ASTROBIOLOGY

Volume 10, Number 4, 2010

ª Mary Ann Liebert, Inc.
DOI: 10.1089=ast.2009.0423

Stromatolites in the *3400 Ma Strelley Pool Formation,

Western Australia: Examining Biogenicity
from the Macro- to the Nano-Scale

David Wacey

Abstract
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The 3426–3350 Ma Strelley Pool Formation (SPF) is a silicified, dominantly sedimentary unit within the Pilbara
Supergroup, Western Australia. It is found widely across the East Pilbara Terrane, and it forms a prominent
marker horizon and separates the largely volcanic 3520–3427 Ma Warrawoona and 3350–3315 Ma Kelly groups.
It has become one of the key formations for study by astrobiologists, following reports of some of the world’s
oldest stromatolites.
Abundant contextural and morphological evidence has been presented over the last decade in support of a
biological role in SPF stromatolite formation. This evidence is reviewed here, and additional data are presented
from recent fieldwork carried out across the *25 km of SPF outcrops in the East Strelley greenstone belt of the
East Pilbara Terrane. In addition to contextural and morphological evidence, a compelling claim for early life
requires geochemical evidence for biological cycling. A potential avenue of approach to obtain such evidence
for the SPF stromatolites (and other ancient examples) is discussed in the context of a pilot study in which
nano-scale secondary ion mass spectrometry (NanoSIMS) was used. Key Words: Biogenicity—Archean—
Stromatolites—Strelley Pool Formation—NanoSIMS. Astrobiology 10, 381–395.

1. Introduction surface of initiation’’ (Semikhatov et al., 1979). This distinction

is vital, especially when considering structures that formed on

O ne of the most intensely studied expressions of mi-

crobial mat activity is that of stromatolites. But what
exactly is a stromatolite? The term stromatolite can mean
different things to different researchers. Many automatically
associate the term stromatolite with the activities of microbial
mats. This is understandable based on present-day analogues
and given that the earliest use of the word stromatolite ap-
peared in a description of laminated organic structures
thought to be produced by ‘‘simple plant-like organisms’’
(Kalkowsky, 1908). This led to a genetic definition of the term
stromatolite: ‘‘organogenic, laminated calcareous rock struc-
tures, the origins of which is clearly related to microscopic life,
which in itself must not be fossilized’’ (Krumbein, 1983),
which was cited as a translation from Kalkowsky (1908) but
was actually a misquote (Riding, 2008). Others understand the
term stromatolite in the nongenetic sense, that is, as a de-
scription of the key textural and morphological characteristics
of the structure without the implication of biological in-
volvement: ‘‘an attached, laminated, lithified sedimentary
growth structure, accretionary away from a point or limited
early Earth or may have formed in extraterrestrial environ-
ments where the participation of biology cannot be assumed.
A comprehensive review of the history and usage of the term
stromatolite can be found in Riding (1999, 2008). Bearing this
in mind, it is not surprising that the earliest stromatolitic
structures in the Archean rock record (reviewed in Hofmann,
2000) have proved contentious in debates concerning the
earliest evidence of life on Earth.
Brasier et al. (2006, and references therein) suggested that
three independent and mutually supporting lines of evidence
are required to substantiate any case for early life and to
falsify the ‘‘null hypothesis’’ of a nonbiological origin for
any candidate structure: (1) evidence for a well-constrained
age and plausible geological context; (2) evidence for a
morphology unique to biology; (3) geochemical evidence
for metabolic cycling. Here, evidence for age, context, and
biological morphology applied to the *3400 Ma Strelley
Pool Formation (SPF) stromatolites is reviewed. New data
are presented from the East Strelley greenstone belt, where
stromatolite diversity and morphology is compared and

Centre for Microscopy, Characterisation and Analysis, and the School of Earth and Environment, The University of Western Australia,
Crawley, Australia.

381
 382
contrasted to that observed at the more widely studied
Trendall locality in the Panorama greenstone belt. Nano-
scale secondary ion mass spectrometry (NanoSIMS) data
from stromatolites of Holocene to Archean age, including an
example from the Trendall locality, are also presented, which
highlight the technique as a promising new tool to obtain
evidence for metabolic cycling within stromatolites.
2. Regional Geology of the Pilbara
The Pilbara Craton of Western Australia (Fig. 1) contains
some of Earth’s best preserved stratigraphic successions of
early Archean rocks, including some of the oldest indisput-
able carbonates and sandstones (e.g., Lowe, 1983). The craton
comprises an Archean proto-continent (Smithies et al., 2005),
which was formed by mantle plume events and consists of
granitic bodies emplaced into and overlain by the Pilbara
Supergroup (Van Kranendonk et al., 2001). The oldest part of
the craton is the East Pilbara Terrane, which comprises
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*3650 to *2850 Ma granitic bodies and an almost contem-
poraneous *3520–3000 Ma volcanogenic carapace and
associated sediments (Pilbara Supergroup) now preserved as
several greenstone belts (Van Kranendonk, 2006).
The Pilbara Supergroup contains four unconformity
bound stratigraphic groups—the *3520–3427 Ma Warra-
woona Group, *3350–3315 Ma Kelly Group, *3240 Ma
Sulphur Springs Group, and the *3200–3000 Ma Soanesville
Group (Fig. 2). Each of these groups consistently dips away
FIG. 1. Geological map of the Pilbara Craton in Western Australia (modified from Van Kranendonk et al., 2007). The best-
preserved stromatolite outcrops are found around McPhee Creek in the East Strelley greenstone belt and around the Trendall
locality in the Panorama greenstone belt. Color images available online at www.liebertonline.com=ast.
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from the domal granitic complexes; dips gradually decrease
with time, which suggests deposition as thickening wedges
adjacent to the growing granitic diapirs (e.g., Hickman, 1984;
Van Kranendonk et al., 2002). The Strelley Pool Formation
previously widely termed the ‘‘Strelley Pool Chert’’ occurs
across 11 greenstone belts and a depositional area in excess
of 30,000 km2 in the East Pilbara Terrane. It comprises a
distinctive assemblage of sedimentary facies within the oth-
erwise predominantly volcanic Pilbara Supergroup, and the
most recent review of mapping and lithostratigraphic evi-
dence established the SPF as an important marker formation
between the Warrawoona and Kelly groups (Hickman,
2008). In addition to the stromatolites of the SPF, older ca.
3490 Ma stromatolites occur in the Dresser Formation of the
Pilbara Supergroup (Walter et al., 1980; Van Kranendonk
et al., 2008). These structures were reviewed by Buick et al.
(1981) in a study that attempted to define universal stro-
matolite biogenicity criteria and concluded that the Dresser
stromatolites were ‘‘probable or possible’’ biogenic stromat-
olites. These structures will not be discussed further here.
3. SPF Stromatolite Pioneers
The SPF stromatolites came to prominence with the de-
scription of conical forms by D. Lowe in 1980 from the
‘‘Strelley West’’ locality (now formally named McPhee Creek,
Wacey et al., 2010) of the East Strelley greenstone belt (ESGB)
(Fig. 3). These structures were originally interpreted as bio-
 STRELLEY POOL STROMATOLITES 383
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FIG. 2. Stratigraphy of the Pilbara Supergroup within the East Strelley greenstone belt (left), showing its relationship with
the East Pilbara Terrane as a whole (right) (data from this study; Van Kranendonk, 2006; Hickman, 2008). Color images
available online at www.liebertonline.com=ast.

logical stromatolites that are very similar to common Prote-
rozoic examples called Conophyton and formed in a shallow
marine, evaporitic setting (Lowe, 1980). Their primary,
synsedimentary origin is not in doubt, and secondary
mechanisms for cone formation, such as post-depositional
deformation structures, can be discounted in the field (dis-
cussion detailed in Hofmann et al., 1999). In 1994 however,
Lowe rescinded his biogenic interpretation for the stromat-
olites, arguing that the extreme continuity of the stromatolite
laminae, the absence of fine-scale crinkly laminae, paucity of
detrital material, absence of an axial zone, lack of fenestrae or
gas bubbles together with the inferred evaporitic setting,
were more consistent with a nonbiological origin through
evaporitic precipitation (Lowe, 1994). There ensued a lively
back-and-forth between Buick et al. (1995a) and Lowe (1995),
by which they debated the biogenicity of the SPF stromato-
lites and other examples older than 3200 Ma. This served to
highlight the challenges that persist when investigating
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FIG. 3. Geological map of the ESGB indicating the position of the logged sections shown in Fig. 4. Mapping and sampling
was performed along approximately 20 km of ridge outcrop of the SPF (white). The four areas of McPhee Creek, Perfect
Stranger, Strelley Pool, and Sulphur Springs Creek are named after geographic features found on the 1:100,000 North Shaw
topographic map and have been formalized by the Western Australia Geographic Naming Committee and the Shire of East
Pilbara. Color images available online at www.liebertonline.com=ast.

heavily silicified carbonates in which any microbial micro-
textures that may once have been present are destroyed.
Discussions concerning the SPF stromatolites were re-
ignited in 1999 by the publication of field observations of
stromatolites from the Trendall locality (see Fig. 1) in the
Panorama greenstone belt (PGB) (Hofmann et al., 1999). This
site was discovered by A.F. Trendall in 1984, and initial in-
vestigations suggested a nonbiological origin for the struc-
tures (Grey, 1984). However, a re-examination of the site,
and especially the discovery of much larger (decimeter-
sized) coniform and branched pseudocolumnar stromatolites
that were more morphologically diverse than any previously
described, lead Hofmann et al. (1999) to reassert a biogenic
origin for these SPF stromatolites. The morphological evi-
dence used to imply biological participation was substantial:
(1) combination of steeply conical forms modified by second-
order corrugate lamination; (2) co-existence of cones with
branching pseudocolumns and the continuation of laminae
across these different forms; (3) juxtaposition of uniform
laminae in the conical forms with uneven wispy laminae in
the inter-cone areas; (4) close comparison to younger forms
of accepted biological origin such as Jacutophyton.
Van Kranendonk et al. (2003) followed up these observa-
tions with geochemical data (REE þ Y) from relatively well-
preserved SPF carbonates, which indicated an anoxic, shallow
marine depositional environment for the SPF in both the
PGB and ESGB. This conclusion was consistent with the
sedimentological observations of Lowe (1983) and Hofmann
et al. (1999), and enabled Van Kranendonk et al. (2003) to
refute an abiotic hydrothermal formation mechanism for SPF
stromatolites (cf. Lindsay et al., 2005). Van Kranendonk et al.
(2003) also provided additional morphological observations
that pointed toward microbial mediation rather than stro-
matolite formation as rigid, presumably abiological, seafloor
crusts (cf. Lowe, 1994). In particular, Van Kranendonk et al.
(2003) cited pockets of thin, flat pebbles in troughs between
the coniform stromatolites that may be reworked microbial
mat material and examples of sand-sized carbonate sedi-
ment, a ready source of loose detritus to be trapped and
bound by microbial mats.
4. Detailed Field Studies—Trendall Locality
In recent years, a series of papers by Allwood et al. (2006,
2007a, 2007b, 2009) described in detail what they interpreted
to be a stromatolite reef developed on a peritidal carbonate
platform in the vicinity of the Trendall locality. Allwood et al.
(2006) described seven distinct stromatolite morphotypes
that occur over several kilometers of outcrop around and
along strike from the Trendall locality. By relating changes in
the morphology of these structures to changes in the sedi-
mentary facies and inferred water depth, they argued that
 STRELLEY POOL STROMATOLITES
the stromatolites were likely biogenic structures. Allwood
et al. (2006) highlighted one of the seven morphotypes for
specific attention (the large complex cones previously de-
scribed by Hofmann et al., 1999). For this morphotype, All-
wood et al. (2006) presented the following geometric,
textural, and geochemical arguments for biogenicity: the
geometry of conical pseudocolumns shows that the stro-
matolites underwent preferential vertical growth that can be
most plausibly explained by upward-migrating microbial
colonies at the sediment-water interface; differences in lam-
inae textures between the cones and the inter-cone areas in-
dicate changes from mechanical deposition (inter-cone areas)
to biologically influenced chemical deposition (cones); a 250-
fold enrichment in rare earth elements in relict carbonate
laminae compared to chert laminae is consistent with
younger microbial carbonates.
Allwood et al. (2007a) repeated and expanded upon ob-
servations made in their 2006 study, which highlighted the
occurrence of multiple stromatolite morphotypes within
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one paleoenvironmental facies, the presence of grains that
were likely trapped and bound by microbial mats, macro-
morphological comparisons to known younger stromatolite
taxa, and a strong ecological control on stromatolite growth.
Additionally, they showed that chemical precipitation played
an important role in SPF stromatolite genesis, highlighted by
abundant evaporite crystal pseudomorphs in close associa-
tion with stromatolite laminae. Allwood et al. (2007a) also
commented on the type of organisms responsible for con-
structing the stromatolites, citing (1) the close spatial corre-
lation of stromatolites and evaporites as evidence that any
organisms must have tolerated high salinity, (2) the lack of
correlation between stromatolites and hydrothermal deposits
as evidence that the organisms were not thermophilic, and
(3) vertical stromatolite growth as evidence for biological
response to either sunlight (phototropism) or some chemical
gradient (chemotropism).
Allwood et al. (2009) continued their investigations of the
Trendall stromatolites at a finer scale and identified micro-
textural evidence from coniform and domical morphotypes
that appear to strengthen the case for a biological component
to SPF stromatolite genesis. In particular, Allwood et al.
(2009) identified organic laminae within the domal mor-
photype that has a texture, Raman spectrum, and micro-
spatial distribution consistent with both a syndepositional
and in situ origin. In the coniform morphotype, Allwood et al.
(2009) again found significant, highly localized contrasts
between the microfabrics in the cone areas (indicative of
deposition of clastic sediment from agitated water) and those
in the inter-cone areas (indicative of precipitated layers),
which they interpreted as microbially mediated. However,
the organic laminae found in the domal stromatolites were
not replicated in the coniform examples. From several years
of detailed field and microtextural work, these authors con-
clude that the data appear much more consistent with a
biological mediation of SPF stromatolites than with purely
physicochemical processes.
5. Detailed Field Studies—East Strelley Greenstone Belt
The author has been part of investigations of the SPF in the
ESGB for the last five years. A summary of the geological
context and stromatolite morphologies in this greenstone belt
385
is given here so that they may be compared to those found
around the Trendall locality in the PGB. Full details can be
found in McLoughlin (2006) and Wacey et al. (2010), and a
further detailed field study of this greenstone belt can be
found in Barnes (1983). A regional map of the greenstone belt
(Fig. 3) highlights the key field areas of McPhee Creek, Perfect
Stranger, Strelley Pool, and Sulphur Springs Creek.* Within
the ESGB, the SPF consists of east-west trending, steeply
southward–dipping, fault-segmented ridges that lie uncon-
formably upon the *3515 Ma Coonterunah Subgroup (Buick
et al., 1995b), a series of basalts, andesites, and intercalated
chert and banded iron formation ridges, and the *3470 Ma
Carlindi granitic complex (Nelson, 1999). The SPF is overlain
by the 3350–3325 Ma Euro Basalt, a thick succession of tho-
leitic and high-Mg basalts, minor felsic tuffs, and thin chert
ridges. The stratigraphy in the ESGB differs somewhat from
that seen in many other greenstone belts in the East Pilbara
Terrane, including around the Trendall locality, PGB; most
notably, much of the Warrawoona Group is absent with the
SPF sitting unconformably on only the lowermost members of
this Group. Throughout the ESGB, the SPF has experienced a
maximum of lower-greenschist–grade metamorphism (Van
Kranendonk, 2000), which allows relatively good preserva-
tion of original sedimentological features.
Detailed mapping of the SPF along some 20 km of strike in
this greenstone belt confirms the subdivision of the forma-
tion into 5 members (cf. Lowe, 1983). In stratigraphic order
these are the basal sandstone (1), which correlates with a
breccio-conglomerate in other greenstone belts—these are
overlain by gray-white cherts (often laminated) (2); then
stromatolite-bearing cherts (3) that are interbedded with
crystal fan pseudomorphs; these cherts are followed by di-
verse upper cherts (4); and finally the upper clastics (5).
Frequent discordant dykes have also been identified but are
not classed as a lithological member of the SPF. Figure 4
highlights the lateral variation in these members along the
*20 km of strike. Members 2 and 3 in the ESGB correlate
with the stromatolite-containing Member 2 of Allwood et al.
(2006) in the PGB.
The environment of deposition can be constrained by field
observations, and several lines of evidence suggest shallow-
water, high-energy conditions for Member 1 in the ESGB
(Wacey et al., 2006). These include cross-bedding, arranged in
truncated sets; relatively shallow scour-and-fill channel
structures, possibly produced by rip currents; textural
maturity shown by moderately sorted to well-sorted, sub-
rounded to well-rounded grains; compositional maturity
shown by the quartz-dominated grains, resulting from pro-
longed mechanical and chemical breakdown; and, lastly,
localized conglomeratic lag deposits that contain large clasts
of the basement rock. The sandstones and conglomerates
appear, therefore, to have been deposited during a shallow
marine transgression across a terrestrial unconformity sur-
face. This depositional interpretation is in broad agreement
with that of Lowe (1983) and the evidence for a rocky
shoreline at the base of the SPF presented by Allwood et al.
(2006, 2007a, 2007b) in the PGB. Putative biogenic features
*These names originate from key geographic features and have
been formally approved by the Western Australia Geographic
Naming Committee.
 386
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FIG. 4. Stratigraphic logs of the SPF from west to east across the ESGB, showing the lateral variations of each of the
members along almost 20 km of strike. Members 2 and 3 in this greenstone belt correspond to Member 2 of Allwood et al.
(2006) in the PGB. Color images available online at www.liebertonline.com=ast.
have been observed in Member 1 (Wacey et al., 2008a, 2008b,
2008c), which indicates that the depositional environment of
the SPF was likely habitable to life prior to deposition of the
carbonate=stromatolite members. There is also evidence
within Member 1 of modern microbial processes, in the form
of microtubular structures that show no substratum prefer-
ence and have been interpreted as endolithic fungal hyphae
(Wacey et al., 2008b). It is likely that similar modern con-
tamination is present in the stromatolite-bearing Member 3,
so great care must be taken not to confuse biological signals
from such contamination with truly ancient signals.
Following deposition of Member 1, clastic sedimentation
ceased, as the ocean inundated the low-lying land surface.
Supersaturation of the water column, perhaps initially aided
by hydrothermal fluids, induced carbonate precipitation
(Members 2 and 3). At the outcrop scale, the laminae of
Member 2 exhibit a spectrum of morphologies from gently
undulose to ripploid surfaces to angular, corrugated sheets.
Disruption of the laminae by brecciated horizons, soft sedi-
ment slumping, and dewatering features is commonly
observed. In places, this disruption results in a massive
megaquartz-rich chert, in which the original laminae are
barely recognizable. These features suggest deposition by bi-
directional currents, with subsequent recrystallization and
dewatering, plus sediment expansion and aggressive silici-
fication. Polished slabs of Member 2 show a complex history
of recrystallization, silicification, and fracturing of the chert.
In thin section, these cherts lack both clay minerals and
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carbonaceous grains. Member 3 in the ESGB contains the
stromatolites and is described in detail below.
Member 4 includes fine-grained black-white-jaspilitic
chert with rare crystal fan horizons that are now pseudo-
morphed by megaquartz. These are primary seafloor pre-
cipitates that lack undulose laminae, which suggests
low-energy conditions and further deepening of the water
column. Overlying gray-green silicified ashes (tuff ) suggest
an increase in the relative contribution of windblown vol-
caniclastics, perhaps due to increased volcanic activity or
chemical inhibition of carbonate precipitation, or both. It is
notable that soft sediment deformation and brecciated hori-
zons are more abundant lower in the SPF section, which
likely record storm events that did not affect the deeper-
water sediments preserved in the upper parts of the SPF. The
upper clastics of Member 5 serve as evidence for a return to
higher-energy conditions and the possible progradation of
sediment fans on the flanks of a volcanic landmass. A green
tuff, often found directly above Member 5 of the SPF, marks
the base of the Euro Basalt and represents an initial explosive
event prior to the extrusion of lava flows.
Discordant black chert veins originate in the underlying
Coonterunah Subgroup and possibly also the Carlindi
granitic complex, and are typically <15 m wide and extend
to over 200 m depth. They often interfinger with the upper
parts of the SPF facies, causing local thickening over strike
distances of *50 m and some local brecciation, but neither
appear large enough nor frequent enough to be feeder
 STRELLEY POOL STROMATOLITES 387

material for the stratiform chert members (Wacey et al.,
2010).
5.1. The stromatolites of Member 3 in the ESGB
Member 3 of the SPF in the ESGB is up to 15 m thick and
comprises gray-white laminated chert and cream-brown
dolomitized carbonate that is best preserved in creek sec-
tions. Coniform structures (Figs. 5b–d, 6) are the defining
feature of this unit and can be traced along strike for over
500 m in the vicinity of McPhee Creek (Fig. 5a; this equates to
‘‘Strelley West’’ of Lowe, 1980), where they are best devel-
oped. The most regularly spaced coniform structures, or so-
called ‘‘egg carton’’ horizons (Figs. 5b, 5d, 6 upper image) are
best exposed in the upper part of Member 3 here. Previous
reports of conical stromatolites (Lowe, 1980) require some
qualification, as not all the structures that appear peak-
shaped in cross section are conical in three dimensions. Ra-
ther, the majority of the SPF structures in the ESGB are
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relief and are unbranched, with apical angles of 70–808, and
those that occur in bedding plane exposures (n ¼ *50) are
ovoid in plan section with their long axes parallel to strike
(Fig. 6 top image). The laminae are between 1 and 3 mm thick
and show no fine-scale crenulations. Near McPhee Creek,
these coniform structures appear to grade into linear current
ripples and undulose bedding (e.g., Fig. 6 middle and lower
images), both along strike and down section, although lack
of continuous outcrop prevents detailed analysis of this
pattern. The cones tend to be best developed higher in
Member 3 with inheritance of coniform shapes over 50 cm or
more. The cones lack a distinct axial zone and sometimes
display a small trough at the base. They may be upright or
inclined by up to 208 to the paleo-vertical, although the
majority are not strongly inclined.
In contrast, when traversing east toward Strelley Pool
(type locality for the SPF) and Sulphur Springs Creek, we
find undulose chert laminae, but coniform structures are
rare. Throughout the mapped area, the Member 3 cherts

noncircular in plan section, have inclined axes, and are thus also contain linguoid ripples (e.g., Fig. 6 lower images)
described as coniform. The cones have 20–60 mm of synoptic and extensive undulatory laminae. The dome-shaped and

FIG. 5. Stromatolites of the ESGB at McPhee Creek. (a) View looking east from the McPhee Creek area showing east-west
trending ridges of the SPF and occasional large black silica dykes perpendicular to them. The McPhee Creek area contains the
most extensive outcrops of SPF Member 3 in this greenstone belt. (b) Plan view of a typical ‘‘small coniform laminite’’
stromatolite morphotype (cf. Allwood et al., 2006) from McPhee Creek, showing linked conical pseudocolumns only a few
centimeters in height. Pencil for scale is 14 cm long. (c) Vertical cross-section view through a typical ‘‘large complex cone’’
stromatolite morphotype (cf. Allwood et al., 2006), McPhee Creek. Some of the coniform shape towards the bottom of the
image has been obliterated by secondary silicification. This is common throughout the greenstone belt. (d) Horizontal cross
section through a typical ‘‘egg carton laminite’’ stromatolite morphotype (cf. Allwood et al., 2006), showing closely spaced
circular to elliptical cones <5 cm in width. Color images available online at www.liebertonline.com=ast.
 388
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pseudocolumnar structures that are best exposed in the vi-
cinity of the Trendall locality, where the largest and most
morphologically diverse stromatolites occur (Hofmann et al.,
1999; Allwood et al., 2006, 2007a, 2007b, 2009), are not
present in the ESGB.
The stromatolite-bearing cherts in the ESGB are inter-
bedded with crystal fan pseudomorphs (and cavity fills) that
occur both as beds up to 2 m high and extend hundreds of
meters laterally, and as pods or lenses less than 0.5 m across.
The laterally extensive crystal fan beds appear to represent
primary seafloor precipitates, because they are commonly
draped by ripploid black cherts, and some of the crystal
terminations show evidence of localized dissolution (Wacey
et al., 2010). Occasionally, they can be preserved in ash but
are most commonly preserved as megaquartz. These are
distinguished from the smaller lenses that represent sec-
ondary diagenetic precipitates formed within the sediment
pile and are widely associated with solution collapse, soft
sediment deformation features, and agate-like mega-quartz
(Wacey et al., 2010).
6. Comparing Stromatolites of the ESGB
and PGB—Implications for Biogenicity
A summary of the key differences between ESGB and PGB
stromatolites is given in Table 1. Evaluations of the bio-
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FIG. 6. Line drawing and ac-
companying photographs to show
the range of bedforms from ripploid
to coniform preserved in Members
2 and 3 of the SPF. These are ar-
ranged stratigraphically to show
the transition in the predominant
bedforms observed up section. Pen
for scale in lower photograph—all
photographs at same scale. Line
drawing courtesy of Martin Brasier.
Color images available online at
www.liebertonline.com=ast.
genicity of the Strelley Pool stromatolites have largely fo-
cused upon the rheology of the primary sediment as inferred
from the laminae geometries and especially the relative
contribution of precipitated carbonate crusts (Riding, 2008)
versus trapped and bound detrital sediment in their con-
struction. For the coniform morphotypes at the Trendall lo-
cality, a principal feature that has been argued to support a
biogenic origin is that many of the coniform flanks are
steeper than 458 and up to 758, which far exceeds the angle of
repose of detrital sediment. From this, chemical or biological
binding of the sediment, or both, has been inferred (e.g.,
Hofmann et al., 1999; Allwood et al., 2006). Previous inves-
tigators have also emphasized the absence of any literature
that describes nonbiological, chemically precipitated cones
and have therefore invoked cohesive microbial mats in the
formation of these SPF structures (Buick et al., 1995a; Hof-
mann et al., 1999; Van Kranendonk et al., 2003; Allwood et al.,
2006).
The presence of a detrital ‘‘trapped and bound’’ sediment
component associated with the SPF stromatolites at the
Trendall locality was shown by Allwood et al. (2007a, 2007b,
2009), but this component is absent in the ESGB. Putative
microbial mat rip-up clasts are also reported from the
Trendall locality (Van Kranendonk et al., 2003). No such
microbial mat rip-ups have been found in the ESGB; there
are lenses of silicified carbonate breccia within the laminated
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Table 1. A Summary of Stromatolite Occurrence in the East Strelley Greenstone Belt (ESGB) Compared to the Panorama Greenstone Belt (PGB)

ESGB stromatolites PGB stromatolites

Paleoenvironmental setting Shallow marine, periodically hypersaline. Stromatolites are associated
with linguoid ripples and silicified evaporite crystals.
Morphological diversity Low: Unbranched conical columns, mostly equivalent to ‘‘egg carton’’
and, rarely, ‘‘large complex cone’’ morphotypes in the PGB. (Flat
and undulose laminite is common, but no evidence exists to support
classification as stromatolite morphotypes).
Geographical extent Common over *500 m around McPhee Creek; rare over remaining
*25 km of greenstone belt.
Size Small, mostly <10 cm width, <10 cm of synoptic relief, with vertical
inheritance over a maximum of *70 cm.
Mineralogy Chert with subordinate carbonate.
Laminae Fine, even and continuous across cone and inter-cone areas; cones
show no obvious axial zones indicative of upwards motility of
filaments; no second-order crinkly or wavy modification of laminae.
Peritidal carbonate platform. Stromatolites occur in three
regressive cycles within an overall shallow marine
transgressive sequence.
High: Seven morphotypes recognized:
(1) Large complex cones; (2) Coniform ‘‘egg carton’’
laminite; (3) Small crested=coniform laminate; (4)
Encrusting=domical laminite; (5) Coniform cuspate swales;
(6) Wavy laminite; (7) Iron-rich laminite.
Environmental distribution indicates some ecological control.
Span *10 km along strike from the Trendall locality. Multiple
morphotypes found within one paleoenvironmental facies.
1–20 cm width, up to 20 cm of synoptic relief, up to *1.5 m high
(coniform morphotypes). Domical morphotypes reach over
2 m in height. Other morphotypes show little synoptic relief.
Partly silicified dolomite.
Often anisopachous; uniform laminae in cones, uneven wispy
laminae in inter-cone areas; some modification by second-
order corrugate lamination; axial zones absent (coniform
morphotypes). Crinkly laminae with several orders of
curvature (domical morphotype).
Continuous laminae across different, closely adjacent

389
morphotypes.
Detrital component Rare to absent. Sand-sized carbonate grains and flat pebbles of possible
reworked microbial mat in the inter-cone areas (coniform
morphotypes); granular sediment possibly within laminae
(domical morphotype).
Fenestrae No. Yes (domical morphotype).
Organic component Rare to absent. Rare laminae (domical morphotype) and carbonaceous
particles (coniform morphotypes).
Biogeochemistry No. Co-occurrence of carbon, nitrogen, and sulfur with black
organic particles aligned with laminae. 250-fold enrichment
of rare earth elements in carbonate over chert laminae
(coniform morphotypes).
Younger analogues Coniform stromatolites such as Conophyton. Jacutophyton, Conophyton, and Thyssagetes (coniform
morphotypes).
Conusella (for domical morphotype).
Irregularia (for wavy laminite morphotype).
Thesaurus (for cuspate swale morphotype).
Likely formation mechanism Chemical precipitation, possibly with some biological mediation. Combination of mechanical deposition, chemical
precipitation, biological trapping/binding, and biologically
induced precipitation. Evidence for biology is strongest
in coniform and domical morphotypes.

PGB data from Hofmann et al. (1999), Van Kranendonk et al. (2003), Allwood et al. (2006, 2007a, 2007b, 2009), and this study. ESGB data from Lowe (1980, 1983, 1994), Wacey et al. (2006), McLoughlin
(2006), and this study.
 390
horizons but no morphological or textural evidence that
these are anything other than brittle fracturing of carbonate
crusts (Wacey et al., 2010). Furthermore, the reported rip-up
clasts of Van Kranendonk et al. (2003) are straight rather than
curved, so these too may represent intraclasts of precipitated
crusts rather than microbial mats.
Many of the stromatolite laminae from the PGB have been
reported to be finer and more uniform within the cones or
columns and coarser, less regular, in the inter-cone areas,
which enhances the view that more variable environmental
conditions and detrital sediment prevailed in the inter-
stromatolite areas (Hofmann et al., 1999; Van Kranendonk
et al., 2003; Allwood et al., 2009). Precisely the converse ob-
servation of remarkably continuous laminae within and be-
tween the stromatolites at the McPhee Creek site was argued
by Lowe (1994) to support their nonbiological origin. Dif-
ferential silicification, which modifies the preservational
windows between the two greenstone belts, cannot account
for these differences alone.
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One of the criteria outlined by Buick et al. (1981) by which
to establish the biogenicity of a putative stromatolite was
that the stromatolite should exhibit wavy or crinkly laminae
with several orders of curvature. This has been shown, in part,
in the case of one very-well-preserved domical morphotype
from the Trendall locality (Allwood et al., 2009). Regionally,
however, wavy or crinkly laminae (and indeed organic lam-
inae) are absent. This has been attributed to poor preservation
and extensive recrystallization (Van Kranendonk et al., 2003),
yet such laminae are absent in coniform morphotypes close to
the Trendall locality (Allwood et al., 2009) and at McPhee
Creek, which are equally well preserved as the domical
example of Allwood et al. (2009).
In the ESGB, there is a closer relationship between ripploid
bedforms and coniform stromatolites (Fig. 6) than in the
PGB. There are rare surfaces near McPhee Creek on which
linguoid ripples and small coniform stromatolites co-occur.
There is also a broad transition from corrugated and ripploid
cherts in the lower SPF up into cherts with coniform struc-
tures (again, best displayed near McPhee Creek where the
silicified carbonate units are thickest). In some instances, it
appears that the coniform structures inherit the topography
of underlying ripploid bedforms and that their subsequent
distribution and accretion were also strongly controlled by
multidirectional currents.
The so-called ‘‘egg carton’’ horizons of regularly spaced
cones accrete in many cases shortly above (a few laminae
above) crystal fan horizons in the ESGB. These cones inherit
the topography and spacing of the underlying crystal fan ar-
rays, and the ‘‘egg carton’’ regularity may reflect the nucle-
ation pattern of the underlying crystal fans. Unfortunately,
this relationship is somewhat obscured by the thin chert layers
that drape the crystal fans; nonetheless, the comparable
spacing and predominance of both crystal fans and ‘‘egg
carton’’ cone horizons toward the top of Member 3 are con-
sistent with this hypothesis. Elsewhere, in the PGB, the con-
verse relationship is also observed with crystal fans capping
coniform stromatolites, which confirms that carbonate su-
persaturation and chemical precipitation were prevalent
during accretion of the stromatolites (Van Kranendonk et al.,
2001; Allwood et al., 2007a).
Viewed in isolation, evidence for biological participation
in stromatolite formation in the ESGB is equivocal. The
WACEY
stromatolite morphologies are much less diverse than they
are around the Trendall locality in the PGB, there is no evi-
dence for granular sediments or trapping and binding by
microbial mats, and there is a close association with struc-
tures that indicate strong physical (ripploid bedforms) and
chemical (crystal fans) controls on growth. In laboratory
experiments, McLoughlin et al. (2008) highlighted the role
that diffusion-limited aggregation of colloids in a turbulent
flow regime, a purely physicochemical process, plays in the
formation of wrinkle structures and domal and columnar
stromatolites. Such colloids, in the form of silica and car-
bonate gels, may have been widespread in the early Archean
environment; it is notable, however, that coniform stromat-
olite morphologies have not been reproduced in this manner.
It is tempting to invoke a biological formation mechanism for
all SPF stromatolites based on morphological similarities in
closely adjacent greenstone belts. However, in the early Ar-
chean caution must be advised, and the null hypothesis (cf.
Grotzinger and Rothman, 1996) of a nonbiological mecha-
nism for the stromatolites of the ESGB, and some of the
morphotypes in the PGB, cannot yet be falsified.
7. Preservation of Biochemical Signals
in Stromatolites
Regardless of the conclusions drawn from morphological
and contextural studies, the case for biogenicity of the SPF
stromatolites would become more compelling with geo-
chemical evidence for biological cycling. This evidence may
be in the form of specific isotopic signatures that indicate
fractionation by biological metabolisms, or enrichments=
specific patterns of suites of elements that are essential for
biological function. NanoSIMS is one technique that pos-
sesses both the spatial resolution and detection sensitivity to
advance our knowledge in this area; the ability of NanoSIMS
to detect biologically important elements and isotopic frac-
tionations in micron-sized objects has been shown in a series
of recent reports in the field of Archean paleobiology (Ras-
mussen et al., 2008; Wacey et al., 2008a, 2008b; Kilburn and
Wacey, 2010).
7.1. Materials
In the present study, three samples were chosen to inves-
tigate the capability of NanoSIMS to map biologically im-
portant elements within stromatolites: (1) a modern (<1500
years before present) lacustrine stromatolite from Lake Thetis
in Western Australia (Fig. 7a) to act as a baseline where
biological participation in the formation of the stromatolite is
undoubted (Grey et al., 1990); (2) an ancient stromatolite from
the 2720 Ma Tumbiana Formation of Western Australia
(Fig. 7b), an example of a well-preserved ancient stromatolite
where a biological formation mechanism is almost univer-
sally accepted (Walter, 1972; Buick, 1992; Lepot et al., 2008);
(3) A moderately well-preserved *3430 Ma coniform
stromatolite from the PGB (Fig. 7c) that most closely corre-
sponds to the ‘‘large-complex-cone’’ morphotype described
by Allwood et al. (2006).
7.2. Methods
Details of NanoSIMS methodology for qualitative
elemental mapping are given in Wacey et al. (2008a) and
Kilburn and Wacey (2010). Briefly, a primary ion beam (Csþ
 STRELLEY POOL STROMATOLITES 391

FIG. 7. Samples used to investi-

gate the applicability of NanoSIMS
elemental mapping to stromato-
lites. (a) Modern (<1500 years be-
fore present) domical stromatolites
from Lake Thetis, approximately
200 km north of Perth in Western
Australia (photograph courtesy of
Ruth Ellison). A disc, 10 mm in
diameter, is required for Nano-
SIMS analysis. This was cut from a
standard thin section, and lami-
nations within a microstromatolite
(arrowed) were targeted for Na-
noSIMS investigation. (b) Small
columnar stromatolites from the
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Meentheena Carbonate Member of

the *2720 Ma Tumbiana Forma-
tion, Western Australia. Again,
specific laminations (arrowed)
within a 10 mm diameter disc
were targeted for NanoSIMS
analysis. (c) Coniform stromatolite
from the *3430 Ma Strelley Pool
Formation, Western Australia.
Due to sampling restrictions in this
heritage-listed area, NanoSIMS
analysis was conducted on a thin
section from a loose piece of ma-
terial pulled off of a fixed outcrop.
Nevertheless, it is clear that this
material is identical to that found
in situ. NanoSIMS analysis tar-
geted small black carbonaceous
grains that occur along poorly
defined laminations (arrowed).
Color images available online at
www.liebertonline.com=ast.

or O
) is scanned across the surface of the sample, and the
sputtered ions are extracted to a double focusing mass
spectrometer. Lateral resolution under 50 nm is achievable
for elemental mapping with use of the Csþ primary beam. In
the Cameca NanoSIMS 50, five elements or isotopes can be
mapped simultaneously with one fixed and four moveable
detectors. Fortuitously, elements commonly representative of
organic material—C, N, O, S—have relatively high negative
secondary ion yields when sputtered with a Csþ primary ion
beam due to their high electron affinity. Although N does not
readily ionize during SIMS sputtering, it does have a par-
ticularly strong emission when coupled to C as the CN
ion.
Images were typically acquired over a 50–60 mm field of view
with a 2–4 pA beam current, an acquisition time of 20 min-
utes, and an image resolution of 256256 pixels. In a 50 mm
field of view, each square pixel measures 195 nm in width, so
for images produced via the Csþ primary beam, the maxi-
mum resolution is defined by the pixel size (as the primary
ion beam is smaller than the pixel). Samples were coated
with a thin (5 nm) layer of gold to provide conductivity, and
each analysis area was presputtered for at least 5 minutes to
remove surface contaminants and implant Csþ ions into the
sample matrix.
7.3. Results
NanoSIMS ion images of the Lake Thetis stromatolite
(Fig. 8a) show very closely correlated distributions of carbon
(12C
), nitrogen (26CN
), and sulfur (32S
), both spatially and
in terms of qualitative intensity (i.e., a number of very bright
areas in the carbon map correlate with very bright areas in
the nitrogen and sulfur maps). These elements pick out
 392 WACEY
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FIG. 8. NanoSIMS elemental maps from modern and ancient stromatolites. (a) Top row, elemental maps of 12C
, 26CN
,
and 32S
from the modern Lake Thetis stromatolite. (b) Center row, elemental maps of 12C
, 26CN
, 32S
, and 28Si
from the
*2720 Ma Tumbiana Formation stromatolite. (c) Bottom row, elemental maps of 12C
, 26CN
, 32S
, and 28Si
from the
*3430 Ma Strelley Pool Formation stromatolite. See text for details; dashed ellipses refer to areas highlighted in the text.
Color images available online at www.liebertonline.com=ast.

organic laminations that run from top left to bottom right
across the field of view. The chemical laminations correlate
with dark laminations visible in optical light microscope
images (Fig. 7a). Given that the dark laminations persist
below the sputtered area of sample surface, they are unlikely
to be an artifact of sample preparation. Since modern stro-
matolites are known to form from microbial trapping and
binding of sediment or microbially induced precipitation, or
both (e.g., Walter, 1976; Riding and Awramik, 2000), the
logical conclusion is that these organic laminations are bio-
logical and the covariance of carbon, nitrogen, and sulfur
represents a biochemical signal that can be preserved during
stromatolite lithification. The NanoSIMS maps also indicate
that these laminations are not simple structures; they com-
monly comprise networks of anastomosing microbial mate-
rial together with occasional larger clumps. Individual
microfossils cannot be identified in these chemical images. It
is also worth noting that NanoSIMS images give only the
relative concentrations of each of the elements within each
pixel; it is not possible to obtain an absolute concentration.
Additionally, it is very difficult to obtain meaningful ratios of
elements present in a sample. For example, the N=C ratio
of the microbial material would be of great interest in order
to provide information about the parts of microbes that are
preserved in stromatolites. However, the apparent N=C ratio
of a sample in SIMS is controlled not only by its true N=C
ratio but also by SIMS instrumental parameters that control
CN
and C
ion yields and by the matrix that the N- and
C-rich material lies in. These variables are not yet fully
understood, so quantitative N=C ratios were not attempted
here.
The 2720 Ma Tumbiana Formation stromatolite was ana-
lyzed under identical conditions to that of the Lake Thetis
example. Once again, results for carbon, nitrogen, and sulfur
show a very close correlation and highlight, in particular,
a number of laminations running from left to right across the
field of view (Fig. 8b). As with Lake Thetis, the laminations
are not simple structures; of particular interest here is the
relationship of the organic material to silica (28Si
), especially
in the center of the images (dashed oval in 8b), where an area
 STRELLEY POOL STROMATOLITES
rich in silica is enclosed by organic material. This could
represent original trapping and binding of a detrital silica
grain, most likely volcanic ash (Buick, 1992), by a microbial
mat. Alternatively, it may represent a fenestral gas escape
structure that has been subsequently infilled with silica (cf.
Allwood et al., 2009), although it is somewhat smaller than
typical fenestrae.
7.4. Discussion
Although the elemental patterns observed here are very
similar to those in the modern Lake Thetis example, it is not
straightforward to attribute them to biology. The presence of
nitrogen alone cannot be used to infer biogenicity, but a close
correlation on a submicron scale of nitrogen with carbon
has been used as an indicator of biological processing in well-
preserved carbonaceous microfossils from both the *1900 Ma
Gunflint Chert of Canada (Robert et al., 2005) and the
*850 Ma Bitter Springs Chert of central Australia (Oehler
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et al., 2006), where the carbon and nitrogen chemical maps
show direct correlation with optical images of the microfos-
sils. An abiogenic source of nitrogen in organic matter is
possible but requires specific conditions and reactants, for
example, high-temperature Fischer Tropsch synthesis, which
requires CO, H2, NH3, and a metal catalyst (Hayatsu et al.,
1972; Kung et al., 1979). Field observations of the Tumbiana
stromatolites indicate deposition in a lacustrine or shallow
marine environment with little hydrothermal influence (e.g.,
Lepot et al., 2008), which is inconsistent with conditions re-
quired for such abiogenic nitrogen synthesis. Sulfur is also
commonly concentrated by biological fixation processes
(Frausto Da Silva and Williams, 2001); and, while its mere
presence may, of course, be due to both biological and
abiological concentration, a positive correlation with carbon
and nitrogen supports a biological formation mechanism in
this case.
The SPF stromatolite is much more difficult to evaluate
because much of the original carbonate mineralogy has been
replaced by silica, and any organic material present is almost
750 Ma older and more metamorphically altered than even
the Tumbiana example. Again, carbon, nitrogen, and sulfur
can be mapped in this sample (Fig. 8c), and they correlate with
black carbonaceous particles observed in transmitted and
reflected light microscope images dotted sporadically along
laminations (Fig. 7c). In the SPF sample, carbon is more
prevalent than nitrogen or sulfur, as may be expected in pu-
tative biological material of this age and metamorphic grade.
While the correlation between carbon, nitrogen, and sulfur is
less well defined than in the younger stromatolites, there are,
nevertheless, numerous areas (dashed ovals in Fig. 8c) where
close spatial and intensity correlations are observed. Overall,
preservation of the organic material shows an inverse rela-
tionship to the degree of silicification (Fig. 8c). Whether these
elements represent degraded microbial material in rocks of
this age depends very much on the interpreted environment
of deposition. Carbon and nitrogen have been shown to co-
occur in organic material from hydrothermal veins in the SPF
(Oehler et al., 2009), presumably synthesized by high-
temperature Fischer-Tropsch–like processes. In contrast, the
coniform stromatolites of the SPF are interpreted to have
formed on a shallow marine carbonate platform without
hydrothermal influence (Allwood et al., 2006, 2007a). In this
393
case, biology represents the most likely mechanism for co-
occurrence of these elements in organic material.
8. Summary
The study of SPF stromatolites in two adjacent greenstone
belts (ESGB and PGB) highlights significant differences in
stromatolite morphology and microstructure (Table 1).
Stromatolites in the PGB comprise seven morphotypes that
occur over several kilometers of semi-continuous outcrop
(Allwood et al., 2006), with coniform and domical morpho-
types possessing microstructural features consistent with
biological mediation (Allwood et al., 2009). In contrast,
stromatolites in the ESGB are less morphologically diverse,
comprising mostly small coniform varieties, lack obvious
biologically mediated microstructures, and highlight the in-
fluence of both chemical and physical sedimentary processes.
NanoSIMS was used to investigate the possibility of ob-
taining evidence for biological cycling from stromatolites of
different ages. NanoSIMS elemental mapping by no means
offers conclusive proof as far as determining the biogenicity
of stromatolites is concerned. It does, however, reveal ele-
mental patterns that are consistent with biology at a scale
and sensitivity ideally suited to the search for very miniscule
traces of life. These signals can be traced from the youngest
to some of the world’s oldest stromatolites. Future work will
incorporate in situ isotopic analysis at the same scale to in-
crease our understanding of the micro- to nano-scale pro-
cesses responsible for the formation of stromatolites still
further. As with any new technique, caution must be exer-
cised, and limitations must be recognized. Chemical signals
should, whenever possible, be matched to morphological
information from petrographic thin sections to guard against
contamination, and careful attention must be paid to stan-
dardization and instrumental artifacts when attempting to
obtain any quantitative chemical information.
The weight of paleoenvironmental, morphological, mi-
crotextural, and geochemical evidence points toward bio-
logical participation in the formation of the PGB coniform
stromatolites and likely also the PGB domical stromatolites.
For the ESGB and remaining PGB stromatolite morphotypes,
the evidence for biogenicity remains equivocal.
Acknowledgments
The author is grateful to the Geological Survey of Western
Australia for extensive logistical support of the ongoing field
program and, in particular, to Arthur Hickman, Martin Van
Kranendonk, and Kath Grey for generous advice and assis-
tance; to Owen Green, Cris Stoakes, and the late John
Lindsay for field assistance; and to Matt Kilburn for assis-
tance with NanoSIMS analysis and image processing. Martin
Brasier and Nicola McLoughlin are thanked for provision of
Tumbiana and SPF stromatolite samples and numerous
discussions of previous versions of this manuscript. NERC
and the Geological Society of London provided financial
support for fieldwork. The author was supported by NERC
grant NE=C510883 and is now supported by a Postdoctoral
Research Fellowship from The University of Western
Australia. The author also acknowledges the facilities and
scientific and technical assistance of the Australian Micros-
copy & Microanalysis Research Facility at the Centre for
Microscopy, Characterisation & Analysis, The University of
 394
Western Australia, a facility funded by The University, State
and Commonwealth Governments.
Competing Interests Statement
No competing financial interests exist.
Abbreviations
ESGB, East Strelley greenstone belt; NanoSIMS, nano-scale
secondary ion mass spectrometry; PGB, Panorama green-
stone belt; SPF, Strelley Pool Formation.
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Address correspondence to:
Dr. David Wacey
Centre for Microscopy, Characterisation and Analysis (M010)
The University of Western Australia
Crawley, WA 6009
Australia
E-mail: David.Wacey@uwa.edu.au
Submitted 2 September 2009
Accepted 19 February 2010
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