sides of
the rift basin (Legarreta et al., 1992; Kokogian
et al., 1993; Barredo et al., 2011). The master
fault is located to the east in the northern area
of the basin and to the west in the southern area
(Ramos & Kay, 1991; Legarreta et al. 1992; Bar-
redo, 2005). The Las Pe~ nas sub-basin is located
in the northern area of the rift (Fig. 1), where
Triassic strata crop out in the East Precordillera
(Mendoza province) overlying Devonian marine
deposits of the Villavicencio Formation (Koko-
gian et al., 1993).
Basin (Fig. 1) at base contain coarse clastic
facies of the R
ıo Mendoza Formation, which
interfingers with the sandstones, mudstones and
tuffs of the Cerro de las Cabras Formation (Koko-
gian et al., 1993). This succession is associated
with an initial phase of rapid tectonic subsi-
dence (syn-rift phase) in the basin (Kokogian &
Mancilla, 1989; L
opez-Gamund
ı et al., 1994;
Kokogian et al., 2001). The R
ıo Mendoza–Cerro
de las Cabras succession was developed under
markedly seasonal conditions (wet and dry peri-
ods) in agreement with global palaeoclimatic
models for the Triassic (Sellwood & Valdes,
2006; Preto et al., 2010; Holz, 2015). Also, in the
Potrerillos sub-basin (passive margin), topo-
graphic highs to the west are thought to have
generated local rain-shadow effects (Benavente
et al., 2015). The age of the succession is con-
strained to the Anisian (Middle Triassic) by an
absolute date U-Pb (SHRIMP) of 243
4.7 Ma

Fig. 1. Location map of the sedimentary Triassic Cuyana rift basin of the west-central region of Argentina with
the Las Pe~nas and Potrerillos sub-basins. Modified from Stipanicic & Marsicano (2002). Generalized stratigraphic
section with approximate thickness and palaeoenvironment of the Cuyana basin infill. Dates in millions of years
are from the Geologic Time Scale (Ogg, 2012). *243.3
4.7 Ma radioisotopic age of the Cerro de las Cabras Forma-


tion (Avila et al., 2006); *239.7
2.2, 239.2
4.5 and 230.3
2.3 Ma radioisotopic age of the Potrerillos Forma-
tion (Spalletti et al., 2008).

© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology

4 A. C. Mancuso et al.

from Cerro de las Cabras levels of the Potrerillos


sub-basin (Avila et al., 2006).
METHODS
Stratigraphic sections were measured at the Val-
let south, Vallet north and Cabrera quarries, and
La Pe~na creek (Fig. 2). The Rock Color Chart of
the Geological Society of America (GSA, 1948)
was used for rock colour descriptions. Hand
samples were collected for mineralogical analy-
sis. The level and position of the ichnological
record was documented in the detailed strati-
graphic sections. The sedimentological analysis
consisted of the determination of facies defined
by sedimentary structures and textures and
recorded using the acronyms of the lithofacies
code proposed by Miall (1996), and facies asso-
ciations were identified to interpret the environ-
ments and sub-environments.
Mineralogical composition of the samples
was analyzed through X-ray diffraction (XRD)
analysis at the Centro de Investigaciones
Geol
ogicas (La Plata) with a Panalytical X0 PERT
Pro equipment (Malvern Panalytical Limited,
Malvern, UK) using Cu Ka (ka = 1.5403 A)  and
filtered radiation at 40 mA y 40 kV and two
detectors, one between 4° and 37° and the other
between 2° and 27° and counting time of 0.04°/s.
Mineral identification was performed on glyco-
lated, not glycolated and heated (500°C) sam-
ples. For mixed illite/smectite the contribution
of illite was determined by the Reichweite value
(R) calculated following Moore & Reynolds
(1997).

Fig. 2. Geological map of the Las Pe~ nas sub-basin, central-western Argentina, showing the study area and general-
ized stratigraphic section of the R
ıo Mendoza and Cerro de las Cabras Formations showing palaeoenvironments
and the stratigraphic location of study track-bearing sections. 1: Vallet south quarry, 2: Vallet north quarry, 3:
Cabrera quarry, 4: La Pe~na creek.

© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology

 Underfilled lake characterized by multi-approach
Taphonomic tetrapod-track modes are evalu-
ated according to physical and morphological (fi-
delity of the trackmaker’s pedal anatomy) traits
within footprints in order to identify the source of
such variation (see Gatesy & Falkingham, 2017).
Thus, the presence or absence of different mor-
phological and extra-morphological features such
as palm-sole pads, digit impressions, claw marks
and marginal rims, among others, are documented
(Mancuso et al., 2016). These taphonomic modes
include the Preservation Scale recently defined by
Marchetti et al. (2019).
Ichnofacies analyses are based on the archety-
pal ichnofacies models proposed for continental
environments (Seilacher, 1963, 1964; Frey et al.,
1984, Buatois & M
angano, 1995, 2004, 2009,
2011; Genise et al., 2000; Ekdale et al., 2007;
Hunt & Lucas, 2007; Krapovickas et al., 2016).
The concept of Taphonomic Pathways is used
according to Zhang et al. (1998), and the Biotur-
bation Index (BI) follows Taylor & Goldring
(1993) for cross-sections and Miller & Smail
(1997) for bedding planes. The BI is categorized
from 1 to 6 for cross-sections and 1 to 5 for bed-
ding planes. Herein the BI is followed by the
range of strata removed by biological activity
expressed as a percentage for each case study.
PALAEOENVIRONMENTS OF THE R
IO
MENDOZA–CERRO DE LAS CABRAS
SEQUENCE
The R
ıo Mendoza and Cerro de la Cabras formations
have been studied separately; however, they form a
continuous depositional succession (Kokogian &
Mancilla, 1989; L
opez-Gamund
ı et al., 1994; Koko-


gian et al., 2001; Avila et al., 2006, Benavente et al.,
2015). In the Las Pe~nas sub-basin, this depositional
succession begins with approximately 300 m of
alluvial fan conglomerates (Fig. 2). The dark red-
dish-brown, structureless, clast-supported conglom-
erates are dominated by porphyry, quartz,
sandstone and granite clasts up to 30 cm in diame-
ter, along with coarse-grained sandstone matrix
with similar composition. The conglomerates occur
in intervals that alternate with subordinate moder-
ate-red sandstones that become dominant up-sec-
tion (Rolleri & Criado Roqu
e, 1968; Carrara, 1970;
Kokogian & Mancilla, 1989; Kokogian et al., 1993).
The following 450 m are a non-rhythmic alternation
of greyish-red-purple and moderate-red, fine to
coarse-grained tuffaceous sandstones with subordi-
nate mudstones. The sandstone and mudstone beds
show ripple cross-lamination, mud-cracks and rain-
© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology
5
drop impressions. The succession is interpreted as
an ephemeral fluvial system with the development
of sandflat deposits (Fig. 2) (Rolleri & Criado Roqu

e,
1968; Carrara, 1970; Kokogian & Mancilla, 1989;
Kokogian et al., 1993). Overlying a 700 m thick suc-
cession of alternating greyish-orange, fine to med-
ium-grained sandstones, tuffs and dark mudstones
show a fining-upward trend. They are dominated by
symmetrical-ripple and asymmetrical-ripple cross-
lamination, horizontal lamination, mud-cracks,
rain-drop imprints, mudstone drapes, wrinkle struc-
tures and tetrapod footprints. These deposits record
sedimentation in sandflat and mudflat sub-environ-
ments developed in the ephemeral distal fluvial and
playa-lake systems (Fig. 2) (Rolleri & Criado Roqu
e,
1968; Carrara, 1970; Kokogian & Mancilla, 1989;
Kokogian et al., 1993). Up-section, there are 150 m
of carbonaceous dark mudstones, fine-grained sand-
stones and tuffs interpreted as perennial-lake depos-
its (Fig. 2) (Rolleri & Criado Roqu
e, 1968; Carrara,
1970; Kokogian & Mancilla, 1989; Kokogian et al.,
1993). The next 350 m thick succession is domi-
nated by greyish-orange pink fine to medium-
grained sandstones in beds 0.5 to 3.0 m thick
(Fig. 2). The sandstone consists of quartz, feldspar,
rhyolite and porphyry grains with cross-stratifica-
tion in the thicker beds (Rolleri & Criado Roqu
e,
1968; Carrara, 1970; Kokogian & Mancilla, 1989;
Kokogian et al., 1993). Finally, 350 m of greyish-
brown sandstones interbedded with carbonaceous
mudstones were deposited in a fluvio-lacustrine
environment (Fig. 2) (Rolleri & Criado Roqu
e, 1968;
Carrara, 1970; Kokogian & Mancilla, 1989; Kokogian
et al., 1993).
The study was concentrated on the tuffaceous
sandstones and mudstones deposited in the sand-
flat and mudflat sub-environments, located
between 450 to 1400 m of the unit (Fig. 2), which
represent the underfilled lake succession domi-
nated by an aggradational stratal stacking pattern.
These levels contain most of the ichnological
record, such as invertebrate traces and tetrapod
track-bearing beds. The detailed sedimentary
and mineralogical features of the sandflat and
mudflat sub-environments are described in the
following sections and summarized in Table 1
and Figs 3 to 8.
Sandflat facies association
The sandflat facies association comprises four
facies (Table 1; Figs 4 to 6): cross-bedded sand-
stone (Sp/St); structureless sandstone (Sm); lam-
inated sandstone (Sh); and ripple cross-
laminated sandstone (Sr).
 6

Table 1. Facies associations (FA) and their characteristics defined for the Cerro de las Cabras Formation (Anisian) at the Las Pe~
nas Sub-basin, North area
of the Cuyana Basin.

FA and sub- Bed Fossil

environment Facies Sedimentary structures fabric geometry Vertical and lateral relations content Interpretation

Sandflat Cross-bedded Medium-grained sandstone of Tabular to Overlies and underlies the – Tractive flows
sandstone moderate-red colour (5R 4/6) and sheet, 20 to laminated sandstone facies (Sh) and
(Sp/St) very pale orange (10YR 8/2), 100 cm thick ripple cross-laminated sandstone
A. C. Mancuso et al.

moderate to good sorting. Rip-up facies (Sr)

mud-intraclastic lag. Planar and
trough cross-bedding. Sp-St cosets of
20 cm. Greyish red (10R 4/2)
mottling and nodules

Structureless Medium-grained sandstone of Tabular to Overlies and underlies the Footprints, Tractive flows
sandstone moderate-red colour (5R 4/6), sheet, 10 to laminated sandstone facies (Sh) and invertebrate bioturbated
(Sm) greenish-grey (5GY 6/1) and very 40 cm thick, ripple cross-laminated sandstone traces
pale orange (10YR 8/2). Rip-up mud- amalgamate facies (Sr)
intraclasts and pebbles in the base. 120 cm
Structureless. Greyish pink (5R 8/2)
mottling. Illite and mixed illite/
smectite

Laminated Fine to medium-grained sandstone of Tabular, 2 to Overlies and underlies the cross- Footprints, Unconfined
sandstone moderate-red colour (5R 4/6) and 150 cm thick bedded sandstone facies(Sp/St), invertebrate tractive flows in
(Sh) very pale orange (10YR 8/2). structureless sandstone facies (Sm) traces, root a low gradient
Horizontal lamination (0.2 to 2.0 mm and ripple cross-laminated traces, plant slope with
thick), convolute structures, mud- sandstone facies (Sr) remains pedogenesis
chips, mottled fabric, mud-drapes,
mud-cracks. Dominantly illite with
subordinate smectite

Ripple cross- Fine to medium-grained sandstone of Tabular, 2 to Overlies and underlies the cross- Footprints, Unconfined
laminated moderate-red colour (5R 4/6) and 50 cm thick bedded sandstone facies (Sp-St), invertebrate tractive flows in
sandstone very pale orange (10YR 8/2). structureless sandstone facies (Sm) traces a low gradient
(Sr) Asymmetrical ripple cross- and laminated sandstone facies (Sh) slope
lamination sets 2 to 4 cm thick,
convolute structures and mud-drapes
with cracks

© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology

 Table 1. (continued)

FA and sub- Bed Fossil

environment Facies Sedimentary structures fabric geometry Vertical and lateral relations content Interpretation

Mudflat Laminated Mudstone yellowish-grey (5Y 7/2), Tabular, 10 Overlies and underlies the Footprints Settle out
mudstone dusky red (5R 3/4), brownish-grey to 200 cm laminated sandstone facies (Sh), invertebrate deposition
(Fl/Fm) (5YR 4/1). Horizontal lamination (0.1 thick structureless sandstone facies (Sm), traces,
to 1.0 mm thick), asymmetrical ripple cross-laminated sandstone wrinkle
ripple on the top of strata, mud- facies (Sr), ripple cross-laminated structures
cracks, mud-drapes, rain-drops. mudstone facies (Fr) and nodular
Dominated by illite limestone facies (Ln)

Ripple cross- Mudstone yellowish-grey (5Y 7/2),
laminated dusky red (5R 3/4), brownish-grey
mudstone (5YR 4/1). Asymmetrical ripple
(Fr) cross-lamination (0.2 mm thick).
mud-cracks, mud-drapes, rain-drops
Tabular, 1 to Overlies and underlies the – Unconfined
20 cm thick laminated mudstone facies (Fl/Fm), tractive flows in
ripple cross-laminated sandstone a low gradient
(Sr), cross-bedded sandstone facies slope
(St), and nodular limestone facies
(Ln)

Nodular Carbonate mudstone, brownish-grey Tabular, Overlies and underlies the – Marginal
limestone (5YR 4/1) nodule fabric, with 10 cm thick laminated mudstone (Fl/Fm) and vadose
(Ln) nodules of 2 to 4 cm diameter. ripple cross-laminated sandstone
Dominated by illite (Sr)

© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology

Underfilled lake characterized by multi-approach
7
8 A. C. Mancuso et al.

Fig. 3. Stratigraphic logs of study track-bearing sections. Vallet south quarry (1) and Vallet north quarry (2) represent
the sandflat facies association, Cabrera quarry (3) represents the transition between sandflat and mudflat facies associa-
tions, and La Pe~ na creek (4) represents the mudflat facies association. The sections include the facies, invertebrate trace
fossils, the track-bearing levels (arrows) and footprint taphonomic modes. St: trough cross-stratification sandstone, Sp:
planar cross-stratification sandstone, Sh: horizontal lamination sandstone, Sm: structureless sandstone, Sr: ripple
cross-lamination sandstone, Fm: structureless mudstone, Fl: horizontal lamination mudstone, Fr: ripple cross-lamina-
tion mudstone, Ln: nodular limestone. Scale bar = 2 m in Sections 1, 2 and 3. Scale bar = 50 m in Section 4.

© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology

 Underfilled lake characterized by multi-approach 9

Fig. 4. Outcrop photographs of the sandflat facies association from the Cerro de las Cabras Formation in Las
Pe~nas sub-basin. (A) Detail of the sandstone showing St from Vallet south quarry. Total length of ham-
mer = 32.5 cm. (B) Detail of the sandstone showing intraclastic lag from Vallet north quarry. Scale bar = 20 cm.
(C) Detail of the sandstone showing carbonate-rich nodules (arrow) from Vallet north quarry. Scale bar = 3 cm. (D)
Detail of the sandstone showing greyish pink mottling from Vallet south quarry. (E) Detail of the sandstone show-
ing horizontal lamination sandstone from Vallet south quarry. (F) Detail of the sandstone showing convolute
structures from Vallet south quarry.

Cross-bedded sandstone lags observed in some beds. The red colour of

Description. Cross-bedded sandstone facies (Sp/
St) is composed of moderate-red (5R 5/4) and
very pale orange (10YR 8/2), medium-grained
sandstone, moderate to well-sorted in 20 to
100 cm thick tabular/sheet strata. They have
mainly trough cross-stratification (Fig. 4A) in
8 cm thick sets and 20 cm thick co-sets. The
planar cross-stratification is recorded only in a
few beds (see Section 1 in Fig. 2); therefore it is
included in this facies. Rip-up mud-intraclast
lags are observed on some lower contacts
(Fig. 4B). Greyish-red (10R 4/2) mottled and car-
bonate-rich nodules (2 to 4 cm in diameter) are
present (Fig. 4C).
Interpretation. The cross-bedded sandstones
(Sp/St) indicate traction currents implying a
highly competent flow producing migration of
sandy bedforms (Fisher et al., 2007a,b, 2008).
The thickness of co-sets points to a waning-
flow regime where flows become unconfined
(Smoot & Lowenstein, 1991). Erosive-based
sandstone sheets are suggested by intraclast
mottling indicates incipient palaeosols with
good drainage and high hematite content. Car-
bonate nodules were formed in the vadose
zone by precipitation induced by root activity
in the rhizosphere (Ghosh, 1997; Owen et al.,
2008). Development of incipient palaeosols and
pedogenic effects are commonly found in
ephemeral sub-environments in which sedi-
mentation is discontinuous (Smoot & Lowen-
stein, 1991).
Structureless sandstone
Description. The sandstone is medium-grained
and is moderate-red (5R 4/6), greenish-grey (5GY
6/1) and very pale orange (10YR 8/2) with grey-
ish-pink (5R 8/2) mottling (Fig. 4D). Occasion-
ally, rip-up mud-intraclasts and pebbles are
present in the base of some beds. The beds have
tabular to sheet geometry, and form structureless
(Sm) layers 10 to 40 cm thick. In some cases,
these beds are amalgamated and can be up to
120 cm thick. Mineralogy from the clay-fraction
of a mottled bed shows a predominant

© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology

10 A. C. Mancuso et al.

Fig. 5. Invertebrate trace fossils preserved in sandflat facies association. Traces preserved at the Vallet north and
Vallet south quarries. (A) Arenicolites isp., Skolithos isp. and Palaeophycus tubularis. (B) Skolithos isp. in cross-
section view. (C) Arenicolites isp. in cross-section view. (D) Skolithos isp. and Palaeophycus isp. in cross-section
view. (E) Arenicolites isp., Palaeophycus tubularis, Scoyenia gracilis, Helminthoidichnites tenuis and Helminthop-
sis tenuis. (F) Taenidium barrette and Helminthoidichnites tenuis. A: Arenicolites, S: Skolithos, P: Palaeophycus,
Sc: Scoyenia, T: Taenidium, D: Diplopodichnus, Ho: Helminthopsis, Hi: Helminthoidichnites. Scale bars = 3 cm.

© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology

 Underfilled lake characterized by multi-approach 11
Fig. 6. X-ray diffraction data of the sandflat and mud-
flat facies associations from the Cerro de las Cabras
Formation in Las Pe~ nas sub-basin. Structureless sand-
stone facies: Illite (I) (9.94 h; 5 h), mixed illite/smec-
tite (IS) (10,61 h). Laminated sandstone facies: Illite (I)
(9.9 h; 4.97 h), smectite (Sm) (16.88 h). Laminated
mudstone facies: Illite (I) (9.97 h; 4.97 h). Nodular
limestone facies: Illite (I) (9.92 h; 4.98 h).

composition of illite and mixed illite/smectite

with Reichwaite (R) = 0 corresponding to 10%
illite (Fig. 6).

Ichnology. Structureless sandstone records a

poorly diverse assemblage composed mostly of
vertically oriented trace fossils such as Arenicol-
ites isp. and Skolithos isp., and horizontal traces
such as Palaeophycus tubularis (Fig. 5A and B).
Tetrapod tracks are recorded at the top surface
of some beds. Bioturbation Index: in cross-sec-
tion BI 2 (16 to 30%); on bedding planes BI 3
(20 to 30%).

Interpretation. The structureless deposits are

the result of high rates of sedimentation in
response to a rapid decrease in flow velocity
(Fisher et al., 2007a). Such rapid deceleration is
likely to result from a waning flow coming onto
a lower slope. The presence of intraclast lags in
some layers supports the interpretation of this
facies as erosive sandstone sheets. These sedi-
ments are interpreted as deposits from a deceler-
ating flow that debouched into a distal sandflat
sub-environment. The mottled fabric indicates
incipient palaeosols (Retallack, 1991; Kraus &
Hasiotis, 2006). The presence of illite points to a
probably enhanced hydrolysis index, which is
consistent with high erosion rates (Chamley,
1989; Do Campo et al., 2010). Since the illite/
smectite contains only 10% layers of illite, it is
interpreted as almost pure smectite and there-
fore as indicator of seasonality in the system
(Chamley, 1989). The ichnological assemblage in
this facies is considered to represent a low
diversity Scoyenia ichnofacies.

Laminated sandstone
Description. This facies comprises fine-grained
to medium-grained sandstone of moderate-red
(5R 4/6) and very pale orange (10YR 8/2) colour.

© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology

12 A. C. Mancuso et al.
This sandstone forms 2 to 150 cm thick tabular
strata and exhibits horizontal lamination (Sh)
with laminae from 0.2 to 2.0 mm thick (Fig. 4E),
in some cases with convolute structures
(Fig. 4F). Mudstone chips, mottled fabric and
mudstone drapes with mud-cracks in them char-
acterize these layers. This facies preserves
remains of fossils, plants and organic material.
Mineralogy of the clay fraction of a laminated
sandstone bed with mudstone chips is domi-
nantly illite with subordinated smectite (Fig. 6).
Ichnology. The laminated sandstone facies
records a diverse trace-fossil assemblage domi-
nated by vertical and horizontal dwelling bur-
rows (Arenicolites isp., Skolithos isp. and
Palaeophycus tubularis) (Fig. 5C to E), and hori-
zontal to subhorizontal grazing and feeding
traces produced by mobile deposit feeders
(Helminthoidichnites tenuis, Helminthopsis
tenuis, Taenidium barretti and Scoyenia gracilis)
(Fig. 5E and F). There are also preserved loco-
motion traces, such as Cruziana problematica
and Diplopodichnus isp., delicate root traces
and, recurrently, tetrapod footprints in the lami-
nated sandstone. Bioturbation Index: cross-sec-
tion BI 2 to 3 (16 to 35%); bedding plane BI 3
(25 to 33%).
Interpretation. The Sh facies indicates deposi-
tion under upper-flow-regime conditions, near
the boundary between sub-critical and super-
critical flows (Miall, 1996). These conditions
usually occur during flash floods (Fisher et al.,
2007a,b, 2008). In this case, the tabular geome-
try of the strata and their lateral extent of tens
of metres supports unconfined tractive flow.
The associated convolute structures are the
result of rapid deposition of material with dif-
ferent densities (mixed loads) that generate dif-
ferential loading and consequent syn-
sedimentary deformation (van Loon, 2009) or
could be attributed to animals walking on and
disrupting such a substrate. Cracked mudstone
drapes show the ephemeral nature of sedimen-
tation (Smoot & Lowenstein, 1991; Gierlowski-
Kordesch & Rust, 1994) with occasional mud-
stone chips resulting from their erosion (Smoot
& Lowenstein, 1991; Fisher et al., 2007a). The
illite content suggests a high erosion rate; its
origin is considered detrital from fine sedi-
ments present in the source areas (Chamley,
1989; F€ ursich et al., 2005; Do Campo et al.,
2010). The presence of subordinate smectite
supports a seasonal component in the rainfall
© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology
regime and is usually abundant in semiarid
regions with alkaline volcanic rocks in the
catchment (Chamley, 1989; Scott et al., 2007;
Wakelin-King & Webb, 2007; Do Campo et al.,
2010). The ichnological assemblages found in
these layers can be associated with both
Scoyenia and Mermia ichnofacies.
Ripple cross-laminated sandstone
Description. This facies comprises fine to med-
ium-grained sandstone of moderate-red (5R 4/6)
and very pale orange (10YR 8/2) colour. It forms
2 to 50 cm thick tabular strata with asymmetri-
cal ripple cross-lamination (Sr) in sets 2 to 4 cm
in thickness. This facies also contains convolute
structures along with mudstone drapes with
mud-cracks.
Ichnology. The ripple cross-laminated sand-
stone facies, similar to the laminated sandstone
facies, records an assemblage of vertical and
horizontal dwelling burrows (Arenicolites isp.,
Skolithos isp. and Palaeophycus tubularis),
horizontal to subhorizontal grazing and feeding
traces (Helminthoidichnites tenuis, Helminthop-
sis tenuis, Taenidium barrette and Scoyenia
gracilis) and locomotion traces (Cruziana prob-
lematica and Diplopodichnus isp.), and tetrapod
footprints on the sandstone layers. Bioturbation
Index: cross-section BI 2 to 3 (16 to 35%); bed-
ding plane BI 3 (25 to 33%).
Interpretation. The Sr facies indicates that rip-
ples formed under tractive low regimen flow
processes, which is common during sheet floods
(Young et al., 2003; Fisher et al., 2008) and is
characteristic of sandstone sheets (Fisher et al.,
2007a,b). Unconfined tractive flows are sup-
ported by the tabular geometry and lateral extent
(tens of metres) of the strata (Fisher et al.,
2007a,b, 2008). As in the laminated sandstone
facies the presence of convolute structures is the
result of rapid deposition of material with different
densities (van Loon, 2009) or an animal walking
on and disrupting a substrate. Mudstone drapes
were the result of suspension settle-out in a quiet
environment. The ichnological assemblages found
in these layers can be associated with both Scoye-
nia and Mermia ichnofacies.
Sandflat facies association interpretation
The geometry and lateral extension of the facies
support the interpretation of unconfined sheet-
flood deposits with superimposed features that
indicate subaerial exposure. Thus, the general
 Underfilled lake characterized by multi-approach 13

features documented in the facies and their rela-
tion allow interpretation of the facies association
as the deposit of a sandflat sub-environment at
the distal areas of alluvial fans, as suggested by
its relation with other facies associations of the
unit (Smoot & Lowenstein, 1991; Benavente
et al., 2015).
mudstone facies is depleted of trace fossils and
only some beds are bioturbated. The tetrapod
tracks are documented on top of laminated mud-
stone beds. Bioturbation Index: bedding plane
BI 2 (10 to 15%). Bioturbation was not observ-
able in cross-section for measuring BI.

Mudflat facies association

The mudflat facies association comprises three
facies (Table 1; Figs 6 to 8): laminated mudstone
(Fl/Fm), ripple cross-laminated mudstone (Fr)
and nodular limestone (Ln).

Laminated mudstone
Description: This facies forms 10 to 200 cm
thick tabular beds (Fig. 7A) with yellowish-grey
(5Y 7/2), dusky-red (5R 3/4) and brownish-grey
(5YR 4/1) colours. The mudstone beds contain
horizontal lamination (Fl) (Fig. 7C) with lami-
nae from 0.1 to 1.0 mm thick or, in some cases,
they are structureless (Fm), symmetrical ripples
are preserved on top of the beds (Fig. 7B). The
mudstone contains mud-cracks (Fig. 7D and E),
rain-drop imprints and wrinkle structures
(Fig. 7F and G). The mineralogical composition
of the clay fraction is dominated by illite
(Fig. 6).

Ichnology. The laminated mudstone facies con-

tains an assemblage dominated by vertically ori-
ented trace fossils, as Arenicolites isp. and
Diplocraterion parallelum (Fig. 8). Scarce hori-
zontally oriented dwelling traces, such as
Palaeophycus tubularis, are also preserved. It is
worth noting that most of the laminated

Fig. 7. Outcrop photographs of the mudflat facies

association from the Cerro de las Cabras Formation in
Las Pe~nas sub-basin. (A) General view of laminated
mudstone from La Pe~ na creek. (B) Detail of the mud-
stone showing symmetrical ripple on the top surface
of bed from La Pe~na creek. (C) Detail of the mudstone
showing horizontal lamination from Cabrera quarry.
(D) Detail of the mudstone showing multiple mud-
cracks from La Pe~na creek. (E) Detail of the mudstone
showing incipient mud-cracks from Cabrera quarry.
(F) Detail of the mudstone showing multiple direc-
tions ripple on the top surface bed from Cabrera
quarry. (G) Detail of the mudstone showing multiple
directions ripple on the top surface bed from La Pe~na
creek. (H) Detail of the carbonate mudstone showing
nodule fabric from Cabrera quarry. Scale bar = 20 cm.

© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology

14 A. C. Mancuso et al.
Fig. 8. Invertebrate trace fossils preserved in mudflat facies association. (A) Diplocraterion parallelum ‘Di’; (B)
Skolithos isp. ‘S’ and Arenicolites isp ‘A’. Scale marked in centimetres.
Interpretation. The thin lamination indicates
suspension settle-out (Smoot & Lowenstein,
1991; Gierlowski-Kordesch & Rust, 1994), and the
structureless mudstone indicates rapid suspen-
sion settle-out. The Fl/Fm characterize mudflat
sedimentation (Hardie et al., 1978; Gierlowski-
Kordesch & Rust, 1994). Desiccation cracks at the
top of the units, as well as the rain-drop imprints
and tetrapod tracks, confirm that the deposit was
formed under ephemeral flow conditions. The
presence of illite is consistent with high erosion
rates (Chamley, 1989) and an enhanced hydroly-
sis index (Do Campo et al., 2010). The ichnologi-
cal assemblage is interpreted as a low diversity
Scoyenia ichnofacies.
Ripple cross-laminated mudstone
Description. This facies forms 1 to 20 cm thick
tabular beds with yellowish-grey (5Y 7/2), dusky-
red (5R 3/4) and brownish-grey (5YR 4/1) colours.
The mudstone beds contain asymmetrical ripple
cross-lamination (Fr) with laminae up to 0.2 mm
in thickness and, in some cases, symmetrical rip-
ples on the top surface. The mudstone contains
mud-cracks and rain-drop imprints.
Ichnology. The ripple cross-laminated mud-
stone facies contains a similar assemblage as the
laminated mudstone with vertical trace fossils
(Arenicolites isp. and Diplocraterion parallelum)
and scarce horizontal dwelling traces (Palaeo-
phycus tubularis) only in some beds. The tetra-
pod tracks are documented on top of these beds.
Bioturbation Index: bedding plane BI 2 (10 to
© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology
15%); however, it was not observable in cross-
section.
Interpretation. The ripple cross-lamination
points to muddy sheet flood deposits (Maroulis
& Nanson, 1996; Wright & Marriott, 2007). Beds
with cracks at the top of the layers, as well as
rain-drop imprints and tetrapod tracks, confirm
that this facies formed under ephemeral-flow
conditions. The ichnological assemblage is inter-
preted as a low-diversity Scoyenia ichnofacies.
Nodular limestone
Description. This facies is characterized by
brownish-grey (5YR 4/1) carbonate mudstone
that forms 10 cm thick tabular strata with nodu-
lar fabric (Fig. 7H). The nodules are sub-
rounded and 2 to 4 cm in diameter. The miner-
alogical composition of the clay fraction is also
dominated by illite (Fig. 6).
Interpretation. The nodules are interpreted as
the result of preferential precipitation of carbon-
ate in the vadose zone of the mudflat sub-envir-
onment above the water table (see Alonso-Zarza,
2003). The presence of illite suggests high ero-
sion rates (Chamley, 1989) and an enhanced
hydrolysis index that might be consistent with
more humid palaeoenvironmental conditions
(Do Campo et al., 2010).
Mudflat facies association interpretation
This facies association is dominated by mud-
stone deposits with a minor occurrence of
 Underfilled lake characterized by multi-approach
limestone layers. The successive beds of mud-
stone with lamination, structureless and ripple
cross-lamination, with tabular geometry and
laterally very extensive, support the mudflat
interpretation for these deposits. Moreover, the
very abundant evidence of subaerial exposure
reinforces the interpretation (Smoot & Lowen-
stein, 1991; Benavente et al., 2015).
Vertical and lateral distribution of facies
associations
The sandflat and mudflat facies associations
are mainly distributed between the 450 to
1400 m interval of the R
ıo Mendoza-Cerro de
las Cabras sequence (Fig. 2), where an aggrada-
tional stratal stacking pattern is recognized
(Rolleri & Criado Roqu
e, 1968; Carrara, 1970;
Kokogian & Mancilla, 1989; Kokogian et al.,
1993). The four sections described in detail in
this contribution [Vallet south (1), Vallet north
(2) and Cabrera (3) quarries, and La Pe~ na creek
(4)] represent part of that succession. The
short sections documented in some of the
quarries and the disconnection between them
enable only partial recognition of trends. Their
location and reference to the general section
(R
ıo Mendoza-Cerro de las Cabras sequence),
however, enable characterization of a general
trend for the succession. The Vallet south and
Vallet north quarries document the sandflat
facies association at different stratigraphic
intervals [(1) around the 500 m in the general
section, and (2) around the 1000 m in the gen-
eral section, see Fig. 2]. The stacking pattern
observed in the Vallet south quarry displays a
potential progradational trend (see Fig. 3), and
the section is located in a marginal position of
the depositional system. The Vallet north
quarry section shows an aggradational stratal
stacking pattern with an intermediate marginal
position in the depositional system (see Fig. 3).
The Cabrera quarry section displays an interval
of the transition between sandflat and mudflat
facies associations located in an intermediate
position of the depositional system (see Fig. 3).
Despite the limited vertical extent of the sub-
section, a general aggradational trend most
likely characterizes the stacking pattern.
Finally, the La Pe~ na creek section, located in a
central position in the depositional system, is
the thickest and documents dominance of
mudflat facies association with a clear aggrada-
tional stratal stacking pattern (see Fig. 3).
© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology
15
TETRAPOD FOOTPRINT TAPHONOMY
Three taphonomic modes of track preservation
are defined at the Cerro de las Cabras succes-
sion. They are based on their fidelity to the
trackmaker’s pedal anatomy, and the presence
or absence of different morphological and extra-
morphological features such as palm-pad, digit
impressions, claw marks and marginal rims,
among others (see Table 2, Fig. 9). The tetrapod
footprints are represented by isolated prints, iso-
lated manus-pes sets, and trackways (a series of
manus-pes sets).
Mode A
Mode A represents the highest fidelity of the track-
maker’s pedal anatomy (Preservational Scale 3,
Marchetti et al., 2019). This mode is typified by
detailed impression of the digits and palms
(Fig. 9). The palm/sole impressions are separated
from those made by digits, which are generally
associated with claw drag marks (Fig. 9); the marks
are usually relatively long, sharply incised grooves
with well-defined borders. The deepness of the
tracks varies between 0.5 cm and 3.0 cm. The foot-
prints included in this category are preserved on
the top of mudstones and fine to medium-grained
sandstones, with horizontal lamination, ripple
cross-lamination, and structureless beds from
sandflat and mudflat facies associations (Fig. 3).
The studied material is in situ at the La Pe~
na creek,
Vallet south, Vallet north and Cabrera quarries
(Fig. 3), and includes tracks and trackways with
good morphological quality, attributed to Chiro-
therium higuerense, Rigalites sp., Dicynodontipus
and cf. Ameghinichnus (Marsicano et al., 2004,
Krapovickas et al., 2015). Where the imprinted
surface also presents ripples and desiccation
cracks, generally, the tracks are printed over the
ripples, whereas the desiccation cracks are on
top of the footprints. When associated with inver-
tebrate traces, footprints most frequently cross-cut
all of the traces and are preserved on top of them.
Interpretation
The good anatomical preservation of this
mode, including both digit and palm impres-
sions, suggests at least a certain plasticity of
the sediment at the time of imprinting.
Although the size of the tracks is related to
the bodyweight of the producer, footprint
depth is related to both the bodyweight of
the animal and the resistance of the
16 A. C. Mancuso et al.
Table 2. Morphological and extra-morphological features of tetrapod footprints from the Cerro de las Cabras For-
mation (Anisian) at the Las Pe~
nas Sub-basin, north area of the Cuyana Basin that characterize the taphonomic
modes.
Mode A
Palm outline Transversely elongated
oval or slightly irregular
Digit impressions Yes
Presence of rim No
Texture Fine to medium-grained
sandstone
Surface Firm
Water content Low
substrate to deformation (substrate consis-
tency) (Laporte & Behrensmeyer, 1980; Marsi-
cano et al., 2010). The footprints here
included in Mode A show a relatively consis-
tent depth, around 2 cm, independent of their
size and thus the bodyweight of the track-
maker. Mode A taphonomy suggests track for-
mation in relatively firm surfaces (sandy and
muddy deposits in the sandflat and mudflat
see Figs 3, 10 and 11), based on the well-
defined and shallow-depth footprint impres-
sions, with the absence of deformational
structures and cross-cutting invertebrate traces
of soft substrate suites.
Mode B
Mode B represents low fidelity to the track-
maker’s pedal anatomy (Preservational Scale 1,
Marchetti et al., 2019.). Tracks are poorly
defined and preserved as oval to somewhat
irregular outline impressions with a relatively
deep shaft (from 0.5 to 6.0 cm) (Fig. 9). Mode B
differs from the proposed preservational scale of
Marchettli et al. (2019) in the deepness of the
impressions included in this taphonomic mode.
There is no clear evidence of digits or consistent
palm/sole dimensions, and they lack deforma-
tional structures. The track-bearing levels con-
sist of horizontally laminated, ripple cross-
laminated and structureless mudstones, and fine
to medium-grained sandstones of sandflat and
mudflat facies associations. The footprints
included in this taphonomic mode are preserved
© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology
Mode B Mode C
Oval or irregular Oval or irregular
impressions impressions
No No
No Yes
Fine to medium-grained Mudstone, fine-grained
sandstone sandstone
Moderate plasticity Higher plasticity
Moderate High
in situ at the Vallet south and Vallet north quar-
ries, and the La Pe~ na creek (Fig. 3). The track
impressions included in this mode show vari-
able sizes according to the producers. In some
track surfaces, desiccation cracks cover the
exposed bedding plane, including the prints,
and in cases where the surface includes ripples
and invertebrate traces, the prints disrupt them.
This cross-cutting relation among the tracks and
the sedimentological features characterize them
as true tracks.
Interpretation
Footprints appear as featureless and relatively
deep impressions suggesting at least moderate
plasticity of the sediment on a soft surface at the
time of imprinting. As mentioned before, the
size and depth of the tracks are related to the
size and weight of the track-maker, as well as
substrate consistency (Laporte & Behrensmeyer,
1980; Marsicano et al., 2010). The footprints
included in Mode B were imprinted in sandy or
muddy deposits corresponding to sandflat and
mudflat sub-environments. They are approxi-
mately 3.5 cm deeper than footprints of similar
size included in Mode A. All of this supports
the interpretation herein of a soft substrate with
moderate plasticity – greater than in Mode A.
Mode C
Mode C represents very low fidelity to the track-
maker’s pedal anatomy (Preservational Scale 0,
Marchetti et al., 2019). The tracks have oval or
 Underfilled lake characterized by multi-approach 17

Fig. 9. Field photographs of the surfaces with footprints recognized as taphonomic Mode A, Mode B and Mode C.
Scale bars = 3 cm and 20 cm.
© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology
18 A. C. Mancuso et al.
irregular outlines surrounded by deep and dis-
tinctly symmetrical marginal rims (Fig. 9). The
footprints lack individual thick impressions. The
rims are gently convex, producing smooth
rounded inner track-walls. The depth of the prints
is around 4 cm, and the thickness of the marginal
rims is variable, between 1 cm and 6 cm, related
to track size. The footprints included in this mode
occur on top of horizontally laminated, ripple
cross-laminated, and structureless mudstone and
fine-grained sandstone beds of sandflat and mud-
flat facies associations. Most of the material is still
in situ at the La Pe~na creek, and Vallet south and
Cabrera quarries (Fig. 3). On the track surface, as
also occurs in the other modes described previ-
ously, the tracks are imprinted over the ripples,
and the exposed surface is disrupted on the top by
desiccation cracks. The footprints mostly lack
physical connection with invertebrate traces but,
when they do, they are cross-cut by grazing traces
of soft substrate suites.
Interpretation
The preservation of the footprints as single
depressions without evidence of detailed anatom-
ical features and associated with marginal rims,
strongly suggests high plasticity conditions of the
sediment during imprinting. Such rims are caused
by the compression of the relatively plastic and
soft sediment by the foot during the step cycle
(Allen, 1997; Brown, 1999; Manning, 2004; Milan
et al., 2004, Mil
an & Bromley, 2007). Another line
of evidence is the cross-cutting relationship of the
footprints and invertebrate traces. Grazing traces
are preserved within the footprints, suggesting
that the substrate was wet at or right after the time
of implantation. As mentioned before, the foot-
print depth and the size of marginal rims can be
related to both the bodyweight of the animal and
substrate consistency (Laporte & Behrensmeyer,
1980; Marsicano et al., 2010). The symmetrical
displacement was probably produced by a nor-
mally directed force to the substrate caused by a
regular walk and a similar moisture content in the
sediment areally. Thus, the footprints included in
Mode C were imprinted in sandy or muddy, soft
substrates deposited in sandflat and mudflats set-
tings with high plasticity.
TAPHONOMIC PATHWAYS OF PLAYA-
LAKE ICHNOFAUNA
A series of taphonomic pathways is identified in
the Cerro de las Cabras palaeolake strata based
© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology
on ichnological content, abundance, cross-cut-
ting relation, and association with sedimentary
structures and facies. The succession of a series
of ichnological suites characterizes each tapho-
nomic pathway, and they reflect a continuum of
ichnological processes. The recognized suites
are: (1) the moderate-diversity suite dominated
by grazing traces such as Helmintoidichnites
and Helmintopsis and arthropod locomotion
traces as Diplopodichnus and Cruziana; (2) the
low-diversity suite dominated by vertical bur-
rows such as Arenicolites, Skolithos and equilib-
rium traces as Diplocraterion; and (3) the low-
diversity suite dominated by meniscate and sim-
ple dwelling traces (Paleophycus, Taenidium
and Scoyenia) (Figs 10 and 11). A general suc-
cessional pattern is evident in almost all out-
crops varying on all sub-environments (sandflat
or mudflat) depending on the grain size and
mineralogy, water table and degree of substrate
humidity, hydrodynamic energy, and time. In
shallow and relatively stable standing waters,
small notostracan crustaceans, insect larvae and
nematodes inhabited the soft substrate at the
bottom of low energy playa lakes producing del-
icate horizontal grazing and locomotion traces
(Diplopodichnus, Helminthoidichnites and
Helmintopsis) (Suite 1, Figs 10 and 11). These
traces were cross-cut and followed by vertical
burrows of dipterous larvae and/or oligochaetes,
emplaced still in soft substrates of shallow and
moderate energy episodic floods (Suite 2,
Figs 10 and 11). With the withdrawal of stand-
ing water and fluctuation of the water content of
the substrate, it was colonized by semiaquatic
insects that produced meniscate (Taenidium
and Scoyenia) and dwelling structures (Palaeo-
phycus) included in Suite 3 (Figs 10 and 11).
The highest trace-fossil diversity of the Cerro
de La Cabras palaeolake is preserved in sand-
flat–facies-association beds at the Vallet south,
Vallet north and Cabrera quarries. They record a
complete pathway that includes the delicate
horizontal grazing and locomotion traces (Suite
1) first produced in fine-grained soft substrates
of low energy shallow lake bottoms with stable
water levels. The laminated mudstone most
likely records fall-out sedimentation in the playa
lake. Tetrapod footprints cross-cut by grazing
invertebrate traces are scarce, but when they
occur, they show irregular outlines and marginal
rims as in taphonomic Mode C. Eventually,
ephemeral floods entered the playa lake, and
Suite 2 was formed. Where these substrates lost
water content slowly, Suite 3 developed.
 Underfilled lake characterized by multi-approach 19

Fig. 10. Sandflat taphonomic pathways. T: Time, E: Hydrodynamic energy, S: Substrate humidity, A: Arenicolites,
S: Skolithos, P: Palaeophycus, T: Taenidium, D: Diplopodichnus, Ho: Helminthopsis, Hi: Helminthoidichnites, G:
mud-cracks, MA: Footprint-taphonomic Mode A, MB: Footprint-taphonomic Mode B.

Locally, traces were produced on firm substrates
recording the delicate scratch marks of Scoye-
nia. Tetrapod footprints with low fidelity to the
trackmaker’s pedal anatomy (Mode B) were fre-
quently preserved in close association with
Suites 2 and 3. Finally, when the substrate was
completely exposed, footprints with high fidelity
to the trackmaker’s pedal anatomy (Mode A)
were imprinted. When dry, its surface was cov-
ered by desiccation cracks on top of mudstone
drapes. With enough time, the firm-grounds
were vegetated, recorded by the presence of
scarce root traces, remains of fossils plants and
organic material. This situation is mainly
recorded at Vallet north quarry, suggesting a
more marginal position within the system
(Fig. 10, T3).
Sandflats also record two other taphonomic
pathways indicating shorter exposure (Fig. 10,
T1 and T2). They are recorded in close strati-
graphic connection with the previously men-
tioned sandflat taphonomic pathways; most
likely implying an areal variation of time of
exposure of the substrate at playa-lake margins.
It comprises delicate traces of Suite 1 estab-
lished on laminated mudstones accumulated
due to fall-out sedimentation in standing waters
that were quickly exposed (Fig. 10, T1). The
suite is usually cross-cut by tetrapod footprints
of taphonomic Mode A suggesting that they
were produced on firm substrate quickly
exposed, and not allowing the generation of
soft-ground traces (Suites 2 and 3). The other
pathway records the playa-lake margin

© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology

20 A. C. Mancuso et al.

Fig. 11. Mudflat and sandflat–mudflat transition Taphonomic pathways. T: Time, E: Hydrodinamic energy, S:
Substrate humidity, A: Arenicolites, S: Skolithos, Di: Diplocraterion, P: Palaeophycus, T: Taenidium, D:
Diplopodichnus, Ho: Helminthopsis, Hi: Helminthoidichnites, G: mud-cracks, MA: Footprint-taphonomic Mode A,
MC: Footprint-taphonomic Mode C.

conditions dominated by high-energy ephemeral
floods. Suite 1 has a low diversity, Suite 2 is
the most abundant and Suite 3 is frequently
absent. Tetrapod footprints of taphonomic
Mode B are produced together with vertical
traces of Suite 2. These suites usually, but not
always, lack desiccation cracks and root traces
(Fig. 10, T2).
The transition between sandflat and mudflat is
best documented at the Cabrera quarry, where
the taphonomic pathway most commonly
recorded involves colonization by delicate hori-
zontal grazing and locomotion traces (Suite 1) in
low energy shallow lake bottoms as the only
recorded association (Fig. 11, T1), occasionally
they are associated with tetrapod footprints with
irregular outline and marginal rims (Mode C).
Locally, ephemeral floods occurred, and Suites 2
and 3 are present on soft substrates. When the
surface was exposed, the firm mud preserved
footprints with high fidelity of the producer’s
pedal anatomy together with desiccation cracks
(Mode A) (Fig. 11, T2).
The mudflat facies association (only recorded at
La Pe~na creek) contains few trace fossils, although
individual beds are bioturbated. Laminated mud-
stone facies records almost exclusively Suite 2
(vertical dwelling Arenicolites isp. and equilib-
rium trace Diplocraterion parallelum) together
with tetrapod footprints of taphonomic Mode C
(Fig. 11). These pioneers had the ability to colo-
nize the muddy soft substrate once the suspended
material settled out. The mudflat, with high sus-
pended-sediment content, was not viable to be
colonized by invertebrate larvae that mostly prefer
sandy substrates under clean water conditions.
DISCUSSION
The R
ıo Mendoza–Cerro de las Cabras sequence
in the Las Pe~
nas sub-basin is dominated by an

© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology

 Underfilled lake characterized by multi-approach
alluvial fan, an ephemeral fluvial system with
development of playa-lake and a perennial-lake
associated with fluvio-lacustrine environment
(Rolleri & Criado Roqu
e, 1968; Carrara, 1970;
Kokogian & Mancilla, 1989; Kokogian et al.,
1993). This contribution is focused on sandflat
and mudflat deposits developed in the
ephemeral distal fluvial and playa-lake system,
where the ichnological record (invertebrate
traces and tetrapod tracks) is concentrated. The
analyzed succession includes abundant sedi-
mentological features such as mud-cracks,
mud-chips and rain-drop imprints that indicate
subaerial exposure, thus, suggesting that the
surface water+sediment supply was ephemeral
(Gierlowski-Kordesch & Rust, 1994; Gierlowski-
Kordesch, 1998). The stacking pattern observed
in the sections is mainly aggradational, a char-
acteristic of an underfilled lake-basin (Bohacs
et al., 2000; Benavente et al., 2019). The sand-
flat facies association (documented mainly in
the sections of the Vallet south and Vallet north
quarries, see Fig. 3) represents a marginal sub-
environment, developed in the ephemeral distal
fluvial system, whereas the mudflat facies asso-
ciation documented in the La Pe~ na creek sec-
tion represents the central sub-environment
developed in the playa-lake system. Further-
more, the transition between sandflat and mud-
flat facies association observed in the Cabrera
quarry section (Fig. 3) constitutes the transition
between the marginal and central sub-environ-
ments in the ephemeral distal fluvial and playa-
lake system.
Footprint preservation
The preservation potential of tetrapod tracks
and invertebrate trace fossils strongly depends
on the rheological condition of the sediment,
such as grain size, mineralogy and moisture con-
tent, at the moment of emplacement (McKee,
1947; Allen, 1997; Zhang et al., 1998; Manning,
2004; Marchetti et al., 2019). Accordingly, the
timescales on which accretion–erosion occur
can produce partial or total infilling or erosion
before final entombment, and/or obliteration by
subaerial processes such as wind, desiccation,
precipitation and trampling. Therefore, preserva-
tion of tetrapod tracks is restricted to a narrow
range of sediment texture and moisture content
with limited potential for permanent burial.
Thus, fundamental steps for footprint preserva-
tion are: a moist substrate where the tracks are
© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology
21
printed, followed by a short period of subaerial
exposure that produces a firm surface able to
resist erosion when the overlying bed is depos-
ited and, finally, the surface is covered by a new
layer of sediment which rapidly entombs the
tracks (McKee, 1947; Allen, 1997; Tucker &
Burchette, 1977; Laporte & Behrensmeyer, 1980;
Manning, 2004).
Moisture content of sediments is one of the
undoubtedly crucial factors at the time of track-
making and affects not only the formation of the
tracks but also the quality of anatomical details
imprinted. When the sediment is underwater,
the saturation of interstitial voids produced dur-
ing the track-making generates liquefied sedi-
ment, the collapse of the deformed sediment
and, in some cases, the disturbance is obliter-
ated (Cohen et al., 1991; Allen, 1997; Paik et al.,
2001). The Cerro de las Cabras succession con-
tains very few levels that had the sediment-
saturation conditions to preserve tracks under
these circumstances. This situation might be
because animals avoided walking over this type
of surface. Convolute structures that could be
attributed to animal disturbance, however, were
recorded, particularly in Section 1 (Vallet south
quarry see Figs 3 and 4F).
High moisture content of sediment is
reflected by potential flow and collapse pro-
duced during the track-making but less promi-
nently than with underwater conditions.
Generally, the sediment preserved the deforma-
tion as a shaft surrounded by a marginal ridge,
without signs of liquefaction. As a result of
deformation, tracks show only very general
anatomical features (Cohen et al., 1991; Allen,
1997; Paik et al., 2001; Manning, 2004). The
Mode C assemblages documented at Cerro de
las Cabras were interpreted as footprints made
on surfaces with high water content (Fig. 12).
Particularly, they show a well-defined oval or
irregular deep shaft without anatomical fea-
tures. The marginal rim that is symmetrically
developed surrounding the shaft indicates simi-
lar moisture content areally in the sediment
and can be related with substrate consistency
more than with animal progression (see Marsi-
cano et al., 2010).
Sediment with moderate moisture content dis-
turbed by animal walking retains the general
shape of impressions, thus preserving some
anatomical detail (Cohen et al., 1991; Allen,
1997; Paik et al., 2001, Manning, 2004). The lack
of pronounced deformation (collapsed layers
and flow) reveals that high moisture and
22 A. C. Mancuso et al.
underwater conditions are absent. Under these
conditions, marginal rims could develop weakly
and, in some cases, were limited to only one
side of the shaft or absent. In the Cerro de las
Cabras succession, the Mode B tracks are inter-
preted to have been produced on a soft substrate
with moderate moisture content (Fig. 12). Most
frequently, the tracks lack anatomical detail of
the trackmaker’s manus/pes (Fig. 9) and the
marginal rim is absent, the degree of deforma-
tion of sediment reveals intermediate water satu-
ration.
A firm surface with underlying sediment of
low moisture content has sufficient plasticity to
record footprints with anatomical detail. The
progression of an animal on such a surface pro-
duced a shallow shaft without signs of collapse,
marginal rims or flow, and preservation of
anatomical detail (Cohen et al., 1991; Allen,
1997; Paik et al., 2001; Manning, 2004). Like-
wise, good preservation with details of the digit
and palm impressions and even claw drag marks
are documented in the Mode A track of the
Cerro de las Cabras succession (Fig. 9). The low
moisture condition (Fig. 12) is reflected by shal-
low depth independent of the size of the track
and, therefore, of the trackmaker’s bodyweight,
© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology
Fig. 12. Summary of the
relationship between the surface
condition and footprint-taphonomic
mode recognized and defined in
this contribution.
the clear impression of morphological features,
and the absence of deformation structures.
Grain size and mineralogy play an important
role in the behaviour of the substrate (e.g.
McKee, 1947; Allen, 1997; Manning, 2004). For
a particular moisture content, variation in the
proportion of mud and sand in the layer domi-
nates the firmness of the surface (Fig. 12) (Cohen
et al., 1991; Paik et al., 2001). A higher propor-
tion of mud introduces high plasticity on the
surface, resulting in tetrapod tracks with larger
deformation structures for the same moisture
content. Regarding clay mineralogy, the swelling
clays that can hold more water generate high
plasticity with same mud/sand proportion, and
layers that contain non-swelling clays have
lower plasticity (Fig. 12) (Gierlowski-Kordesch &
Rust, 1994; Gierlowski-Kordesch & Gibling,
2002; Scott et al., 2010). In the present case, the
Cerro de las Cabras succession reveals a low
proportion of mud in the sandy layers with
Mode A and Mode B preservation, specifically
in Sections 1 and 2 (Vallet south and Vallet
north quarries), despite the presence of mud-
stone drapes (Fig. 9). The opposite situation is
reported in the La Pe~ na creek (Section 4, see
Fig. 3) where the footprints are mainly
 Underfilled lake characterized by multi-approach
preserved as Mode C and, in less proportion, as
Mode B and Mode A, associated with a higher
mud proportion in the sediments at the time of
imprinting (Fig. 3). Section 3 (Cabrera quarry,
see Fig. 3) shows an intermediate situation
where the dominance of Mode A of preservation
is related to the well-developed alternation of
sandy and muddy layers, with sandy layers pro-
viding firm support to the muddy ones (Fig. 3)
(Cohen et al., 1991; Paik et al., 2001). The clay
mineralogy is dominated by illite, a non-swel-
ling clay, forming single beds with a minor con-
tribution of swelling clays (smectite and
interstratified illite/smectite dominated by smec-
tite). This condition suggests low plasticity of
the beds, which is supported by the lack of
deformation structures in all tetrapod track
taphonomic modes, even in the large tracks
imprinted on the muddy layers. Absent in the
succession are potential deformation footprint
features such as collapse or flow due to liquefac-
tion that would be produced by large animals
moving in underwater conditions.
After impression, footprints can be exposed to
such potential processes of destruction as tram-
pling, extreme desiccation, erosion, and also
they can be disturbed by invertebrate and plant
activity (Tucker & Burchette, 1977; Laporte &
Behrensmeyer, 1980; Paik et al., 2001). The fre-
quency, duration and severity of these modify-
ing events all influenced the preservation or
destruction of the footprints (Tucker & Burch-
ette, 1977; Laporte & Behrensmeyer, 1980; Paik
et al., 2001). Features such as the presence of
mudstone chips, mottled fabric, mud-cracks,
rain-drops and wrinkle structures recorded in
the track-bearing levels indicate subaerial expo-
sure, which is consistent with the preservation
state of the footprints and the ephemeral-flow
conditions of the palaeoenvironment. However,
the particularly well-preserved features docu-
mented in each mode, and the absence of pedo-
genic disturbance, trampling and/or evidence of
strong disturbance by wind, desiccation and/or
precipitation, supports short periods of exposure
of the imprinted surface. The entombing flows
of track-bearing levels mostly were not erosive,
in agreement with the type of depositional sys-
tems, where recurrent unconfined inflows
deposited sediments that acted as successive
potential track-bearing levels (Fisher et al.,
2007a,b, 2008), but despite this can still record
some erosive flows. The abundance and distri-
bution of mud-cracks in close relation with the
tracks do not necessitate extended periods of
© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology
23
exposure, because such cracks can form in just a
few hours (Tang et al., 2011). Thus, in the Cerro
de las Cabras succession case, the time between
deposition of the track-bearing layer with
enough moisture to preserve tracks and the
entombment of the layer probably ranged from
hours to days (Scott et al., 2010; Tang et al.,
2011).
To summarize, the factors that enhanced the
preservation of the tetrapod tracks in the Cerro
de las Cabras succession include a moderate
proportion of mudstone containing mostly illite
in the substrate, low to moderate moisture-con-
tent, and relatively rapid final entombment.
These observations allow characterization of
the different areas of the system more specifi-
cally. In a marginal position, such as Sections
1 and 2 (Vallet south and Vallet north quar-
ries), sandy layers dominate with a low pro-
portion of mud, whereas the central position,
represented by Section 4 (La Pe~ na creek), is
dominated almost exclusively by muddy layers
with strongly rhythmic alternation where thin
sandy layers provided firm support. The low
content of mud in the marginal position is
mainly related to recurrent surficial waning
flows that removed the light particles and
deposited them basinward. The potential to
maintain substrate moisture is closely related
to the water-table position and its fluctuations
with climate (Allen, 1997), particularly in
ephemeral systems (Rosen, 1994; Mel
endez
et al., 2009). The general dominance of illite
suggests an enhanced erosion rate consistent
with a high hydrolysis index that provided
illite-rich clays into the Cerro de las Cabras
depositional system from the source areas, per-
haps linked to humid conditions. Humidity is
also in agreement with the low content of the
muddy component of the sandstone layers in
the marginal basin position. This also explains
the frequent occurrence of surficial flows, both
in the sandflat and mudflat sub-environments.
The high frequency of surficial flows would
have enhanced footprint preservation and sug-
gests short dry periods. A relatively high water
table is evidenced not only by well-developed
footprints and invertebrate traces basinward
but also because the development of mud-
cracks progressively needs a continuous supply
of water from within the soil to the evapora-
tion surface (Tang et al., 2011). Moreover, the
relatively high water-table could be evidence
of persistent standing water in the lake devel-
oped in the deepest area of the sub-basin, as
24 A. C. Mancuso et al.
is recorded upward in the Cerro de las Cabras
succession.
Global palaeoclimatic models propose marked
seasonality (wet and dry periods) during the
Triassic for western Gondwana (Parrish, 1993;
Sellwood & Valdes, 2006; Preto et al., 2010;
Holz, 2015). For the Cuyana Basin, smectite-
rich clay assemblages and the presence of verti-
sols in the Cerro de las Cabras Formation
(Potrerillos sub-basin) also support the interpre-
tation of seasonal conditions during its deposi-
tion (Benavente et al., 2015). Smectite can
potentially be the product of basic volcanic
rocks depositing in alkaline lacustrine waters
leading to clay assemblages dominated by zeo-
lites (Chamley, 1989). Nevertheless, no zeolites
have been identified in clay minerals within
the Potrerillos sub-basin. Therefore, most likely
detrital smectite in that area of the rift points
to weathering of poorly drained terrains under
seasonally high rainfall conditions (Singer,
1980; F€ ursich et al., 2005). In that particular
case, the location of the outcrops at the passive
margin of the rift allowed the inference of a
possible local rain-shadow effect as well (Bena-
vente et al., 2015). Evidence recorded in the
Cerro de las Cabras Formation (illite-rich clay
assemblages with smectite contribution and
illite/smectite dominated by smectite) in the
Las Pe~ nas sub-basin does not support such an
effect and points to seasonality in the system,
which is probably related to the main develop-
ment of the highest terrains in the east of the
sub-basin, with a low margin to the west. This
palaeogeographic configuration of the sub-basin
possibly promoted the dominance of wet
periods over drier ones.
Underfilled ichnological preservation model
In lacustrine environments, the Scoyenia and
Mermia ichnofacies have been recorded in
lake-margin and lake-centre sub-environments,
respectively, and are present from overfilled to
underfilled lake-basins (Buatois & M
angano,
2004, 2009). The information available for playa-
lake-centre ichnology of underfilled lake-basins
is quite scarce. It is not surprising that the
Scoyenia ichnofacies is not only preserved on
the playa-lake margins but also in the sandflat
facies association, towards the playa-lake-centre
position. The Mermia ichnofacies is commonly
absent in playa-lakes (Buatois & M
angano, 2004,
2009).
© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology
In the Cerro de las Cabras Formation, the
sandflat facies association is mostly character-
ized by the Scoyenia ichnofacies, typically dom-
inated by low to moderate ichnodiversity and
locally high abundance of traces. Classically, it
records an abundance of meniscate traces of
mobile deposit feeders and locomotion traces
produced by arthropods, the presence of vertical
domiciles, and also the presence of tetrapod
footprints. It characterizes low-energy deposits
periodically exposed to air and then inundated,
situating the ichnofacies as the intermediate
between aquatic and non-aquatic sub-environ-
ments (Frey et al., 1984; Buatois & M
angano,
2011). This soft-ground to firm-ground suite
mostly comprises tetrapod footprints with the
most detailed pedal anatomical preservation
(Mode A) and, less abundant, horizontal dwell-
ing and feeding traces of semiaquatic inverte-
brates (Suite 3; Palaeophycus, Taenidium and
Scoyenia).
However, the soft-ground suite is dominated
locally (spatially and temporally) by vertical
dwelling burrows (Suite 2; Skolithos and Areni-
colites) of mainly subaqueous invertebrates of
the Skolithos ichnofacies. It is mostly recorded
on high-energy ephemeral-flood deposits that
occurred in the sandflat facies association of the
playa-lake margin. The tetrapod footprints pre-
served in the soft-ground suite correspond to the
taphonomic modes B and C and constitute a sec-
ondary element of the suite (Figs 10 and 13).
Other local components of the sandflat facies
association are the delicate grazing
(Helmintoidichnites and Helmintopsis) and
arthropod locomotion traces (Diplopodichnus
and Cruziana) of the Mermia ichnofacies in
shallow-water ponds that rapidly got exposed
and desiccated. The tetrapod footprints pre-
served together with this assemblage correspond
to Mode A and mostly occurred after dewatering
of the sediments (Figs 10 and 13).
In contrast, the basin-centre position is almost
devoid of invertebrate trace fossils, but replete
with tetrapod footprints, probably because the
centre areas were the location of the last rem-
nant waterholes (standing water) during dry
periods. The Skolithos ichnofacies is mainly
recorded towards the basin centre, in the mud-
flat facies association, with vertical dwelling
and equilibrium traces (Suite 2; Arenicolites isp.
and Diplocraterion parallelum) (Figs 11 and 13).
Basin-centre position of a playa lake system
does not represent the profundal, permanently
submerged subaqueous zone of the lacustrine
 Underfilled lake characterized by multi-approach 25

Fig. 13. Underfilled ichnological preservation model. MA: Footprint-taphonomic Mode A, MB: Footprint-tapho-
nomic Mode B, MC: Footprint-taphonomic Mode C, A: Arenicolites, S: Skolithos, Di: Diplocraterion, P: Palaeophy-
cus, T: Taenidium, D: Diplopodichnus, Ho: Helminthopsis, Hi: Helminthoidichnites, G: mud-cracks.

system, at least not at all scales. Tetrapod foot-
prints of large size, such as those produced by
archosaurs and large dicynodont therapsids, are
mostly recorded on the soft muddy substrates
(Mode C). In contrast, small size tracks, pro-
duced mainly by small therapsids and small
archosaurs, are recorded on the firm substrates
(Modes A and B) (Figs 11 and 13).
Sequence stratigraphy
The ichnological record, by means of recording
all sorts of biological activities of plants, inverte-
brates and tetrapods, reinforces the identifica-
tion of non-depositional surfaces that could not
be discerned based uniquely on physical sedi-
mentary structures (Buatois & M
angano, 2004,
2009; Scott et al., 2009; Marsicano et al., 2010).
The record and hierarchal rank of the hiatus rep-
resented by a surface of non-deposition is
related to the kind of non-depositional process
and the time span involved. Non-depositional
© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology
processes include erosion, bypass, down-lap,
omission and firm-ground formation (Mitchum
& Vail, 1977; Bohacs, 1998). Depositional
sequence boundaries are characterized by ero-
sion, bypass and onlap; they record prolonged
abnormal subaerial exposure in proximal areas
(cf. Bohacs et al., 2007) to be distinguished from
surfaces that record ‘normal’ subaerial exposure
of short duration, such as might be found in a
floodplain or lake-plain environments. Sequence
stratigraphy uses the variety of such surfaces
and their hierarchical relations as boundaries of
depositional sequences, and the parasequence
sets that constitute them that, in turn, allow
reconstruction of basin infill patterns (Catu-
neanu et al., 2011).
A depositional sequence boundary is typically
a well-expressed unconformity in basin-margin
areas, whereas towards the basin centre this
might be recorded as a correlative conformity
(Catuneanu et al., 2011). Omission surfaces rec-
ognized via their record of overprinted
26 A. C. Mancuso et al.

biological activity can be correlative conformi-
ties linked with an unconformable part of a
sequence boundary (Buatois & M
angano, 2004,
2009; Bohacs et al., 2007; Marsicano et al.,
2010). Thus, the observation of extensive biolog-
ical activity recorded as trace fossils (inverte-
brate and vertebrate) helps to recognize non-
depositional surfaces and reconstruct the basin
infill patterns (Buatois & M

angano, 2004, 2009;
Bohacs et al., 2007; Marsicano et al., 2010). It is
essential, however, to recognize not only the
non-depositional surfaces but also their hierar-
chy. Thus, the integration of the ichnological
content (invertebrate and vertebrate), within a
detailed taphonomical context is useful as a sen-
sitive indicator of the surface hierarchy. This
integration combined with sequence stratigraphy
helps to refine its evolutionary interpretation
and discriminate autogenic versus allogenic con-
trols in this lake basin history. Table 3 summa-
rizes the steps of this integrated multiproxy
approach, which has a high potential for appli-
cation in other underfilled lake basins that pre-
serve ichnological evidence.
In the Cerro de la Cabras palaeolake, biologi-
cal activity was recorded by tetrapod foot-
prints, with three taphonomic modes and
invertebrate trace fossils characterized by three
suites grouped in multiple taphonomic path-
ways. The mode of footprint preservation,
along with the absence of pedogenic distur-
bance, trampling, or evidence of strong distur-
bance by wind, desiccation or precipitation,
supports the fact that most surfaces record
‘normal’ subaerial exposure of short duration
between successive flood beds. In consequence,

Table 3. Integrated multiproxy approach for evolutionary interpretation and identification of autogenic versus
allogenic controls in underfilled lake-basins history.
Describe sedimentology, mineralogy, stratigraphy and paleogeographic setting of lake strata.
Acknowledge complexities difficult to model in the lake basin (palaeogeography linked to catchment areas and
rain shadows, size of the lacustrine basin versus size of catchment, among others).

Determine lake-basin type:

Characterize lake-basin type from stratigraphy, sedimentology, mineralogy, etc. (Bohacs et al., 2000;
Benavente et al., 2019).

Recognize invertebrate ichnology record in context and analyze:

Recognize the ichnological taxa in a stratigraphic position;
Recognize the ichnofacies;
Analyze the taphonomic pathways to estimate temporal, spatial and facies variability.

Recognize vertebrate ichnology record in context and analyze:

Recognize the ichnological taxa in a stratigraphic position;
Recognize the taphonomic modes;
Analyze the taphonomic modes to estimate temporal, spatial and facies variability.

Integrate the taphonomic modes and the taphonomic pathways in an underfilled ichnological preservational
model and estimate temporal, spatial, and facies variability (Buatois & M

angano, 2004, 2009).

Interpret hydrography and hydrology:

Characterize the table water position and fluctuation from the underfilled ichnological preservational model.

Interpret palaeoclimate:
Use ichnological preservational model to estimate the seasonality of precipitation/temperature;
Use primary clay assemblages as proxies for climate conditions
Compare interpretations with paleoclimate models for the same age and region;
Compare interpretations with coeval units from same general vicinity to differentiate effects of regional
climate from local hydrology.

Interpret sequence stratigraphy:

Use ichnological preservational model to identify non-depositional surfaces;
Use the non-depositional surfaces identified to estimate the hiatus represented and distinguish depositional
sequence boundaries (allogenic) from surfaces that record ‘normal’ subaerial exposure of short duration
(autogenic)

© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology

 Underfilled lake characterized by multi-approach
the studied exposure surfaces in the Cerro de
las Cabras succession do not represent
sequence boundaries, neither as erosional and/
or as omission surfaces. Thus, the portion of
the succession studied represents the evolution
of a playa-lake system (including sandflat and
mudflat deposits) in a continuous subsidence
context. The observed vertical and lateral varia-
tions probably indicate autogenic processes
and, to a lesser extent, allogenic ones. The
main tectonic role is the generation of accom-
modation in the rift basin during the syn-rift
phase, as the allogenic control on basin centre
subsidence and spill-point elevation. The
absence of erosional and/or omission surfaces
suggests that the subsidence was uninterrupted
and without episodic tilting events (Pietras &
Carroll, 2006; Catuneanu et al., 2011; Scott
et al., 2012). The studied sections display a
dominantly aggradational stacking pattern of
sandflat facies association in sections 1 and 2
(Vallet south and Vallet north quarries, respec-
tively), and transition between sandflat and
mudflat facies association in Section 3 (Cabrera
quarry) and mudflat facies association in Sec-
tion 4 (La Pe~na creek), with a potential progra-
dational trend in Section 1 (see Fig. 3). The
aggradational trend observed strongly suggests
an equilibrium between rates of accommoda-
tion increase and sediment supply. The short
periods of exposure recorded by biological
activity not only in the sandflat (basin margin)
but also in the mudflat (basin centre) indicate
that the rate of sediment supply into the basin
was enough to keep the accommodation filled;
thus, the centre of the basin did not experience
prolonged starved conditions. Therefore, the
climate, as another allogenic control, may have
promoted water and sediment supply in the
basin, associated with seasonal conditions with
the dominance of wet periods over dry ones.
CONCLUSIONS
The abundance, diversity and preservation of
tetrapod footprints, and invertebrate trace fossils
in underfilled systems, in particular related to
sandflat and mudflat sub-environments, agree
with the general hypothesis that these settings
commonly produce the narrow range of condi-
tions needed for their preservation. On the other
hand, invertebrate and vertebrate traces, ready
to be interpreted at first sight on the field, pro-
vide insights and detailed characterization of
© 2020 The Authors. Sedimentology © 2020 International Association of Sedimentologists, Sedimentology
27
underfilled systems not available otherwise.
Moreover, both analyses, invertebrate traces
focusing on their ethology and taphonomy (ich-
nofacies models and taphonomic pathways) and
tetrapod-track taphonomic modes, provide
taxon-independent or taxon-free approaches for
multi-age case studies. Sensitive indicators of
the environmental conditions and the timescales
on which accretion–erosion occurs in lacustrine
systems are based on simple taxon-free informa-
tion (fidelity to the trackmaker’s pedal anatomy,
presence or absence of foot marks and marginal
rims, and trampling degree on the surface). The
particular well-preserved features documented
in each mode and the absence of pedogenic dis-
turbance, trampling obliterating the tracks, or
evidence of strong disturbance by wind, desicca-
tion or precipitation, supports short periods of
exposure of the imprinted surface. Thus, the
vertical and lateral variations indicate autogenic
processes. The aggradational trend observed sug-
gests an equilibrium between rates of accommo-
dation increase and sediment supply (controlled
mostly by allogenic processes). The short peri-
ods of exposure recorded by biological activity
not only in the sandflat (basin margin) but also
in the mudflat (basin centre) facies associations
indicate that the rate of sediment supply into
the basin was enough to keep the accommoda-
tion filled; thus the centre of the basin did not
experience prolonged starved conditions. There-
fore, tectonics, as an allogenic control, during
the syn-rift phase, generated accommodation by
subsidence and spill-point elevation. Climate, as
the other allogenic control, promoted seasonal
water and sediment supply in the basin with
wet periods being more significant than drier
ones. The integrated multiproxy approach is
useful for evolutionary interpretation and identi-
fication of autogenic versus allogenic controls in
underfilled lake-basin history.
ACKNOWLEDGEMENTS
We are grateful to Carlos Vallet, Jorge and Gra-
ciela Sterponi, Carlos Cabrera, Juan Carlos Cena-
tore, the owners of the quarries, for their
support during this project. For access per-
misi
on, we thank Elina Albarran (Direcci
on de
Patrimonio Cultural provincial de Mendoza).
Authors thank Gilda Collo for rich discussions
on clay mineralogy. The authors thank two
reviewers Mangano, G. and Bohacs, K., Associ-
ate Editor Veiga, G. and Chief Editor Della Porta
28 A. C. Mancuso et al.
G. for help that improved the quality of the
manuscript. Funding for this research was pro-
vided by projects PICT-2013-0805 (ACM),
UBACyT 20020100100728 (CAM), PICT- 2014-
1921 (VK) and PICT 2014-0489 (CAB), PIP CON-
ICET 2015-00613 (Ottone), PICT-2017-2159 (de
la Fuente). Additional financial support was
provided by the Consejo Nacional de Investiga-
ciones Cient
ıficas y T

ecnicas. This is CAM’s and
VK’s contribution R-297 to the Instituto de Estu-
dios Andinos Don Pablo Groeber (IDEAN, CONI-
CET).
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Manuscript received 14 June 2019; revision accepted
16 March 2020