Marine Geology 352 (2014) 367–380

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Marine Geology
journal homepage: www.elsevier.com/locate/margeo

Modern microbial mats in siliciclastic tidal flats: Evolution, structure and

the role of hydrodynamics
Diana G. Cuadrado a,b,⁎, Gerardo M.E. Perillo a,b, Alejandro J. Vitale a,c
a
CONICET-IADO, Florida 5000, 8000 Bahía Blanca, Buenos Aires, Argentina
b
UNS, Dto. Geología, San Juan 670, 8000 Bahía Blanca, Buenos Aires, Argentina
c
UNS, Dto. Ingeniería Eléctrica y Computadoras, Avda. Alem 1253, 8000 Bahía Blanca, Buenos Aires, Argentina

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

Article history: Microbial mats in coastal environments play an important role in both stabilizing the sediment and preserving
Received 18 June 2013 sedimentary structures. They represent the first biological systems that appeared on Earth, even forming large
Received in revised form 1 October 2013 fossils in carbonate settings known as stromatolites. However, microbial mats have also been recognized for
Accepted 7 October 2013
siliciclastic environments as sandy mat structures. In particular, mats are the primary element of the so-called
Available online 16 October 2013
Microbially-Induced Sedimentary Structures (MISS) and their various types represent very useful proxies for
Keywords:
the interpretation of the environmental conditions in which they were formed.
Sedimentary structures The objectives of this paper are twofold. The first is to provide a general background review of microbial mats in
MISS tidal flats, including their evolution, structure, and morphology. The second objective is to provide a modern case
Wave shear stress study in which we examine the role of storms relative to day-to-day tidal currents. Although tidal currents could
Tidal currents be an important eroding factor, we demonstrate that waves, no matter their magnitude, play a significant role in
Tidal flat biostabilization developing different types of MISS such as wrinkle structures and eroding mats into flipped-over fragments.
Furthermore, we also show the significance of wave-induced pressure onto groundwater in forming other
types of MISS as Kinneya structures and petee ridges. Knowing the environmental conditions to which present-
day microbial mats are subject and how they evolve upon drastic changes in those conditions (i.e., storm events),
represent an important basis for the interpretation of fossil MISS.
© 2013 Elsevier B.V. All rights reserved.

1. Introduction of ancient sedimentary rocks which requires the understanding of

processes that resulted in a particular sedimentary stratum. A very
Geobiology has recently been defined by Knoll et al. (2012) as useful proxy is the study of processes on modern settings as analogues
the scientific discipline in which the principles and tools of biology of ancient environments. Although the processes are similar (not
are applied to studies of the Earth. These parent specialties and identical), they often provide useful insights for the interpretation of
their subdisciplines gather together to produce new discoveries that the processes that occurred in the past.
contribute to a better comprehension of Earth in modern and past A major bridge that joins the past with the present is the world of
times. Moreover, the search of life beyond the Earth (i.e. astrobiology) microbiota such as cyanobacteria, which they are the only oxygenic
is extending and, in that sense, geobiological approaches provide phototrophs known to have existed from before 2 Ga (Knoll et al.,
the basis for studies that focus on the exploration for extraterrestrial 2012) up to the present. In most environments, sediments are densely
life on other planets (Baumgartner et al., 2009; Cady and Noffke, 2009 populated by benthic microorganisms that form biofilms around
and references there-in). As a consequence, a large number of inter- individual sediment grains or develop thick, carpet-like layers termed
disciplinary studies are focused on the documentation and under- microbial mats (i.e., Fig. 1; some examples given in this article are
standing of the evolution of Earth. taken from sites in Argentina, Fig. 1a). Probably, the major forms of
Geobiogists are interested in knowing the ways in which the life on the early Earth, prior to the evolution of algae and higher plants,
organisms affect the Earth and the interactions between biological and were photosynthetic microbial mats. In that sense, Schopf (1996)
physical processes in order to infer their participation in the planet's states that the Precambrian biosphere belonged almost exclusively
history. This interaction is useful for geologists in the interpretation to microbiota due to a different atmospheric composition present in
those periods. Later, when plants and animals evolved and, due to a
rapid growth of grazing metazoan communities (Grotzinger, 1990),
extensive microbial mats became rarer. However, they are still present
⁎ Corresponding author at: CONICET-IADO, Florida 5000, 8000 Bahía Blanca, Buenos in many ecosystems and have a significant impact on global biogeo-
Aires, Argentina. chemical cycles (Seckbach and Oren, 2010). Several studies revealed

0025-3227/$ – see front matter © 2013 Elsevier B.V. All rights reserved.
http://dx.doi.org/10.1016/j.margeo.2013.10.002
368 D.G. Cuadrado et al. / Marine Geology 352 (2014) 367–380

Fig. 1. (a) Maps of the SW portion of the Buenos Aires Province (Argentina) with locations mentioned in the text. Oceanographic Tower (OT) is shown at the mouth of the Bahia Blanca
Estuary. (b)Thick coherent and leathery microbial mat present in a sabkha environment, Paso Seco near San Blas, where the main physical process is wind. The mat was ripped off and
removed, exposing the underlying substrate in wide openings. Note the detached and deformed piece of flexible mat making a fold (mat deformation structures, MDS sensu Bouougri
and Porada, 2012). (c) Initial stage of microbial biofilm according to Stoodley et al. (2002) in Puerto Rosales tidal flat. (The photograph was taken on July, 2006). (d) Biofilm formed in
the lower intertidal zone within a tidal creek in Rio Negro Estuary. Foreground width is 4 m. (e) Epibenthic microbial mat composed of three main bacterial groups differentiated by
multicolored layers. Cyanobacteria at the top, sulfate-reduction bacteria (black layer) and purple sulfur bacteria below. The photo was taken at Puerto Cuatreros, inner Bahía Blanca
Estuary. (f) Bird tracks in the epibenthic mat (formed mainly by M. chthonoplastes) in Puerto Rosales.

the earliest direct evidence for terrestrial microbial mats affecting grazing animals (Pflüeger and Gresse, 1996). Thus, they may be found
siliciclastic sediments in sedimentary rocks of South Africa (Eriksson in peritidal environments such as coastal lagoons (Javor and Castenholz,
et al., 2000; Watanabe et al., 2000; Noffke et al., 2003). Other studies 1981) as well as in extreme environments with high temperatures and
reported microbial mat proliferation during the Mesoproterozoic hypersaline conditions, where competition by higher organisms is absent
(Seilacher, 1999; Eriksson et al., 2000; Altermann, 2004; Schopf, 2004). and grazers are excluded (Seckbach and Oren, 2010).
Today, extensive microbial mats only exist in special places such There is a growing interest in the importance that microbial mats
as tidal flats, where they have little competition from vascular plants or play in coastal environments with regards to their role in stabilizing
 D.G. Cuadrado et al. / Marine Geology 352 (2014) 367–380 369

the sediment, and preserving and being part of the formation of discovered in Shark Bay (a hypersaline environment) in Western
sedimentary structures. The objectives of this paper are thus twofold. Australia (Logan, 1961; Davis, 1970; Hoffman, 1976) and, more recently,
The first is to provide a review of microbial mats in coastal wetlands along the margins of Exuma Sound (in open marine conditions) in the
with special reference to tidal flats, including their evolution, structure, Bahamas (Dravis, 1983; Dill et al., 1986; Reid and Browne, 1991; Reid
and morphology. The second objective is to provide a modern case et al., 1995). Subtidal stromatolites were studied at Highborne Cay,
study in which we examine the role of storms relative to day-to-day Bahamas (Reid et al., 2000; Decho et al., 2005).
tidal currents. Stromatolites provide valuable information on the micro-
biogeochemical processes interaction during their formation and
2. Microbial mats the precipitation of CaCO3 (Stolz, 2000). Visscher and Stolz (2005),
in contrast to previous studies, stated that processes such as photo-
In a recent article, Gerdes (2010) listed several definitions of synthesis and sulfate reduction result in net precipitation, while aerobic
microbial mats, opportunely stated by different authors (Krumbein, respiration and sulfide oxidation produce net dissolution of the
1983; Cohen, 1989; Wachendörfer et al., 1994; Decho, 2000a; Stolz, associated carbonates. These authors also describe the role of microbial
2000; Krumbein et al., 2003). The author suggested a definition of mats as bioreactors producing a characteristic biosignature. However, it
microbial mats as “advanced biofilm stages forming laminae on bedding is important to remark that the main difference between stromatolites
surfaces where they reflect gaps in sedimentation or, in other terms, time and non-carbonate mats is the precipitation of CaCO3 by microbes to
for growth, biomass condensation and biological succession”. form stromatolites, which does not happen in siliciclastic mats, which
In turn, Stoodley et al. (1999, 2002) explained the formation of is the focus of this paper.
microbial mats from the development of a biofilm around individual
sand grains, to its expansion to adjacent grains ultimately forming
4. Microbially-Induced Sedimentary Structures (MISS)
a laterally-continuous organic layer (Fig. 1c). In favorable (nutrient
enriched) conditions well adapted benthic consortia forms complex,
The classification of primary physical sedimentary structures made
multilayered microbial ecosystems in salt marshes and estuaries
by Pettijohn and Potter (1964) includes various types of bedding,
(Krumbein et al., 1977; Stal et al., 1985; Nicholson et al., 1987; Stal,
markings on bedding surfaces and deformational phenomena. They
2010) that may extend their distribution even into the lower intertidal
also involved stromatolites as positive growth structures projecting
zone (Fig. 1d).
from former depositional surfaces, reflecting the fact that the microbes
The initial step in the formation of a microbial mat involves the
construct the structure in conjunction with synsedimentary cementation.
secretion of an exopolymer to allow the microorganism to adhere to
However, Noffke et al. (2001) recognized that there are other
the substratum (Yallop et al., 1994; Wetherbee et al., 1998; de Winder
microbially-mediated structures in siliciclastic or sandy deposits that
et al., 1999; Decho, 2000b). Both, cohesive and non-cohesive sediments
“result from epibenthic cyanobacteria interacting with the physical agents
are commonly embedded in this thick mucilaginous matrix of extra-
of erosion, deposition, transportation or deformation” which these latter
cellular polymeric substances (EPS), which may consist of 90% or
authors named MISS (Microbially-Induced Sedimentary Structures).
more of polysaccharides (Hoagland et al., 1993), forming sticky coatings
Because these structures differ in appearance and origin from other
on individual sediment particles. The rheology and stability of the
physical sedimentary structures, MISS were proposed to have their
sediment bed is thus influenced by EPS (Paterson, 1989, 1997; Dade
own category in the existing classification scheme for primary
et al., 1990; Ruddy et al., 1998). Microbial cells typically present a high
sedimentary structures (Noffke et al., 2001). Thus, the scholarly
potential for resiliency during periods of physical stress (e.g., changes
definition of MISS is: “… are primary sedimentary structures that rise
in salinity and temperature, UV irradiation, desiccation, inundation)
syndepositionally from the interaction of biofilms and microbial mats
and may overall enhance their physiological activities (Decho, 2000b).
with the physical sediment dynamics in siliciclastic aquatic environments”
This resiliency can also be extended to direct physical processes such
(Noffke et al., 2001), being the siliciclastic counterparts to stromatolites
as water flow and wave activity.
that occur in carbonate settings. Although we know that cyanobacteria
All trophic groups (i.e., primary producers, consumers and
are the main prokaryotes responsible for present-day MISS, during the
decomposers) are present in microbial mats, which is of great impor-
Archean other prokaryotes of similar morphology and behavior than
tance from an ecological perspective. Their populations are organized
cyanobacteria may have also been involved in the formation of MISS
into specific communities interacting with each other and their envi-
(Noffke, 2003a, 2009).
ronment (Stolz, 2000). The large diversity of species within microbial
mats support a wide spectrum of metabolic processes (Visscher and
Stolz, 2005) resulting in coupled reactions (e.g., reduction and oxidation 4.1. Categories of MISS
of essential elements such as C, S, N; or, alternatively, stepwise oxidation
or reduction of a compound) that support robust biogeochemical cycles On the basis of modern tidal flat studies (Noffke et al., 1997; Noffke,
(Fig. 1e). The result can be the formation of trace gases and mineral 1998, 1999, 2010; Noffke and Krumbein, 1999) defined 17 individual
precipitates. MISS morphologies ranging in size from m to mm scale (Fig. 2).
Noffke (2009) distinguished 5 categories of MISS based on the microbial
3. Microbial mats in carbonate environments activity that develops the mats: structures induced by biostabilization,
baffling, growth, trapping and binding, as well as structures that are
For a long time, the microbial influence on sediments was directly induced by the interference of those parameters.
related to biogeochemical processes that enhance precipitation of The stabilization of the sediment is the effect caused by the
carbonate from seawater causing stromatolites. They are mainly microbiota to sandy substrata in shallow-marine settings, evidencing
formed by cyanobacteria and associated heterotrophic bacteria, which a response to physical dynamics caused by waves and currents.
developed widespread microbial reefs throughout shallow waters This response is due to the highly mobile organisms that can move
of Precambrian oceans (Awramik, 1992). They can be easily preserved actively through the grains (Golubic and Knoll, 1999). Filamentous
so they are abundant in the fossil record and they represent the earliest microorganisms, such as cyanobacteria, entangle the mineral particles
macroscopic evidence of life forms (Schopf, 1996). Thus, numerous like an organic meshwork and the EPS glue the sediment particles
studies have been made on these rigid stacks of layers produced by together resulting in the biostabilization (sensu Paterson et al., 1994)
microbial organisms in carbonate settings (Awramik, 1992; Hoffman, of the sediments. On the other hand, Noffke (2010) has proposed that
2000; summary in Altermann, 2004). Also, modern stromatolites were the filaments (trichomes) of cyanobacteria are oriented horizontally
370 D.G. Cuadrado et al. / Marine Geology 352 (2014) 367–380

Fig. 2. Classification of microbially induced sedimentary structures extracted from Noffke, 2010 (with permission from Springer-Verlag). The diagram shows the formational relationship
of all MISS. The structures are induced by growth, biostabilization, baffling, trapping, or binding. The center of the diagram includes MISS caused by the interference of all biotic–physical
interactions.

within the grains. The presence of EPS affects the chemical structure of amount of sediment input into the system. In the same way, tides and,
the filaments resulting in a layer that is more resistant to erosion. specifically, tidal currents are dominant along the tidal channels and
Baffling occurs during low energetic periods when cyanobacteria at the lower intertidal zones. Normally, in the upper intertidal zone,
(mainly the filamentous Microcoleus chthonoplastes) move up the currents tend to be weaker and they are more prone to deposit any
sediment in order to reach optimal light conditions (phototaxis), suspended sediment they may carry up to the edge of tidal inundation.
orienting their filaments vertically. The vertical filaments act to slow Depending on their location, either within restricted embayments
sediment transport and so, suspended silty sediments deposit and (i.e., enclosed estuaries, coastal lagoons) or directly connected to the
mats become enriched in silt-sized particles. Growth of the microbial open sea (e.g., Wadden Sea, Hangzhou Bay) the circulation over the
mat is due to the biomass enrichment by means of cell replication, flats is dominated by tidal currents alone or the interaction between
which in turn results from favorable sunlight, temperature, humidity waves and currents (Piccolo et al., 1993). Wave action may also be
and available nutrients (Noffke, 2010). On the other hand, the effect important within estuaries if they have a large-enough fetch along
of the sticky EPS gluing small-sized particles to the mat surface is the predominant wind direction (i.e., Bahía Blanca and St. Lawrence
termed trapping. Binding is the formation of a mat fabric by the active estuaries, Chesapeake Bay).
movement of cyanobacteria generating a mesh-like network during Regardless of the importance of establishing circulation patterns
calm conditions. over tidal flats, there has been much less work done in that field,
In fact, the biostabilization counteracts erosion, while baffling and compared to those measurements done within tidal courses (sensu
trapping generate deposition of sediment. Binding and growth take Perillo, 2009) that are associated to coastal wetlands. Pioneering,
place during latencies, which are time periods of no- or low sediment preliminary studies were made by Postma (1967) for the Wadden Sea
reworking. Inorganic mineral precipitation does not play a role in the tidal flats. Later on, Groen (1970, 1974) described the deformation
formation of MISS but, in some cases, the biological activity produces processes affecting tidal currents as they move from the open sea
geochemical changes in the microenvironment that enhance authigenic to channels to the intertidal area and back. While the currents are
mineral precipitation. reversible in the channels, in the open ocean and on the flats, a vector
diagram for a tidal cycle normally shows a rotational pattern. However,
5. General hydrodynamics on tidal flats Piccolo et al. (1993) demonstrated after a month long data set of tidal
currents made on an extensive tidal flat in Minas Basin (Canada), that
Coastal wetlands are ecosystems found within an elevation gradient they can also present a reversing pattern. These authors showed that
that ranges between subtidal depths to which light penetrates to the local geomorphology plays a major role in defining the circulation
support photosynthesis of benthic plants and the landward edge pattern of the flat, even when, as in the case study, the flats were
where the hydrologic influence of the sea transition into groundwater covered by as much as 8 m of water. Modern studies have also included
and atmospheric processes. At the seaward margin, biofilms, benthic the influence of waves in sediment transport processes (i.e., Lee, 2004;
algae, and seagrasses are representative biotic components. At the Pratolongo et al., 2010; Shi et al., 2012).
landward margin, vegetation boundaries range from those located on The upper wetland and supratidal zones are flooded only during
groundwater seeps or fens, in humid climates, to relatively barren salt spring tides, or by water set up due to wind influence. In particular
flats in arid climates (Wolanski et al., 2009). Coastal wetlands can be when wind plays a major role in the dynamics of a wetland, waves
found in any tidal range, although their size is directly proportional to become increasingly more important in the various processes of
tidal range but inversely proportional to coastal slope, given the same erosion, transport and deposition of sediment in the system. In open
 D.G. Cuadrado et al. / Marine Geology 352 (2014) 367–380 371

coast wetlands, oceanic waves can penetrate as far as permitted by Accordingly, the predominant type of microbial mat directly affects
factors such as the flooding and geomorphology, but their energy can the biostabilization of sediments. Epibenthic microbial mats have a
be significantly reduced due to friction (correlated with distance into “slippery” surface, probably because they are EPS-rich (Noffke, 2010),
the estuary). However, local waves in areas with enough fetch can be which reduces the frictional flow forces and rises the erosional current
determinant in tidal flats although their influence is reduced on velocity to 1.60 m s−1 (in 3–5 cm water depths). Estimations of the
marshes (i.e., Moller, 2006; Koch et al., 2009; Pratolongo et al., 2010) magnitude of this erosional current velocity range from being 9 times
where vascular plants dampen waves and currents. higher than for similar grain size, uncolonized sediments (Noffke,
Therefore, in the places where microbial mats normally occur, tidal 2010), to 12 times higher (Yallop et al., 1994; Paterson, 1997; Noffke,
currents are normally low and have little effect on the mats and the 1998). Type I biostabilization (Noffke, 2010) protects the sedimentary
geomorphology. Nevertheless, contrary to that indicated by Noffke surface from erosion and can only be mechanically disrupted or
(2010), we observed that ebb currents are much more effective in destroyed by wave activity during strong storms.
modifying the sediment surface than flood ones. Ebb currents acquire Contrarily, in endobenthic microbial mats, the cyanobacteria (mainly
the highest velocities even reaching supercritical conditions due to the O. limosa) form a mesh between the sand grains and the filaments,
small depths and the large slope over which the water flows. Piccolo resulting in a rough sedimentary surface, turning the smooth laminar
et al. (1993) showed that ebb currents in tidal flat of the Minas Basin flow into a rough turbulent flow. The erosional current velocity can
where more concentrated in direction and stronger than flood currents. reach 60 cm s−1, which is also higher than in loose sterile sand
On the other hand, waves are much more effective in acting over the (Noffke, 2010). This type of biostabilization is named Type II by Noffke
plain. For instance, Pratolongo et al. (2010) demonstrated that tidal (2010). Geoforms can develop in these conditions, but they are smaller
currents on the edge of a marsh of the Bahía Blanca Estuary, which than those of non-colonized surfaces, hiding the actual hydraulic
gets inundated on every tidal cycle, were basically negligible in contrast conditions. For field measurements of the initial erosion shear stress,
to the strong effect produced by waves generated by local winds. They Noffke (2010) used a Manzenrieder flume chamber (Führböter and
even pointed out the formation of fluid mud by injection of water by Manzenrieder, 1987).
the waves during storm conditions which resulted in significant Moreover, results of flume experiments (Hagadorn and McDowell,
changes in the topography of the bordering mudflat. 2012) demonstrated that microbial mat communities growing over
Furthermore, in cores taken on tidal flats it is common to observe the sediment increase the required flow velocity to produce grain
relatively thin layers of sand between layers dominated by silt and movement with age of the mat (Fig. 3). Under such conditions, a
clay (i.e., Ginsberg and Perillo, 1990). The sandy layer corresponds to flow velocity at least ca. 50 to 100% higher is necessary to develop
a lag material when the finer sediments are removed by the action the known sequence of bedforms (ripple-dune-plane-bed lamination)
of the locally generated waves with small periods but high steepness obtained from experiments with sterile sands (Hjulstrøm, 1935;
(H/L, where H is wave height and L is wavelength) (Perillo and Shields, 1936; Southard and Boguchwal, 1990; Souslby, 1997). A sig-
Sequeira, 1989; Perillo, 2003). Contrarily, the fine sediment layer is nificant result of the experiments by Hagadorn and McDowell (2012)
the result of the deposit of suspended sediments during fair conditions. is the correlation between the age of the microbial mat and the
When present, microbial mats can play an important role in the unidirectional flow velocity (u), which combine to produce various
preservation of the different layers (i.e., biolaminites), provided they types of modification on the mats. With sterile sands, grains move
have enough time to develop (Cuadrado et al., 2013). basically with current velocities N 20 cm s−1 and, at u N30 cm s−1, the
formation of ripples has been observed even in mats a few days old
6. Hydrodynamics over microbial mats (Fig. 3). The condition of no movement reaches the required maximum
current velocity at about 30 days of age but, as mats grow older, their
One of the criteria for biogenicity of MISS (Noffke, 2009) states capacity to avoid sediment movement is considerably reduced.
that they reflect the average hydraulic pattern that governs a defined Experimentally increasing velocity further, flip-overs (see below) are
area (tidal, lagoonal or shallow paleoenvironments). In ancient shelves the next step once flow velocity (u) is higher than the threshold. The
dominated by fair weather and storm waves, the characteristic structures presence of flip-overs increases for the younger mats and with increasing
are transparent and non-transparent wrinkles formed by endobenthic velocities, but their upper boundary is more stable with older mats.
microbial mats. Similar structures can be found in lagoons, typically However, at about 30 days of age and for velocities 35 cm s−1, the flip-
thin bedded (fine sandstones separated by fine siltstone layers). overs transform into roll-ups, which does not occur for lower velocities.
However, the more complex hydraulic systems are tidal flats where Rip-ups (Fig. 3) follow the general trend of the flip-overs, but are
different structures can be found along a transect from the mean low to
high tide levels (Bose and Chafetz, 2009). This fact results in different
types of microbial mats as a function of the area of inundation, being
a primary control factor for mat growth in conjunction with local
topographic differences.
This lateral succession is called biofilm-catena and is typically
characterized by a dominance of cyanobacteria which are best adapted
to dynamic conditions (Noffke and Krumbein, 1999; Noffke, 2003b). In
temperate siliciclastic tidal flats, the lower intertidal zone is colonized
only by a biofilm. In the upper intertidal zone, which gets inundated
twice every day, the endobenthic mat is formed in just a few hours
and the main builder is the cyanobacteria Oscillatoria limosa. Single
trichomes interweave with sand particles in the upper mm of the
sedimentary surface (Fig. 1d). In contrast, the lower supratidal zone,
inundated periodically by spring tides and sporadically during storms
events, is characterized by planar epibenthic mats, arising from the Fig. 3. Correlation between age of microbial mats (in days) vs. critical velocities based on
construction of a coherent network by M. chthonoplastes filaments in laboratory experiments by Hagadorn and McDowell (2012). Bedforms and sediment
transport are produced with sterile or very young mats. As mats grow older there is an
presence of abundant EPS on top of the sedimentary surface (Fig. 1f). increase in the critical velocity which is maximum for initiation at about 30 days. Once
The extent of these types of microbial mats is different in relation to the mats are affected, they show flip-overs and with higher velocities rip-ups are observed
the tidal range (Noffke, 2010; see her Fig. II.14). (extracted from Hagadorn and McDowell, 2012).
372 D.G. Cuadrado et al. / Marine Geology 352 (2014) 367–380

associated with higher velocities. In other words, these experiments However, after performing a detailed inspection of these structures,
could provide significant information on the dynamic conditions we were able to observe an arrangement of elongated domes and
present during the evolution of sediments covered by microbial mats irregular folds among the upturned-margins cracks (Figs. 4c, 5a).
as specific forms can be well correlated to a particular range of Furthermore, some of the elongated domes showed an initial crack at
velocities. However, a drawback to this approach is that asserting the the top of the dome (Fig. 5a). This is the origin of most of the cracks
age of mats in the fossil record is not possible. which progressively shrink due to tensile shear under desiccation
Microbial mats clearly modify the erodibility of the original periods forming a polygonal pattern. The ridged dome features may
sediments. The erodibility can be defined as an erosion threshold, be associated with soft sediment deformation. The cohesive microbial
which is the bed shear stress needed to erode the surface of the bed mat forms a zone of low permeability, separating the sediment below
(Andersen and Pejrup, 2011). However, as indicated by Tolhurst et al. the mat (mainly sandy sediments, Fig. 5c–d) which is water saturated
(2000), the diversity of instruments and field vs. laboratory experiments like a “confined aquifer” (Porada et al., 2008; Fig. 6).
considered for its estimation, makes comparison of results difficult. It is important to consider mechanical aspects of tidal flat at Puerto
Andersen and Pejrup (2011) sustain another important variable, which Rosales. The water filling the pore spaces of the substratum flows to
is also complex to obtain and seldom reported in the biofilm literature: downslope under pressure. This hydraulic pressure is greatest in the
the erosion rate. As for the other parameters above-mentioned, diffi- supratidal zone where the confining mat begins and has the same
culties in providing sound measurements, based on common criteria magnitude along equipotential lines. It acts as an uplift force leading to
arise from major differences in the type of instrumentation employed the detachment of the mat (Fig. 7). The tidal oscillation superimposed
and in local field characteristics. Another way to represent the impact on the hydrostatic pressure (according to tidal period, over the intertidal
of biofilms is the stabilization index proposed by Tolhurst et al. (1999), and sometimes over the supratidal zone), increases the total pore water
which is calculated as the ratio between the erosion threshold for a pressure on the sediment. This liquefied sand moves laterally (Eriksson
colonized- and a biologically-sterilized sediment bed. Neumeier et al. et al., 2007 and references therein; Porada et al., 2007). On the other
(2006) summarized studies for mudflat sediments and found that hand, when evaporation occurs there is a vertical hydraulic gradient
the stabilization index ranged from 3 to 10 which clearly show the that replaces the evaporated water near the surface. This happens during
importance of biofilms in reducing sediment erodibility. summer or during low tide, replacing the water consumed by organisms
(Porada et al., 2007). The Bahía Blanca Estuary area is characterized by
7. Modern case study: Bahía Blanca Estuary dry and windy weather (mean precipitation is 650 mm, mean humidity
72%, Piccolo and Diez, 2004), therefore, the exposed flats have high
7.1. Tidal flats colonized by microbial mats. Study area evaporation rates (Pan et al., 2013a) resulting in a vertical water flow
from the lower sediment levels.
The microbial mats present at Puerto Rosales (middle zone of Bahía Other factors, like various endobenthic fauna (e.g., crabs, worms,
Blanca Estuary—BBE, Fig. 1a) have been studied for a few years insect larvae) may play a role in the dynamics of groundwater
now (Cuadrado and Pizani, 2007). The low gradient supratidal zone even though the original sediment lacks the adequate porosity and,
which is inundated occasionally, during spring tides or storm events, especially, permeability. For instance, Perillo et al. (2005) observed an
provide the essential physical conditions necessary to allow an initial increase on the groundwater dynamic and its outflow along a tidal
colonization and a progressive establishment of a coherent network channel margin due to high density burrows made by the burrowing
of microbial mats (Pan et al., 2013a,b). Other conditions necessary crab Neohelice granulata in the adjacent salt marsh. In the Bahía Blanca
to colonize the tidal flat are low sedimentation and fine sand to silt Estuary, this crab plays a major role in the geomorphologic evolution
sediment granulometry (Cuadrado et al., 2011). of salt marshes (Perillo and Iribarne, 2003a,b) and in the formation
At this location, the modern microbial mats preferentially grow of tidal courses. Burrows made by this crab are active in the study
in zones with low hydrodynamic energy. However, the resistant area. In fact, since the beginning of microbial mat studies in the area,
epibenthic microbial mats of ˜1–2 cm-thick can tolerate high current crabs have advanced constantly up the flat and have reduced the area
velocities and waves such as those severe storm events occurring at available for mat development.
least once a year. Such events can be preserved in sediments and
evidenced by biolaminites (Cuadrado et al., 2013). Moreover, the plastic 7.3. Hydrodynamic conditions under a storm event in BBE and its
nature of the microbial mat can sustain biogenic structures such as bird- consequences
footprints (Fig. 1f) that bear a rise in energy, as those produced by
severe storms (Carmona et al., 2011). Such biogenic structures can To demonstrate the degree of influence of waves over the upper
ultimately be preserved in sedimentary rocks due to the precipitation intertidal microbial mats, we analyzed wave records during a storm.
of authigenic minerals as a consequence of biogeochemical processes Wave data were taken at an Oceanographic Tower (OT) located just
in the mat (Carmona et al., 2012). The presence of authigenic minerals outside the BBE (Fig. 1a) and corresponded to the period July 20–30,
that would enhance the preservation was documented for the Puerto 2009. They consisted of values of significant height (Hs) and period
Rosales tidal flat (Cuadrado et al., 2012). (Ts) calculated every 30 min estimated from the spectrum of a fast
Fourier transform applied over a series of 18 min pressure estimates of
7.2. Sedimentary structures under normal energy conditions the water level at a frequency of 2 Hz.
Provided actual wave data was not available at the study site, we
Several MISS have been recognized and described for the supratidal compared a three month wave dataset with a similar period of data
zone of the Bahía Blanca Estuary (Cuadrado et al., 2011; Pan et al., taken at a nearby tidal flat (Vitale, 2010), and corrected the Hs of the
2013a,b). The general appearance of the tidal flat during 2009 showed OT for refraction and friction effects on wave propagation, due to the
soft sediments characterized mainly by the presence of straight to geomorphology of the estuary. Based on this correlation, we estimated
slightly-curved microbial shrinkage cracks, forming a polygonal pattern that the H at the Puerto Rosales tidal flat as the Hs/5.05. No correction
on the supratidal zone (Fig. 4a), starting at the end of the Austral for the period was made. Maximum corrected wave heights for July
summer. Polygons had 4 to 6 sides and their dimensions were variable, 2009 ranged from 0.02 to 0.52 m whereas periods were from 4 to 15 s.
between 6 and 20 cm in width, and the majority of the polygons tended At the study site, there is a pressure sensor that estimates water level
to be equidimensional. They evolved to cracks with upturned margins every 2 min which allows a precise determination of water depth (d).
(Fig. 4b), a characteristic of microbial shrinkage cracks due to prolonged Considering the possible errors involved in the estimation of the local
aerial exposure (Porada and Druschel, 2010) (Fig. 4e). Hs by applying the indicated correction, calculation of the wave
 D.G. Cuadrado et al. / Marine Geology 352 (2014) 367–380 373

Fig. 4. (a) General view of the tidal flat in May 7th, 2009. The tidal flat is covered by shrinkage cracks. (b) General view of the tidal flat in July 9th, 2009 after periods of desiccation.
(c) Close-up from (a). Some cracks are sealed-up due to subsequent hydration and recolonization (black arrows). Other cracks have their margins separated due to shrinkage (open
arrows). Some elliptic bulges can be observed (white solid arrows). The reference square is 20 × 20 cm. (d) The same reference square as (c) two month later. The crack margins
(same arrows shown in c) are opened and upturned showing the substrate below, due to desiccation. See the small domes as a product of photosynthesis by underlying cyanobacteria
called “blister mat” (Gerdes, 2007) formed in patches (black arrows). (e) Tidal height (above 3.90 m) that inundate the supratidal flat. The storm was marked with a circle.

parameters were done using the Airy linear theory; therefore, the The bed shear stress was calculated for each wave data when the
horizontal wave orbital velocity (uw) was calculated as water depth was greater than 0.05 m as suggested by Noffke (2010).
πHs During the period of the storm activity there were 15 high tides that
uw ¼ ð1Þ flooded the area colonized by microbial mats. This was an unusual
T s sin hðkdÞ
situation for this site, in which typically there are periods of several
where k = 2π/L and L is the local wavelength estimated iteratively from days on which the water level does not reach the area (Fig. 4e). The
maximum values of uw ranged between 0.55 and 3.6 m s−1, and τw
gT 2 ranged from 0.7 to 11.6 N m−2.
L¼ thðkdÞ ð2Þ
2π Considering that the typical sediment grain size at the tidal flat is
and g is the acceleration of gravity. D = 0.3 × 10−3 m, we calculated the value of Shields parameter (θ)
The bed shear stress produced by the waves (τw) was calculated from generated by the waves as

1 2 τw
τw ¼ ρf u ð3Þ θ¼ ð6Þ
2 w w ðρs −ρÞgD
where ρ is the water density (1027 kg m−3) and fw is the wave friction
factor which was estimated following Whitehouse et al. (2000) as where ρs (= 2650 kg m−3) is the sediment density. For each cycle the
−0:5 maximum value of θ was plotted as a function of the friction Reynolds
f w ¼ 2R ð4Þ
number over the Shields graphic proposed by Noffke (2010) (Fig. 8).
where R is the Reynolds number which was also calculated from In her original graphic θ values were estimated in the field employing
a portable erosion meter (Führböter and Manzenrieder, 1987) to
uw d determine the erosion stability of both endobenthic and epibenthic
R¼ ð5Þ
ν microbial mats. The wave shear stress exerted over the plain is greater
where ν is the cinematic viscosity (1.36 × 10−6 m2 s−1) than the magnitudes obtained by Noffke (2010) for the stabilization of
374 D.G. Cuadrado et al. / Marine Geology 352 (2014) 367–380

Fig. 5. (a)Aspect of the tidal flat showing elongated bulges (black arrows) in several directions and initial formation of cracks on the crest of bulges (white arrows). (b) Wrinkle structures
as sinuous crest (Kinneyia structures) after severe wave energy over the supratidal flat. A ridged bulge present in a piece of mat. The arrow shows the direction of the flood-tide.
(c) Liquefied sand underneath the microbial. Note the crack on the upper border of the opening and the elongated bulges on the top the picture. (d) Sand layer below the microbial
mat. (e) Piece of mat which is above the liquefied sand (not shown). The ridged bulge shown in (b) in plan-view is coincident to the crack in cross section. The crack is formed from
the underneath sand to the top of the mat due to hydraulic pressure. View text for explanation.

microbial mats. In the next section the response of the microbial mat definition as “wrinkle structures may reflect morphological surface and
against waves during a high-energy event is presented. subsurface features, mat deformation patterns, or be structures arising
from subsurface processes, e.g. gas trapping”. The structures recognized
7.4. Mat structures found after a severe storm event in this study have amplitudes of 2–3 cm in an orthogonal disposition
to the tide- and wave-advancement, and they are presented in a patchy
About 10 days before the storm, the conditions on the tidal flat were distribution. It is important to note than during the storm a rainfall
rather different from those after the July storm event. Periods of drying occurred (30.6 mm precipitation between July 20th and July 22th),
conditions make curled-up edges of microbial cracks that may lose their influencing the hydraulic pressure.
attachment to the sandy substrate (Fig. 4b, d). The storm occurred As a result of the increased energy during the storm event, polygonal
between 22nd and 31st July, 2009 and was characterized by strong fragments of flipped-over mat edges were formed (commonly named
winds from SW direction that caused the inundation of the supratidal “flip-overs”) along weakness of the cracks network (Fig. 10a). They
sector twice a day during several consecutive days (Fig. 9). So, there were completely turned over at the edges, some of them in the
was a high level of energy over the tidal flat which supported high direction of the tidal current (Fig. 10b). Thus, part of the substratum
wave energy as represented by the high values of θ (Fig. 8). After is exposed producing localized erosional pockets (Noffke, 1999) or
that episode, the lower supratidal zone exhibited a large number of erosional “windows” (Eriksson et al., 2007). Ten days after the storm,
sinuous ridges displayed in the cohesive and flexible mat surface a recolonization began with the formation of a biofilm (Fig. 10c), similar
as deformations of the microbial mats (Fig. 10b). These features might to the younger cultures obtained by Hagadorn and McDowell (2012).
be called “wrinkle structures”. After the thorough review made by However, that recolonization was not efficient in protecting the tidal
Porada and Bouougri (2007) about these structures, we adopted their flat and two months later the erosional pocket remained probably

Fig. 6. Hydraulic conditions over Puerto Rosales tidal flat modified from Porada et al. (2007) and Gardner and Wilson (2006). Open arrows = hydrostatic pressure; solid arrows =
hydraulic pressure. MLW = mean low water; MHW = mean high water; STHW = spring tide high water.
 D.G. Cuadrado et al. / Marine Geology 352 (2014) 367–380 375

Fig. 7. Examples of floating microbial mats. In the bare spaces of a Saccocornia perennis high marsh there is significant colonization of mats. The vascular plants have relatively low density.
(a) (b) Microbial mat covered by water. The combined effect of groundwater (pushed upward by the incoming tide) and the gases generated by the microorganisms induce the mat to float
in the typical ponds of the area. (c) Healed-up wide cracks. Note the curled crack margins. The bubbles in the water are due to photosynthetic activity of microorganisms forming the mat.
(d) Sieve-like surface as the impression of photosynthetic gas bubbles formed in the thin mat. Scale (coin) is 23 mm in diameter.

uncolonized (Fig. 10d, e). This patch is the source of sandy sediment that play a significant role in defining the various properties of sediments.
might be transported by tidal currents and waves. Accordingly, the activities of organisms can be generally divided into
bioturbation and biostabilization.
8. Discussion Bioturbation considers all the biological activities that modify
sediment packing properties by burrowing, sediment displacement,
Originally thought as bare environments, tidal flats are an excellent modification of the level of oxidation–reduction, etc. Whereas,
example of multiple and complex physical–biological interactions biostabilization deals with biological processes (i.e., generation of
which are tied up in an ecological cascade (Daborn et al., 1993). pellets, sediment surface alteration, gluing, etc.) that make more difficult
Due to growing field research on intertidal environments in the last for the physical processes (i.e., currents, waves, wind) to dislodge the
20–30 years, there are now clear evidences that biological activities surface sediment by increasing the critical shear stress required to
mobilize sediment particles. When present, microbial mats represent
one of the most significant and complex of biological systems that
intervene in the latter. Within mats there is interplay between
cyanobacteria, diatoms and other groups of protists which, besides
their proper presence, generate EPS that act as a very effective “glue”
among both inorganic and organic sediment particles.
The fact is that an action exerted by the same biota is capable of
perturbing the sediment in an occasion and is also capable of producing
exactly the reverse effect. A valid analogy would be that of a person
walking on a beach, who disturbs the sediment packing by displacing
surface sediment into de rim of the crater made with every step taken,
but also, due to the pressure exerted, extrudes water and the sediment
becomes more packed (Perillo, 2003). Similarly, macrofaunal benthos
(e.g., crabs, worms) have the same effect on tidal flat. As they burrow,
they change the oxidation–reduction conditions of the underlying
layers of sediment, they modify the packing by either disturbing the
Fig. 8. Shield's diagram for initiation of grain sediment transport which includes
the threshold for suspended load proposed by Bagnold (1966). The circles data points sediment particles or developing different types of pellets in the same
correspond to field experiments made by Nofke and Manzenrieder in 2002–2005 using a process, but simultaneously generate mucus that increases the inter-
portable unidirectional erosion meter designed by Führböter and Manzenrieder (1987). particle bonding.
Grey circles represent measurements made over endobenthic microbial mats over a Although studies related to microbial mats date back to the early
sandy substrate and black circles corresponds to epibenthic microbial mats (both reported
by Noffke, 2010). Red squares are the maximum values of θ for the 15 cases in which the
1960s, the recognition of MISS on the last ten years, added a completely
tidal flat at Puerto Rosales was inundated and having wave activity over epibenthic new perspective to the interpretation and analysis of physical–
microbial mats. biological interactions, and the resulting sediment transport in coastal
376 D.G. Cuadrado et al. / Marine Geology 352 (2014) 367–380

Fig. 9. Tide height and wave height during July when the severe storm occurred. During the storm (between July 22nd and 31st) the measured tide was higher than the forecasted tide and
also the wave height was greater. The plot shows only the tides that inundated the supratidal zone (tide height N3.9 m). Note that the exertion of water and waves over the plain was twice
a day during consecutive days.

wetlands in general, and tidal flats, in particular. As previously key mechanism involved in the formation of Kinneyia structures is
described, microbial mats interact with the sediment in different ways an instability induced in a viscoelastic film under flowing water.
and, depending of the species, they participate in the formation of After laboratory experiments, they confirmed that the ripple has a
various morphologic structures mentioned before which were classified wavelength roughly three times the thickness of the film, independently
by Noffke (2010) (Fig. 2). of the viscosity of the film and the flow conditions. They also confirmed
The lower supratidal zone of tidal flats is commonly colonized by that storm events can deform an established microbial mat in an
epibenthic mats mostly composed by M. chthonoplastes which forms undulate manner. In the present study we propose that the action of
a coherent microbial mat network. Sand particles are covered by hydrostatic and hydraulic pressures over the microbial mat is the origin
the ubiquitous EPS (Noffke, 2010). The latencies in this zone depend of the wrinkle structures observed after the storm event considered. In
on the occurrence of storms and the direction of wind-generated this case, the repetitive variation in tidal levels twice a day for several
waves, which influence how much high tides reach on the flat. The consecutive days, created a unique direction of formation of wrinkles
cohesiveness of the microbial meshwork has the capability of inhibiting (Fig. 5b, white arrow in Fig. 10b). These wrinkle structures which are
the escape of gas produced by complex biogeochemical reactions during similar to Kinneyia-type structures (the most typical structures on
bacterial degradation into air or water (Gerdes, 2010) forming gas ancient siliciclastic sediments since Archean; Porada et al., 2008) have
domes (Cuadrado et al., 2011). This strong cohesive layer is also capable been presented in different studies (Porada and Bouougri, 2007 and
of confining water and inducing greater hydraulic pressure in the upper references therein; Porada et al., 2008).
slope of the tidal flat. The latter fact produces an uplift force, favoring Deformation structures as wrinkle structures are a consequence of
the rise of sediment in the mat substratum. This force alternatively the balance between hydrostatic and hydraulic pressures. Additionally,
counteracts the hydrostatic pressure made by the sea level during during storms events the tidal plain is also affected by the action of
flood and ebb tides. waves which can have effects on the supratidal zone and, even though
In regular conditions, the genesis of MISS is related to the action they have very small wave heights, the high values of shear stress
of the hydrostatic pressure over the sealing microbial mat which is produce erosional signatures. On average, wave action over the plain
under hydraulic pressure (Porada et al., 2007, 2008). The variation in is much more effective in generating destructive structures as flipped-
tides over the microbial mat during several days (see the high frequency overs. In our case study, these erosional features occurred only under
of tidal inundation in Figs. 4e, 10f) originates elongated domes with high energy level, once a year; in agreement with the highest wave
different orientations over the plain (Fig. 5a). The pressure of the heights over the tidal plain. The analysis of storms made for the
water column during high tide produces the formation of a crack supratidal flat at Puerto Rosales considered an unusual period during
below the mat, uplifting the underneath liquefied sand into the crack, which the plain was flooded 15 times in less than 10 days. Under such
while the flexible upper mat remains intact (Fig. 5e). These structures conditions, the microbial mats in the tidal flat were able to withstand
can be compared to “petee” or “petee ridge” (Bouougri et al., 2007). very high energetic conditions which were much higher than the critical
The first definition (Gavish et al., 1985) established that this structure values for both sediment and microbial mat dislodging (Fig. 10).
is related to mat growth and gas pressure from underlying mat decay. Although the flexible characteristic of the microbial mat, flip-overs
This condition is responsible for detaching the cohesive mat and start along pre-existing crack margins in the microbial mat, formed by
deforming it into domes and irregular folds (petee sensu stricto by both hydraulic upward pressure and desiccation processes. Although
Gehling, 1999, 2000), a definition that is still current (Porada and not measured in our study, the excess pressure induced by waves on
Druschel, 2010). Schieber (2004) associated the genetic definition the groundwater may represent a potential force that helps dislodge
with mat cracking and used the term “petee ridges” where the pattern the mat during the formation of a flip-over. On the other hand, the
may relate to desiccation cracks (petee sensu lato; Eriksson et al., 2007). presence of the microbial mats could potentially prevent water from
Parizot et al. (2005) discussed these structures in the fossil record, being fully injected into the sediment to develop a fluid mud, as
which are similar to those recognized in the present study. described by Pratolongo et al. (2010). These structures were recognized
We propose the term wrinkle structures, especially Kinneyia in the clastic rock record as destruction features (Schieber, 1999).
structures to those found after the storm (Figs. 5b, 10b). The origin of Once upturned, the mat behaved as a mat fragment and a few
wrinkle structures has been controversial. Hagadorn and Bottjer months later microbial recolonization around their edges could be
(1997, 1999) had suggested the interrelation between microbial activity verified (white arrow in Fig. 10e). The erosional pocket left by the mat
and wrinkle structures formation, including in this terminology both fragment turned into a source of sediment that might be transported
Kinneyia and elephant skin structures. Pflüeger (1999) recognized a by tidal currents and waves over the tidal flat. This fact produces a slight
pattern similar to “Kinneyia”, by experimentally generating it with accumulation over the microbial mat near the erosional pockets that
a fluidized sand and gas pressure and related the presence of these can be recognized as siliciclastic biolaminite (Bouougri and Porada,
structures to storm waves in the fossil record. In spite of Noffke 2007) in vertical cross-section of the plain (Fig. 11a). The process
(2010) stating that they only occur in the fossil record and Porada and that controls the deposition of siliciclastic biolaminites includes periods
Bouougri (2007) declared that these structures have not been observed of sedimentation in which the cyanobacteria can move upward
in modern mat, a recent study (Thomas et al., 2013) proposed that the establishing a new mat at the top, over old mat layers. The fine sand
 D.G. Cuadrado et al. / Marine Geology 352 (2014) 367–380 377

Fig. 10. Tidal flat after the severe storm. (a) Fragments of flip-overs showing the uncolonized sediment underneath the microbial mat. Note the initial formation of thin mat over
the uncolonized sediments in the pocket left by the flip-over (bottom of the figure). (b) Flip-overs turned in the direction of flood current pointed by the black arrow. Petee-ridges
(white arrow) are orthogonal to the food direction over an un-erosional sector at the bottom of figure. (c–e) Comparison of the flip-over during two months. (c) There is a thin mat
“skin” forming in the pocket left by the flip-over. (d) 20-days after the previous picture the thin mat was taken-off by natural processes. (e) Subsequent rehydration and aerial exposure
made the flip-over fragment re-colonized and adhesion to the mat (white arrow). Note sand grains over the mat transported from the sand pocket to the left of the pocket (references
squares were 20 × 20 cm). (f) Height of the tide (higher than 3.90 m) that produces the inundation of supratidal zone after the storm occurred in July (indicated by a circle). Letters
refer to the pictures date.

lamination recognized in the cross section taken in 2010 indicates the that were not recolonized by microbial mat and also between the
sporadic high energy event that occurred the year before. cracks margins (Fig. 11 b). Gerdes et al. (2000) declared that the
Crabs in this area (N. granulata) typically become very active in late burrowing marine species are excluded from the zones colonized by
spring and throughout summer (Escapa et al., 2008); therefore during M. chthonoplastes because they must expend a great deal of energy to
this period crabs advanced over the area making use of the open patches penetrate through such thick filamentous surface layers. Evidently
378 D.G. Cuadrado et al. / Marine Geology 352 (2014) 367–380

Fig. 11. (a) Vertical cross section of the sedimentary surface reveals the biolamination. The photograph was taken on May 2010. The thin sand layer (shown by an arrow) would reflect the
strong storm happened the year before. Note the EPS layers in the layers below reflecting periods of no or low sedimentation (ns). (b) Colonization of crabs between the crack margins
(thin or absent microbial mat) marked with an arrow. Note the bioturbated loose sediment (darker) extracted from underlying layers over the microbial mat. The square dimensions are
20 × 20 cm.

they choose special weaker zones and this crab colonization advancing range of stressors, they contribute to retention and consolidation of
from the lower intertidal toward the upper supratidal flat has been sediments and, even prevent their erosion; while on the other hand,
recognized in previous studies (Cuadrado et al., 2011). Thus, this when perturbed by desiccation or by physical and/or biological
significantly contribution to bioturbation must be considered in future processes, the unprotected sector can be easily eroded and sediment
balances of biostabilization–bioturbation rates. It is reasonable to assume transported elsewhere. Nevertheless, microbial mats have proven
that, as the crabs are neutralizing the formation of mats, the area will to be an excellent indicator of environmental conditions as they are
produce more sediment to be transported over the plain, what would an important mechanism in the preservation of structures in the
ultimately affect the presence and evolution of microbial mats. stratigraphic record.

9. Conclusions Acknowledgments

Microbial mats have played a significant role in stabilizing the This work was partly funded by grants from ANPCyT, CONICET and
sediment, being part of the formation of sedimentary structures and SECyT-UNS of Argentina. The authors would like to thanks John T.
preserving them along the Earth history but their importance has only Wells and two anonymous reviewers for their helpful comments that
been recognized in the last decades. Although we focus in tidal flat improved the manuscript. We appreciate the English by J. Pan.
environments, MISS observed in the stratigraphic record of different
types of environments may give a clue about the physical, biogeo-
chemical and sedimentary processes. Therefore, knowing in higher Appendix A. Supplementary data
detail the processes involved in modern microbial mat colonization
over tidal flats result in an essential tool for geological interpretation. Supplementary data to this article can be found online at http://dx.
Low sedimentation is needed for the survival of microorganisms and doi.org/10.1016/j.margeo.2013.10.002.
the presence of microbial mats could be connected mostly to relatively
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