Characterization of modern dolomite stromatolites from

hypersaline Petukhovskoe Soda Lake, Russia

OLGA S. SAMYLINA AND LYUBOV V. ZAYTSEVA

Samylina, O. S. & Zaytseva, L. V. 2019: Characterization of modern dolomite

stromatolites from hypersaline Petukhovskoe Soda Lake, Russia. Lethaia, Vol. 52, pp.
1–13.

Microbialites occur in a number of modern soda lakes, but the formation of dolomite
stromatolites in these environments is rare. Microbial metabolic activities are consid-
ered to influence precipitation of carbonate minerals (including dolomite) in marine,
hypersaline and slightly alkaline environments due to their capacity to buffer pH of
the local environmental settings. Hydrochemical features of hypersaline alkaline Petu-
khovskoe Soda Lake are already favourable for the precipitation of carbonate minerals,
and the metabolic activities of microorganisms may not be significant in this type of
environment. Stromatolites from this lake provide an excellent opportunity for the
study of recent natural dolomite formation in the highly saturated soda environment.
To understand the genesis of these stromatolites and to provide insight into environ-
mental aspects of its formation, we have made a detailed structural and mineralogical
description with the use of light and scanning electron microscopy, infrared spec-
troscopy and X-ray analysis. Due to the special fluctuating conditions of sedimenta-
tion in the lake, stromatolites contain both layers with chemogenic precipitation of
minerals and layers with fossilized algal, cyanobacterial and coccoid bacterial forms.
The presence of dolomite, calcite and Mg calcites with different content of magnesium
is detected. The primary independent binding of Mg and Si with the following binding
of Ca in the pericellular and released exopolysaccharides is shown. Extracellular poly-
meric substances (EPS) and fluctuating hydrological regimen of the Petukhovskoe
Soda Lake are considered to play the key role in the precipitation of Ca–Mg carbonates
including dolomite. □ Ca–Mg carbonates, dolomite, extracellular polymeric substances,
soda lake, stromatolite.

Olga S. Samylina ✉ [olga.samylina@gmail.com], Laboratory of Relict Microbial Commu-

nities, Winogradsky Institute of Microbiology, Research Center of Biotechnology of the
Russian Academy of Sciences, Moscow 119071, Russia; Lyubov V. Zaytseva [l.zaytseva@
mail.ru], Department of Analytical Instruments, Borissiak Paleontological Institute of
Russian Academy of Sciences, Moscow 117647, Russia; manuscript received on 14/12/
2017; manuscript accepted on 13/05/2018.

Non-marine carbonate build-ups originate in large
spectrum of terrestrial subaerially exposed and sub-
merged aquatic environments, including different
kinds of lakes (fresh water, saline and alkaline).
There are different types of carbonate build-ups
within lakes depending on depositional settings: sal-
ine and alkaline lake’s margin carbonate build-ups,
sublacustrine spring mounds and pinnacles,
hydrothermal travertines and fluvial tufas (Della
Porta 2015). A combination of organomineralization
(s.l. Dupraz et al. 2009) and inorganic processes dri-
ven by physicochemical properties of environments
leads to the formation of carbonate build-ups.
Saline alkaline and soda lakes are characterized by
deposition of different carbonates. The common
carbonates described in microbialites from the well-
studied soda lakes of the East African Rift (Magadi,
Natron, Bogoria and others), Turkey (Van, Salda),
USA (Mono Lake), China (lakes Nuoertu and Huhe-
jaran) and Mexico (Alchichica) are calcite, aragonite,
low-Mg calcite and hydromagnesite (Kempe et al.
1991; Casanova 1994; Arp et al. 1998; L
opez-Garc
ıa
et al. 2005; Souza-Egipsy et al. 2005; G
erard et al.
2013). Precipitation of dolomite in soda lakes is also
a well-known fact (Dapeng 1987; Renaut & Jones
2011; Brady et al. 2013; Della Porta 2015). The soda
lakes of Kulunda Steppe, formed in cryo-arid conti-
nental climate, are understudied regarding the pres-
ence of microbialites. But it was shown that calcite
and dolomite predominate among the insoluble car-
bonate minerals in salt sediments of Kulunda Steppe
lakes of different hydrochemistry, including soda
lakes (Lebedeva (Verba) et al. 2008). A small margin
carbonate build-ups with the predominance of stoi-
chiometric dolomite and Ca dolomite were found in
the Petukhovskoe Soda Lake. These build-ups repre-
sented laminated crusts formed in shallow brine
conditions with the participation of algo-cyano-bac-
terial communities. It allowed us to define them as
modern stromatolites (Samylina et al. 2016), but the
detailed description of the peculiarities of different
layers was not done.

DOI 10.1111/let.12286 © 2018 Lethaia Foundation. Published by John Wiley & Sons Ltd
2 O. S. Samylina & L. V. Zaytseva
Epicontinental soda lakes are considered to be
widely spread in Archaean and Proterozoic. They
could be significant for the early evolution of life
as assumed in the hypothesis of ‘soda continent’
(Zavarzin 1993, 2003; St€ ueken et al. 2015). Dolo-
mite-depositing soda lakes can be considered as
modern representatives of ancient types of lake
basins, forming a ‘hydrological relic’ that aids in
understanding the specific features of sedimenta-
tion in mineralized basins of arid regions. Despite
a numerous discussions on the importance of
microbial metabolic activity for precipitation of
carbonate minerals in different environments (Pet-
rash et al. 2017), the specific role of microorgan-
isms in deposition of minerals in highly saturated
soda lakes is still poor understood. The problem of
precipitation of stromatolite-like sedimentary lay-
ered structures is widely discussed in Precambrian
publications (e.g. Grotzinger 1989; Bartley et al.
2000; Sharma & Sergeev 2004; Knoll et al. 2016).
It is argued that in some Precambrian-restricted
basins with high carbonate alkalinity, the totally
inorganic precipitation of carbonates occurs. The
example of Petukhovskoe Soda Lake is unique for
possibly total inorganic carbonate precipitation
and could be compared to some Precambrian-
restricted basins with high saturation of dissolved
inorganic carbonate. Thus, the purpose of this
study is a detailed description of structural and
mineralogical peculiarities of modern stromatolites
from Petukhovskoe Soda Lake and discussion of
possible role of microorganisms in deposition of
carbonate minerals in the highly saturated soda
environment.
Materials and methods
Geographical setting of the sample site
The Kulunda Steppe is located in the southern part
of the West Siberian Plain between the rivers Ob and
Irtysh, along the Kazakhstan–Russia border. The
area contains numerous of salt lakes with varying
salinity and chemical composition, including chlo-
ride–sulphate and soda types. The soda lakes are
located in its southern part.
The Petukhovskoe Soda Lake is located in the
Klyuchevskoi area of Altai Region (Russia), at coor-
dinates 52°60 20.52″N, 79°90 22.19″E. Although
known as Petukhovskoe Sodovoe in Russian, it is
variously referred to by other names in English-lan-
guage articles: Petukhovskoe Soda Lake (Lebedeva
(Verba) et al. 2008), Petukhovo (Kompantseva et al.
2009, 2010) or Cock soda lake (Foti et al. 2007,
LETHAIA 52 (2019)
2008; Sorokin et al. 2010, 2011a,b; Samylina et al.
2014, 2016).
The first published description of the lake was
made in 1927 with respect to its trona deposits
(Gebler 1927). It is an undrained shallow lake
formed in a depression, subjected to periodic drying
and irrigation. The main source of feeding is
groundwater, but some contribution comes from
surface run-off and atmospheric condensation. The
lake brines belong to the carbonate (soda) type. The
sediments of the lake are represented by grey clays
with a high content of sand on the northern coast
and arkosic sands on the south coast. The subsurface
sediments are highly reduced and contain molar
concentrations of acid labile sulphides (HS + FeS).
The lake is surrounded by sand dunes with conifer-
ous forest. The diversity of eukaryotic organisms in
the lake is relatively low: there are few species of
green algae and diatoms, higher plants are absent,
and invertebrates are represented by Ephydra larvae
and Artemia sp. (Samylina et al. 2014).
Hydrochemical measurements
Salinity and pH of the brines were measured using a
WTW field potentiometer–conductometer. For the
hypersaline conditions, the average pH values
between the native brine and its 1:5 dilution was
used. The soluble carbonate alkalinity was deter-
mined in the field by a two-step titration with 1 M
HCl. The content of Si, Mg and Ca was measured on
atomic emission spectrometer Optima-4300 DV
(‘Perkin-Elmer’, USA).
Structure and composition of the stromatolitic
crusts
Study of the surface and polished cross-sections of
stromatolitic crusts was performed using a light
stereomicroscope LEICA M165C. Examination of
the samples by scanning electron microscope (SEM)
with EDX (energy dispersive X-ray spectral micro-
analysis) was carried out using EVO-50 Zeiss elec-
tron microscope equipped with the microanalyser
INCA Oxford 350 at 15–20 kV. Part of the sample
was treated with 0.1 M HCl for the dissolution of
the surface layer of microsections, and another part
was examined as an intact stromatolitic crust. All
samples were gold-coated (15 nm thickness).
Semi-quantitative elemental analysis of micron size,
and smaller sites was carried out using EDX
spectrometry.
Determination of mineral composition of the
stromatolitic crust by Fourier transform infrared
spectroscopy (FT-IR) was conducted on a FT/IR-
LETHAIA 52 (2019) Petukhovskoe Soda Lake stromatolites 3

4100 spectrometer using the method of pressed Table 1. Salinity, pH and alkalinity of Petukhovskoe Soda Lake
brines, measured in July–August of each year (data for 2002–
tablets with potassium bromide and the Universal 2010 obtained from D. Yu. Sorokin, 2011–2016, own data
Sampling Archaean PIKE MIRacle. The IR-active obtained in collaboration with D. Yu. Sorokin).
compounds were identified by comparing the
obtained spectrum with the spectra of reference Alkalinity (M)
minerals. The characteristic bands of calcite are 1430 Year Salinity (g/l) pH Total CO2
3
(m3), 873 (m2) and 712 (m4)/cm. Dolomite is charac-
terized by following bands: 1440 (m3), 882 (m2) and 2002 54 9.93 0.9 n/d
2003 63 10.1 0.8 n/d
729 (m4)/cm. Magnesium calcite with different con- 2005 65 10.13 0.44 0.31
tent of Mg is identified by the position of the spec- 2006 60 10.3 0.9 0.85
trum band m4 from 712 to 728/cm, which correlates 2008 120 10.36 1.16 1.0
2009 120 10.3 0.82 0.62
with the amount of Mg. Silicates are characterized 2010 70 10.1 0.68 0.34
by the following position of the characteristic 2011 100 10.18 1.11 0.96
absorption bands: 1100–900, 830–740 and 500–400/ 2012 200 9.8 2.7 2.4
2013 50 9.6 0.65 0.5
cm. Amorphous silicates exhibit broad diffuse 2014 65 10.03 0.45 0.35
absorption bands without distinct maxima. Cryp- 2015 85 10.1 0.6 0.56
tocrystalline silica species, including opal, can repre- 2016 55 9.92 0.62 0.56
sent a mixture of a-quartz, a-cristobalite and a-
tridymite with a noticeable presence of molecular
water (Sklyarov et al. 2010).
X-ray analysis (XRD) was held on diffractometer
Advance D8. Detailed mineralogy of carbonates
Structure and mineralogical composition of the
assumes the determination of the position of most
stromatolites
intense reflection (interplanar distances hkl = 104) Stromatolites found in Petukhovskoe Soda Lake in
of the trigonal carbonates in the diffraction patterns. 2011–2012 represent the perforated and cavernous
The magnesium calcites hkl are located between the crusts with alternation of thin mineralized layers of
corresponding values of calcite and stoichiometric different colour and structure (Fig. 1). From one to
dolomite. These values are in the range from d104 several alternations of light and dark layers can be
3.036 (calcite) to d104 2.889 (stoichiometric dolo- distinguished in different samples. The development
mite) and can serve as a semi-quantitative measure of algo-cyano-bacterial community occurs on its
of Mg content in carbonates (after Lumsden 1979). surface (Fig. 2). The usual for Kulunda Steppe soda
lakes’ dominant filamentous benthic cyanobacteria
belonging to the genera Geitlerinema and Nodosi-
Results linea (Oscillatoriales) and the green filamentous
algae Ctenocladus circinnatus are identified in the
community (Fig. 2) (Samylina et al. 2014). The
Hydrochemical features of Petukhovskoe Soda
presence of mineralized cyanobacteria, bacteria and
Lake
algae in the interior of stromatolites was shown pre-
The lake is hypersaline and alkaline. The total salin- viously (Samylina et al. 2016). Visually, the stroma-
ity of the water, monitored from 2002 to 2016 (in tolites consist of intermittent layers of different
July–August of each year), varied from 50 to 200 g/l colour and texture: dark grey and light brown layers,
wherein the pH remained highly alkaline (9.6– white thin layers and a bottom layer with predomi-
10.36). Stable alkaline pH values are due to high nance of clastic material.
total soluble carbonate alkalinity which varied from The use of EDX analysis allowed identifying the
0.45 to 2.7 M HCO 3 þ CO3 during the same per-
2
distribution of chemical elements across the profile
iod (Table 1). In the 2011–2012, when the samples of stromatolite and in cross-sectional area that also
were collected, the lake was at the stage of maximum includes all of the layers (Fig. 3). The dominant ele-
salinity: TDS 100 g/l, 1.11 M HCO 3 þ CO3 , pH
2
ments are Ca, Mg and Si. Fe and Al are also found in
10.18, in 2011 and TDS 200 g/l, 2.7 M a small amount, especially in the layer of the cemen-
HCO 3 þ CO3 , pH 9.8, in 2012. The content of Si
2
ted clastic material. The high content of Si is charac-
(3.02 mg/l), Mg (18.5 mg/l) and Ca (1.89 mg/l) was teristic for soda lakes as a result of leaching from
measured in 2016 when the lake brine was relatively clastic material at highly alkaline conditions (Shvart-
diluted (TDS 55 g/l, pH 9.9, 0.62 M sev et al. 2014). The Ca and Mg supply to the brine
HCO 3 þ CO 2
3 ). most probably occurs through groundwater seepages
4 O. S. Samylina & L. V. Zaytseva
Fig. 1. Modern stromatolites from Petukhovskoe Soda Lake. A, surface view. B, microsection of a stromatolite as a whole. [Colour figure
can be viewed at wileyonlinelibrary.com]
Fig. 2. Algo-cyano-bacterial community that develops on the surface of a stromatolite from Petukhovskoe Soda Lake. A, SEM micro-
graph. B, cyanobacteria Geitlerinema sp. (G) and Nodosilinea sp. (N). C, filamentous algae Ctenocladus circinnatus.
and may have seasonal quantitative fluctuations as it
has been shown for the Tanatar soda lake system in
Kulunda Steppe (Telentyuk 1952). Chlorophylls and
bacteriochlorophylls originated from the decompos-
ing phototrophic biomass which accumulates in sig-
nificant quantities in Petukhovskoe Soda Lake
(Samylina et al. 2014) may also serve as a source of
Mg during the formation of stromatolites.
The distribution of Mg, Ca and Si across the pro-
file and in cross-sectional area of stromatolite seems
to be intriguing. Mg is uniformly distributed over
the cross-sectional area at an average value of 6–7
atomic % (Fig. 3). In contrast, the distribution of Ca
is clearly zonal localizing in the white layers defined
as calcite (approximately 1.4 and 3.2 mm). Within
the range of 1.6–3 mm, an almost synchronous
change of Mg and Ca peaks is observed, and the
atomic ratio of these elements is close to 1:1. This
region corresponds to a dark grey layer, where
numerous mineralized microorganisms are detected
(see below). Asynchronism of Mg and Ca peaks in
the area of 0.5–1.2 mm can be explained by the abil-
ity of these elements to interact differently with EPS
localized within the bottom layer of clastic material
(Fig. 3).
Silicon is present throughout the profile because
of silicification of biota and organic matrix. The dis-
tribution of Si has a number of features. High con-
tent in the bottom of the crust (0–2 mm) is due to
LETHAIA 52 (2019)
the presence of large amounts of clastic material. In
the upper part of the profile, the amorphous silica
may be present.
The presence of Ca–Mg carbonates with different
ratios of Ca and Mg and silicates of different compo-
sition and degree of crystallinity was established in
mineralogical composition of stromatolite by IR
spectroscopy and X-ray diffraction.
According to the IR spectrum of Petukhovskoe
Soda Lake stromatolite (Fig. 4), the presence of
dolomite, calcite and magnesium calcites with differ-
ent content of Mg was determined, as well as various
forms of cryptocrystalline silica. The cryptocrys-
talline silica is possibly representing some forms of
opal or chalcedony, which occur in plenty in Pre-
cambrian shallow water dolomites as lenses and
noodles of cherts and contain abundant remains of
cyanobacteria. But the presence of clastic silicate
inclusions of different composition in the crusts
under study makes it difficult to identify accurately.
The XRD spectroscopy showed the presence of fol-
lowing carbonate minerals: stoichiometric dolomite
(d104 2.889), Ca dolomite (d104 2.897), high-magne-
sium calcites with varying content of Mg (d104 2.908
and 2.935) and low-magnesium calcite (d104 3.011–
3.004) (Table 2). The presence and positions of the
principle dolomite ordering reflections (101, 015,
021) confirm that dolomite in the studied stromato-
lite was ordered (Table 3). The presence of aragonite
LETHAIA 52 (2019) Petukhovskoe Soda Lake stromatolites 5

Fig. 3. Distribution of Mg, Ca and Si across the profile and in cross-sectional area of stromatolite. Quantification of the elements in
cross-sections is given in the brightness gradient from black (absence) to white (maximum presence).

in trace amounts cannot be excluded (d111 3.38–
3.42). Quartz and feldspars, as a part of clastic mate-
rial, were also detected.
SEM investigation with EDX analysis allowed to
study the composition and structure of each layer at
a micrometre scale.
Layers with clastic material
Clastic material can be observed in a certain amount
throughout the vertical profile of stromatolite, but it
is mainly concentrated in the bottom part (Fig. 1).
This layer includes quartz and feldspar grains of
varying degrees of roundness (particle size ranging
from 25 to 300 lm).
Clastic material is cemented by brown matrix
(Fig. 5). This matrix is a complex precipitate, and its
formation is due to both chemogenic and biogenic
processes. The participation of biota in the matrix
formation is proved by the presence of mineralized
cyanobacterial filaments entangling grains in this
layer (Fig. 5D–F). The EPS released by microbial
community is exposed to mineralization impact of
the lake brine, and subsequently penetrates through
6 O. S. Samylina & L. V. Zaytseva LETHAIA 52 (2019)

Fig. 4. The IR spectrum of a stromatolite (a powder of the crushed stromatolite). [Colour figure can be viewed at wileyonlinelibrary.
com]

Table 2. XRD spectrum parameters of Petukhovskoe Soda Lake content in a penetrating mass at the base of
stromatolite (powder) in the range 29.00–32.57 2Θ°CuKa. microfracture (Fig. 5H–I, point No 2) may serve as
Peak evidence of the later accumulation of Ca. High silica
d104 intensity content in the matrix is due to solubilization of the
Carbonates N° 2Θ°CuKa (
A) (%) underlying Si-containing minerals by the alkaline
Stoichiometric dolomite 1 30.95 2.889 100 brine of soda lake. The formation of interstices in
Ca dolomite 2 30.86 2.897 93 the sand grains caused by this process is illustrated
High-magnesium calcites 3 30.74 2.908 71 in Figure 5G.
4 30.46 2.935 23
Low-magnesium calcites 5 29.74 3.004 11
6 29.67 3.011 11
Light brown and dark grey layers
The stromatolite body mainly consists of alternating
light brown and dark grey layers. The EDX analysis
microfractures and interstices, filling them (Fig. 5G– showed that the dark grey layers contained more
I). The chemical composition of the penetrating total carbon (approximately 2%) than the brown
mass contains an abundance of Mg and Si, but lacks ones. We consider this indicates the higher organic
Ca (Fig. 5H–I, point No 3). The illustration of such content in dark grey layer compared with brown.
process can be also seen in the range from 0.5 to The difference in colour is probably due to high con-
1.2 mm in the profile (Fig. 3). High Ca content in tent of organic matter in combination with silica in
the cement (Fig. 5H–I, point No 1) outside of the dark grey layer and microimpurities of iron hydrox-
quartz grain (Fig. 5H–I, point No 4) and its slight ides in the brown layer.

Table 3. Unit cell and XRD information for dolomite from Petukhovskoe Soda Lake stromatolite in comparison with dolomite stan-
dards (CuKa radiation).

Dolomite standards

Petukhovskoe Soda Lake stromatolite PDF 36-0426 Ideal dolomite

Hexagonal unit
cell (hkl) 2Θ° d(
A) I 2Θ° d(
A) I 2Θ° d(
A) I

101 21.972 4.040 4.5 22.022 4.033 1.0 22.04 4.03 <5
012 24.176 3.681 5.5 24.038 3.699 4.0 24.10 3.69 5
104 30.851 2.898 100.0 30.938 2.888 100.0 30.96 2.886 100
006 33.459 2.678 4.9 33.536 2.670 4.0 33.54 2.670 10
015 35.267 2.544 13.4 35.321 2.539 3.0 35.31 2.540 10
110 37.447 2.402 13.1 37.376 2.404 7.0 37.36 2.405 10
113 41.147 2.193 34.3 41.127 2.193 19.0 41.15 2.192 30
021 43.717 2.071 3.1 43.804 2.065 3.0 43.78 2.066 5

Abbreviations: PDF 36-0426, Powder diffraction file PDF 36-0426; ideal dolomite, according to Kaczmarek et al. 2017; 2Θ°, XRD peak
location; d(
A), d-spacing (angstroms); I, relative XRD peak intensity.
LETHAIA 52 (2019) Petukhovskoe Soda Lake stromatolites 7

Fig. 5. Layer with the inclusion of clastic material. A, light micrograph; B, SEM micrograph; C, elemental composition of clastic material
(No. 1–feldspar grain, No. 2–quartz grain) and mineralized matrix (No. 3); D, imprint of the grain with mineralized cyanobacterial fila-
ments (E) and the diagram of its elemental composition (F) at the point marked with an arrow; G, area of the dissolution of clastic mate-
rial (indicated by arrows); H, penetration of mineralized exopolysaccharide through microfractures in the clastic material and elemental
composition of penetrating mass (I) at the points Nos. 1–4.

The massive presence of mineralized biota repre-
sented primarily by fossilized cyanobacterial fila-
ments, Ctenocladus circinnatus, and coccoid forms
most likely of bacterial origin was revealed in the
dark grey layer during SEM investigation (Fig. 6).
This algo-cyano-bacterial community secretes
EPS, presented in two forms: either as pericellular
structures (sheaths and capsules) or as unstructured
diffuse mucilage, which is found in all layers of stro-
matolites as mineralized matrix. Pericellular EPS are
mineralized primarily by Mg and Si and are depleted
in calcium: Si:Mg:Ca (atomic %) = 4.8:4.0:0.1
(Fig. 6A), 5.7:4.3:0.3 (Fig. 6C) and 8.5:6.8:0.0
(Fig. 6D). SEM also revealed the formation of rhom-
bohedral crystals in contact with the mineralized
EPS. Figure 6D shows the formation of crystals in
contact with the mineralized sheaths of cyanobacte-
ria. The fact that the crystals in the zone of contact
with EPS often skirt filaments of cyanobacteria and
maintain connection with EPS (indicated by black
arrows) allows suggesting that crystal growth begins
by interacting with EPS. The formation of crystals in
contact with mucous matrix of bacterial colonies is
illustrated in Figure 6D. Border contact is also
clearly visible (arrows).
Rhombic crystals are characteristic for Ca–Mg
carbonates. Ca–Mg carbonates of different composi-
tion are determined in contact with EPS in the sam-
ple of stromatolite by EDX analysis: high-Mg calcite
(4–40% MgCO3), Ca dolomite (40–50% MgCO3)
and dolomite (50% MgCO3) (Fig. 7). This agrees
with the results of IR spectroscopy and XRD
analysis.
White layers
From one to four white layers with a variable thick-
ness (from a few lm to 200–400 lm) can be present
in the microsections of stromatolitic crusts (Fig. 1).
White layers are wavy and may interlace with one
another. A zone with high density of Ca was detected
by EDX mapping (Fig. 3). The SEM investigation
revealed a layer containing elongated radial axial
crystals (Fig. 8). The mineral composition of this
layer identified with EDX analysis is defined as cal-
cite with low Mg and Sr content: 0.22 and 0.69
8 O. S. Samylina & L. V. Zaytseva LETHAIA 52 (2019)

Fig. 6. Mineralized microorganisms and algae in the stromatolite from Petukhovskoe Soda Lake. A, cyanobacteria; B, Ctenocladus circin-
natus; C, bacterial colony; D, E, formation of rhombohedral crystals in contact (indicated by black arrows) with EPS of cyanobacterial cell
wall (D) and bacteria (E). White arrows indicate the points in the pericellular EPS where the ratio Si:Mg:Ca is measured (see text).

Fig. 7. SEM micrograph and elemental composition (in the table) of Ca–Mg carbonates in the contact with the EPS: 1, 2–magnesium
calcite, 3–Ca dolomite, 4–dolomite.

atomic %, respectively. Strontium is a common minerals (biologically induced organomineralization

companion of calcium in geochemical processes and
can be included as an isomorphous impurity in the
crystal lattice of calcium minerals. The presence of
biota is not detected in this layer. This may indicate
its chemogenic origin.
Discussion
Dolomite precipitation in marine, hypersaline and
slightly alkaline environments is widely discussed.
Microorganisms inhabiting microbial mats are con-
sidered to mediate dolomite and different Mg cal-
cites formation through an influence of their
metabolic activity on precipitation of carbonate
sensu Dupraz et al. 2009). Different types of meta-
bolic pathways are discussed to be related to dolo-
mite precipitation: sulphide oxidation, sulphate
reduction, phototrophic CO2 assimilation, methano-
genesis coupled to anaerobic oxidation of methane
and even aerobic heterotrophic metabolism (Oppen-
heimer & Master 1965; Vasconcelos et al. 1995; Vas-
concelos & McKenzie 1997; Moreira et al. 2004;
S
anchez-Rom
an et al. 2008, 2009; Kaczmarek et al.
2017; Petrash et al. 2017).
We have found fossilized microorganisms in
different layers of the Petukhovskoe Soda Lake
stromatolites. The possibility of morphological iden-
tification of these microorganisms is limited mainly
by large forms, such as cyanobacteria and
LETHAIA 52 (2019)
Fig. 8. SEM micrograph (A, B) and elemental composition (C) of white layers in the Petukhovskoe Soda Lake stromatolite.
filamentous algae (Fig. 2). However, it should be
understood that the microbial community inhabit-
ing lake brines and sediments and probably able
to participate in the formation of stromatolites
includes phylogenetically and functionally diverse
microorganisms, which are studied quite well
(Sorokin et al. 2014, 2015b). The phototrophic
prokaryotes are represented by oxygenic cyanobac-
teria (dominated by Geitlerinema, Nodosilinea)
and anoxygenic Chromatiaceae (Thiorhodospira,
Thiorhodovibrio) and Ectothiorhodospiraceae (Halorho-
dospira and Ectothiorhodospira) (Kompantseva et al.
2010; Samylina et al. 2014).
The rates of oxygenic and anoxygenic photosyn-
thesis in the soda lakes of Kulunda Steppe (0.62 and
0.12 g C/(m2 per day), respectively, for Petukhovs-
koe) are close to the moderately saline soda lakes of
the Transbaikal Region and Mongolian Plateau and
much inferior to the eastern African and American
soda lakes (Kompantseva et al. 2009; Namsaraev
et al. 2015). The sulphur cycle is also active (Sorokin
et al. 2011a). It involves microorganisms using vari-
ous inorganic sulphur compounds either as electron
donors (lithotrophic sulphur-oxidizing prokaryotes,
anoxygenic phototrophic bacteria) or as electron
acceptors (sulphate-reducing prokaryotes). The sul-
phur-oxidizing bacteria from Kulunda Steppe soda
lakes belong to three genera within the Gammapro-
teobacteria: Thioalkalimicrobium, Thioalkalivibrio
and Thioalkalibacter. These genera include mainly
obligately alkaliphilic and some facultatively alka-
liphilic strains. Sulphate-reducing bacteria (SRB)
and their activity in the Petukhovskoe Soda Lake
were also studied (Foti et al. 2007, 2008; Sorokin
et al. 2011b). The sulphate reduction rates (SRR)
attained 116 lmol SO2 3
4 (dm per day) in the upper
2 cm of sediments. Culturable SRB from the Petu-
khovskoe Soda Lake belong to species Desulfona-
tronum thioautotrophicum, Desulfonatronovibrio
thiodismutans and Desulfonatronovibrio magnus.
They are obligately alkaliphilic with a pH optimum
at 9.5–10 and moderately salt tolerant. They are not
Petukhovskoe Soda Lake stromatolites 9
able to grow when pH <8–8.5 (Sorokin et al.
2011b). The measurements of methanogenesis
potential activity in Petukhovskoe Soda Lake
revealed a non-competitive methylotrophic, litho-
trophic and acetoclastic pathways. All pathways were
functioning exclusively within the alkaline pH range
between 8 and 10.5 (Sorokin et al. 2015a). Thus,
most types of microbial activities usually related to
dolomite precipitation are present in Petukhovskoe
Soda Lake.
It is considered that the influence of microbial
metabolic activity is due to the direct or indirect
increase in total alkalinity and pH which creates an
environment favourable for dolomite formation.
The peculiarity of Petukhovskoe Soda Lake hydro-
chemistry is that the brine has a high pH buffer
capacity. Despite the significant oscillation of total
alkalinity (0.45–2.7 M HCO 3 þ CO3 ), the brines
2
2
are always of soda type with CO3 as the main
anion. Even during the years with the lowest salinity
in the lake, the concentration of CO2 3 anion
remains high (>300 mM) (Table 1). It allows main-
taining stable alkaline pH values which were never
observed below 9.6. Furthermore, most of the cul-
turable microorganisms inhabiting this lake are obli-
gately alkaliphilic and natronophilic. The concept of
natronophily assumes that such microorganisms are
not able to survive without elevated concentrations
of soluble carbonates and at the pH below 8–8.5
(Sorokin et al. 2011a). It means that the hydrochem-
ical features of the lake brines are already favourable
for the precipitation of carbonate minerals and the
metabolic activities which may cause elevation of pH
and producing bicarbonate may not be significant in
this type of environment. The similar ideas regard-
ing the impossibility of direct influence of such
microbial activities as photosynthetic CO2 assimila-
tion and sulphate reduction on precipitation of car-
bonate minerals in alkaline environments were
discussed earlier applied to hypothesis of Precam-
brian soda ocean with assumed total alkalinity about
22 mM (Arp et al. 1999, 2001, 2003; Bosak &
10 O. S. Samylina & L. V. Zaytseva
Newman 2003; Gallagher et al. 2013; Meister 2013,
2014), which is much less then in Petukhovskoe
Soda Lake (Table 1).
Obviously, hydrological and hydrochemical fea-
tures of the brines play an important role in dolo-
mite precipitation in the Petukhovskoe Soda Lake.
The same conclusion was made by Deelman (2011)
based on analysis of literature data: the formation of
dolomite is typical for the ‘dynamic’ lakes (subjected
to variations in pressure, temperature, salt concen-
tration) but not for the ‘static’ ones. The Petukhovs-
koe Soda Lake is a typical representative of the
‘dynamic’ lake. The ancient lacustrine build-ups
with dolomite precipitation also represent ‘dynamic’
saline alkaline/soda lake environments with fluctuat-
ing salinity and alkalinity: Green River Formation
(Eocene), Ries Crater Lake (Middle Miocene) and
Qaidam Basin (Eocene) (Della Porta 2015; Huang
et al. 2015). So it is possible to suggest a wide distri-
bution of dolomite formation in saline–alkaline or
soda lakes in the past (including Precambrain) if this
type of inland waterbodies was also spread widely
(St€ueken et al. 2015).
We consider that biologically influenced (sensu
Dupraz et al. 2009) formation of Ca–Mg carbonates
(including dolomite) occurs in Petukhovskoe Soda
Lake stromatolites. All the microorganisms inhabit-
ing this environment produce exogenous organic
matter: EPS and different metabolites. The role of
soluble exometabolites is still unclear although there
are some considerations about the influence of
chelating effect on this process (Zaitseva et al.
2006). The effect of EPS which serve as nucleation
site for mineral growth is discussed much wider
(Braissant et al. 2007; Decho 2009; Krause et al.
2012; Bontognali et al. 2013; Paulo & Dittrich 2013;
Perri et al. 2018; Petrash et al. 2017). It is also dis-
cussed that Mg calcite and disordered Ca dolomite
can be precipitated in association with EPS in the
absence of active metabolism of SRB (Bontognali
et al. 2013). The components of microbial cell walls
can also be involved in Ca–Mg carbonates precipi-
tation. For example, the archaeal protein S layers
can also provide nucleation sites for the formation
of disordered dolomite (Qiu et al. 2017). Thus, in
the environment where the microbial metabolic
influence on the physicochemical conditions may
be insignificant, EPS are considered to play the key
role in the precipitation of Ca–Mg carbonates,
including dolomite. EPS in the Petukhovskoe Soda
Lake, produced mainly by cyanobacteria, were dis-
cussed earlier (Samylina et al. 2016). And the cell
walls of green alga Ctenocladus circinnatus were also
suggested to be involved in precipitation of Ca–Mg
carbonates.
LETHAIA 52 (2019)
Despite the fact that the structure and composi-
tion of EPS differ depending on the microorganism
producing it (Pereira et al. 2009), there are possibly
some common peculiarities in the structure of EPS,
synthesized by different cyanobacteria in soda lakes,
which affect their role as nucleation sites. Only few
cyanobacteria from alkaline saline lakes (Cyanospira
capsulata from lake Magadi, Kenya, and two strains
of Cyanothece sp. from lake Abijata, Ethiopia) were
studied for the structure of their EPS, and the high
content of uronic acids was established there (Gar-
ozzo et al. 1995; De Philippis et al. 1998). Uronic
acids are those components which mostly provide
the acidic property of EPS and play an important
role in the binding of divalent cations. It was esti-
mated by Arp et al. (1999) that acidic polysaccha-
rides act as a Ca2+ ‘buffer’ in alkaline salt lakes,
thereby preventing precipitation at first, and there-
fore, the nucleation occurs only after the saturation
of binding sites. Ca2+ supply is the limiting factor to
achieve this saturation in soda environments as most
of the Ca2+ cations are precipitated inorganically as
calcite, what is clearly seen in Petukhovskoe Soda
Lake stromatolites as white layers (Figs 1, 8). The
following mechanism can be assumed: Ca enters the
lake water with the groundwater seepages. In the
case when a porous stromatolitic crust is positioned
above the site of groundwater leakage, the supersatu-
rated calcium carbonate solution penetrates into the
voids inside the stromatolite and crystallizes there,
forming thin uneven intersecting calcite layers.
The sequence of Mg, Ca and Si precipitation in
EPS is discussed for various halophilic and slightly
alkaline environments. It was argued that initially
Mg calcite permineralization of the EPS occurs and
the replacement by amorphous Mg–Si phase is sub-
sequent during microbialite formation in Satonda
Crater Lake with increased alkalinity (Arp et al.
2003). On the other hand, initial Mg–Si precipita-
tion is known in microbialites from different mod-
ern saline and alkaline lakes (Souza-Egipsy et al.
2005; Zeyen et al. 2015; Pace et al. 2016). In some
cases the coexistence of calcification and silicification
of cyanobacterial EPS in alkaline lakes is discussed
(Kremer et al. 2012). According to our data, the pre-
cipitation of Mg and Si in pericellular and diffuse
EPS in the Petukhovskoe Soda Lake stromatolite is
the primary process and the precipitation of Ca
occurs in a second step (Figs 5 and 6). It is assumed
that microbial metabolic activity controls the
sequence of Mg, Ca and Si precipitation in slightly
alkaline environments where it can be a sufficient
force to influence the pH values. Depending on the
specific microbial activities, pH can be lowered or,
conversely, raised, thereby affecting the local
LETHAIA 52 (2019)
precipitation or dissolving of carbonate and silicate
phases. The ability of microorganisms from Petu-
khovskoe Soda Lake to influence pH value at micro-
scale in the hyperalkaline strongly buffered
conditions is still not understood. But as Ca supply
is a limiting factor (because of inorganic precipita-
tion in calcite layers), primary binding of Mg and Si
seems logical without microbial influences.
We have not found FT-IR or IR evidence that Mg
and Si are precipitated as any separate mineral phase
(amorphous and/or crystalline-like kerolite), as dis-
cussed in the literature (Zeyen et al. 2015). We con-
sider Mg and Si are precipitated simultaneously, but
independently. The initial precipitation of Mg and Si
associated with endolithic cyanobacterial biofilms
was observed in sandy tufas of soda lake Mono
(Souza-Egipsy et al. 2005). It was shown that the liv-
ing cell envelopes were characterized by the presence
of Mg, with little or no Si, while the decaying cells
contained an increased amount of Si, often equal to
or greater than the amount of Mg. It also argues in
favour of an independent Mg and Si precipitation in
EPS. The low-Mg calcite precipitation in sandy tufas
of Mono Lake was confined to the areas around sand
particles and permineralized biofilm remains and
considered to be a periodic process (Souza-Egipsy
et al. 2005). The same is possible for Petukhovskoe
Soda Lake as the supply of divalent cations to the
brine may have seasonal quantitative fluctuations
(Telentyuk 1952).
The specific processes that occur during EPS min-
eralization and formation of rhombohedral crystals
of carbonates in Petukhovskoe soda lake stromato-
lites are still not clear. We can assume that the fine
structure of EPS is favourable for nucleation and
growth of Ca–Mg carbonates and silicates enabling
the non-replacive coexistence of dolomite and
silicates by mechanisms probably similar to those
discussed in the literature (Sanz-Montero et al.
2008; Perri et al. 2018). The limitation in Ca2+ sup-
ply can be a reason of the specific sequence of
Mg–Si–Ca precipitation in Petukhovskoe soda lake
stromatolites.
Conclusions
The Petukhovskoe soda lake (Kulunda Steppe, Rus-
sia) can be considered as a possible environmental
counterpart for Precambrian carbonate shallow
water deposits with cherts containing numerous
cyanobacterial remains. We have made a detailed
description of structure and mineralogical composi-
tion of different layers of the modern stromatolites
from this lake. Despite the hydrochemistry of
Petukhovskoe Soda Lake stromatolites 11
Petukhovskoe Soda Lake is unique for possibly total
inorganic carbonate precipitation, we found out two
types of processes which occur there: biologically
influenced organomineralization and inorganic min-
eralization (sensu Dupraz et al. 2009). Studied stro-
matolites contain both layers with chemogenic
precipitation of calcite and layers with fossilized
algal, cyanobacterial and coccoid bacterial forms
with numerous Ca–Mg carbonate crystals attached
to EPS. It is not quite clear whether microbial meta-
bolic activity can directly influence carbonate pre-
cipitation and whether it is possible to change local
pH or influence CO2/HCO3/CO32 equilibrium,
but the influence of microorganisms through the
release of EPS is surely of utmost importance for the
deposition of carbonate minerals including dolomite
in the highly saturated soda lake.
Acknowledgements. – This work was financially supported by the
RAS Presidium Program ‘The evolution of the organic world and
planetary processes’ (subprogram II), Russian Academy of
Sciences and Federal Agency of Scientific Organizations (project
0104-2018-0030), and RFBR grants 16-04-00758 and 17-04-
00324. We also thank Dr D.Yu. Sorokin (Winogradsky Institute
of Microbiology RAS) for the opportunity to participate in expe-
ditions to Petukhovskoe Soda Lake and RFBR for the financial
support of field work.
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