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From the sea to the land: How microbial mats dominated marine and
continental environments in the Ediacaran Camaquã Basin, Brazil

Ilana Lehn , Paulo Sergio Gomes Paim , Farid Chemale Jr.

PII: S2772-8838(24)00033-5
DOI: https://doi.org/10.1016/j.geogeo.2024.100283
Reference: GEOGEO 100283

To appear in: Geosystems and Geoenvironment

Received date: 6 November 2023

Revised date: 20 February 2024
Accepted date: 29 February 2024

Please cite this article as: Ilana Lehn , Paulo Sergio Gomes Paim , Farid Chemale Jr. , From
the sea to the land: How microbial mats dominated marine and continental environments
in the Ediacaran Camaquã Basin, Brazil, Geosystems and Geoenvironment (2024), doi:
https://doi.org/10.1016/j.geogeo.2024.100283

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Highlights
 Geochemical proxies indicate mainly oxic conditions during Camaquã Basin deposition.

 Ediacaran well-preserved microbial mats associated with organic-walled microfossils.

 Clay particles associated with EPS sealed the organic matter and avoided their decay.
From the sea to the land: How microbial mats dominated marine and

continental environments in the Ediacaran Camaquã Basin, Brazil

Ilana Lehn*, Paulo Sergio Gomes Paim, Farid Chemale Jr.

Universidade do Vale do Rio dos Sinos, Geology Graduate Program, Av. Unisinos, 950, Cristo Rei 93022-000, São

Leopoldo, Brasil

*Corresponding author. E-mail address: ilanalehn@gmail.com

Handling Editor: Arubam C. Khelen

Abstract

Precambrian sedimentary successions exposed worldwide record microbial deposits within

both marine and continental strata. The Camaquã Basin records the transition from the Ediacaran

to Cambrian periods in strata that comprise marine to lacustrine environments. The Camaquã Basin

fossils comprise microbial mats and organic-walled microfossils in very close association.

Carbonaceous laminae are common within the siltstone and mudstone layers and represent

remarkably well-preserved Ediacaran microbial mats. These mats occurred along shallow marine

through deep and shallow lake settings. The excellent preservation of these structures seems to be

the product of calm hydrodynamic conditions and floodings. These hydrodynamic undisturbed

environments allowed microbial mat growth. Besides, episodic floodings brought clay particles,

which deposits guaranteed isolation from organic decomposition of mats. In general, the siltstones

and mudstones record microbially induced sedimentary structures (MISS) that are typical from
Ediacaran strata, such as wrinkle marks and elephant skin structures. Our research revealed that

the occurrence of these structures is not limited to marine settings, as common in records from this

period, but also present in lacustrine environments. Field studies were combined with optical and

scanning electron microscopy plus energy-dispersive X-ray spectroscopy (SEM-EDS) analyses to

demonstrate the vast occurrence of microbial mats along the deposits that registered the Camaquã

Basin development. Geochemical data (whole-rock and total organic carbon (TOC) analyses)

contribute to improving sedimentological descriptions, bringing light regarding the depositional

environment where these microbial communities have developed. The excellent preservation state

of organic matter on these marine and lacustrine siliciclastic sequences represents an impressive

record in a Proto-Gondwana basin.

Keywords: Microbial mats; Organic-walled microfossils; Lacustrine deposits; Geochemical

proxies; Ediacaran; Proto-Gondwana basin

1. Introduction

The supercontinent Gondwana has its origin associated with ocean closures and

rearrangement associated with collision and suturing of the older continental crust parts from the

Archean-Mesoproterozoic cratons during the Neoproterozoic Pan-African-Brazilian Orogeny

(Trompette, 1994; Unrug, 1997). As a result of the later stages of this orogeny, several small basins

were formed along the outer part of the Brasiliano belts of the Mantiqueira Province (eastern

Brazil; Philipp et al., 2016).

The Camaquã Basin was formed along the Late Ediacaran/Early Cambrian, associated with

late orogenic stages of Brasiliano/Pan-African Orogeny. Several works have been carried out on
stratigraphy (Paim et al., 2000; Janikian et al., 2003; Paim and Scherer, 2007; Paim et al., 2014),

sedimentology (Fambrini et al., 2006; Borba et al., 2008; Marconato et al., 2014; Lehn et al., 2018),

and volcanic events (Wildner et al., 2002; Janikian et al., 2012), but most of them focusing on

specific units rather than the entire succession. Information about life in the Camaquã Basin is

scarce, limited to a few works focusing on ichnology (Netto, 2000, 2012) and, more recently,

microfossils (Lehn et al., 2019, 2022). This is the first time that microbial mats from both, marine

and continental strata, of the Camaquã Basin are shown in detail and supported by geochemical

data.

Evidence suggests that microbial mats have inhabited both marine and terrestrial

environments since the Archean Eon. The oldest microbially induced sedimentary structure

(MISS), preserved on siliciclastic sequences, includes wrinkle structures and roll-up structures in

tidal flats of 3.2 Ga Moodies Group, South Africa (Noffke et al., 2006). However, the biogenicity

of some of these structures are still under discussion (Saitoh et al., 2021; Homann and Heubeck,

2021), which demonstrate the necessity of research on this subject. Microbial mats are also

reported in Precambrian (2.7 and 2.6 Ga) paleosoil (Watanabe et al., 2000) and fluvial deposits

(Homann et al., 2018) in South Africa. Following ancient world examples, the Camaquã Basin

presents remarkably well-preserved examples of Proterozoic MISS within marine and lacustrine

facies.

A wide morphological variety of MISS can be found in the Precambrian fossil record. These

structures result from distinct variables, such as different sedimentary environments, substrate

particle size, types of interaction between the organisms and the sediment, biofilm presence, and

taphonomic processes (Noffke et al., 2003).

 Geochemical proxies provide hints about the depositional environment conditions. Shales and

mudrocks, specifically, are enriched with suites of major elements and trace metals that reflect

their depositional environment, provenance, and diagenesis (Ross and Bustin, 2009). Clues about

the depositional environment can be assessed through total organic carbon (TOC) content, clay

components, and radioactive elements, emphasizing physical and chemical properties (Ibrahim et

al., 2020). The hostess rocks geochemical characterization represents a fundamental tool to

understand the environment that the studied strata represent and, consequently, the fossil

preservation on them.

Ediacaran strata are well known for the presence of microbes and microbial textures

preserved in sandstones and siltstones/mudstones. The exceptional preservation of microbes in

Ediacaran sediments suggests a unique taphonomic window (Newman et al., 2016). The

Ediacaran-type preservation of microbial textures is explained by some hypotheses that propose

elevated silica concentrations in Precambrian oceans (Siever, 1957) or even a shallowing of the

Ediacaran seafloor redox boundary that reduced the degradation of organic matter and enhanced

microbial preservation (Callow and Brasier, 2009). Aluminum- and iron-rich clay minerals

preclude decay bacteria and increase decay resistance through clay-organic matter interactions

(McMahon et al., 2016; Naimark et al., 2016; Anderson et al., 2020). A silica- and aluminum-rich

system seems suitable to explain the preservation of microbial forms in the Ediacaran Camaquã

Basin record; with clay layers preserving the organic mats and associated structures.

The Camaquã Basin is part of Proto-Gondwana basins that host primitive microorganisms.

Understanding these environments help us to improve the knowledge about life dispersion in both

continental and marine settings on the Earth. Therefore, this research uses organic and inorganic

geochemical data coupled with optical microscopy and SEM/EDS analysis of mudstones,
sandstones, and organic matter to constrain paleoenvironmental conditions and test previous

sedimentological-based paleoenvironmental interpretations. Additionally, this paper highlights the

remarkable preservation of organic matter in siliciclastic rocks for the first time within a Proto-

Gondwana basin.

2. Geological background

The Camaquã Basin is located in southernmost Brazil (Fig. 1) and records late to post-

collisional stages of the Brasiliano/Pan-African Orogeny in the Dom Feliciano Belt (Philipp et al.,

2016 and references therein). The geological record (Fig. 2) crops out along Piquirí–Arroio Boici,

Santa Bárbara, Guaritas, and Ramada-Taquarembó structural blocks (Paim et al., 2000). They

include a partial record of four successive tectonic-volcanic-sedimentary episodes (or groups) (Fig.

3) evolving from compressive through transpressive and transtensive to distensive tectonic regimes

recorded as four unconformity-bounded units (Paim et al., 2014).

The Maricá Group has been ascribed to a retroarc foreland basin (Paim et al., 2000). The

sedimentary succession comprises a siliciclastic package about 2,000 m thick (Paim et al., 2000;

Borba et al., 2008) at the base of the basin and includes the Arroio América, São Rafael, and Passo

da Promessa formations (Pelosi, 2003).

The Bom Jardim Group is a volcano-sedimentary succession that comprises a second

unconformity-bounded unit of the Camaquã Basin (Fig. 3b). It records the infill of two late- to

post-collisional basins (East and West Bom Jardim transcurrent basins) developed within a back-

arc context (Paim et al., 2000, 2014). The Bom Jardim Group is about 4,000 m thick and includes

intermediate to basic volcanic rocks of shoshonitic to high-K, calc-alkaline affiliation, as well as

volcanoclastic strata of the Hilário Formation (Wildner et al., 2002). The siliciclastic succession is

subdivided into the Picada das Graças and Cerro da Angélica formations (Janikian et al., 2003).

The Santa Bárbara Group is the third unconformity-bounded unit of the Camaquã Basin

and records the filling of two basins (Santa Bárbara East and West rifts; Paim et al., 2014)

developed in transtensional regime during post-collisional stages (Paim et al., 2000; Borba and

Mizusaki, 2003). At its base, it contains alkaline, mostly acidic volcanic rocks (Acampamento

Velho Formation; Wildner et al., 2002). The siliciclastic package that overlies the Acampamento

Velho Formation includes the Santa Fé, Serra dos Lanceiros, and Pedra do Segredo formations.

The uppermost unconformity-bounded unit of the Camaquã Basin is the Guaritas Group.

It is around 800 m thick and includes, at its base, volcanic rocks (Rodeio Velho Formation),

interlayered with and overlain by the aeolian strata of the Pedra Pintada Formation. A

disconformity delineates the boundary between the Pedra Pintada and Varzinha formations (Paim

and Scherer, 2007).

2.1. Stratigraphy of Camaquã Basin

Each unit of the Camaquã Basin comprises different siliciclastic sedimentary facies,

according to the depositional systems, which lead to different depositional environment

interpretations (Table 1). Petrographic and geochemical data will be used to scrutinize and analyse

the current interpretations present in the literature.

The Maricá Group sedimentary succession is divided into three major units, which denote

linked depositional intervals: Passo da Promessa, São Rafael, and Arroio América formations

(Pelosi et al., 2003). The base of the group (Passo da Promessa Formation) is composed of trough

cross-bedded, lenticular conglomeratic sandstones and conglomerates and tabular, either ripple-
cross laminated or massive sandstones ascribed to a braided fluvial depositional system grading

upwards to a delta front setting (Fig. 3). The intermediate interval (São Rafael Formation)

comprises tabular siltstones and mudstones interlayered, tabular very fine-grained sandstones and

sandstones with hummocky cross-stratification and wave-ripples, interval interpreted as distal, low

concentration turbidites deposited a fluvial- and wave-influenced marine environment (Borba et

al., 2008; Paim et al., 2014). MISS is very common in these fine-grained facies. These facies host

small and unornamented acritarchs as the only member of this microfossil association, suggesting

a restricted, highly influenced by river discharge marine environment (Lehn et al., 2019). The last

interval (Arroio América Formation) is composed of lenticular and sigmoidal, massive to trough-

cross bedded conglomeratic sandstones, interpreted as a fluvio-deltaic system (Fig. 1; Borba et al.,

2008; Paim et al., 2014).

The proximal facies of the Bom Jardim Group comprise tabular graded, coarse-grained

sandstones and conglomerates ascribed to sheet floods taking place on delta plains of fan deltas,

developed along the active faulted margins of the transcurrent basins (Paim et al.,

2000). Centimeter- to decimeter-scale, tabular beds of fine- to very fine-grained sandstones and

mudstones interlayered are interpreted as thin-bedded turbidites deposited along the delta-front of

the fan deltas (Paim et al., 2014). MISS are present in the fine-grained facies, which seldom may

present desiccation cracks. Microfossils were recovered from siltstones and mudstones, and as

expected for lacustrine settings, acritarchs show simple morphology, mostly

from Leiosphaeridia genera (Lehn et al., 2019). The microfossil content in fine-grained facies

indicates that during periods of low alluvial discharges, microbial mats and acritarchs development

was favored.
 The Santa Bárbara Group presents three major sedimentary depositional sequences. In

general, they encompass lenticular beds of through cross-bedded conglomerates or coarse- to very-

coarse-grained sandstones ascribed to braided fluvial systems at their bases. The intermediate

intervals are composed of tabular and sigmoidal sandstones interbedded with siltstones/mudstones,

with abundant mud cracks, deposits related to mouth-bar (sigmoidal bodies), hyperpycnal turbidity

currents, and cohesionless debris flows deposited on a delta front to prodelta setting subjected to

recurrent exposures (Paim et al., 2014; Lehn et al., 2018). The upper intervals present lenticular,

massive, or cross-stratified sandstones and/or conglomerates that were associated with delta plain

deposits of braidplain deltas (Paim et al., 2000). MISS, sphaeromorphs, and acanthomorphs

acritarchs are very common in fine-grained facies, mainly associated with the maximum flooding

interval of each sequence. Like in the previous unit, acritarchs present simple morphology,

recorded by the Leiosphaeridia sp. and Lophosphaeridia sp. genera, which is consonant with other

lacustrine examples in the world (Horodyski, 1994; Arouri, 1999; Prave, 2002; Strother et al.,

2011). The shallow lakes were able to lodge life for some periods, while the microbial mats would

grow and establish a substrate (Lehn et al., 2019). Recurrent desiccation mud cracks indicate that

these conditions were sustained until large parts of the lakes were drained and the substrate

exposed. On the other hand, thin-bedded turbidites and associated debrites record the humid phases

associated with high-frequency climatic cycles, a common aspect of lacustrine systems (Carroll

and Bohacs, 1999).

The Guaritas Group rests on an angular unconformity above the Santa Bárbara Group. The

Pedra Pintada Formation is intercalated with and rests on the andesitic rocks of the Rodeio Velho

Formation. This unit records successive palaeoergs bounded by super surfaces (Paim and Scherer,

2007). The Varzinha Formation includes fine- to coarse-grained, trough cross-bedded sandstones,
ascribed to braided fluvial systems at its base, and sigmoidal and tabular beds of very fine- to fine-

grained climbing- and wave-rippled sandstones interlayered with mudstone, above the fluvial

deposits. This facies association was related to delta front to prodelta facies developed on shallow

ephemeral lakes, frequently subaerial exposed (abundant mud cracks; Paim et al., 2000). MISS are

rare but occur in the fine-grained facies related to ephemeral lakes, from which microbial mat

fragments were recovered through palynological extraction.

3. Material and methods

Petrographic analyses were conducted in forty-seven thin sections of fine-grained rocks

from all units of the Camaquã Basin (Fig. 2) using optical and scanning electron microscopy. A

palynological preparation technique was used for the extraction of acid-insoluble microfossils.

Following Grey’s method (1999), raw samples were mechanically disaggregated. After, they were

digested with HCl and HF for carbonate and silicate removal, respectively. Boiling HCl was used

for the removal of clay minerals. After filtration (10 µm filter size) and swirling to separate heavy

minerals, strew slides were prepared and examined under a transmitted light microscope. Optical

microscopy was performed at Universidade do Vale do Rio dos Sinos (UNISINOS, Brazil). Three

scanning electron microscopes with energy dispersive detector (SEM-EDS) were used, a first one

(Quanta 650FEG model) at Centro Nacional de Pesquisa em Energia e Materiais (CNPEM, Brazil),

a second one (Zeiss EVO/MA15 model) at Itt fossil (UNISINOS, Brazil) and a third one (Quanta

400FEG model) at University of California at Santa Barbara (UCSB, United States).

Samples were analyzed for total organic carbon (TOC) at the Geochemical Analysis

Laboratory of Pontificia Universidade Católica do Rio Grande do Sul (PUCRS) in Brazil, using a
LECO elemental analyzer (TRUSPEC model). This involved first pulverizing the samples and

digesting them with hydrochloric acid (3.24 M) to release inorganic carbon (CO₂ generated from

carbonate decomposition). The amount of CO₂ generated during combustion was then measured.

Additionally, eight samples were analyzed for major, trace, and rare earth elements (REE) at the

SGS Geosol Laboratory in Brazil. Major oxides were analyzed using X-ray fluorescence after

sample fusion. Trace elements and REE were analyzed using Inductively Coupled Plasma Mass

Spectrometer (ICP-MS) after sample fusion with lithium metaborate (LiBO2) and digestion with

an acid solution (C4H6O6 and HNO3; Tomascak et al., 2012).

4. Results

4.1. Microbial induced sedimentary structures

Different kinds of MISS were reported in each unit of Camaquã Basin (Table 2), mostly

related to transgressive portions of rock successions. Typically, there is an upward decrease in

MISS diversity in the rock succession. The Maricá Group has a diverse range of MISS, including

bubbles, wrinkle marks, pitted texture, ridge reticulated, grain prison + wrinkle, elephant skin,

Arumberia, ridges and mounds, gas escape, and reticulated marks (Fig. 4). The Bom Jardim Group

also showed a significant variety of MISS, such as Arumberia-like/leveling, leveling, palimpsest

(bubbles and reticulated marks), bubbles, grain prison, setulfs, and pitted structures (Fig. 5). The

Santa Bárbara Group shows only five types of MISS: cuspate mounds, bubble marks, Arumberia-

like, and wrinkle marks (Fig. 6). At last, only wrinkle marks were found in the Guaritas Group

(Fig. 6).
4.2. Petrography of Ediacaran mats

Thin sections show the presence of microbial mats as fragments (centimetric to

micrometric pieces) or in situ. The mats may occur on bedding planes, either surrounding the

grains with organic filaments or composing the layer itself or even penetrating the siliciclastic

layer (Fig. 7). Structures such as aligned mica grains parallel to bedding and elongated organic

grains (size of fine-grained sand) within an organic-rich matrix indicate the presence of ancient

microbial mats on fine-grained facies trapping the particles.

In the Camaquã Basin, microbial mats are preserved in siliciclastic rocks, mainly siltstones,

and mudstones. Occasionally, organic fragments also occur as mat chips in sandstones. Microbial

mat fragments were discovered alongside microfossils using the palynological technique for

extracting acritarch and other acid-insoluble microfossils (Grey, 1999; see Fig. 8).

Microbial mats can be found as small, organic-rich films mixed with fine-grained sediment

or as fragments within sandstones. These dark layers contain the original organic matter that made

up the biofilm on marine and lake substrates during the development of the Camaquã Basin. In

some cases, the cellular structure has been preserved in the carbonaceous material (Fig. 9). The

composition of carbon in the mats has been confirmed through EDS results (Fig. 10).

4.3. Eodiagenetic cement and microbial mats

The Maricá Group facies often contain early diagenetic carbonate cement, as shown in Fig.

11a. In contrast, the Bom Jardim Group has iron oxide coating that was formed before eodiagenetic

calcite cementation, also shown in Fig. 11b. The Santa Bárbara Group has more iron coating, but

less eodiagenetic calcite cement (Fig. 11c). Finally, in the Guaritas Group, the only eodiagenetic

cement present is iron oxide coating (Fig. 11d). As we move upwards along the Camaquã Basin
succession, there is a decrease in calcite cement and an increase in iron coatings. Additionally, the

diversity of MISS and the occurrence of microbial mats also decrease.

4.4. Geochemical proxies

Trace element ratios can provide information about paleoredox environment conditions.

The use of proxies as Th/U, Ni/Co, V/Cr can be used for organic-rich mudrocks to investigate the

role of bottom water oxygen levels and the water-column productivity reflected in inorganic

geochemical variations (Demaison and Moore, 1980; Tyson, 1987; Hatch and Leventhal, 1992;

Jones and Manning, 1994; Rimmer et al., 2004; Ross and Bustin, 2009; Zhao et al., 2016). To

better interpretate each different basin portion (or unit), which reflect different environments,

analyses was conducted in each unit separately. Zhao et al. (2016) suggests that some geochemical

proxies such as (Th/U, Ni/Co, and V/Cr) can indicate paleoredox environment conditions. The

work proposes that Ni/C ratios between 0-5 indicate oxic conditions, between 5-7 dysoxic and 7-

16 anoxic conditions; V/Cr ratios indicate oxic (0 to 2), dysoxic (2 to 4), or anoxic (4 to 12) states;

and Th/U ratios <2 signalize anoxic conditions. Based on our results (Table 3), it is possible to

indicate that oxic to slightly dysoxic conditions dominated during the deposition of the entire basin

(Fig. 12).

Just like some previously mentioned ratios that show the environmental oxidation state,

such as Ni/Co, V/Cr, and Th/U (Ross and Bustin, 2009; Zhao et al., 2016), certain elements can

also indicate bioproductivity (P), the presence of organic compounds (Co), and whether the biota

is marine or terrestrial (Sr), as suggested by Felitsyn et al. (1998). In this study, the CB samples

showed consistently low P content (less than 0.1 ppm), and Co content decreased from the bottom
to the top of the succession (Fig. 13). Only the Maricá Group sample from the CB had Sr values

higher than 500 ppm, indicating a marine deposition setting.

Table 5 displays the TOC content of 16 samples, consisting of siltstones, claystones and

mudstones, taken from all units of the CB. The mean values are generally low, varying from 0.08

to 0.35. However, the larger values are associated with the Bom Jardim Group.

5. Discussion

5.1. Camaquã Basin MISS

According to this study, the marine strata (attributed to the Maricá Group) display a higher

number of MISS types compared to the lacustrine ones (Bom Jardim, Santa Bárbara and Guaritas

groups). This can be attributed to the hydrodynamic and depositional conditions prevailing in

lakes, such as sedimentary discharge and subaerial exposure, thereby leading to erosion and

decomposition of microbial mats. Thus, in terms of environmental conditions, a marine setting

would be expected to preserve more MISS than a lacustrine one.

Predominantly, wrinkle marks are the most recurrent MISS type on CB, recorded in every

unit, even in the Guaritas Group (Fig. 6d). The term “wrinkle structure” has been used as a

collective term for a variety of small-scale crinkly marks on bedding surfaces, evoking the

impression of microbial mats during sedimentary deposition (Porada and Bouougri, 2007; Mata

and Botjer, 2009; Hagadorn and Botjer, 1997). The CB deposits present different sizes of wrinkle

marks, probably representing a colony response in the growth pattern to different ecological

conditions on the bottom (e.g., depth of water column and consequent hydrodynamic responses to

flooding).
 Reticulated structures occur frequently in Maricá and Bom Jardim outcrops (Figs. 4 and 5).

This structure, similar to a network pattern, occurs widespread in the Precambrian strata and is

usually interpreted as a primary feature related to mat growth (Hagadorn and Bottjer, 1999; Gerdes,

2007). Mat growth also causes leveling structures, common at Bom Jardim Group. Elephant skin

texture also presents a reticulated growth pattern and appears commonly in Bom Jardim and Santa

Bárbara groups.

Petrographic analyses demonstrate recurrent grain prison (Noffke et al., 2011) structures

caused by microbial binding of sediments on the substrate (Fig. 7a). Other common structures

associated with grain imprisonment are setulfs, i.e., a shadow of grains oriented parallel to

paleoflow and deposited at the down-current side of small obstacles (Friedman and Sanders, 1974).

According to Kolesnikov et al. (2012), the interaction between microbial mats and fluids also

explains the origin of the Arumberia structure, which is very recurrent at CB strata (Fig. 4a).

In the Maricá and Bom Jardim groups, the CB fine-grained facies display recurring pit and

mound structures, as well as bubble marks (Figs. 4 and 5). These structures are formed by trapped

gases that emerge from beneath a microbial mat. The gases are usually produced by decomposing

older microbial communities, causing doming of the mat surface (Gerdes et al., 1993). The gas

then escapes to the surface, resulting in small and shallow pits (<1 cm diameter) and creating pit

and mound structures (Reineck and Singh, 1980; Rindsberg, 2005; Davies et al., 2016).

Millimetric rounded marks found in fine-grained sandstones and siltstones of Camaquã

Basin (specifically in the Santa Bárbara Group) were associated either with raindrops (Paim, 1994)

or with possible late Precambrian discoidal fossils, like Aspidella terranovica (Netto, 2000, 2012).

Raindrops and Aspidella marks produce particular morphologies, such as partially superposed
discs with external rims, for the first one and discs with radial ridges and central hollow or

concentric ridges with no radial ornamentation for the fossils. Detailed analyses of transversal,

polished cuts of hand samples, and transversal or parallel to bedding thin sections, show no

similarity with the above-mentioned morphologies. The lack of these characteristic structures (Fig.

14a) indicates another origin for those marks. Also, polished hand sample and thin section from

the same portion indicate the presence of microbial mats on the surface of the sample (Fig. 14a-

b). The parallel and transverse to bedding thin sections demonstrate the existence of microbial

mats with the same macro-scale morphology. Although the possibility of raindrop marks is not

ruled out because they do occur in the Camaquã Basin (Fig. 14d), we propose that microbial

activities produce some of these rounded marks. Similar structures were described for siliciclastic

lacustrine sequences on the ~1000 Ma Diabaig Formation, northwest Scotland, and associated with

MISS (Callow et al., 2011; Strother and Wellman, 2016). The main difference between microbial-

induced marks and raindrops is the mark’s juxtaposition. The observed rounded structures occur

side by side, while the raindrops could fall one above another, causing a juxtaposition of forms.

5.2. Taphonomic issues

Samples show microbial mats frequently present in fine-grained substrates, suggesting that

they grew during periods of low energy and, consequently, low accumulation. The mats would

grow during the deposition of fine-grained particles, as evidenced by the prison structures, or when

there was no deposition, which prevents disturbance by strong currents that would erode the mats.

Additionally, mat fragments, such as chips and organic matter, are often found in sandstones of a

medium to fine grain size, indicating that the substrate was reworked, and the mats were deposited

as bioclasts.
 Optical and scanning electron microscopy show microbial mats surrounded by fine-grained

matrix in detail (Figs. 8, 9, and 10). EDS results reveal organic particles wrapped by an aluminum-

and silica-rich clay matrix with minor potassium and iron content (Fig. 10). The organic matter

remains preserved within the fine-grained rocks of CB. Furthermore, whole-rock geochemical

analyses (Supplementary Data, Tables S1-S2) indicate a high content of silica, aluminum, iron,

calcium, magnesium, sodium and potassium oxides.

Clay minerals can improve organic preservation by physically stabilizing substrate

molecules within their interlamellar lattice or modifying the active sites on the compound

(Butterfield, 1990). Additionally, fine-grained particles have valuable mechanical properties and

serve as excellent packing material, isolating and supporting delicate structures as observed in

macrofossil preservation. The presence of very fine-grained sediments can form a sealing effect,

preventing heterotrophic microbes or their respective electron acceptors from accessing the

organic matter (Alexander, 1965; Butterfield, 1990; Anderson et al., 2020). A study by Anderson

et al. (2020) found that the silicon- and aluminum-rich clay matrix can surround and preserve

organic components. Therefore, the fine-grained matrix found in CB samples plays a crucial role

in preserving organic matter (Fig. 10).

The Torridon Group, in Scotland, is an immature siliciclastic sequence (1.2-1.0 Ga) that

presents lacustrine strata preserving organic-walled microfossils by combining clay minerals and

phosphate (Strother et al., 2021). Also, experiments conducted by Newman et al. (2016)

demonstrated that clay minerals can coat cyanobacterial cells under oxic conditions without

interruption of cell growth or photosynthetic processes. Besides clay capacity of organic matter

(OM) preservation, experiments demonstrate the importance of exopolymeric substances (EPS) in

OM fossilization. According to Pacton et al. (2007), the highly hydrated nature of EPS can protect
organic matter from degradation, sealing the OM with macromolecules resistant to strong acids

and acid hydrolyses (Goth et al., 1988; Tagelaar et al., 1989; Pacton et al., 2007). The CB samples

demonstrate the recurrent presence of organic layers and fine-grained particles, which indicates

that clay minerals and possible EPS production represent the OM preservation mode for these

Ediacaran strata.

5.3. Eodiagenetic cement and microbial mats

Petrographic analyses reveal eodiagenetic types of cement along the entire succession. But

there is an upward change in terms of early diagenesis processes. The lower units present carbonate

as the dominant eodiagenetic cement, whereas iron oxide coating was commonly observed as the

cement in the upper levels. It is noteworthy that the reduction in carbonate cementation occurred

concomitantly with the decrease in microbial mats and the occurrence of MISS, which suggests

that the early diagenetic process of carbonate cementation was more favorable in deposits that

were rich in microbial mats.

Apparently, early diagenetic processes were related to organic matter content in the

sediments. Biochemical reactions during bacterial degradation of mats can be responsible for

carbonate (calcite, aragonite) precipitation in eodiagenetic processes (Krumbein and Cohen, 1977).

In experiments with modern bacterial mats, Cohen et al. (1977) concluded that carbonates mimic

filament structures and form 1-2 mm wide oncoidal structures. This process involves the fixation

of CO₂ by bacteria and an increase in pH because of bicarbonate production, which creates a

microenvironment around the cells that readily favor carbonate precipitation (Défarge et al., 1994).

Maisano et al. (2020) propose that the EPS can act as nuclei for in situ CaCO3 precipitation. The

presence of calcium carbonate as an eodiagenetic cement, as well as the MISS abundance, are

widespread in the basal unit of Camaquã Basin and decreases from the base to the top of the basin.
The abundant record of MISS associated with the register of early precipitated carbonate cement,

associated with the decrease of both for records to the top of the basin, seems to indicate a relation

between the microbial activity and the carbonate precipitation.

5.4. Geochemical proxies and the depositional environment

Based on previous research (Demaison and Moore, 1980; Tyson, 1987; Hatch and

Leventhal, 1992; Jones and Manning, 1994, Rimmer et al., 2004; Ross and Bustin, 2009; Zhao et

al., 2016), trace element ratios were used to achieve information about paleoredox environment

conditions. According to Zhao et al. (2016), Ni/C ratios between 0-5 indicate oxic conditions,

between 5-7 dysoxic and 7-16 anoxic conditions; V/Cr ratios indicate oxic (0 to 2), dysoxic (2 to

4), or anoxic (4 to 12) states; and Th/U ratios <2 signalize anoxic conditions. The analyzed

geochemical proxies (Ni/Co, V/Cr and Th/U) point to oxic to slightly dysoxic bottom conditions

throughout the entire succession. Based on sedimentological models and petrographic data, all

units display evidence of a strong influence of alluvial and/or fluvial discharge and common

hyperpycnal flows. And some of them present common features related to subaerial exposures.

Besides, widespread iron oxide coatings occur as the first cement around grains along the upper

three units of CB corroborating oxygenated bottom waters. Altogether, these characteristics are

compatible with the geochemical proxies’ results.

Results indicating oxic and dysoxic environments seem coherent with the low phosphorus

content of the analyzed samples. During the Proterozoic, limited phosphorus input and low primary

production are supposed to be a result of the low levels of oxygen in the atmosphere (Laakso and

Schrag, 2014). The results of P analysis pointed to contents between 0.03 and 0.09%, which are

similar to the P content found by Laakso et al. (2020) on Late Ediacaran siliciclastic samples that

presented a mean value of 0.06%.

 The cobalt content found in CB samples presents an upward decrease from 13 ppm in

Maricá Bom Jardim groups to around 4 ppm in Santa Bárbara/Guaritas groups. A similar decrease

trend is also perceptible in MISS types and microbial mats abundance in the basin records. The

presence of Co is related to organic compounds in Upper Vendian shales that contain around 14-

16 ppm, according to Felitsyn et al. (1998). The higher Co values were found in the Maricá and

Bom Jardim groups, which are supposed to record marine and relatively deep-lacustrine settings,

respectively, based on purely sedimentological criteria. These environments are characterized by

sedimentation of fine-grained particles under low energy conditions, therefore more suitable for

microbial mat growth.

Geochemical results indicate low and relatively constant strontium concentrations in all

analyzed samples, indicating no variation of available Sr during the CB evolution. Due to the rock

weathering, isolated use of strontium in ppm concentration is not recommended for marine and

terrestrial differentiation. Nevertheless, the observation of Sr content remains valid as an additional

reference point, offering insights about the environments of a complex Ediacaran basin, as

Camaquã Basin. According to Felitsyn et al. (1998), Sr concentrations are higher in marine (about

500 ppm) than in terrestrial biota because oceanic water comprises much higher Sr concentrations

than fresh water. The analyzed samples present Sr values much lower than 500 ppm, hence

suggesting continental water, exceptionally by one sample of the Maricá Group, which comprises

the only unit assumed as marine, reflecting a possible marine influence.

According to Tribovillard et al. (2006), redox-sensitive trace metals can be used as

paleoredox proxies due to association of some elements (U, V, Mo, Ni Cu, Zn and Cd) with organic

matter. Those elements are more soluble under oxidizing conditions and less soluble under

reducing conditions, producing authigenic enrichments in oxygen-depleted sedimentary facies.

Molybdenum enrichment occurs under anoxic-sulfidic conditions, but in oxic environments Mo

enrichment is limited (Zheng et al., 2000; Algeo and Tribovillard, 2009). Uranium and

molybdenum present similar geochemistry in aqueous conditions, however uranium enrichment

occurs under less intense reducing conditions and shallower depths within the sediment column

that Mo (Morford et al., 2009). Under oxic-suboxic conditions U is soluble as U (VI), chemically

unreactive, and present a limited enrichment (Algeo and Tribovillard, 2009; Morford et al., 2009).

For Precambrian samples, the enrichment factors for Mo and U enrichment were obtained using

the formula XEF = [(X/Al)sample/(X/Al)PAAS], and X and Al stand for the weight concentrations

of element X and Al, respectively (Rico et al., 2019). Samples are normalized using the post‐

Archean average shale compositions of Taylor and McLennan (1985), using average background

values of Mo (0.25 ± 0.05 ppm), U (1.02 ± 0.15 ppm), and Al (3.1 ± 1.1 wt.%; after Chappaz et

al., 2008). The samples from all units of CB Basin shows values of MO and U enrichment between

2 and 0.2 (Table 4). According to Rico et al. (2019) low values indicate oxic to suboxic

environments. The MO and U enrichment values obtained in CB samples indicate an oxic to

dysoxic depositional environment for every CB interval.

Total organic carbon (TOC) content results (Table 5) are similar to values found in other

siliciclastic Precambrian successions (Anbar et al., 2007; Pehr et al., 2018; Duda et al., 2020). The

organic matter is mainly composed of microbial mats and secondary by organic-walled

microfossils. The highest values appear on mudstones from Bom Jardim (1.15%) and Santa

Bárbara (0.55%) groups. The low TOC content indicates oxic conditions during the deposition in

the Camaquã Basin, which are not favorable for organic matter preservation. However, relatively

higher TOC values were found in the Bom Jardim and Santa Bárbara fine-grained samples that

also showed proxies values indicating dysoxic depositional conditions. The organic matter
conservation appears to be higher on clay-rich rocks, supporting the hypothesis that clays act as a

seal, isolating the organic matter from decomposition in CB deposits.

Overall stressing conditions, an upward decrease in life-supporting conditions, and a

marine setting restricted to the lowermost Maricá Group, are geochemical-based readings

compatible with previous sedimentological interpretations. These interpretations indicated an

overall continentalization along the Camaquã Basin, with basal marine deposits changing from

deep to shallow lakes, always associated with significant freshwater discharge. The higher values

of organic matter content in the Bom Jardim Group reinforce the idea of a relatively deeper

lacustrine setting for this unit. Even the deepest portion of the basin, the Maricá Group, presents a

higher quantity and variety of MISS, which suggests that the photic zone reaches the substrate,

suitably to the bibliography, presenting a shallow marine environment.

6. Conclusions

Throughout its record, the Camaquã Basin has impressively maintained the preservation of

microbial mats and related MISS. However, a noticeable decrease in the amount and types of

fossils and sedimentary structures can be observed upwards. This decrease can be attributed to an

increase in subaerial exposure events, which has been proposed in previous sedimentological

literature.

The well-preserved organic matter (microbial mats), associated with the presence of

organic-walled microfossils, is possibly related to the lacustrine depositional conditions. The

commonly described low-concentration turbidity currents and possibly hypopycnal plumes

derived from fluvial discharge could easily bring fine-grained particles to usually quiet water
regions, where microbial mats were fully developed. These mud and clay particles associated with

EPS could seal the organic deposits and avoid their decay.

Geochemical proxies indicate oxic to dysoxic conditions during the deposition of all units.

Oxic conditions are concordant with the sedimentological-based models previously proposed for

the Camaquã Basin as well as petrographic pieces of evidence. Sedimentological models point to

a strong alluvial and/or fluvial influence in all units, hence bringing oxygenated water into the

subaqueous realms. They also indicate an upward increase in the frequency of subaerial exposure

events, which also enhances oxic conditions. Early diagenetic iron coatings are also present in

almost all units and also corroborate oxygenated lake floors.

Understanding microbial life's interaction with the continental environment requires the

study of Precambrian lacustrine rocks that house well-preserved microbial communities. The

Camaquã Basin, which developed during a time of significant changes in the complexity and

diversity of life, provides valuable insight into this field. The petrographic and geochemical

conditions that preserved organic forms in the basin indicate an oxygen-rich environment, despite

the challenges posed by microscopic life spreading in such conditions. The clay content and EPS

presence play a crucial role in preserving organic matter. The microbial mats discussed in this

context depict ancient life occupying stressful continental niches of the Proto-Gondwana during

the Proterozoic/Phanerozoic transition.

Acknowledgements

The first author thanks CAPES for her scholarship (grant number #88887.150701/2017-

00). The second author acknowledges CNPq for the long-term support (grant number
#311175/2019-8). The third author acknowledges the National Council for Scientific and

Technological Development – CNPq for financial support (grant #301441/2019-7). The authors

are grateful to UNISINOS, CNPEM and UCSB for providing lab facilities (SEM and X-ray) and

field equipment. We thank Dr. Susannah Porter (UCBS) for her valuable assistance and review of

this MS.

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Figure Captions

Fig. 1. Geological map of the Sul-Riograndense Shield, indicating the Camaquã Basin location. The circles indicate

the sites where samples were collected. Modified from CPRM (1995), according to Philipp et al. (2016).
Fig. 2. Outcrops of Camaquã Basin where the samples were collected. (a) Thin layers of intercalated claystone and siltstone from

Maricá Group; (b) mudstone of Bom Jardim Group with MISS on the top; (c) mudstone layers of Santa Bárbara Group; (d) layers

of mudstone between sandstone layers on Guaritas Group.

Fig. 3. Sedimentary logs of the Maricá (a), Bom Jardim (b), Santa Bárbara (c), and Guaritas (d) groups indicating

stratigraphic divisions and facies association for each unit. Log b based on Janikian et al. (2003) and log c based on

Paim et al (2000) and Lehn et al. (2018), and log d based on Paim and Scherer (2007).
Fig. 4. Hand samples with examples of different types of MISS found in the Maricá Group. (a) Arumberia-type; (b)

elephant skin-type; (c) reticulated marks; (d) gas escape marks.

Fig. 5. Hand samples with examples of different MISS types identified in the Maricá and Bom Jardim groups. (a)

Reticulated ridges (Maricá Group); (b) leveling-type (Bom Jardim Group); (c) grain prison + setulfs (Bom Jardim

Group); (d) ridges and mounds (Bom Jardim Group).

Fig. 6. Hand samples from Santa Bárbara and Guaritas Group demonstrating examples of different MISS types found

in these groups. (a) Bubbles (Santa Bárbara Group); (b) cuspate mounds (Santa Bárbara Group); (c) wrinkle structure

(Santa Bárbara Group); (d) wrinkle structure (Guaritas Group).

Fig. 7. Photomicrographs of fine-grained facies with microbial mats. (a) Example from the Maricá Group containing

microbial filaments surrounding grains, photomicrograph under natural light. Green arrows indicate microbial

filaments. (b) Example from the Bom Jardim Group showing a microbial colony around siliciclastic grains and other

organic forms, obtained using natural light. A red arrow points to a possible acritarch. (c-d) Microbial mat on a layer

from the Santa Bárbara Group on polarized light (c) and natural light (d). € Microbial mats as intraclasts in fine-

grained sandstones of Santa Bárbara Group. Blue arrows indicate microbial fragments. (f) Filaments (indicated by

green arrows) and fragments (indicated by blue arrows) of mats in fine-grained sandstones of the Guaritas Group,

photomicrograph under natural light. Photomicrographs a and b modified from Lehn et al. (2019).
Fig. 8. Photomicrographs of microbial mat fragments recovered through palynological extraction with a view under

natural light. (a) Maricá Group; (b) Bom Jardim Group; (c) filamentous mat fragment from Santa Bárbara Group; (d)

Guaritas Group.
Fig. 9. SEM images of microbial mats in thin sections (a-c) and as isolated fragments of palynological extraction (d-

f). (a) Detail of an organic-rich layer, interpreted as a microbial mat from Maricá Group; (b) detail of organic layers

understood as a microbial mat from Santa Bárbara Group; (c) feature of an organic-rich layer interpreted as a microbial

mat in a thin section from Santa Bárbara Group; (d) organic fragment interpreted as a microbial mat fragment from

Bom Jardim Group; (e) small scale, roll-up clast from the microbial mats of Maricá Group; and (f) microbial mat

fragment from Maricá Group.

Fig. 10. SEM images and EDS elemental mapping of two samples of fine-grained rocks, including siliciclastic material

and microbial mats. (a) Note the composition of the microbial mat and oxygen, siliceous, and aluminum associated

with siliciclastic particles; (b) In situ, parallel to the bedding plane microbial mat. Notice the carbon content related to

the microbial mat and oxygen, aluminum, silicon, and iron content related to the clay particles.
Fig. 11. Photomicrographs from thin sections showing the most common cement in each unit from CB. (a) Claystone

thin section from Maricá Group in natural light (NL). Yellow arrows pointing to calcite cement. (b) Same as (a) in

polarized light (PL). (c) Siltstone from Bom Jardim Group; viewed in NL. (d) Same image as (c) in PL. (e) Fine-

grained sandstone from Santa Bárbara Group, viewed in NL. Red arrows indicate iron coating as the first eodiagenetic

cement. (f) Same as (e) in PL. (g) Very fine-grained sandstone from Guaritas Group. Red arrows indicate iron coating

cement. (h) Same as (g) in PL.

Fig. 12. Crossplots of paleoredox proxies. (a) Ni/Co vs. Th/U; (b) V/Cr vs. Ni/Co; (c) V/Cr vs. Th/U. Ranges of redox

proxies from Hatch and Leventhal (1992), Jones and Manning (1994), and Zhao et al. (2016).
Fig. 13. Elements content in CB samples. (a) P content on each sample; (c) Sr content on each sample; (c) Co content

on each sample.
Fig. 14. Discoidal structures from Santa Bárbara Group. (a) Detail of bubbles-type of MISS, located right above the

organic-rich layer. Sample from Maricá Group. (b) Photomicrography of bubbles-type of MISS of Maricá Group.

Thin section parallel to bedding showing the circular morphology of filamentous mats. (c) Raindrop and hailstone

impression forms. Rd – rain-print depth; RD – rain-print diameter; Hd – hailstone imprint depth; HD – hailstone

imprint diameter; ERh – external rim height; ERw – external rim width (modified from Remin et al., 2014). (d) Sample

of very fine-grained sandstones with raindrops from Santa Bárbara Group.

Fig. 15. Examples of MISS and palynological material. A comparison between (a) discoidal features in positive relief

from siltstone top surface of Diabaig Formation (Callow and Brasier, 2009) and (b) discoidal structures in mudstone

surface related to gas escape from Bom Jardim Group. (c-f) Examples of similar microbial fragments recovered with

palynological extraction. (c) large colony of E.lacustrina from Nonesuch Shale (Strother and Wellman, 2016); (d)

Extracted organic material associated with E. lacustrine, from siltstones of Maricá Group; (e) distributed colony of

E.lacustrina in a flat organic fragment from Nonesuch Shale (Strother and Wellman, 2016); (f) Santa Bárbara Group

sample of bacterial fragment associated with samples of Nonesuch Shale.

Table Captions

Table 1 Camaquã Basin facies and facies association. Maricá Group facies adapted from Pelosi et al, 2003; Bom

Jardim Group facies adapted from Janikian et al. (2005). Santa Bárbara Group facies adapted from Lehn et al.

(2018); Guaritas Group facies adapted from Paim and Scherer (2007).

Unit Facies association Facies

Trough-cross bedded conglomerate
Normal graded conglomerate
Fluvial facies Massive conglomerate
Trough-cross bedded conglomeratic sandstone
Trough-cross-bedded sandstone
Ripple cross-bedded sandstone
Climbing-ripple cross-bedded sandstone
Laminated sandstone
Horizontally bedded sandstone
Marine facies
Maricá Group Massive sandstone
Hummocky cross-bedded sandstone
Laminated mudstone
Heterolytic sandstone and siltstone
Massive conglomerate
Trough-cross bedded conglomeratic sandstone
Delta front to delta plain/braided Trough-cross bedded sandstone
fluvial facies Horizontally bedded sandstone
Massive sandstone
Laminated mudstone
Ripple cross-bedded sandstone
Climbing-ripple cross-bedded sandstone
Fan-delta (delta plain) facies
Horizontally bedded sandstone
Trough-cross bedded sandstone
Ripple cross-bedded sandstone
Laminated sandstone
Distal delta front to prodelta facies
Laminated siltstone and claystone
Bom Jardim Group
Massive claystone
Trough-cross bedded sandstone
Horizontally bedded sandstone
Massive sandstone
Delta front facies
Convoluted sandstone
Laminated siltstone and claystone
Massive siltstone
Trough-cross bedded conglomerate
Fluvial facies Trough-cross bedded conglomeratic sandstone
Trough-cross bedded sandstone
Trough-cross bedded sandstone
Santa Bárbara Graded sandstone
Group Inverse- to normal-graded sandstone
Delta front facies Undulated sandstone
Massive sandstone
Laminated sandstone
Mudstone
 Laminated sandstone
Horizontally bedded sandstone
Lacustrine facies Silt-rich sandstone
Clay-rich sandstone
Mudstone
Low-angle cross-bedded sandstone
Trough cross-bedded sandstone
Fluvial-lacustrine facies Wave-rippled sandstone and siltstone
Current-rippled sandstone and siltstone
Guaritas Group Massive sandstone
Dune Large-scale trough cross-bedded sandstone
Trough cross-bedded sandstone
Dry and wet interdune and damp
Inverse graded sandstone
interdune facies
Claystone
Maricá Group facies adapted from Pelosi et al. (2003); Bom Jardim Group facies adapted from Janikian et al. (2005); Santa
Bárbara Group facies adapted from Lehn et al. (2018); Guaritas Group facies adapted from Paim and Scherer (2007).
 Table 2. MISS types on each group of the Camaquã Basin

Maricá Group Bom Jardim Group Santa Bárbara Group Guaritas Group
Bubbles Arumberia-like/leveling Cuspate mounds Wrinkle
Wrinkle Leveling Bubbles marks
Pitted texture Reticulated marks Arumberia-like
Ridges reticulated Bubbles Wrinkle
Grain prison + wrinkle Grain prison
Elephant skin Setulfs
Arumberia Pitted texture
Ridges and mounds
Gas scape
Reticulated structures

Table 3. Geochemical proxies

Sample Unit Lithology Ni/Co V/Cr Th/U La/Sc Sc/Th Cr/Th Co/Th
P156 Guaritas Sandstone 2.50 2.22 4.08 2.80 0.8 2.45 0.84
P145 Guaritas Mudstone 2.29 2.8 5.29 4.95 0.35 0.80 0.5
SB06 SB Sandstone 2.45 2 4.62 2.93 0.76 3.58 1.05
SB17 SB Mudstone 3.23 1.08 4.27 4.59 0.88 6.60 1.11
I53A BJ Sandstone 2.41 1.76 4.03 4.86 0.94 3.79 1.84
PC02 BJ Mudstone 3.04 1.91 4.48 4.68 0.89 2.96 1.11
MD1 Maricá Sandstone 1.92 2.31 6.12 2.55 0.93 4.43 1.64
CG96 Maricá Mudstone 3.17 1.87 4.70 3.79 1.36 4.43 1.56
Table 4. Molybdenum and uranium enrichment values

Sample Unit MoEF UEF

ID

P145 Guaritas Gr. 0,975 0,671

P156 Guaritas Gr. 0,712 0,723

SB06 Santa 1,987 0,422

Bárbara Gr.

SB17 Santa 0,887 0,331

Bárbara Gr.

153A Bom Jardim 1,887 0,671

Gr.

PC02 Bom Jardim 2,011 0,860

Gr.

MD1 Maricá Gr. 0,75 0,197

CG96 Maricá Gr. 0,612 0,498

 Table 5. TOC content

TOC (%)
Sample ID Unit
(%) Range Average
P145 Guaritas Gr. 0.07
P145B Guaritas Gr. 0.07
0.06 to 0.11 0.08
P148 Guaritas Gr. 0.06
P156 Guaritas Gr. 0.11
SB01 Santa Bárbara Gr. 0.12
SB04 Santa Bárbara Gr. 0.09
0.06 to 0.55 0.21
SB06 Santa Bárbara Gr. 0.06
SB17 Santa Bárbara Gr. 0.55
PC02 Bom Jardim Gr. 0.08
153A03 Bom Jardim Gr. 0.09
0.08 to 1.15 0.35
153A04 Bom Jardim Gr. 1.15
153B Bom Jardim Gr. 0.09
CG33 Maricá Gr. 0.10
CG45 Maricá Gr. 0.10
0.08 to 0.10 0.09
CG96 Maricá Gr. 0.09
MD1 Maricá Gr. 0.08
Graphical Abstract

Credit authorship contribution statement

Ilana Lehn: Conceptualization, Project administration, Methodology, Investigation, Software, Writing –

Original draft preparation. Paulo Sérgio Gomes Paim: Methodology, Writing – Reviewing and Editing,

Supervision, Validation. Farid Chemale Jr: Methodology, Supervision, Validation, Funding acquisition,

Writing – Reviewing and Editing.

Declaration of interests

☒ The authors declare that they have no known competing financial interests or personal relationships
that could have appeared to influence the work reported in this paper.

☐ The authors declare the following financial interests/personal relationships which may be considered
as potential competing interests: