ELSEVIER Sedimentary Geology 126 (1999) 147–157

The role of sulphate-reducing bacteria and cyanobacteria in dolomite

formation in distal ephemeral lakes of the Coorong region,
South Australia
David T. Wright *
Department of Earth Sciences, University of Oxford, Parks Road, Oxford OX1 3PR, UK
Received 4 May 1998; accepted 3 March 1999

Abstract

The dolomitic distal ephemeral lakes of the Coorong region of South Australia are dynamic systems in which the
concentrations of solutes generally increase during late spring and summer when the rate of evaporation greatly exceeds
aqueous input. During late evaporation, active bacterial sulphate reduction occurs in uppermost, anoxic lake sediments as
rising salinity and then H2 S-toxicity kills off the macrobiota. These hypersaline lakewaters are characterised by high pH
and elevated concentrations of carbonate and sulphate ions. Magnesium concentrated in cyanobacterial cells and sheaths
is released into the strongly-electrolytic, alkaline lake brines from lake-margin stromatolites and mats during desiccation,
and may explain the remarkable increase in Mg levels recorded at this time. Continued evaporation of the lakewaters
produces a halite crust overlying the dolomite: sulphate-free residual puddles and the absence of gypsum indicates that all
sulphate is removed by sulphate-reducing bacteria. Dolomite forms in situ as a primary precipitate from these microbially
mediated lake porewaters, and both the morphology and size distribution of dolomite grains is identical to bacterial cells.
It is not proven exactly when dolomite precipitation occurs, but the submicron-sized grains and multiple defects in a
disordered lattice indicate that precipitation is rapid and follows bacterial removal of sulphate, which is known to inhibit
dolomite formation. Bacterial sulphate reduction and cyanobacterial degradation may thus lead to the removal of all kinetic
inhibitors to dolomite formation, and provide a mechanism for dolomite precipitation around bacterial cells in shallow
subsurface sediments during the late stages of evaporation. In adjacent non-dolomitic lakes, sulphate is present throughout
evaporation, so that other minerals, principally aragonite and hydromagnesite, are precipitated.  1999 Elsevier Science
B.V. All rights reserved.

Keywords: dolomite; bacterial sulphate reduction; kinetic inhibitors; cyanobacteria

1. Introduction 1959; Von der Borch, 1965, 1976; Von der Borch and
Lock, 1979; Lock, 1982; Warren, 1988). It occurs
The Coorong area of South Australia is well in several of a string of small, distal ephemeral
known for the occurrence of modern dolomite, and lakes, characteristically elevated a minimum of 1 m
has been the subject of many studies (Alderman, above sea level, located to the north of the Coorong
Lagoon (Fig. 1). The dolomite is not replacive, and
Ł E-mail: davew@earth.ox.ac.uk has been interpreted as a primary precipitate (Von

0037-0738/99/$ – see front matter  1999 Elsevier Science B.V. All rights reserved.
PII: S 0 0 3 7 - 0 7 3 8 ( 9 9 ) 0 0 0 3 7 - 8
148 D.T. Wright / Sedimentary Geology 126 (1999) 147–157

Fig. 1. Locality map showing distal ephemeral lakes of the Coorong region near Salt Creek, South Australia. Dolomitic study lakes
appear in bold type. After Warren (1988).

der Borch and Lock, 1979; Rosen et al., 1988;
Wright, 1999), but the proposed hydrological and
evaporative models for dolomite formation, though
popular, are inconsistent with empirical data and do
not explain the contrasting mineralogy of the most
recent sediments in adjacent lakes in the chain.
The size and shape of the dolomite grains re-
semble bacterial bodies, suggesting that they become
mineralised during precipitation of dolomite. Benthic
microbial communities, especially sulphate-reducing
bacteria, dominate the lake ecosystem during the
latest stages of evaporation, and their metabolic ac-
 D.T. Wright / Sedimentary Geology 126 (1999) 147–157
tivities can have profound effects on ambient water
chemistry. This paper provides morphological and
geochemical evidence for an essential role for mi-
crobes in dolomite formation in distal ephemeral
lakes.
2. Geological setting and lake stratigraphy
In the early Holocene, the site of the present dis-
tal ephemeral lakes was connected to the sea and
formed part of an estuarine to lagoonal environment.
Deposition of calcareous sediments began to choke
the waterway as coastal beach ridges formed a bar-
rier, and the lagoon began to break up into the chain
of ephemeral lakes seen today. Lake deposits, which
accumulated throughout most of the Holocene in
interdunal depressions, comprise a stratigraphically
differentiated range of mostly non-skeletal carbonate
minerals, reflecting changing depositional environ-
ments during continuing marine regression. Cored
sections from dolomitic and non-dolomitic lakes all
record a shallowing-upwards vertical sequence, be-
ginning with a 6500 year old basal lagoonal facies
of mixed aragonite and calcite muds with a restricted
marine fauna sharply overlying calcretised marine
Pleistocene sands (Von der Borch and Lock, 1979;
Warren, 1988). Isolation of the lakes led to the depo-
sition in each of a stratigraphically correlatable suite
of carbonate minerals with similar textures; however,
mineralogical variability is observed in lakes above
the lagoonal carbonate muds. In dolomitic lakes, the
lagoonal muds are succeeded by proximal ephemeral
lacustrine calcitic and protodolomitic facies, which
are capped by a ca. 1 m thick dolomite=magnesite or
pure dolomite unit of the distal ephemeral lacustrine
facies (Fig. 2). It is this capping dolomite which
forms the basis of this study. In the non-dolomitic
distal ephemeral lakes near Salt Creek, this upper-
most unit comprises aragonite and hydromagnesite
(Von der Borch, 1965; D.T. Wright, unpubl. data).
3. Lake ecology
The distal ephemeral lakes are fed by rainfall and
by seaward-flowing, continental groundwaters from
a regional unconfined aquifer which rises above the
149
lake floor in winter, but become isolated during des-
iccation in the dry summer months as the water
table falls (Von der Borch, 1976). The plant Ruppia
maritima and an aquatic macrofauna including the
brine shrimp Paratemia zietziena, ostracods, cerithid
gastropods and nematode worms colonise the lakes
during winter and early spring, but are progressively
killed off by the rising salinity of increasingly con-
centrated brines, and toxicity of H2 S. Abundant tests
of small gastropods, which thrive when lakewaters
are brackish, are blown lagoonwards as the lakes dry
out, thus removing fossils (and calcium) from the
lake (Von der Borch, 1965; Warren, 1988). Microbes
flourish during these late evaporative stages (Von der
Borch, 1976; Wright, 1999); important benthic mi-
crobial components include surficial cyanobacterial
mat and stromatolites, and sulphate-reducing bacte-
ria within the sediments, the latter including Desul-
fovibrio desulfuricans (L.L. Robbins, pers. commun.,
1996; see Section 4). Small, isolated, filmy, conical
stromatolites 2–3 cm high, occur in Dolomite and
McFaiden lakes, but are completely degraded during
desiccation and do not participate directly in min-
eralisation processes. Single and colonial forms of
the cyanobacterium Synechococcus dominate surface
waters, but decline in abundance with depth. Other
autotrophs, and the green bacteria Chloroflexus, were
observed. Desiccation cracks may be colonised dur-
ing later stages of evaporation by cyanobacteria, sup-
ported by capillary-driven water supply, and=or dew.
4. Methods
Core samples were obtained from the capping
sediments of dolomitic Dolomite, Pellet and Mc-
Faiden lakes, and from non-dolomitic South Stro-
matolite Lake, using 4-cm-diameter plastic piping
and=or sample tubes. Most were immersed in 4%
glutaraldehyde, which was also added to lakewater
samples. Lakewater samples obtained by the author
were filtered on site using 0.45 mm filters and du-
plicates were analysed at CSIRO (Adelaide). pH was
measured in situ in the field, thus avoiding sam-
ple contamination or alteration during conventional
laboratory analysis, and in the laboratory for com-
parison, using an Orion A290 pH meter and Ross
‘sureflow’ electrode.
150 D.T. Wright / Sedimentary Geology 126 (1999) 147–157

Fig. 2. Simplified carbonate mineral stratigraphy of a typical dolomitic distal ephemeral lake, as determined by XRD analysis. Carbonate
mineralogy is related to changing environments, reflected in successive facies changes during progressive isolation of the lakes; modern
dolomite occurs only in ephemeral lakes which experience seasonal desiccation (after Von der Borch and Lock, 1979).

Mg and Ca concentrations were obtained by microscopy, by Prof. L.L. Robbins, University of

atomic absorption, sulphate by gas chromatography,
and carbonate alkalinity by titration. Mineralogy
was determined by XRD: carefully disaggregated
powders were mounted and run on a Philips HT
Generator Model PW 13 20=00 at the Department
of Earth Sciences, University of Oxford, using a
Diffractor PW 1050. Microbes were identified from
sediments by culture using Bergey’s manual of deter-
minative bacteriology (Holt, 1993); and by electron
South Florida, Tampa, Florida, USA.
5. Dolomite textures in uppermost lake sediments
The top few centimetres of the capping dolomite
of the dolomitic distal ephemeral lakes has a gelati-
nous consistency reminiscent of yoghurt, and is as-
sociated with organic detritus from degraded Ruppia
 D.T. Wright / Sedimentary Geology 126 (1999) 147–157
Fig. 3. SEM photograph of dolomite sediment from Dolomite
Lake. Grains are mostly submicron-sized, with larger grains re-
sulting from aggregation of smaller grains. Close inspection of
the smallest grains reveals a subspherical or elliptical, bacte-
rial-shaped core often with an overgrowth of white dolomite.
Grains are interpreted to be the result of mineral encapsulation
of bacterial cells.
plants and cyanobacteria. The dolomite, which be-
comes more plastic and stiffer with depth, has a
structureless, aphanitic texture comprising submi-
cron-sized grains and spherular aggregates which
range in size from <5 µm to >63 µm, though pel-
lets in Pellet Lake may be >500 µm. Under the
scanning electron microscope (SEM), subspherical
and rod-shaped individual dolomite grains (Figs. 3
and 5) are seen to be morphologically identical to
bacteria (e.g. Défarge et al., 1996; Folk, 1997; Gour-
nay et al., 1997), whereas aragonite and hydromag-
nesite sediments from adjacent non-dolomitic lakes
comprise elongate crystalline prisms and platelets
(Fig. 4). Both the shapes and size distributions of
the dolomite grains and clusters are typical of bacte-
rial precipitates (Buczynski and Chafetz, 1991; Folk,
1993). Discrete grains show very limited morpho-
logical variation and no regular crystal habit. This
suggests that the bacteria have been mineralised
by encapsulation while alive (e.g. Castanier et al.,
1989; Chafetz and Buczynski, 1992). The smallest
spheres are identical in shape and size to so-called
‘nannobacteria’ (Folk, 1993). Dolomite collected at
5 cm depth from Lake McFaiden has apparently
undergone recrystallisation, with individual rhombic
crystals typical of crystalline dolomite apparently
151
forming by the coalescing of the submicron-sized
grains (Fig. 5).
6. Nature and timing of dolomite formation
Radiocarbon age dating of surface dolomites is
300 š 250 years (Von der Borch, 1976), proving that
they are recent. The absence of evidence for replace-
ment of calcium carbonate by dolomite supports the
contention that the dolomite is a primary precipitate.
The dolomite was not found to be intergrown with
any other carbonate phase. Dolomite forms in situ
— there is no dolomite present in interdune flats or
calcareous dunes — and must therefore be related in
some way to lakewater and=or porewater chemistry.
In the dry season, the lakes are perched above the
water table, and apart from rare showers, there is
no input of water. This seasonal hydrological iso-
lation of the lakes means that chemical evolution
of lakewaters is driven by evaporation and internal
factors: the lakes form dynamic systems in which
the concentrations of solutes generally increase as
evaporation continues, but changes in alkalinity must
be related to internal processes, of which sulphate
reduction is known to be important. When exactly
dolomite precipitation occurs is still questionable. It
is highly probable that precipitation is rapid, as sug-
gested by the micron-sized grains, lattice disorder
of cations and closely spaced lattice defects (Rosen
et al., 1988; Wright, 1999). Chemical analysis of
lakewaters (Tables 1 and 2), together with experi-
mental evidence that sulphate inhibits precipitation
of dolomite (Baker and Kastner, 1981; Slaughter and
Hill, 1991), suggests that the dolomite precipitates
when all sulphate is removed, just prior to total desic-
cation. Analytical test-strips detected sulphate levels
at between 0 and 200 mg=l in residual wet patches in
Dolomite Lake just prior to complete evaporation.
7. Results and discussion
Tables 1 and 2 show that all lakes record high pH,
carbonate alkalinity and sulphate concentrations rel-
ative to surface seawater. Given the calcareous nature
of the surrounding dune aquifers (Von der Borch,
1965), it is surprising that Ca levels are so low (be-
152 D.T. Wright / Sedimentary Geology 126 (1999) 147–157

Fig. 4. (a, b) SEM photographs of euhedral aragonite crystals and platelets of hydromagnesite from sediment North Stromatolite Lake. In
the general view (a) the abundant larger grains are non-microbial in form, and few submicron grains reminiscent of bacteria are present
(note presence of rare possible organism at centre-left). Enlarged detail of the sediment comprising prisms and platelets appears in (b).
 D.T. Wright / Sedimentary Geology 126 (1999) 147–157 153

Table 1
Chemical analyses of Coorong distal lakewaters (late October)

Lake=ion Mg2C Ca2C Mg=Ca CO23 SO24 pH Mineralogy

(mg=l) (mg=l) (mg=l) (mg=l)
Pellet 1730 66 26 23.24 5,000 8.470 dolomite
Dolomite 957 115 8 34.67 3,500 9.260 dolomite
S. Stromatolite 134 54.1 2.5 28.12 35,788 8.780 aragonite=hydromagnesite

Table 2
Chemical analyses of Coorong distal lakewaters (early December)

Lake=ion Mg2C Ca2C Mg=Ca CO23 SO24 pH Mineralogy

(mg=l) (mg=l) (mg=l) (mg=l)
Pellet 5,280 165 32 21.68 9650 8.461 dolomite
Dolomite Dried out, except for residual wet patches <200 8.512 dolomite
S. Stromatolite 28,720 112 256 12.4 73400 7.863 aragonite=hydromagnesite
Seawater 1,313 400 3.2 16.2 2688 7.9

low seawater concentrations). A possible reason for
this may be the precipitation of calcium carbonate
as biotic skeletal elements in Pellet and Dolomite
lakes, where dolomite is the sole non-skeletal sedi-
ment, rather than deficiency of supply. Comparison
of Tables 1 and 2, and reference to seasonal chemical
analyses of several distal lakes of varying mineral-
ogy in the Coorong chain reported by Von der Borch
(1965), show that concentrations of Mg dramatically
increase late in the evaporative cycle. This peak
cannot be attributed solely to concentration by evap-
oration in Pellet Lake because Mg=Ca ratios increase
significantly, and the only carbonate phase present
is dolomite. Cyanobacteria are known to concentrate
Mg preferentially in their sheaths by up to four times
its abundance in seawater, and in experiments were
shown to contain 60% MgO (Gebelein and Hoffman,
1973). In South Stromatolite Lake, the dramatic
increase in Mg=Ca ratio follows desiccation of lake-
margin stromatolites, suggesting that Mg stored in
stromatolitic cyanobacteria throughout their growth
is released rapidly during late evaporation. Data from
the non-dolomitic South Stromatolite lake (Table 2)
clearly show that high Mg=Ca ratios alone cannot be
a controlling factor in dolomite formation.
These modern, massive dolomites and other car-
bonates in the lacustrine surficial sediments have
been interpreted as typically evaporative in origin
(e.g. Warren, 1988). However, there are a number
of problems associated with this view. Experiments
involving evaporation of the ephemeral lakewaters
under laboratory conditions have consistently failed
to produce dolomite (Von der Borch, pers. com-
mun., 1995, 1996). The dominant carbonate mineral
precipitated in such experiments was in fact arago-
nite, which is in line with experiments performed by
Borchert and Muir (1964). Aragonite is not found
in any of the lake sediments where modern dolomite
occurs. Furthermore, although elevated sulphate con-
centrations were recorded during evaporation, gyp-
sum is not found in any of the studied lakes. The pre-
cipitation of halite crusts in desiccated lakes negates
the possibility that gypsum was somehow flushed
from the system: it never formed. Lippmann (1973)
argued that evaporation would not favour dolomitisa-
tion despite consequent high Mg=Ca ratios, because
of the accompanying decrease in the CO23 -free ion.
The already low activity of the CO23 ion would fall
further, due to its complexing to form MgCO03 neu-
tral ion pairs while Mg2C would also complex with
sulphate, giving and MgSO04 , thus further enhancing
kinetic inhibitions. The apparent interdependence of
dolomite with evaporation is therefore misleading.
Clearly, other processes must be involved in the
precipitation of dolomite.
Modern evaporitic environments are often colo-
nised by benthic microbial communities (BMCs),
principally cyanobacteria, and a genetic link be-
154 D.T. Wright / Sedimentary Geology 126 (1999) 147–157

Fig. 5. (a, b) SEM photographs of dolomite sediment at 5 cm depth from Lake McFaiden. The uppermost sediment comprises
submicron-sized grains of bacterial affinity overlying more structured grains with developing crystal faces (a). There appears to be an
intimate relationship between clusters of these mineralised, bacterial-shaped nanno-grains and crystalline dolomite: where the two grain
types are in physical contact, lower surfaces of nanno-grains can be seen partially incorporated into crystalline dolomite. Remnant outer
surfaces of nanno-grains can be seen within crystal faces where they become almost totally enveloped by the larger crystal — clearly
seen at higher magnification in (b). The loss of identity of the nanno-grains at the expense of the larger crystals may represent diagenetic
self-organisation of mineral grains driven by neomorphic crystal growth from encapsulated, mineralised bacterial spheres.

tween BMCs and dolomite has been observed in both
modern and ancient sediments (e.g. Compton, 1988;
Wright, 1993, 1994, 1997, 1999; Vasconcelos et al.,
1995; Vasconcelos and McKenzie, 1997). Cyanobac-
teria, in the form of stromatolites and as free-floating
autotrophs, were recorded in all the studied lakes,
and the yoghurt-like consistency of the carbonate
muds is due in part to microbial binding (see also
Rosen et al., 1988). Microbial populations are thus
abundant in the dolomitic muds of the Coorong distal
ephemeral lakes, and sulphate-reducing bacteria are
particularly active in shallow subsurface lake sed-
iments during the later stages of evaporation. The
spherular dolomite resembles the grains produced in
microbial experiments by Vasconcelos et al. (1995)
through the mineralisation of organic material in a
sulphate-reducing medium. This suggests a similar
origin for the Coorong dolomite in shallow subsur-
face sediments: anoxic organic degradation driven
by sulphate-reducing bacteria can cause profound
chemical changes to porewaters, and by diffusion
to overlying lakewaters, leading to elevated pH and
carbonate concentrations. The removal of sulphate
from overlying lakewaters may proceed by diffusion
to exhaustion, so that dolomite may then precipi-
tate both authigenically within pre-existing dolomitic
sediments and from modified lakewaters.
Within the zone of sulphate reduction, organic
matter is consumed, sulphate removed, and sulphide
released into ambient waters with metabolic CO2
(Berner, 1980):
2CH2 O C SO24 ! H2 S C 2HCO3
The sulphide may combine with available iron
to produce pyrite and=or pyrrhotite (e.g. Last and
De Decker, 1990), and pyrite-rich laminae occur in
sediments of the north shore of Pellet Lake. Present-
day cyanobacteria contain nitrogen-rich proteins and
fatty acids (Huc, 1980). Sulphate-reducing bacte-
ria and other microbes oxidise cyanobacterial and
 D.T. Wright / Sedimentary Geology 126 (1999) 147–157
Fig. 5 (continued).
other organic matter to support their metabolism,
producing ammonia from enzymatic breakdown of
proteins, which is rapidly absorbed by ambient wa-
ters, thereby increasing pH and carbonate alkalinity
to levels necessary for dolomite formation (Berner,
1980; Durand, 1980; Slaughter and Hill, 1991):
2
2NH3 C CO2 C H2 O ! 2NHC
4 C CO3
NH3 C H2 O ! NHC
4 C OH
OH C HCO3 ! H2 O C CO23
These reactions help explain the elevated pH val-
ues obtained from lakewaters and sediment porewa-
ters. Sulphate reduction may also lead to the release
of free Mg2C ions (Slaughter and Hill, 1991):
2CH2 O C MgSO04 ! 2HCO3 C H2 S C Mg 2C
The sharp increase in lakewater Mg concentra-
tions concomitant with desiccation of cyanobacteria
indicates that cyanobacterial degradation may be an
important source of unhydrated Mg ions (Murata
et al., 1972; Gebelein and Hoffman, 1973; Lyons
et al., 1984). Sulphate-reducing bacteria in associa-
tion with cyanobacterial degradation can thus create
155
conditions favourable to dolomite precipitation, by
increasing the activity of carbonate and magnesium
ions and removing sulphate. The chemical data from
Dolomite and Pellet lakes would appear to be con-
sistent with this hypothesis. In South Stromatolite
Lake on the other hand, sulphate concentrations are
much higher, and persist throughout evaporation. It
is known that calcite as well as dolomite is inhib-
ited by the presence of sulphate, which may explain
why aragonite is the preferred mineral carbonate
phase (e.g. Bathurst, 1975; Walter, 1986), while the
remaining Mg is precipitated as hydromagnesite.
8. Conclusion
Sulphate-reducing bacteria and other microbes
play an essential role in dolomite precipitation in
distal ephemeral lakes of the Coorong, by mod-
ifying ambient waters as a consequence of their
metabolism so that kinetic inhibitors to dolomite for-
mation are overcome. Bacterial sulphate reduction
in shallow subsurface sediments of the dolomitic
lakes reaches a peak during late evaporation, and Mg
156 D.T. Wright / Sedimentary Geology 126 (1999) 147–157
previously concentrated in cyanobacterial sheaths is
released rapidly during desiccation of lake-margin
stromatolites into the highly alkaline, strongly elec-
trolytic, sulphate-depleted brines. This combination
of biochemical processes facilitates dolomite precip-
itation around the numerous bacterial cells found
in ambient waters and sediments during this time.
Carbonate mineralogy is probably controlled by the
degree of biochemical modification of ambient wa-
ters, so that in non-dolomitic lakes, where sulphate
remains in porewaters even in very low concentra-
tions throughout evaporation, aragonite along with
hydromagnesite or magnesite will precipitate rather
than dolomite.
Acknowledgements
I am greatly indebted to Lisa Robbins (Univer-
sity of South Florida, USA) for advice and technical
support in salty waters and to Christ Von Der Borch
of Flinders University, Adelaide, for logistical and
scientific support during field data acquisition in the
Coorong. I would like to record my sincere thanks
to the editor, Gilbert Camoin, for his encouragement,
and to my reviewers, one of whom was Jean-Paul
Perthuisot, for their constructive role in the comple-
tion of this paper. I also acknowledge with gratitude
the support of the Royal Society for travel to Aus-
tralia.
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