Palaeogeography, Palaeoclimatology, Palaeoecology 273 (2009) 75–86

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Palaeogeography, Palaeoclimatology, Palaeoecology

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

Ichnology and sedimentology of a tide-influenced delta, Lower Miocene Chenque

Formation, Patagonia, Argentina: Trace-fossil distribution and response to
environmental stresses
Noelia B. Carmona a,⁎, Luis A. Buatois b, Juan José Ponce a, María Gabriela Mángano b
a
Laboratorio de Geología Andina, CONICET — Centro Austral de Investigaciones Científicas (CADIC), B. Houssay 200, C.P. 9410, Ushuaia, Tierra del Fuego, Argentina
b
Department of Geological Sciences, University of Saskatchewan, 114 Science Place, Saskatoon, Canada SK S7N 5E2

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

Article history: Combined sedimentologic and ichnologic analysis of the Lower Miocene Chenque Formation allows
Received 2 July 2008 recognition of a tide-influenced deltaic succession exposed along the coast of Caleta Olivia city, Patagonia,
Received in revised form 2 December 2008 Argentina. Two main subenvironments were identified, prodelta and delta front, stacked forming a
Accepted 2 December 2008
progradational coarsening-upward succession, up to 10 m thick. The prodelta deposits are mainly
characterized by heterolithic strata (lenticular and wavy bedding), with low bioturbation intensity and
Keywords:
Ichnology
sporadic distribution of trace fossils. The trace-fossil assemblage is dominated by deposit-feeder structures
Tide-influenced delta (e.g. Planolites montanus, Protovirgularia isp., and Teichichnus rectus), constituting an impoverished expression
Miocene of the Cruziana ichnofacies, with respect to their fully marine counterparts. A transition zone between the
Patagonia prodelta and the delta front is discontinuously distributed along this outcrop. This interval consists mainly of
Chenque Formation flaser-bedded sandstone, almost completely obliterated by equilibrium/adjustment trace fossils of large
bivalves (Atrina), and subordinately, by the trace fossils Nereites missouriensis, Phycosiphon incertum, T. rectus,
Thalassinoides isp., and Schaubcylindrichnus freyi. The delta-front deposits consist of sigmoidal cross-stratified
sandstone with mud drapes. The trace-fossil assemblage is dominated by large Rosselia socialis and Macar-
onichnus segregatis in the sandier beds, whereas the mud drapes blanketing the sandstone foresets
commonly contain N. missouriensis and Protovirgularia isp. Ichnologic characteristics (e.g. shallow-tiered
communities, impoverished trace-fossil assemblages, dominance of deposit-feeder structures, and inhibition
of suspension-feeder elements) suggest that different paleoenvironmental stresses, such as changes in
salinity, water turbidity, and fluctuations in energy and in sedimentation rates, affected the infaunal
communities of these tide-influenced delta settings.
© 2008 Elsevier B.V. All rights reserved.

1. Introduction diversity of biogenic structures in these Miocene successions, mainly in

Trace fossils constitute valuable tools in the recognition of stresses
in marine settings. During the last years there has been an increased
interest in the ichnology of deltaic environments (McIlroy, 2004;
MacEachern et al., 2005; McIlroy, 2007). These analyses were
principally focused on describing the trace-fossil content of river-,
wave-, and storm-influenced deltaic deposits. However, the ichnology
of tide-influenced deltas remains mostly understudied with the
exception of McIlroy (2004, 2007).
Shallow-marine environments of the Lower Miocene Chenque
Formation of Patagonia, Argentina, are characterized by superbly
preserved trace fossils. Recent studies (e.g. Buatois et al., 2003; Carmona,
2005; Carmona et al., 2008a) documented the high abundance and
⁎ Corresponding author. Fax: +54 2901 430644.
E-mail addresses: carmonanb@yahoo.com.ar (N.B. Carmona), luis.buatois@usask.ca
(L.A. Buatois), ponce_juanjose@yahoo.com.ar (J.J. Ponce), gabriela.mangano@usask.ca
(M.G. Mángano).
shallow-marine deposits developed under normal-marine salinity
conditions. In addition, these studies also reflected the occurrence of
impoverished trace-fossil suites in estuarine and deltaic deposits. In
particular, outcrops along the coast of Caleta Olivia city have been
interpreted as formed in a deltaic environment strongly influenced by
tides. Therefore, the aims of this study are two-fold: (1) to describe and
interpret the sedimentologic and ichnologic characteristics of these
deltaic deposits, and (2) to evaluate how these trace-fossil suites reflect
paleoenvironmental stresses typical of deltas.
2. Geological setting
The Lower Miocene Chenque Formation crops out in the east
region of Central Patagonia, Argentina (Fig. 1). This formation
comprises a wide variety of shallow-marine and marginal-marine
strata that were deposited during two major transgressions (the
Leonense and Superpatagoniense transgressions), which occurred
during the Early Miocene (Bellosi, 1987, 1995). The Chenque Formation

0031-0182/$ – see front matter © 2008 Elsevier B.V. All rights reserved.
doi:10.1016/j.palaeo.2008.12.003
76 N.B. Carmona et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 273 (2009) 75–86
Fig. 1. Map showing the distribution of the Chenque Formation, and the studied succession at Caleta Olivia city.
comprises five shallowing-upward sequences. The first two sequences
(Sequence I and II) were deposited during the Leonense transgression,
and comprise marine successions formed in shelf environments, as
well as in shallower-water settings (e.g. estuarine and deltaic
environments). The upper three sequences (III–V) were deposited
during the Superpatagoniense transgression (late Early Miocene) and
comprise mainly very shallow, tide-dominated deposits (Bellosi,
1995). The present study focuses on a succession belonging to the
lower sequence (Bellosi, 1987), cropping out along the Atlantic coast of
Caleta Olivia city, Santa Cruz province (Fig. 1).
3. Sedimentology, trace-fossil distribution, and depositional
environments
Two main facies associations have been identified in the studied
succession: tide-influenced prodelta, and tide-influenced delta-front
(Fig. 2). The tide-influenced prodelta association includes heterolithic
facies, encompassing distal- (facies 1) and proximal- (facies 2)
prodelta environments, as well as a transition zone between prodelta
and delta front (facies 3). The tide-influenced delta-front association
includes two sandstone-dominated facies, recording deposition in a
distal (facies 4) to proximal (facies 5) delta-front setting.
3.1. Prodelta facies association
3.1.1. Facies 1: Lenticular-bedded mudstone and sandstone
This facies consists of regular alternations (millimeter- to cen-
timeter-thick) of very fine-grained sandstone interbedded with
mudstone (Figs. 2 and 3). This heterolithic facies is dominated by
massive or parallel-laminated mudstone and sandstone showing
lenticular bedding. Sandstone lenses contain current-ripple cross-
lamination, asymmetrical ripples, and mud drapes (Fig. 3A). Although
individual beds are commonly lenticular and show lateral thickness
changes, bedsets are laterally persistent. Soft-sediment deformation
structures and synaeresis cracks are abundant (Fig. 3A–C). Paleocur-
rent measurements indicate a bipolar pattern with flows directed
either to the southeast and northwest (Fig. 3A). Facies 1 shows
alternation of unburrowed intervals and moderately bioturbated (BI
0–2) beds. The ichnofauna is mostly composed of deposit-feeder
structures, mainly Planolites montanus and Protovirgularia isp.
(Fig. 3D), although some isolated specimens of Thalassinoides isp.
also occur (Fig. 3A).
Interpretation: This facies records deposition in a low-energy
setting, with dominance of mud fallout and fluid muds, punctuated by
tractive sand deposition. The presence of mud drapes in the cross-
 N.B. Carmona et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 273 (2009) 75–86
Fig. 2. Schematic representation of an idealized section at the studied locality, integrating the sedimentologic and ichnologic information.
laminated sandstone lenses reflects mud settling during periods of
slack water, and the common bipolar paleocurrent orientation suggest
tidal processes. Fluctuations in salinity are inferred by the presence of
synaeresis cracks. The impoverished trace-fossil suites also suggest
salinity stress, and probably the occurrence of soupgrounds. On the
contrary, the more bioturbated intervals may reveal periods of
relatively normal-marine salinity conditions, and slower sedimenta-
tion rates. These characteristics suggest that facies 1 was deposited as
distal bottomsets of a tide-influenced distal prodelta.
3.1.2. Facies 2: Wavy-bedded sandstone and mudstone
Facies 2 consists of fine-grained sandstone interbedded with
mudstone, displaying wavy and, more rarely, flaser bedding (Figs. 2
and 4A–B). These heterolithic beds show a gradational contact with
facies 1. As with facies 1, individual layers display common lateral
variations in thickness, but bedsets show tabular geometries. Synaer-
esis cracks occur in thin mudstone intervals (Fig. 4A–C). The sandstone
beds display asymmetrical ripples with mud drapes at the base of this
77
facies, being generally replaced in some levels by symmetrical ripples
at the top of these beds. Soft-sediment deformation structures occur
locally (Fig. 4B). Paleocurrent measurements indicate a bidirectional
pattern with flows directed to the east and west. This facies records an
increase in bioturbation intensity (BI 1–3), and the trace-fossil
assemblage is dominated by deposit-feeding burrows (e.g. Astero-
soma isp., Planolites montanus, Protovirgularia isp., and Teichichnus
rectus) (Fig. 4A, C–D). Subordinate and rare elements include Nereites
missouriensis, Phycosiphon incertum, Schaubcylindrichnus freyi, Sko-
lithos isp., and Thalassinoides isp.
Interpretation: Dominance of wavy bedding indicates alternation
of bedload transport and suspension fallout during slack-water
periods. In addition, fluid muds may have also been responsible for
accumulation of fine-grained material. Tidal action is also indicated by
the bipolar paleocurrent orientation, whereas wave action is only
recognized locally by the presence of symmetrical ripples. The
presence of synaeresis cracks indicates salinity fluctuations, although
these variations are apparently less pronounced than in facies 1.
78 N.B. Carmona et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 273 (2009) 75–86

Fig. 3. Facies 1. Distal prodelta. A and B. Cross-section view of the thinly interbedded sandy mudstone, with lenticular bedding. This facies displays abundant synaeresis cracks (sy),
soft-sediment deformation structures (def), and low bioturbation intensities, with isolated Planolites montanus (Pl) and Thalassinoides isp. (Th). Note also the two opposite current
directions indicated by arrows. C. Bedding-plane view showing the mudstone intervals with synaeresis cracks (sy). D. Bedding-plane view with a specimen of Protovirgularia isp. (Pr).
Note the poorly defined outline of this trace fossil, suggesting that the substrate was relatively fluid.

Increase in grain size suggests a more proximal position than in facies
1, although the heterolithic nature of these deposits still indicates a
position within the prodelta setting (i.e. bottomsets of a proximal
prodelta). The increase in both bioturbation intensity and ichnodiver-
isty may reflect more marine conditions and shorter periods with
salinity stress. An increase in ichnodiversity from the distal to the
proximal prodelta was also observed by McIlroy (2007) in the Jurassic
Lajas Formation, Neuquén Basin, Argentina. He considered that this
pattern was possibly an artifact of preservation potential, although he
also mentioned that this increment could be linked to a greater range
of food resources in the proximal prodelta, and therefore a wider
range of behavioral types (McIlroy, 2007).
3.1.3. Facies 3: Flaser-bedded sandstone
Facies 3 consists of muddy sandstone heterolithics displaying
mainly flaser bedding, and subordinately wavy bedding (Figs. 2 and 5).
Sandstone beds are separated by thin mudstone units. This facies
occurs transitionally from facies 2, and is laterally discontinuous.
Sedimentary structures are almost completely obliterated by biotur-
bation, especially by equilibrium/adjustment trace fossils produced by
large bivalves (Atrina) (BI 3–5) (Fig. 5B–C). Subordinate trace fossils
include Nereites missouriensis (Fig. 5A, C–E), Phycosiphon incertum
(Fig. 5A), Teichichnus rectus (Fig. 5D), Thalassinoides isp., and Schaub-
cylindrichnus freyi.
Interpretation: Presence of equilibrium structures reflects moder-
ate to high rates of deposition. In general, the assemblage shows
dominance of trace fossils made by deposit-feeding organisms (except
for the Atrina structures). Similar bivalve equilibrium trace fossils in
other deltaic successions worldwide have been occasionally referred
to the ichnogenus Siphonichnus. Siphonichnus comprises vertical
backfilled structures, with concave-downward menisci, and the
laminae that form the backfill are cut through centrally by a vertical
tube (Stanistreet et al., 1980). However, the general morphology of the
Caleta Olivia equilibrium trace fossils have a V-shaped retrusive
spreiten, without the central tube and, therefore, are closer to Sca-
lichnus than to Siphonichnus (Hanken et al., 2001). Ichnodiversity is
moderate, although bioturbation intensities are relatively high. The
reduced mud content and the inferred moderate to high depositional
rates, together with the ichnologic characteristics, indicate that this
facies was deposited in a more proximal position than facies 1 and 2.
More precisely, this facies is interpreted as formed in the transition
zone between the prodelta and the delta front.
3.2. Delta-front facies association
3.2.1. Facies 4: Sandstone with mud drapes
This facies consists mainly of decimeter-thick, fine- to medium-
grained sandstone beds with trough and planar cross-stratification
(Figs. 2 and 6). Abundant mud drapes, a few millimeters thick, are
preserved along the foresets (Fig. 6A, D–E). Individual beds are mostly
tabular, although some small channelized geometries have been
recognized. The cross-stratified sandstone shows diffuse and irregular
erosive surfaces, and commonly grades upwards into an interval with
asymmetrical current ripples. Symmetrical and near-symmetrical ripples
are also present on top of the sandstone beds. Reactivation surfaces and
mud clasts are also common (Fig. 6A–C). Bioclastic fragments (e.g.
bivalves, gastropods) commonly occur at the base of this facies. Intensity
of bioturbation is commonly low (BI 1–2), and the trace-fossil assemblage
is dominated by Macaronichnus segregatis and large Rosselia socialis in
the sandstone intervals (Fig. 6B, E–F), while Nereites missouriensis and
Protovirgularia isp. reworked the mud drapes (Fig. 6G–H). Sand dollars
in life position also occur in these mudstone levels.
 N.B. Carmona et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 273 (2009) 75–86 79

Fig. 4. Facies 2. Proximal prodelta. A–B. Cross-section view of the wavy-bedded facies. A. Heterolithic facies with Teichichnus rectus (Te) in cross section, and synaeresis cracks (sy).
B. Heterolithic interval with synaeresis cracks (sy), and soft-sediment deformation. Isolated specimens of Planolites montanus (Pl) also occur. C. Bedding-plane view of Protovirgularia
isp. (Pr), associated to synaeresis cracks (sy). Note the clearly defined outline of this specimen in contrast to the one shown in Fig. 3D. D. Bedding-plane view of Asterosoma isp. (As)
with horizontal bulbs.

Interpretation: Dominance of sandstone with cross-stratification
and ripples suggests bedload transport. The bimodal ripple cross-
stratification, as well as the presence of mud drapes, indicates
variations in flow velocity and direction (Willis et al., 1999), which is
typical of settings with strong tidal-action. Additionally, although not
exclusive of tidal settings, the presence of mud clasts also suggests
tidal influence (Dalrymple and Choi, 2007). Subordinate wave action is
indicated by the presence of symmetrical and near-symmetrical
ripples, the latter interpreted as produced by combined flows. This
facies represents deposition in a relatively high-energetic setting, with
strong tidal influence. Occurrence of biogenic structures adapted to
cope with moderate to high energy, such as the ichnogenera Macar-
onichnus and Rosselia, supports this interpretation. The presence of
deposit-feeding structures, such as Nereites missouriensis and Proto-
virgularia isp. in the mud drapes, indicates opportunistic colonization
during intervals of low-energy. This facies is interpreted as deposited
in a distal delta-front environment.
3.2.2. Facies 5: Sigmoidal cross-stratified sandstone
Facies 5 consists of large-scale (meter-thick) medium-grained,
cross-stratified sandstone with slightly erosive base (Figs. 2 and 7).
The foresets are sigmoidal and locally covered by mud drapes.
Paleocurrent measurements from cross-beds indicate a predomi-
nantly westward direction, suggesting dominance of flood-currents.
Thickness of individual cross-strata is variable. Bioturbation intensi-
ties are low (0–1), and the only recognized biogenic structures are
Macaronichnus segregatis and isolated Rosselia socialis specimens.
Interpretation: Lower abundance of mud drapes in facies 5 most
likely reflects stronger currents than those acting during deposition of
facies 4. The low-diversity trace-fossil suite, with predominance of
deposit-feeding structures and absence of suspension-feeding trace
fossils, reflects strong heightened water turbidity conditions (Buatois
and López-Angriman, 1992; Gingras et al., 1998). Additionally,
relatively strong currents and salinity stress may have also affected
the infaunal suites. This facies is interpreted as formed in a proximal
delta-front setting.
3.3. Depositional environment
3.3.1. Evidence of tidal influence
Several lines of evidence suggest that the studied succession has
been deposited under strong tidal influence. (1) Oppositely dipping
ripple cross-lamination and dune cross-stratification are common
throughout the whole succession, indicating flood and ebb flows
(Klein, 1977; Willis, 2005). (2) There is an abundance of mud drapes
mantling ripple and dune foresets, reflecting fluctuations in current
velocities and mud fallout during slack water (Klein, 1977). (3) The
lower interval is strongly heterolithic and displays flaser, wavy and
lenticular bedding, indicating alternating traction sedimentation and
suspension fallout (Klein, 1977; Willis, 2005). (4) Sigmoidal contacts in
cross-bedded strata, which are typical of tidal bodies (Mutti et al.,1985;
Willis, 2005), occur in the delta-front deposits. (5) Abundant mud
intraclasts occur at several sandstone beds, and are regarded as
common in tide-influenced settings (Dalrymple and Choi, 2007).
3.3.2. Evidence of other subordinate processes
The Caleta Olivia succession is overwhelmingly dominated by tide-
generated structures. However, evidence of other subordinate
processes is present, albeit locally. Synaeresis cracks are thought to
be formed due to freshwater input, and reveal the participation of
river processes. Similar cracks were experimentally produced by
varying the concentration of salt in the fluid (Foster et al., 1955; Weiss,
80 N.B. Carmona et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 273 (2009) 75–86

Fig. 5. Facies 3. Transition zone. A. Cross-bedding view of the interbedded muddy sandstone, with Phycosiphon incertum (Ph) and Nereites missouriensis (Ne) specimens. B–C.
Heterolithic interval containing abundant equilibrium structures (eq) of Atrina. Planolites (Pl) and Nereites missouriensis (Ne) also occurs in this facies. D. Cross-section view of the
heterolithic interval, specimens of Teichichnus rectus (Te) and Nereites missouriensis (Ne). E. Bedding-plane view of facies 3, with Nereites missouriensis (Ne) specimens.

1958, Donovan and Foster, 1972). Soft-sediment deformation struc-
tures suggest high rates of sedimentation in relatively steep, unstable
slopes. Although large-scale deformation features are more typical of
river-dominated deltas (e.g. Bhattacharya and Davies, 2004), smaller-
scale structures, such as those in the Caleta Olivia succession, may also
occur in strongly tide-influenced deltaic settings. Evidence of wave
influence is restricted to wave and combined-flow ripples. Hummocky
cross-stratification is notably absent.
3.3.3. Evidence of deltaic deposition
Although tidal influence can be established on the basis of physical
sedimentary structures and associated facies, identifying if tidal facies
were formed in a delta, estuary or open-marine setting is not
straightforward because these facies may occur in more than one
depositional environment. Careful documentation of stratal stacking
pattern are required also (Willis and Gabel, 2001; Dalrymple et al.,
2003; Dalrymple and Choi, 2007). In the studied progradational
succession, a tide-influenced delta is preferred over other tidal
scenarios, such as an estuary or an open seaway.
The documented ichnofauna clearly points to stressed conditions
typical of marginal-marine environments rather than fully marine
settings. Recognition of brackish-water ichnofaunas requires knowl-
edge of the open-marine expression of the basin to act as a template for
comparison (Buatois et al., 2005). In this case, coeval deposits of the
Chenque Formation formed under normal-marine conditions contain
intensely bioturbated deposits having highly diverse ichnofaunas
(Buatois et al., 2003; Carmona et al., 2008a). Accordingly, the stressed
nature of the Caleta Olivia trace-fossil suites can be firmly established.
However, stressed conditions due to dilution of normal-marine
waters can be produced in various depositional settings influenced by
freshwater discharge. The Caleta Olivia ichnofauna, although depau-
perate with respect to its fully marine counterparts in the basin, shows
some departures with respect to classic brackish-water ichnofaunas,
such as those present in estuaries or interdistributary bays (e.g.
MacEachern and Pemberton, 1994; MacEachern and Gingras, 2007).
Some common ichnotaxa in the Caleta Olivia succession (e.g. Phyco-
siphon) are unusual in more stressed brackish-water settings, but may
occur in prodelta environments, in which freshwater dilution
alternates with periods of normal-marine salinity conditions (e.g.
Buatois et al., 2008). Also, comparison with coeval estuarine deposits
in the Chenque Formation shows that the Caleta Olivia ichnofauna is
more diverse.
In addition, the stratal stacking pattern indicates that the succession
reflects deltaic progradation rather than the backstepping pattern
typically displayed by transgressive estuarine deposits (Dalrymple and
Choi, 2007). A progradational stacking pattern occurs also in estuarine
valleys due to progradation of the bay-head delta during the
subsequent highstand (Zaitlin et al., 1994). However, ichnodiversity
levels in the Caleta Olivia succession are higher than those of a bay-head
delta, which usually experiences strong salinity dilution (MacEachern
and Pemberton, 1994; MacEachern and Gingras, 2007).
3.3.4. Paleoenvironmental reconstruction
Although the studied succession is clearly progradational, it
reflects a short-term regressive event within an overall transgressive
trend within the basin (Leonense and Superpatagoniense Atlantic
transgressions; Bellosi, 1986, 1995). Tidal dominance occurs in areas
with high tidal range and low influence of waves, and is commonly
associated to transgressive intervals (Willis et al., 1999; Nummedal
et al., 2003; Willis, 2005; Ta et al., 2005; Porębski and Steel, 2006).
Amplification of tidal range may also occur in structurally controlled
basins independent of the sea-level cycle (Martinius et al., 2001;
McIlroy, 2004; McIlroy et al., 2005), but tectonically generated
confinement does not seem to be present in the case studied. Low
 N.B. Carmona et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 273 (2009) 75–86 81

Fig. 6. Facies 4. Distal delta front. A. Cross-stratified sandstone with mud drapes, dipping in opposite directions. Note also the presence of mud clasts and reactivation surfaces marked
by white dash lines. B. Interval with mud drapes, flaser and abundant mud clasts (mc). Macaronichnus segregatis (Ma) commonly reworks sandstone beds. C. Bedding-plane view of
facies 4, showing a mud-clast lag (mc) along an erosive surface. D. Dune-scale strata with mud drapes in rippled beds. E. Cross-section view of facies 4, showing an inclined specimen
of Rosselia socialis (Ro) occurring in the rippled and cross-stratified beds. F. Bedding-plane view of facies 4, with abundant sand dollars (sd) and Rosselia socialis (Ro). Lens cap is
5.5 cm. G–H. Ichnofauna associated with mud drapes. G. Nereites missouriensis (Ne). Coin is 2.4 cm. H. Epichnial preservation of Protovirgularia isp. (Pr). Coin is 1.8 cm.
82 N.B. Carmona et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 273 (2009) 75–86

Fig. 7. Facies 5. Proximal delta front. A. General view of facies 5. B. Line drawings of bedding in A. Paleocurrent measurements from cross-stratified strata indicate dominance of flood
currents (black arrow).

wave influence and high tidal range were most likely present in the
low-gradient platform San Jorge Basin during deposition of the
Chenque Formation (Bellosi, 1986, 1995).
In modern tide-influenced deltas, tidal channels and adjacent tidal
flats are common (Coleman and Wright, 1975). Interestingly, in the
studied succession we only recognized delta-front and prodelta
deposits, with no representation of delta-top tidal facies or evidence
of subaerial exposures (Fig. 8). The delta-top facies generally have low-
preservation potential and are normally eroded during transgressive
ravinement (Bhattacharya and Willis, 2001), and accordingly, very few
ancient examples of these more proximal deposits have been recorded
(McIlroy et al., 2005; McIlroy, 2007). Alternatively, Willis et al. (1999)
explained that absence of tidal-flat facies and tidal channels may
reflect the fact that the tide-influenced delta-front deposits can be
generated several kilometers basinward of the subaerial deposits,
particularly in low-accommodation settings (see McIlroy, 2007). In the
present example, absence of lag deposits produced by ravinement or
other evidence of erosion at the top of the delta front, most likely
suggests that the studied deposits were formed seaward from the
delta-top tidal flats and channels.

Fig. 8. A. Correlation scheme of the sections measured along the Atlantic coast in Caleta Olivia city, with facies distribution. Sections are oriented mostly parallel to the inferred
paleocoast. Facies stacking pattern displays a coarsening-upward trend. B. Paleocurrent diagrams from facies 1, 2 and 4, showing bipolar currents both in the prodelta and in the
delta-front facies. C. Correlation panel with less vertical exaggeration, showing the dominantly tabular geometries that characterize these deposits.
 N.B. Carmona et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 273 (2009) 75–86
The Caleta Olivia deposits are different from those of the Fly River
delta recently described by Dalrymple et al. (2003). The Fly River delta
deposits are mud-dominated and, while having a heterolithic delta
front, the prodelta is essentially muddy. In contrast, the Caleta Olivia
succession contains a larger proportion of sand.
The Caleta Olivia succession is similar in scale and sedimentologic
characteristics to that of Eocene tidal bodies from the Baronia
Formation of Spain documented by Mutti et al. (1985). The Baronia
tidal bodies form coarsening-upward successions that were attrib-
uted to progradation from bottomsets to bar-slope and bar-crest
facies. The Baronia tidal deposits have resulted in a tidal-bar model
that became popular during the nineties (e.g. Dalrymple, 1992;
Johnson and Baldwin, 1996). Although Mutti et al. (1985) regarded
these deposits as formed in the distal portion of an estuary or a tide-
dominated delta, this interpretation has been disputed by other
authors. Dalrymple et al. (2003), in their study of the tide-dominated
Fly River delta, noted that deltaic tidal bars generate lateral-accreted
deposits instead of forward-accretion deposits. These authors
suggested that the Baronia deposits are most likely compound
dunes rather than tidal bars. Willis (2005) suggested that the Baronia
deposits may represent tidal bars of a prograding delta or dunes in
waters 20–25 m deep of a shallow shelf. Dalrymple and Choi (2007)
revised the diagnostic sedimentologic features of tide-dominated
environments, and concluded that the Baronia sandbodies are more
likely the deposits of compound dunes or deltas. Preliminary results
of a study conducted by Olariu et al. (2008) suggest that the Baronia
sandbodies were formed by subtidal compound dunes within a strait
or a seaway. The Baronia ichnofauna seems to be diverse and
illustrates an archetypal Cruziana ichnofacies, which is consistent
with an open-marine setting. In contrast, the Caleta Olivia deposits
show evidence of stress conditions, as indicated by their more
restricted and depauperate ichnofauna, suggesting that this tidal
sandbody was not emplaced under fully marine conditions (i.e. open
sea), but within a deltaic system.
4. Organism responses to environmental stresses
Recently, MacEachern et al. (2005) analyzed in detail the most
important controls on the distribution of trace fossils in deltaic
systems. Because tide-influenced deltas are still understudied, the
main paleoenvironmental stresses acting on their infaunas are poorly
known (McIlroy, 2004; Brandsæter et al., 2005; MacEachern et al.,
2005; McIlroy et al., 2005; McIlroy, 2007). Consequently, in the
following paragraphs we discuss the role played by each environ-
mental parameter on the development of the infaunal communities in
this Miocene tide-influenced delta.
4.1. Salinity
Reduced salinity can be inferred from the general low diversity of the
ichnofauna, particularly in more proximal settings. The presence of
echinoid sand dollars in these proximal positions, which are usually
considered stenohaline organisms, seems to contradict this view.
However, modern clypeasteroids (e.g. Mellita) are known from brackish
habitats, with salinities as low as 20‰ (Smith, 1984). Furthermore, sand
dollars have also been documented by Kuehl et al. (2005) in the foreset
beds of the tide-dominated Ganges-Brahmaputra delta. Interestingly,
McIlroy (2007) found biogenic structures attributed to echinoderms (e.g.
Asteriacites, Scolicia) in tide-dominated deltaic deposits of the Jurassic
Lajas Formation, suggesting tolerance of these organisms to lowered
salinities. West and Ward (1990), and Mángano et al. (1999) also
mentioned the occurrence of ophiuroid trace fossils in lowered salinity
environments. In prodelta deposits, fluctuations in salinity are
evidenced from the alternation between moderately-burrowed and
unburrowed intervals. The presence of synaeresis cracks in these
deposits also suggests fluctuations in salinity (MacEachern and
83
Pemberton, 1994). However, significant size reductions in the ichnotaxa
were not observed, except for a few intervals of the distal-prodelta facies
(e.g. Planolites).
4.2. Sedimentation rates
Sedimentation was moderate to high, and certainly played a major
role controlling the distribution of trace fossils. This seems to have
been particularly important in the more proximal settings, where
biogenic structures show equilibrium strategies (e.g. trace fossils of
the bivalve Atrina, Fig. 5B–C). High sedimentation rates were also
observed by Ta et al. (2005) for the tide-dominated interval of the
Holocene Mekong River Delta. These authors suggested that during
the evolution of the Mekong River delta, the tide-dominated interval
recorded higher accumulation rates (40 m/kyr) than in the subsequent
mixed wave- and tide-dominated period (4.7–7.1 m/kyr). Additionally,
the maximum accumulation rate (39 mm/yr) during the tide-
dominated period was recorded in the delta-front slope (Ta et al.,
2005). This agrees with our observations that the highest sedimenta-
tion rates occurred in the transition zone between the delta-front and
the prodelta.
4.3. Hydrodynamic energy and water turbidity
Moderate energy most likely affected the more proximal settings,
especially the proximal and distal delta fronts, with large Rosselia
socialis and Macaronichnus segregatis as dominant elements of these
suites (Fig. 6B, E–F). Interestingly, there are no elements typical of the
Skolithos ichnofacies in the proximal settings, probably indicating that
water turbidity precluded suspension-feeding behavior. Particularly in
the delta-front settings the water turbidity is high, and this commonly
reduces ichnodiversity and bioturbation intensities, as well as
produces restriction of suspension- and filter-feeder organisms
(Buatois and López-Angriman, 1992; Gingras et al., 1998). These
characteristics (e.g. impoverishment of Skolithos ichnofacies struc-
tures, dominance of elements of the Cruziana ichnofacies) are
considered typical of deltaic conditions (e.g. Moslow and Pemberton,
1988; Gingras et al., 1998; MacEachern et al., 2005).
4.4. Hyperpycnal flows
These flows are produced by direct fluvial discharges, and are
generally related to river floods (Mulder and Syvitski, 1995).
Although hyperpycnal conditions are inferred to be common in
deltaic deposits (MacEachern et al., 2005), there is no evidence of
these flows in the studied sections. The seaward distance with
freshwater influence directly depends on the intensity of tidal-
mixing and the amount of river discharge (Willis, 2005). In
particular, tides can modify depositional patterns imposed by the
river, by modulating flow of channel-mouth plumes, changing the
rates of river- and basin-water mixing, and grain sorting within and
away from distributaries, and can also rework sediments between
river floods (Willis, 2005). Therefore, if there were hyperpycnal
discharges in the analyzed succession, their deposits would have
been probably completely reworked by the tidal action. Additionally,
there is no evidence of phytodetrital content in the analyzed interval.
Absence of phytodetritus seems to be reasonable with our inter-
pretation that these deposits formed in the seaward end of the tide-
influenced delta, far from any direct influence of the river, as it is
suggested by the absence of tidal channels and mud flats in the
upper part of the succession.
4.5. Fluid muds
McIlroy (2004), and MacEachern et al. (2005) observed that in
tide-influenced environments, clay flocculation and fluid muds
84 N.B. Carmona et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 273 (2009) 75–86
constitute two important factors that affect infaunal communities.
In particular, these mud-prone settings are characterized by high
depositional rates and important accumulations of flocculated fluid
mud, generating soupground conditions (MacEachern et al., 2005).
These soupgrouds are commonly only colonized by mobile deposit-
feeders, excluding large endobenthic deposit-feeder and suspen-
sion-feeder organisms. Interestingly, in the prodelta facies (facies 1
and 2) and in the mud drapes observed in the delta-front facies
(facies 4), structures interpreted as the locomotion of deposit-
feeder bivalves (Protovirgularia isp.) are common (Figs. 3D, 4C
and 6H). Furthermore, the morphologic analysis of Protovirgularia
isp. reveals important differences in substrate consistency in each
of these facies (Carmona et al., 2008b). For instance, those
specimens of Protovirgularia isp. in muddier beds (e.g. in facies 1)
show irregular and poorly defined chevrons. This type of sediment
allows bivalves to penetrate easily, but offers poor anchorage for
the foot (Trueman et al., 1966; Mángano et al., 1998), resulting in a
highly deformed trace-fossil morphology where the substrate is
too fluid. Conversely, the specimens of Protovirgularia isp. which
occur in relatively coherent, sandier beds (e.g. facies 4) show sharp,
closely-spaced chevrons, revealing that the tracemaker experi-
enced some friction while advancing through this more resistant
sediment.
4.6. Oxygen
Oxygen conditions do not seem to have been very restricted in the
studied succession, mainly in the proximal facies. Trace fossils typical
of low-oxygenated settings (e.g. Chondrites) are not common.
Additionally, most of the biogenic structures display normal sizes.
However, the ichnofauna is impoverished and it does not show
important penetration depths, particularly in the distal facies. Low
diversity and shallow tiers are probably more related to deposition of
fluid muds and generation of soupgrounds than to a decline in oxygen
content, although generation of fluid muds may also lead to temporal
dysoxic or anoxic conditions, so both factors may be interdependent
(Wignall and Pickering, 1993; MacEachern et al., 2005). In any case,
the redox boundary layer most likely was relatively close to the
sediment surface.
5. Comparisons with other deltaic ichnofaunas
There are few ichnologic studies of tide-dominated deltas that
serve for comparison. We restrict this discussion to prodelta and
delta-front deposits, which are the ones represented by the Caleta
Olivia succession. McIlroy (2004) documented the ichnology of the
Jurassic Ile Formation of Norway. This formation is interpreted as a
tide-dominated delta with strong riverine influence, deposited in a
microtidal setting. He noted overall lower ichnodiversities than in
wave-dominated deltas and poor development of the Skolithos
ichnofacies. The prodelta deposits contain a moderately diverse
trace-fossil assemblage, including Palaeophycus, Phoebichnus, Pla-
nolites, Rhizocorallium, Chondrites, Phycosiphon, Teichichnus, Taeni-
dium, Thalassinoides, and Schaubylindrichnus. The delta-front
deposits show the highest ichnodiversity and intensities of biotur-
bation in the Ile Formation. Skolithos, Ophiomorpha, Diplocraterion,
Arenicolites, Rosselia, Asterosoma, Lockeia, Teichichnus, Chondrites,
Trichichnus, Planolites, Palaeophycus, Gyrochorte, and Taenidium are
present in delta-front mouth bars. Collectively both ichnofaunas
illustrate an archetypal Cruziana ichnofacies. Ichnodiversity is higher
than in the Caleta Olivia succession. This may be due to the fact that
the succession is microtidal and higher-energy large-scale bedforms,
such as those that characterize settings with high tidal range, may
not have been formed. Colonization of large tidal bedforms is
commonly inhibited or only possible during short-term windows
(Pollard et al., 1993).
McIlroy (2007) documented the ichnology of a tide-dominated
succession in the Jurassic Lajas Formation of Argentina. The prodelta
deposits are intensely bioturbated, and contain Teichichnus,
Schaubcylindrichnus, Chondrites, Rhizocorallium, and Parahaentzche-
linia, illustrating an impoverished Cruziana ichnofacies. The delta-
front mouth-bar deposits show the highest ichnodiversity, including
deposit-feeding/infaunal predator trace fossils (e.g. Asterosoma,
Helminthorhaphe, Macaronichnus, Palaeophycus, Planolites, Rosselia,
Schaubcylindrichnus, Teichichnus), suspension-feeding trace fossils
(e.g. Diplocraterion, Parahaentzschelinia, Siphonichnus), and biogenic
structures assigned to facultative suspension or deposit feeders (e.g.
Ophiomorpha, and Thalassinoides). Delta-front ichnofaunas illustrate
an archetypal Cruziana ichnofacies. McIlroy (2007) considered that
the observed increase in ichnodiversity from the prodelta to the
delta front may be an artifact of preservational potential, although he
also noted that the combination of episodically strong currents with
periods of slack-water may have provided optimal conditions for
both suspension- and deposit-feeder organisms in the delta-front
setting. Although the Lajas and Caleta Olivia successions seem to
have been deposited in a similar setting, overall ichnodiversity is
higher in the former. This may be linked to the fact that the salinity
stress affected mainly the ebb-tidal channel deposits (McIlroy,
2007), and not the whole succession, as revealed by the analysis of
the Caleta Olivia deposits.
Also, a number of sedimentologic studies have documented the
facies and stratal stacking pattern of tide-influenced deltas (e.g.
Willis et al., 1999; McIlroy et al., 1999; Bhattacharya and Willis, 2001;
Martinius et al., 2001; McIlroy 2004; Brandsæter et al., 2005;
McIlroy et al., 2005), and their ichnologic content has been
subsequently addressed by McIlroy (2004), MacEachern et al.
(2005). Willis et al. (1999), and Bhattacharya and Willis (2001)
discussed tide-dominated deltaic deposits of the Cretaceous Frontier
Formation of Wyoming (see also review in MacEachern et al., 2005).
Prodelta deposits of the Frontier Formation are unburrowed to
weakly bioturbated and trace fossils are sporadically distributed.
The prodelta ichnofauna consists of Planolites, Piscichnus, Teichich-
nus, Thalassinoides, Chondrites, “Terebellina”, escape trace fossils,
and bivalve escape/adjustement structures, illustrating an extre-
mely impoverished Cruziana ichnofacies. Delta-front deposits are
unburrowed or very weakly bioturbated and ichnofossils are
extremely sporadically distributed. The delta-front ichnofauna
consists of Arenicolites, Ophiomorpha, “Terebellina”, Thalassinoides,
Planolites, Cylindrichnus, Diplocraterion, Lockeia, Zoophycos, Macar-
onichnus, Palaeophycus, Piscichnus, Skolithos, escape trace fossils,
and bivalve escape/adjustement structures, illustrating the Cruziana
ichnofacies. Size reduction was noted and most of the bioturbated
zones are associated with pauses in deposition (MacEachern et al.,
2005).
Martinius et al. (2001) analyzed tide-dominated deltaic deposits of
the Jurassic Tilje Formation of Norway (see also review in MacEachern
et al., 2005). Prodelta deposits of this unit are unburrowed or show
low degrees of bioturbation. They contain Planolites, Teichichnus, Di-
plocraterion, Thalassinoides, Bergaueria, Siphonichnus, Palaeophycus,
Skolithos, Rhizocorallium, Cylindrichnus, Chondrites, Taenidium, Phyco-
siphon, and escape trace fossils illustrating an archetypal Cruziana
ichnofacies. Delta-front deposits are unburrowed to sparsely biotur-
bated and contain similar ichnogenera, with the addition of Gyro-
chorte. The delta-front ichnofauna represents a proximal expression of
the Cruziana ichnofacies. Although overall ichnodiversity is moderate,
trace fossils are sporadically distributed and commonly concentrated
at pause planes. Fluid muds and freshets seem to have been the most
important stress factor in the Tilje Formation (Martinius et al., 2001;
MacEachern et al., 2005).
Although it is premature to propose some general ichnologic
model for tide-influenced deltas, several common features are
emerging. First, tide-influenced delta fronts and prodeltas seem to
 N.B. Carmona et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 273 (2009) 75–86
be more stressed than wave-dominated ones but less stressed than
river-dominated ones. Accordingly, tide-influenced delta-front
and prodelta deposits tend to be more sparsely bioturbated, and
ichnofaunas are less diverse than in wave-dominated deltas.
Intensity of bioturbation and ichnodiversity is typically higher
than in river-dominated prodeltas and delta fronts. However, high
ichnodiversity and intense bioturbation have been noted in some
tide-influenced deposits (e.g. McIlroy, 2004, 2007). Also, ichnodi-
versity levels of tide-influenced deltas seem to be higher than in
tide-dominated estuaries (McIlroy, 2007). Second, bioturbation is
commonly associated with bedform mantling, suggesting coloniza-
tion along pause planes (e.g. MacEachern et al., 2005). Third, bivalve
equilibrium trace fossils seem to be common, reflecting vertical
accretion of the sea floor. Fourth, suppression of suspension-feeding
elements is apparently quite common as a response to high content
clay in the water column. An expanding dataset is needed in order to
further evaluate how tide-influenced deltaic ichnofaunas are shaped
by the relative influence of the different stress factors.
6. Conclusions
The lower Miocene succession outcropping along the Atlantic coast
of Caleta Olivia city was deposited in a tide-influenced delta
environment, and consists of two main facies associations: delta-
front and prodelta facies. The ichnologic analysis reveals that the
trace-fossil suites developed in these environments show low
diversities and low to moderate abundance of trace fossils, poor
development of tiering structure, and sporadic distribution.
In particular, the suites developed in the delta-front settings show
reduced ichnodiversity, dominance of deposit- and detritus-feeding
structures (mainly those considered to be tolerant to energetic
conditions), and suppression of suspension-feeding elements. Dom-
inance of structures typical of the Cruziana ichnofacies, and impover-
ishment of elements attributable to the Skolithos ichnofacies are
considered typical of deltaic environments, and probably reflect major
stressful conditions, generated not only by freshwater input, but also
by tidal action (e.g. fluctuation in sedimentation rates and in current
velocities, flocculated muds, and water turbidity).
On the other hand, the trace-fossil suites developed in the prodelta
settings display low to moderate diversity, dominance of deposit-
feeding structures (mainly facies-crossing forms), and sporadic
distribution. These assemblages reflect an impoverished expression
of the Cruziana ichnofacies. Most likely, occurrence of fluid muds
(with the subsequent reduction in oxygenation), as well as salinity
fluctuations, commonly precludes development of diverse commu-
nities in this tide-influenced prodelta environment.
Although there is still a poor understanding of the organism
responses to tidal action in deltaic settings, this study shows that tidal
dynamics generate distinctive effects on the infauna. These effects
relate not only to fluctuations in energy and water salinity, but also
(and probably more significant), to changes in substrate consistency,
water turbidity, and sedimentation rates.
Acknowledgments
We thank Eduardo Olivero, Jeremías González, and M. Isabel
López-Cabrera for discussion in the field. Alvar Sobral is thanked for
preparation of polished samples. Duncan McIlroy and an anonymous
reviewer provided useful criticism. Financial support for this study
was provided by a Post-Doctoral Grant from the Argentinean Research
Council (CONICET), an International Association of Sedimentologists
Postgraduate Grant Scheme, and PICT 840 awarded to Carmona.
Additional funds were provided by the Canadian Natural Sciences and
Engineering Research Council (NSERC) Discovery Grants 311727-05
and 08 and 311726-05 and 08 awarded to Mángano and Buatois,
respectively.
85
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