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Great Salt Lake ooids: insights into rate of formation, potential as paleoenvironmental archives, and biogenicity

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Content i
GREAT SALT LAKE OOIDS: INSIGHTS INTO RATE OF FORMATION, POTENTIAL AS
PALEOENVIRONMENTAL ARCHIVES, AND BIOGENICITY
by
Olivia P. Paradis
A Dissertation Presented to the
FACULTY OF THE USC GRADUATE SCHOOL
UNIVERSITY OF SOUTHERN CALIFORNIA
In Partial Fulfillment of the
Requirements for the Degree
DOCTOR OF PHILOSOPHY
in
GEOLOGICAL SCIENCES
May 2019
ii
Table of Contents
LIST OF FIGURES AND TABLES ............................................................................................. v
ACKNOWLEDGMENTS .......................................................................................................... vii
CHAPTER 1: OOIDS AND THEIR PROBLEMS .................................................................... 1
Figures ......................................................................................................................................... 9
References ................................................................................................................................. 14
CHAPTER 2: RADIOCARBON CHRONOLOGY OF OOIDS FROM GREAT SALT
LAKE, UTAH INDICATE SLOW RADIAL OOID GROWTH ........................................... 29
Abstract ..................................................................................................................................... 29
Introduction ............................................................................................................................... 30
Great Salt Lake ...................................................................................................................... 32
Methods ..................................................................................................................................... 33
Sample collection................................................................................................................... 33
Raman spectroscopy .............................................................................................................. 34
Dissolved inorganic carbon age ............................................................................................ 34
Bulk inorganic and organic carbon ooid ages ...................................................................... 35
Sequential ooid acidification ................................................................................................. 35
14
C analysis ............................................................................................................................ 36
Results ....................................................................................................................................... 37
14
C analyses in the Great Salt Lake ....................................................................................... 37
Bulk ooid
14
C results .............................................................................................................. 38
Serial
14
C ooid record ........................................................................................................... 38
Discussion ................................................................................................................................. 39
Comparison to marine ooid chronologies ............................................................................. 41
Ooids and the history of the Great Salt Lake ........................................................................ 43
Conclusions ............................................................................................................................... 43
Acknowledgments ..................................................................................................................... 44
Figures ....................................................................................................................................... 45
References ................................................................................................................................. 54
CHAPTER 3: GREAT SALT LAKE OOIDS AS HIGH-RESOLUTION RECORDS OF
HOLOCENE STABLE ISOTOPE HISTORY AND LAKE LEVEL CHANGE ................. 69
iii

Abstract ..................................................................................................................................... 69
Introduction ............................................................................................................................... 70
Ooids as lake level indicators ................................................................................................ 71
Methods ..................................................................................................................................... 72
Sequential stable carbon isotope record of ooids ................................................................. 72
Results ....................................................................................................................................... 73
Discussion ................................................................................................................................. 74
Conclusions ............................................................................................................................... 78
Acknowledgments ..................................................................................................................... 79
Figures ....................................................................................................................................... 80
References ................................................................................................................................. 85
CHAPTER 4: MICROBIAL COMMUNITIES OF LACUSTRINE OOIDS AND
CORRESPONDING NUCLEI REVEAL RELICT DNA MAY BE AN IMPORTANT
SOURCE OF DNA IN STUDIES OF BIOGENICITY ......................................................... 100
Abstract ................................................................................................................................... 100
Introduction ............................................................................................................................. 101
Biogenicity of ooids in marine settings ............................................................................... 102
Biogenicity of lacustrine ooids: Great Salt Lake, Utah ...................................................... 104
Methods ................................................................................................................................... 106
Study site and sampling ....................................................................................................... 106
Microbial community composition analysis ........................................................................ 106
Carbonate geochemistry ...................................................................................................... 108
Results ..................................................................................................................................... 108
Discussion ............................................................................................................................... 110
Carbonate chemistry of Great Salt Lake ............................................................................. 111
Implications of relict DNA in studies of biogenicity ........................................................... 113
Conclusions ............................................................................................................................. 115
Acknowledgments ................................................................................................................... 116
Figures ..................................................................................................................................... 117
References ............................................................................................................................... 121
SUPPLEMENTARY INFORMATION .................................................................................. 136
iv

Appendix A ............................................................................................................................. 136
Appendix B ............................................................................................................................. 170
Appendix C ............................................................................................................................. 174

v
LIST OF FIGURES AND TABLES
Chapter 1
Figure 1: Main microfabrics of ooid cortices 9
Figure 2: Examples of modern and ancient ooid microfabrics 10
Figure 3: Lake Bonneville and Great Salt Lake hydrograph 11
Figure 4: Map of Great Salt Lake, Utah 12
Figure 5: Great Salt Lake ooid microfabrics 13
Chapter 2
Table 1: Inorganic and organic
14
C ages from bulk ooids 45
Table 2: Inflowing river and well water
14
C ages 45
Figure 1: Grain mount of Great Salt Lake ooids 46
Figure 2: Map of Great Salt Lake with sample locations 47
Figure 3: Petrographic images of Great Sale Lake ooids depicting the
presence of organic matter using Raman spectroscopy 48
Figure 4: Grain size distribution of GSL ooids 49
Figure 5: Inorganic
14
C chronologies from north and south arm ooids 50
Figure 6: Nuclei composition in northern vs southern arm GSL ooids 51
Figure 7: Spherical vs rod shaped geometry in ooid growth curve integration 52
Figure 8: Monthly average water temperatures of Great Salt Lake (2010 –
vi
2016) 53
Chapter 3
Figure 1: Relationship between δ
13
C and δ
18
O in closed-basin lakes 80
Figure 2: Lake Bonneville and Great Salt Lake hydrograph 81
Figure 3: Stable carbon isotope (δ
13
C) record of inorganic carbonate carbon
from ooids vs. time (cal yr BP) 82
Figure 4: Great Salt Lake ooid microfabric examples 83
Figure 5: Ooid ray microfabric occurrence through time 84
Chapter 4
Figure 1: Artemia fecal pellet vs Great Salt Lake ooid with pellet nucleus 117
Figure 2: Relative abundance of bacterial and archaeal phyla from water,
brine shrimp pellets, and ooids 118
Figure 3: Principal coordinate analysis (PCoA) of variance between
microbial communities from GSL water, brine shrimp pellets, and both 119
unsorted and sieved ooids
Figure 4: Saturation state of calcium carbonate (aragonite) in the southern
arm of GSL in the winter and summer 120
vii
ACKNOWLEDGMENTS
This work was made possible through the support and efforts of a multitude of people. Of
the many, the first and undoubtedly most important is my advisor Frank Corsetti, who has
provided scientific direction and constant enthusiasm and encouragement every step of the way.
He has taught me how to be a better scientist and an effective science communicator. I owe a
huge portion of my scientific achievements and professional growth to him and can’t imagine a
better advisor.
I thank Will Berelson for sparking an unending excitement for carbonate geochemistry,
and Doug Hammond for helping me to become a more confident and capable isotope
geochemist. Frank Will, and Doug, together with the other members of my qualifying
committee, Dave Bottjer and John Heidelberg, have shaped this dissertation.
I want to thank my academic family: the Corsetti lab (past and present) and all my
colleagues (both in the Department of Earth Sciences at USC and beyond). Specifically, I would
like to thank Dylan Wilmeth, Jeff Thompson, Joyce Yager, Audra Bardsley, and Taleen
Mahseredjian who have been field assistants, scientific sounding boards, collaborators, and
above all—amazing friends throughout the entirety of this work.
Thank you to my family for their endless love, support, and enthusiasm. Specifically, I
want to thank my husband, Brendan Paradis, who has been my biggest supporter, an astute field
assistant, honorary isotope geochemist, and a source of endless love and support throughout the
duration of this work and beyond.
1

CHAPTER 1: OOIDS AND THEIR PROBLEMS
The ability of carbonate minerals (i.e., calcite, aragonite, dolomite) to record geochemical
signatures has been exploited in sedimentary rocks in deep time and the modern to garner
information about environments, the evolution of early earth, biological processes, mass
extinctions, and the evolution of life, to name but a few. The following dissertation aims to
investigate the formation of ooids, defined as laminated, carbonate coated grains that grow with
increasing sphericity around a nucleus of some kind (Figure 1). Ooids are abundant in the
geologic record (Wilkinson, 1985; Opdyke & Wilkinson, 1990; Sandberg, 1983), with marine
and lacustrine ooids dating back to the Paleoproterozoic (Beukes, 1983) and Archaean (Awramik
& Buchheim, 2009), respectively. Modern ooids are commonly found in shallow agitated
environments where the local geochemistry is favorable for their development (e.g.,
supersaturation with respect to calcium carbonate). Though modern ooids are predominantly
found in tropical marine environments, ooids are also found in terrestrial environments including
freshwater and hypersaline lakes (Halley, 1977; Kahle, 1974; Plee, Ariztegui, Martini, &
Davaud, 2008; P. Sandberg, 1975), caves (Donahue, 1969), and other hypersaline environments
(Freeman, 1962).
The carbonate cortex of the ooid precipitates from water and thus, records information
about the environment from which they grew. Nuclei are commonly detrital grains (quartz,
feldspar, etc.), peloids, or even other ooids. In the marine realm, ooids are a common sediment
during periods of transgression and regression but are less common during highstands and
lowstands (Wilkinson, 1985). Ooid cortices are most commonly composed of carbonates such as
calcite and aragonite, but other ooid mineralogies (e.g., Fe, phosphate etc.) are known (Heikoop,
2
Tsujita, Risk, Tomascik, & Mah, 1996; Sturesson, Heikoop, & Risk, 2000; Sturesson & Bauert,
1994)
The crystal orientation and microfabric of ooid cortices vary depending on environment
and chemical and physical conditions. Cortex crystal orientations include tangential, radial, and
randomly oriented crystals (Figure 1). Throughout earth’s history, ancient ooids have displayed a
variety of microfabrics including tangential (Figure 2A) and radial (Figure 2B) ooid cortices,
while most modern marine ooids have tangentially oriented concentric aragonite laminations
(e.g., Joulter’s Cay, Bahamas, Figure 2C). While most work on ooid formation focuses on
modern marine examples, we investigate modern ooid formation in Great Salt Lake in Utah, a
well-known site of primary radial ooids (Figure 2D), and thus may represent both an analog for
understanding ancient radial ooid formation and a repository of environmental change in a
fluctuating terminal lake.
The Great Salt Lake
Great Salt Lake (GSL) is a terminal lake in the Great Basin, a basin that has been home to
several deep lakes throughout the Pleistocene, including Lake Bonneville, the predecessor to
GSL between ~30 – 12,000 years ago (Benson et al., 1990; Oviatt, Madsen, Miller, Thompson,
& McGeehin, 2015; Oviatt, Thompson, Kaufman, Bright, & Forester, 1999). Lake Bonneville’s
size fluctuated with climatic perturbations in western North America which resulted in several
transgressions and regressions (Figure 3). The hydrologic history of Lake Bonneville is divided
into three major sections: the transgressive phase, the overflowing phase, and the regressive
phase (Oviatt, 2015). Throughout the transgressive phase (30,000 – 18,000 cal yr), Lake
Bonneville was in a hydrologically closed basin, with the only outflow to the lake being
evaporation. Lake Bonneville fluctuated during the transgressive phase as closed-basin lakes are
3
particularly sensitive to climatic shifts. The Bonneville shoreline marks the shift from the
transgressive phase to the overflowing phase, which lasted until ~15 cal ka. The overflowing
phase includes the Bonneville flood, which resulted from the collapse of the alluvial-fan which
had dammed the basin rim. The Provo shoreline, lower than the Bonneville shoreline, represents
the new spillover level once the lake emptied during the flood. The regressive phase began after
the lake level dropped below the Provo shoreline, and the lake returned to closed-basin
hydrology (Oviatt, 2015). The Gilbert episode occurred next and represented a brief
transgression that culminated around 11 cal ka (Figure 3). Lake Bonneville regressed to average
historic GSL levels (near 1280m above sea level) after the Gilbert episode, after which brine
shrimp cysts and pellets appeared in lake sediment cores (Oviatt et al., 2015), the implication
being the Bonneville through-Gilbert lake was too fresh for brine shrimp, and the lake did not
become salty until after the Gilbert episode, post 11 ka. Little is known about the remainder of
Holocene lake level aside from estimations and extrapolations from sparse paleo shorelines, and
thus the Holocene record represents a target for further investigation.
Today, the Great Salt Lake is a terminal lake, which means water flows into the lake, but
the only outflow is evaporation. With no outflow, GSL is susceptible to lake level variability due
to fluctuations in evaporation and precipitation which result in times of drought and flooding,
respectively. Salinity varies inversely with lake level, with an all-time low lake level occurring in
2016 due to recent drought—the lowest lake level since the 1963, when the lake recorded the
previous historic all-time-low low (Stephens, 1990). In the late 1950s, a rock-fill railroad
causeway was constructed that separated the north and south arms of GSL, restricting the north
arm from inflow and circulation because the significant inflowing rivers (Jordan, Weber, and
Bear) enter the lake in the southern arm (Figure 4). The construction of the causeway
4
led to a salinity gradient whereby the north arm water is more saline (280 ppt) at or above
saturation for halite, and has a lower elevation than relative to the south arm, which currently has
a salinity near 120 ppt. (“National Water Information System data available on the World Wide
Web (USGS Water Data for the Nation),” 2016; Rupke & Mcdonald, 2012; Stephens, 1990).
Calcium bicarbonate river inflow from the Bear, Jordan, and Weber rivers and sodium chloride
dominated springs and groundwaters dominate the solute inputs to the lake (Jones, Naftz,
Spencer, & Oviatt, 2009). Carbonate deposition is common in the lake (Jones et al., 2009), with
GSL carbonates including mud, tufa-like “microbialites” (Bouton, Vennin, Boulle, et al., 2016;
Newell, Jensen, Frantz, & Vanden Berg, 2017; Pace et al., 2016), and ooids (Halley, 1977;
Kahle, 1974; Reitner, 1997; P. Sandberg, 1975).
Great Salt Lake Ooids
Ooids are found as shoreline deposits around the entirety of GSL (Baskin, 2014; Eardley,
1938). GSL ooids are composed of radial carbonate cortices, with nuclei that include detrital
grains and peloids. Typically elongate, the peloids have been demonstrated to originate from the
fecal pellets of the brine shrimp, Artemia that thrive in GSL’s hypersaline water (Eardley, 1938).
The radial crystal fabric was inferred to be the result of recrystallization and likely therefore
calcite in mineralogy until it was demonstrated that the ooids are indeed aragonite (Kahle, 1974)
and that the radial fabric was depositional (Sandberg, 1975). It has been suggested that GSL
ooids may be the result of organomineralization and a local alkalinity pump (possibly tied to
strong sulfate reduction in sediments) (Reitner, 1997), although the evidence is equivocal. GSL
ooids have high organic matter content, which is dominated by terrestrial plant organics and
insect remains (likely Ephydra brine fly). Some suggest the growth and fabric of GSL ooids is
mediated by organic matter adhering to the surface of ooids (Reitner, 1997).
5
GSL ooids have shapes ranging from spherical to rod shaped, typically corresponding to
the shape of the nucleus—rod shaped ooids often have elongated brine shrimp pellet nuclei
(Figure 5A). The radial aragonite concentric cortex fabrics can have a variety of thicknesses and
include large crystal “rays” (Figure 5B), a finer crystalline banded radial aragonite, and thin
clean radial concentric laminae (Figure 5C). The most common crystal fabric to initiate from the
nucleus is the large aragonite crystal ray, and this fabric is often greater than 50 µm thick. Some
ooids display an alternation of these fabrics while others simply have one cortical layer (often the
aragonite “rays”). Tangentially oriented aragonite crystals in GSL ooids are rare (Sandberg,
1975).
Biogenicity of Ooids
Recent studies focusing on modern marine ooid formation have concluded that
microbiota play an active role in ooid formation, thus qualifying ooids as biogenic structures;
however, much of the evidence is circumstantial. Possible evidence for ooid biogenicity
includes: 1) the presence of organic matter trapped within cortex layers (Folk & Lynch, 2001;
O’Reilly et al., 2017), 2) the presence of a common lipid assemblage across ooids from disparate
localities (Summons et al., 2013), 3) high microbial diversity in marine ooids compared to
nearby stromatolites (Diaz, Piggot, Eberli, & Klaus, 2013; Diaz et al., 2014), and 4) the presence
of a functional gene assemblage which is linked to certain microbial metabolisms that have the
potential to foster calcification (Diaz et al., 2014). Microbial mediation has also been invoked in
the formation of the oldest known radial calcite ooids from the 2.72 Ga Meentheena Member,
Tumbiana Formation (a carbonate lake system) based on the presence of organic matter in close
association with primary and early diagenetic mineral phases within the ooids (Flannery et al.,
6
2018). Much of the evidence surrounding ooid biogenicity is circumstantial and disentangling
the attribution of microbes that are building the carbonate vs. residing in it remains challenging.
Ooid “Problems”
Our lack of knowledge on the rate of ooid formation as accretionary structures and
repositories of paleoenvironmental information, the role of microbiota in ooid formation, and the
isotopic heterogeneity of ooids represent a gap in our understanding of a major carbonate
sediment throughout earth’s history. In the following chapters, isotopic, petrographic,
geochemical, and molecular data are combined in order to determine the conditions under which
modern lacustrine radial ooids form, to assess the growth rates of ooids in hypersaline
environments, and to analyze the microbial communities in lacustrine ooids and the implications
for using certain molecular tools in studies of biogenicity in the future.
Chapter Roadmap
Chapter 1 aims to assess the growth rate and lifespan of ooids from Great Salt Lake
(GSL), and thus establish a chronology for ooids in this environment. The growth rates of
modern marine ooids in a few locations have been characterized; however, the growth rates of
lacustrine ooids remains elusive. Modern marine ooids exhibit tangentially-oriented crystal
fabrics, while many ancient ooids exhibit primary radial crystal fabrics. Great Salt Lake ooids are
one of the best examples of modern lacustrine ooids and exhibit a primary radial aragonitic
fabric, thus affording us the opportunity to exhibit another end-member for ooid formation in
comparison to modern marine ooids. This chapter employs radiocarbon chronology along with a
novel carbonate dissolution vessel to sequentially dissolve GSL ooids, collect CO2 from the
reaction with acid, and date the carbon using radiocarbon analysis. We were also able to compare
7
this high-resolution chronologic data with carbon data one might measure from a bulk ooid
sample in the same environment to illustrate the offset in ages that is observed, and highlight
both the need for finer scale analysis, and caution for bulk analysis of accretionary structures
such as these. The results highlight the distribution of organic matter throughout ooid cortices,
cautioning against the use of bulk carbon (organic) age dating to establish a lifespan. The results
also highlight the disparity between lifespan and growth rate between modern tangential marine
ooids and radial lacustrine ooids from GSL and establish an analog for understanding the
occurrence of radial ooids in ancient rocks.
Chapter 2 seeks to utilize GSL ooids as repositories of high-resolution environmental
archives in a terminal lake system that is highly sensitive to climatic change. By coupling stable
carbon isotopes with the established radiocarbon chronology in Chapter 1, we can establish a
record of isotopic change over several thousand years in GSL. Because stable carbon isotopes
covary with stable oxygen isotopes in closed basin lakes (Talbot, 1990), δ
13
C may be used to
evaluate change in lake level over time. This chapter couples the stable carbon isotope record
with fine scale petrographic analysis, which reveal that most GSL ooids have a common
nucleating crystal fabric consisting of large aragonite crystal rays that coincide with a relatively
constant carbon record during that time. The study also highlights that GSL ooids have most
likely been forming after the construction of the rock-fill railroad causeway in 1959, which is the
first confirmation of recent ooid formation in modern times.
Chapter 3 seeks to address the biogenicity of GSL ooids by first establishing the
microbial potential of ooids using 16S rRNA sequencing. The microbial communities of GSL
ooids, brine shrimp pellets (which serve as dominant nuclei in GSL), and water were analyzed.
Most ooid samples had a microbial community that strongly resembles those from brine shrimp
8
fecal pellets, highlighting the importance of relict DNA in this study of biogenicity. The
carbonate chemistry of GSL water supports the slow growth rates and low saturation state of the
system, and thus does not necessitate the intervention of microbial activity to form.
9

FIGURES

Figure 1: The main microfabrics of ooid cortices include tangentially, radially, and randomly
oriented crystals that make up concentric carbonate laminae. Randomly oriented ooid cortices
also include micritized ooids.

10

Figure 2: Examples of ancient and modern ooid microfabrics. A) Neoproterozoic tangential
ooids from the Beck Springs formation. B) Radial ooids from the Neoproterozoic Johnnie
formation. C) Modern ooids from Joulter’s Cay, Bahamas display tangential concentric laminae
that are characteristic of many modern marine ooids. D) Modern ooids from Great Salt Lake,
Utah have a primary radial crystal orientation.

11

Figure 3: Hydrograph depicting lake level fluctuation in Lake Bonneville and Great Salt Lake
(modified from Murchison, 1989; Newell et al., 2017; Oviatt, 1997, 2015; Oviatt et al., 2015).
The transgressive phase (T), the overflowing phase (O), and the regressive phase (R) are marked
with gray boxes on the top of the figure. Lake Bonneville fluctuated from ~30 cal ka until ~15
cal ka when it regressed to Great Salt Lake levels until present.

12

Figure 4: Map of Great Salt Lake in Utah highlighting the previous historic low lake level of
1963 (1277 m above sea level) and the historic average level of 1280 m above sea level (left).
The railroad causeway constructed in 1959 divides the lake in two distinct arms and isolates the
north arm from inflowing river water. Halobacteria and halophilic algae thrive in the resulting
hypersaline north arm water, tinting the water red with their red pigments (right).

13

Figure 5: Great Salt Lake ooid with pellet nucleus (A). Large aragonite crystal rays are often the
initiating crystal fabric against the nucleus (B). The other common crystal fabrics include banded
tightly-packed radial aragonite crystals (white arrow), and thin radial-concentric aragonite
crystals (black arrow).

14

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CHAPTER 2: RADIOCARBON CHRONOLOGY OF OOIDS FROM GREAT SALT LAKE,
UTAH INDICATE SLOW RADIAL OOID GROWTH
Paradis, O., Corsetti, F., Bardsley, A., Hammond, D., Berelson, W., Xu, X., Walker, J., Celestian,
A.
ABSTRACT
Ooids (calcium carbonate coated grains) are common in carbonate environments
throughout geologic time, but the mechanism by which they form remains unclear. In particular,
the rate of ooid growth remains elusive in all but a few modern marine environments. Furthermore,
modern marine ooids display tangentially-oriented aragonite crystals within their cortices, whereas
many ancient ooids have radially-oriented cortices. In order to investigate radial ooid formation,
we used
14
C to date ooids from Great Salt Lake, Utah, a well-known site of primary radial
aragonitic ooids.
14
C dating of bulk ooids obtained from the northern shore of Antelope Island and
the northeast shore of Great Salt Lake were sieved into different size fractions and indicate a range
of dates between 2728±15 and 4373±20
14
C yr BP. Larger ooids were older than smaller ooids,
implying that larger ooids grew in the environment for a longer duration, with the caveat that bulk
age dating integrates the growth history of an ooid. To surmount this effect, ooids from the coarse
fraction were sequentially dissolved and
14
C ages were obtained for each dissolution step to create
a time series of ooid growth. The results of the sequential dating indicate that the formation of
coarse Great Salt Lake ooids began between 5800-6600 ± 60
14
C yr BP while their outer cortices
are nearly modern. Sequentially dated ooids from Antelope Island (southern part of Lake) record
a nearly linear growth history with linear growth rates of ~ 0.01 – 0.015 µm/yr, whereas ooids
from Spiral Jetty (NE part of Lake) record somewhat faster growth between ~6000 and 4000 years
ago (0.03 – 0.06 µm/yr) followed by a slower growth history for the remainder of their lifespan
30

(0.003 – 0.008 µm/yr). The lifespan of Great Salt Lake radial aragonitic ooids ranges is two and
six times longer than those from modern marine environments, and thus provides a unique end
member for understanding the mechanisms behind radial ooid formation. The antiquity of the
ooids would suggest that geochemical parameters measured from bulk ooid dissolution would
integrate over ~6000 years and thus do not represent geochemical snapshots in time as some
previous studies suggest.
INTRODUCTION
Ooids are small (generally < 2 mm) laminated, coated grains, with a calcium carbonate
cortex surrounding a nucleus. Ooids are ubiquitous in the geologic record in marine and lacustrine
settings, and as accretionary structures, may serve as repositories of high resolution aqueous
evolution, preserving both biogeochemical (Diaz et al., 2015, 2013; Summons et al., 2013) and
isotopic (Duguid et al., 2010) information. Despite their ubiquity, ooid formation remains
enigmatic. Both abiogenic and biogenic modes of formation have been proposed (Diaz et al., 2013,
2015, 2017; O’Reilly et al., 2017; Pacton et al., 2012; Summons et al., 2013), and even the rate of
ooid accretion remains elusive for the majority of ooid occurrences. Without a better understanding
of how rapidly ooids form, their utility as paleoenvironmental indicators is hindered and the
question of biogenicity remains unclear.
Radiocarbon (
14
C; half-life = 5730 ± 40 yr) has been successfully used for the step-wise
dating of marine ooids from the Bahamas (Beaupré et al., 2015; Duguid et al., 2010), Australia
(Beaupré et al., 2015; James et al., 2004), and Hawaii (Hearty et al., 2010). Regardless of location,
14
C ages decrease from the ooid nuclei toward their outer surfaces with the exception of a
14
C
anomaly of unknown origin in ooids from Highborne Cay, Bahamas (Beaupré et al., 2015). Using
the radiocarbon chronology, Beaupré et al., 2015 argued “modern” marine ooid net growth rates
31

were slow and relatively constant, with mean lifespans ranging from 800 ± 135 to 1470 ± 280
14
C
years and growth rates ranging from 0.36 ± 0.03 to 2.2 ± 0.3 ng C-CaCO3/ooid/year. However,
calculated net growth rates from these radiocarbon dating experiments on ooids are likely
underestimating gross carbonate precipitation due to abrasion, as lab experiments with seawater
have shown growth can be four orders of magnitude faster than radiocarbon net growth rates
(Trower, Lamb, & Fischer, 2017).
While most modern marine ooids have carbonate crystals tangentially oriented to the
nucleus or micritic cortices, many ancient ooids display crystals that radiate from the nucleus (See
Figure 2 from the Introduction Chapter, p. 9). Thus, there is a need to investigate the controls on
radial ooid formation to both understand how to interpret their occurrence in ancient and modern
environments and compare their growth dynamics to tangentially oriented ooids. Although not a
marine environment, the Great Salt Lake (GSL) in Utah provides a unique opportunity to assess
the net growth rate of primary radial aragonitic ooids that texturally resemble many ancient ooids,
both marine and lacustrine (Figure 1). In addition to their utility in understanding radial ooid
formation, GSL ooids may be targets for understanding the history of GSL, which is particularly
sensitive to climatic shifts, as there is no outflow from the lake (besides evaporation). The GSL
has also been subjected to environmental alteration by human activity, especially notable is the
division of the lake by a railroad causeway that occurred in 1959 which created a northern and
southern salinity gradient. However, like marine ooids, the utility of lacustrine ooids in
reconstructing paleoenvironmental changes is dependent on their placement within a proper
temporal framework. The aim of this study is to use
14
C as a chronometer to sequentially date ooids
from Great Salt Lake, and thus constrain modern radial ooid formation in this setting and provide
32

necessary chronological context so their potential as paleoenvironmental indicators may be
explored.
Great Salt Lake
1) Great Salt Lake environmental setting
Great Salt Lake (GSL) is a terminal lake in northern Utah with circumneutral pH that is
currently meromictic. GSL represents the present phase (since 11.5 ka BP) that resulted from the
transition of the larger and deeper Lake Bonneville (30-11.5 ka BP) to a shallow, hypersaline lake.
(Oviatt, Currey, & Sack, 1992; Oviatt et al., 2015) The north arm of GSL is currently separated
from the south arm by a rock-fill railroad causeway that was constructed in 1959. Because the
three rivers that feed the lake (Bear, Jordan, and Weber rivers) enter the south arm, the north arm
water is more saline (28%) at or above saturation for halite, and has a lower elevation relative to
the south arm, which currently has a salinity near 12% (“National Water Information System data
available on the World Wide Web (USGS Water Data for the Nation),” 2016; Rupke & Mcdonald,
2012; Stephens, 1990)
2) Great Salt Lake Ooids
Great Salt Lake contains one of the best examples of modern lacustrine ooids. Previous
studies on Great Sale Lake ooids describe their morphology and petrology (Eardley, 1938; Halley,
1977; Kahle, 1974; Sandberg, 1975). The shoreline and bottom lake deposits of GSL have been
mapped, and ooids are found as shoreline deposits around the entirety of GSL (Baskin, 2014;
Eardley, 1938). Eardley (1938) also demonstrated that the fecal pellets of the brine shrimp,
Artemia, likely served as the dominant peloidal nuclei due to morphological similarities. Eardley
also inferred ooid cortices were calcitic, and that their radial texture was the result of
33

recrystallization. The assumption of calcitic mineralogy in GSL ooids prevailed until Kahle (1974)
demonstrated that GSL ooids are in fact aragonite, and their cortical fabric is depositional.
However, Kahle concluded aragonite-aragonite inversion had taken place. Sandberg (1975)
confirmed the aragonitic mineralogy of GSL ooids, demonstrated that the radial aragonite fabric
is depositional, and found no evidence that aragonite-aragonite inversion had taken place.
Subsequently, Reitner (1999) suggested that organic matrices on the surface of GSL ooids could
be important in the mineralization of the aragonite. More recently, (Mcguire, 2014) attempted
serial dissolution of unsorted ooids from 15cm water depth in the modern south arm of GSL that
resulted in
14
C ages from 2024±36 yr BP (outermost composite sample) to 8144±29 yr BP
(innermost composite sample), indicating that the ooids were quite old versus modern marine
examples but the coarse sampling resolution could not discern whether modern precipitation took
place. As part of a large-scale survey of the tufa-like carbonate mounds many refer to as
“microbialites”, Bouton, Vennin, Boulle, et al., 2016 measured the bulk
14
C age of unsorted GSL
ooids from the shoreline of the south arm of the lake. Their results yielded a composite ooid
14
C
age of 3.3 ka BP. Thus, while some constraints regarding the age of the ooids exist, many questions
remain.
METHODS
Sample collection
Ooids were collected by hand from the southern half of Great Salt Lake in Bridger Bay on
Antelope Island, and near Spiral Jetty in the north arm of the lake in March 2014 (Figure 2). Ooids
were collected at the sediment-water interface in less than 10 cm water depth. Samples were rinsed
with deionized water, dried in an oven at 50°C, and sieved to partition the ooids into discrete size
fractions (125-250 µm, 250-355 µm, 355-500 µm). Ooids were massed from each size fraction
34

relative to the beginning sample to establish a grain size distribution (Figure 4) Unfiltered lake
water was sampled from the shore of the northern tip of Antelope Island in the south arm of GSL
and the beach at Spiral Jetty in the north arm of GSL in September 2016 for
14
CDIC analysis.
Unfiltered river and well water were sampled in May 2017 from Bear, Jordan, and Weber rivers
as well as a well in Ogden, Utah (Weber State University). At each site, one liter of water was
collected in 1000ml size glass bottles (Fisher #06-414-8) which had been previously rinsed three
times with deionized water, soaked in 10% HCl, and rinsed three more times with deionized water.
The bottles were field rinsed three times before water was sampled with no head space and
immediately poisoned with 100 µl of saturated HgCl2 in the field.
Raman spectroscopy
Raman spectra of GSL ooids were obtained using a Horiba XploRa+ micro-Raman
spectrometer. Specimens were measured using an incident wavelength of 532 nm, laser slits of
200 um, 1800 gr/mm diffraction grating, and a 100x (0.9 NA) objective. Laser spot size was
approximately 2 micrometers in diameter. The laser was projected through individual ooids from
a grain mount. Data were collected for 0.5 sec/exposure at maximum laser power of 532 nm at the
thin section, which achieved sufficient signal-to-noise ratio without altering the spectrum during
data collection. Component spectra were chosen to represent organic material based on the
acquired Raman spectra, and the presence of organic material was mapped on photomicrographs
of ooids based on principal components analysis (PCA). The heat maps represent the presence or
absence of organic material, and not relative abundance.
Dissolved inorganic carbon age
Lake, river, and well water samples were prepared using the headspace-extraction method
(Gao et al., 2014).
35

Bulk inorganic and organic carbon ooid ages
Total organic and inorganic C was extracted from each sieved ooid sample (125-250 µm,
250-355 µm, 355-500 µm) and an unsorted ooid sample. The extracted organic and inorganic
carbon was analyzed for
14
C at the Keck Carbon Cycle Accelerator Mass Spectrometer
(KCCAMS) facility at the University of California, Irvine (Beverly et al., 2010; Southon et al.,
2004). Details regarding methodology for the bulk organic and inorganic carbon extractions may
be found in the supplementary information (Appendix A, Table 1).
Sequential ooid acidification
To assemble an ooid chronology, we measured the
14
C ages of fractions of CO2 collected
during sequential acid addition to sieved ooids (355-500 µm) from Spiral Jetty and Antelope
Island. Ooids (~50g) and 150ml of deionized water (DIW) were placed in a reaction vessel
constructed from a 500ml graduated round media storage bottle (VWR cat. # 89000-238) and a
suspended magnetic stir rod (Appendix A, Figure 1). The reaction vessel was purged with N2 that
was scrubbed with Ascarite-II while a stir bar spun at 700 rpm to drive off any dissolved CO2 in
the water for a total of 30 minutes. The sample was acidified by injecting 60 ml of 3.3M HCl at a
flow rate (acid) of 10ml/min. Gas was shunted for the first 5 seconds of acidification to off-gas
any residual N2 before collecting the sample in Tedlar bags which had been rinsed with ultra-high
purity (UHP) helium scrubbed with Ascarite-II. Gas was collected in 3 Tedlar bags per each
acidification step. The first Tedlar bag collected gas for the first 30 ml of acid added, the second
Tedlar bag collected gas during the second 30 ml acid addition, while the third Tedlar bag collected
remaining CO2 that evolved after all 60 ml acid had been added and was left to sit for 3 minutes
before pulling it off the vessel. Four discrete acidification steps were performed, with a subsample
of 5-10 ooids removed from the acidification vessel between each. The subsample of ooids was
36

examined using a Hitachi TM-1000 environmental scanning electron microscope (SEM) to
confirm dissolution was occurring from the outside to the inside (Appendix A, Figure 2). Between
acidifications, ooids were rinsed three times with deionized water (DIW) and dried overnight. The
reaction vessel and its components were rinsed in 10% HCl and dried between each acidification.
The DIW in the reaction vessel was replaced, and the reaction vessel was purged for 30 minutes
with ascarite-scrubbed N2 to remove any atmospheric carbon. The acidification using 60 ml of 3.3
M HCl, DIW rinse, acid wash, and 30-minute purge was repeated for each acidification (four times
total). Following the final acidification, the remaining nuclei were rinsed three times with DIW,
dried overnight, and reserved for
14
C analysis of the organic carbon fraction. Some calcium
carbonate remained on the oolitic nuclei at the end of the experiment to ensure ancient carbonate
nuclei were not dissolved which would skew the oldest inorganic carbon age.
14
C analysis
For
14
C analysis, gas samples from the sequential leach were cryogenically purified through
a dry ice/ethanol trap and collected in a liquid nitrogen trap. Residuals from ooid dissolution of
bulk ooids and from the sequential leach were combusted at 900°C for 3 hours to obtain CO2. All
purified CO2 samples were graphitized using a sealed-tube zinc reduction method (Xu et al., 2007).
Graphite was pressed into aluminum target holders and analyzed for
14
C at the Keck Carbon Cycle
Accelerator Mass Spectrometer (KCCAMS) facility at the University of California, Irvine
(Beverly et al., 2010; Southon et al., 2004). Data were normalized and background corrected using
both modern coral and radiocarbon-dead reference carbonates acidified in the same reaction vessel.
14
C data are presented according to the conventions presented in Stuiver & Polach, 1977. Non-
calibrated ages (given in BP) are presented in this paper to be consistent with ages reported in the
literature.
37

RESULTS
14
C analyses in the Great Salt Lake
It is important to consider the behavior of
14
C in the GSL, as the
14
C age of lacustrine
carbonates may be subject to a “reservoir effect”, whereby lakes can accumulate old dissolved
inorganic carbon over time. Lakes acquire some of this carbon from inflowing water that travels
over ancient limestones that reside in their catchment, causing their dissolved inorganic carbon
pool to have an apparent age that would deviate from the atmospheric value. Any calcium
carbonate that precipitates from that lake water would record an apparently older
14
C age than
coeval atmospheric
14
C. Our analyses reveal that reservoir effects represent the largest source of
uncertainty in our data. Paired U-Th and
14
C ages from lacustrine cave carbonates suggest the
reservoir effect for Lake Bonneville (from 25 to 13 ka) was 200 years or less (Mcgee et al., 2012),
which agrees with Oviatt’s previous estimates of Lake Bonneville’s reservoir effect (Oviatt et al.,
1992). To date, no one has measured the age of the dissolved inorganic carbon in Great Salt Lake
directly. Surface water enters the lake via three rivers: Bear, Jordan, and Weber, all of which enter
the south arm of GSL. The radiocarbon age of the river water and water from a well in Ogden,
Utah was measured in May 2017, and indicate the water is delivering ancient inorganic carbon to
the lake (Table 1). Modern lake water from the south arm of GSL has a
14
C age of 295 ± 20 yr
BP, and the modern north arm water has a
14
C age of 115 ± 20 yr BP. The difference likely reflects
the inflow of high
14
C river water into the south arm, with atmospheric gas exchange subsequently
supplying an atmospheric CO2 component to the north arm. Two anthropogenic changes may have
influenced the lake reservoir age in contrasting ways. The causeway has reduced the north arm
surface area by a factor of two, reducing the rate of atmospheric exchange in this region
proportionally. However, bomb testing has increased
14
C/
12
C in the atmosphere by an average of
38

50% during the past 50 years. Because those two effects should largely negate one another, we
assume the modern south arm reservoir effect of 295± 20 yr BP is likely more representative of
pre-causeway homogeneous lake conditions and therefore more applicable to this dataset.
However, there remains uncertainty in how the reservoir age may have varied through the past
6000 years.
Bulk ooid
14
C results
Bulk unsorted and sieved ooids from each site yield inorganic and organic
14
C ages that
are a result of averaging between much older and younger carbon in the samples and thus do not
represent a unique age for the ooids. However, bulk ages can help bracket the general age of the
ooids and provide some indication of their antiquity. In general, bulk ooid analyses produced ages
that ranged from 2728±15 to 4373±20 yr BP, whereas bulk organics produced slightly younger
ages, between 1935±15 and 4200±15 yr BP (Table 2). Ooids of finer grain size have younger
average
14
C ages, which is reflected in both inorganic and organic carbon. Total organic carbon of
bulk ooids from both sites varies from 0.43% to 1.34%; however, ooids from the north arm of GSL
have more than double the organic carbon of south arm ooids (Table 2). Raman spectroscopy of
ooid cross sections shows that organic matter is distributed both in the micritic nuclei of ooids
(when the nuclei are peloids) and incorporated throughout the carbonate cortices (Figure 3). Grain
size analysis reveals both north and south arm ooids are skewed toward finer grain sizes, between
63 and 355 µm. North arm ooids are slightly dominated by the 63 – 355 µm size class (45%), while
southern arm ooids are strongly dominated by the 63 – 355 µm size class (73%) (Figure 4).
Serial
14
C ooid record
14
C ages of CO2 that was released during acidification of 355 – 500 µm diameter ooids
from Spiral Jetty increased in a non-linear manner from and age of 660±15 yr in the first layer
39

dissolved to 5830±60 in the last layer dissolved. The
14
C ages of ooids from Antelope Island
increased linearly from 460±20 yr to 6600±60 (Appendix B, Table 1) (uncorrected for reservoir
effect).
14
C ages of organic matter combusted from the nuclei remaining at the end of the
experiment were 5975±15 and 6210±20 yr for Spiral Jetty and Antelope Island ooids, respectively.
As dissolution progresses and ooids become smaller, each successive sample taken
represents a thicker width of ooid cortex dissolved (assuming that sample size is constant), thus
homogenizing the
14
C over a larger range of radial cortex depths. To account for this, ages were
integrated over ooid cortices ranging from 355 – 500 µm in diameter assuming they had a spherical
geometry and constant net growth (precipitation - abrasion) (Figure 5A, 5B). To summarize, our
experiments indicate that 355-500 µm ooids from GSL began precipitating around 5830-6600 ±
60
14
C years BP with a continuous chronology to near modern ages (when corrected for reservoir
effect). Organic carbon extracted from the nuclei material left at the end of the experiment yield
nearly contemporaneous ages with the oldest inorganic carbon samples, lending credence to the
presumed onset of ooid formation and our methods.
DISCUSSION
Bulk
14
C ages from sieved ooids reveal that smaller ooids are younger than larger ooids,
with age disparities between the coarse and fine ooid fractions of 1645 and 789
14
C years for the
north and south arm, respectively. The younger average bulk
14
C ages of finer sized ooids may be
attributed to a more recent onset in formation and implies that an ooid factory is active in Great
Salt Lake. Therefore, ooid size scales with age rather than some later physical sorting mechanism
indicating that ooids did in fact continue to grow in the GSL over at least the past several thousand
years. The grain size distribution is skewed toward finer grain sizes in both arms of the lake, which
is also suggestive of an active ooid factory when combined with bulk ooid age data (Table 2, Figure
40

4). Bulk ooid ages also indicate that the Great Salt Lake ooids are significantly older than the
modern marine ooids from Carbla Beach, Australia and the Bahamas (Beaupré et al., 2015;
Duguid et al., 2010). However, some significant caveats require exploration while interpreting
bulk
14
C ages. Bulk ooid ages do not allow for the differentiation between relic ooids that formed
thousands of years ago versus modern ooid formation. In addition, Raman spectroscopy
demonstrates that organic carbon is not exclusively found in peloidal nuclei but is also
incorporated throughout the ooid cortex. Thus, bulk ooid organic carbon ages represent a mixture
of organic carbon from Artemia pellet nuclei and younger organic carbon incorporated at various
points in the growth of the aragonitic cortex. Ooids from the northern arm of GSL have older bulk
organic carbon ages (Table 2) for each size fraction, including unsorted ooids. Because the total
organic carbon content in north arm ooids is twice that of south arm ooids and the bulk organic
carbon ages are older, we expect this age disparity is attributed to a higher occurrence of ooids
with organic-rich brine shrimp pellet nuclei in the north arm of the lake. Petrographic investigation
of 100 ooids in thin section from the northern and southern arm of GSL confirm this hypothesis,
with 83% pellet nuclei in the north compared to 56% pellet nuclei in the south (Figure 6). The
distribution of organic matter throughout ooid cortices coupled with the need to resolve a
chronology from the carbonate fraction, highlight both the problems with interpreting bulk age
data from ooids and the need for serial dissolution.
Our serial dissolution experiments present a chronology from modern lacustrine ooids that
demonstrate the ancient onset of ooid formation over ~6,000 years ago and once corrected for
reservoir effect, the youngest inorganic carbon ages suggest ooids continued to form until very
recently. We hypothesize ooid formation may still be occurring, as any modern
14
C would be
homogenized with slightly older
14
C in our youngest sample (this is considered more carefully in
41

Ch. 3). The
14
Corg from Antelope Island ooids is slightly younger than the oldest inorganic carbon
sample, and this may be attributed to 1) residual organics of younger origin, 2) partial leaching of
ancient carbonate material in the center of the ooids, and/or 3) a reservoir effect yielding inorganic
carbon which is apparently older by hundreds of years.
The age of the onset of ooid growth from the north and south arm of GSL is similar, as
indicated by the oldest inorganic and organic carbon ages of ooids, but the growth curve of their
chronologies varies. For example, the south arm ooids appear to have a near-constant growth rate
(between ~0.01 – 0.015 µm/yr) within the resolution of the data and assumptions. In contrast, the
growth of the north arm ooids appear to have been initially more rapid (~0.03 – 0.06 µm/yr) and
then slowed somewhat throughout their growth history (0.003 – 0.008 µm/yr). The differences in
slope may be attributed to local site-specific variations affecting carbonate precipitation or
abrasion in each part of the lake, or assumptions made when calculating dissolution depth (i.e.,
constant net growth rate, spherical geometry). To determine whether the assumption of spherical
geometry in age integration is responsible for the difference in the slope of the ooid growth curves,
we integrated the ages over assumed ellipsoidal ooid geometries. The resulting slope differences
were exacerbated when we assumed 100% ellipsoidal geometry (Figure 7), suggesting that there
are likely other effects (environmental, geochemical, or physical) during the ooid growth history
causing their differences in slope. It is intriguing that the north arm ooids fit the prediction that
ooid growth should be rapid at first and then slow as they reach hydrologic equilibrium and spend
more time as bedload versus suspended load (Trower et al., 2017) but coevally, the south arm
ooids display a linear growth trend.
Comparison to marine ooid chronologies
42

The GSL ooid growth histories raise some unexpected questions with respect to how ooids
form in the GSL and thus how ooids grow in general. The lifespan of 355-500 µm radial ooids
from Great Salt Lake is between two and six times longer than most modern marine ooids from
the Bahamas Archipelago and Australia (Beaupré et al., 2015; Duguid et al., 2010). The ooids are
very old compared to modern marine examples, yet sequential dating reveals they experienced
continuous net growth for over 6000 years while existing within the GSL environment. Trower et
al., 2017 note that the balance of precipitation versus abrasion are key components in the formation
of ooids. On the one hand, the GSL has a very different chemical environment versus the marine
settings. For example, in marine settings where ooids grow, the seasonal water temperature
variations are low, whereas the Great Salt Lake experiences comparatively large temperature
fluctuations (Figure 8). The solubility of aragonite increases as temperature decreases, thus the
favorable window for aragonite precipitation in the GSL likely only exists over a short window in
the summer when the lake water is significantly warmer, whereas marine settings are likely to be
supersaturated with respect to aragonite year-round. Additionally, we expect abrasion is less
intense in the GSL than in marine systems as GSL is a significantly lower energy environment
than marine examples.
It is unclear how the balance between abrasion and precipitation should be reconciled given
how slowly net ooid growth appears to be in the GSL system. Do they experience rapid growth
then significant abrasion on a yearly basis, thus accounting for such a slow net growth rate, or, do
they simply grow very slowly? Petrographic investigation reveals what appear to be relatively
delicate ray-like aragonite crystals that we speculate would not survive intense abrasion,
supporting the premise that that perhaps the GSL ooids simply grow very slowly. Furthermore,
how might the radial fabric affect or indicate growth rate versus the tangential fabric in modern
43
marine ooids? Future work, including finer scale sequential dating, may help resolve the
unanswered questions surrounding the GSL ooids.
Ooids and the history of the Great Salt Lake
Throughout the duration of ooid formation, the north and south arms of Great Salt Lake
would have been in communication with one another as part of one large body of water (rather
than two arms separated by a railway), thus the generally similar chronologies from ooids from
each arm of the lake agree with the lake’s history. Furthermore, 10,000
14
C yr BP marks the end
of the Gilbert episode of Great Salt Lake, where the lake experienced a brief 15m transgression
during which the lake had freshened enough to support ostracods and possibly fish (Broughton,
Madsen, & Quade, 2000; Oviatt et al., 2015). After the Gilbert episode, GSL regressed to average
historic GSL levels (near 1280m) and brine shrimp cysts and pellets appeared in lake sediment
cores. It is thought that Great Salt Lake did not transgress higher than modern lake levels during
early parts of the Holocene (11.5-10.2 cal ka BP; 10-9
14
C ka BP), but little is known about the
remainder of Holocene lake level because Holocene sediments on the floor of GSL have been
largely reworked. On the one hand, a bulk geochemical analysis of ooids would represent an
integrated signal over ~6000 years and provides at least one end member for the duration of
geochemical integration that ooids may represent, with relevance to other systems where ooids are
analyzed as paleoenvironmental indicators. On the other, sequential dissolution of the ooids
preserved in GSL ooids has the potential to resolve some of the finer scale lake level variations in
GSL during the last ~6,000 years and potentially longer.
CONCLUSIONS
44

The high-resolution
14
C chronology of GSL ooids demonstrates that:
14
C is a robust tool
for dating ooids in GSL, and GSL ooids have a lifespan between two and six times longer than
modern marine ooids. The long ooid lifespan confirms the need to temporally resolve accretionary
structures like ooids before interpreting bulk geochemical data. The
14
C ages obtained from
organics in ooid nuclei corroborate the timeframe of onset of aragonite precipitation. Additionally,
Raman spectroscopy coupled with
14
C ages from bulk unsorted and sieved ooids shed light on the
importance of sequentially derived chronologies due to the fact that bulk ages underestimate the
maximum age of ooids by thousands of
14
C years. This study highlights the disparity in net growth
rate, lifespan, and seasonality in precipitation between radial ooids from Great Salt Lake and
modern marine ooids. Ancient radial ooids, which serve as textural analogs to GSL ooids, may
have been subject to growth rates more like Great Salt Lake than modern marine examples.
ACKNOWLEDGMENTS
We thank Bonnie Baxter and Jaimi Butler for their support in organizing field sampling.
We also thank Carie Frantz, Dylan Wilmeth, and Joyce Yager for their helpful assistance in the
field and Nick Rollins for help in the lab. This work was supported in part by the SEPM student
research award.

45

FIGURES
Table 1:
14
C and δ
13
C composition of Weber, Bear, and Jordan rivers as well as well water sampled
in Ogden, Utah. Water was treated with HgCl2

Table 2. Inorganic and organic
14
C ages from bulk ooids
Locality
Grain Size
(µm)
Inorganic C
Age
(
14
C yrs BP)
Organic C
Age
(
14
C yrs BP)
% Total
Organic Carbon
Spiral Jetty -- North
Arm 1.34
Unsorted 3872±15 3490±15
355-500 4373±20 4200±15 1.3
250-355 3759±15 3520±20 0.97
125-250 2728±15 2335±15 1.19

Antelope Isl -- South
Arm

0.46
Unsorted 3556±15 2175±20
355-500 3947±15 2680±20 0.43
250-355 3834±15 2250±15 0.48
125-250 3158±15 1935±15 0.55
Water Source δ
13
C (‰) ±
fraction
Modern
±
∆
14
C
(‰)
±
14
C age

(BP) ±
Graphite Size
(mg C)
DIC
(mM)
Well -15.5 0.1 0.8377 0.0014 -169.1 1.4 1425 15 0.67 6.6
Weber River -10.2 0.1 0.9332 0.0015 -74.3 1.5 555 15 0.71 3.2
Bear River -8.4 0.1 0.8348 0.0016 -171.9 1.6 1450 20 0.69 4.1
Jordan River -9.7 0.1 0.8416 0.0014 -165.2 1.4 1385 15 0.70 4.1
46

Figure 1: Grain mount of ooids from Great Salt Lake, Utah showing primary radial aragonitic
fabric. Nuclei range from elongate peloids to quartz or feldspar grains.

47

Figure 2: Map of Great Salt Lake, modified from Currey et al., 1984. Ooid samples were collected
from the sediment water interface from: Spiral Jetty in the north arm of GSL, and Bridger Bay on
Antelope Island in the south arm of GSL.

48

Figure 3: Presence of organic matter (blue) within an ooid from the north arm (A) and south arm
(B) of Great Salt Lake acquired from Raman spectroscopy. A survey of 30 ooids was carried out
to confirm the distribution of organic matter within ooid cortices (Appendix A, Table 2). Scale
bars equal 100 µm. Organic matter is distributed within peloidal nucleus (A) and throughout
carbonate cortex (A and B).

49

Figure 4: Grain size distribution of Great Salt Lake ooids sampled from the north and south arms
of the lake. Ooids from both arms of the lake are dominated by finer sized ooids (63 – 355 µm),
though south arm ooids are more heavily skewed toward fine grain sizes.

50

Figure 5: Inorganic
14
C chronologies from north arm ooids (A) and south arm ooids (B) were
integrated over ooid cortical ranges of 355-500 µm in diameter (represented by shaded region),
assuming spherical geometry and constant net growth (see Appendix B, Table 2 for all ages and
yields).
14
C ages from organic carbon (dotted line) from remaining ooid nuclei were 5975±15 and
6210±20 yr for north arm (A) and south arm (B) ooids respectively.

(

C )

(

C )

51

Figure 6: Nuclei composition in north vs south arm GSL ooids as observed in thin section. North
arm ooids have 83% brine shrimp pellet nuclei, 15% detrital grains such as quartz and feldspar,
and 2% other ooids as nuclei. Southern arm ooids have 56% brine shrimp pellet nuclei and 44%
detrital grain nuclei.

52

Figure 7: Comparison of ooid chronologies from the northern arm of GSL (7B, 7D) to the southern
arm of GSL (7A, 7C). Figures 7A and 7B use the radiocarbon chronology and integrate the ages
assuming spherical geometry of ooids (using mean ooid radius of 213 µm). The resulting slope of
the north arm ooid growth curve (B) is steeper (more rapid growth) during the first several
thousand years of ooid growth and slows down during the alst several thousand years, while the
southern arm ooid growth curve (A) is largely linear. Because north arm ooids have a larger
occurrence of ooids with peloidal nuclei (roughly cylindrical geometry), the radiocarbon
chronology was integrated over an assumed cylindrical geometry using V = L * π r
2
, where V is
volume, L is length (6 * radius), and r is radius (213 µm) in micrometers. The slope of the north
arm ooid growth curve remains exponential even after assuming cylindrical geometry, suggesting
there are other factors (environmental or otherwise) accounting for the difference in slope.

53

Figure 8: Monthly average water temperature in north (circle) and south (square) arms of Great
Salt Lake as measured by USGS from 2010-2016. (U.S. Geological Survey, 2016)

54

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CHAPTER 3: GREAT SALT LAKE OOIDS AS HIGH-RESOLUTION RECORDS OF
HOLOCENE STABLE ISOTOPE HISTORY AND LAKE LEVEL CHANGE
Paradis, O., Corsetti, F., Bardsley, A., Hammond, D., Berelson, W., Xu, X., Walker, J., Celestian,
A.
ABSTRACT
Like many other accretionary structures, ooids (laminated coated grains) have the
potential to record a time series of hydrologic/geochemical change in marine and lacustrine
environments. In Great Salt Lake (GSL), Utah, lake level indicators are sparse between ~10,000
ybp and the onset of historical times, so the Holocene lake variation is somewhat poorly known.
Here, we evaluate the
14
C age and stable carbon isotope composition of ooids that grew in the
lake over the last 7,000 years. Since δ
13
C and δ
18
O covary in closed basin lakes, changes in the
δ
13
C through time can be interpreted to represent changes in lake volume if other assumptions we
make hold true. Ooids were examined by sequentially dissolving layers to create a radiocarbon
chronology and δ
13
C profile with depth in the ooid. Petrographic analysis reveals that, for the
majority of ooids, the initial precipitation upon the nucleus is represented by a comparatively
large aragonite crystal ray texture, followed by alternations of finely crystalline radial aragonite
and thin radial-concentric layers. Net growth rates derived from radiocarbon ages indicate the
large crystal ray fabric corresponds to 7498 to 2073 cal yr BP with δ
13
C between 4.4±0.15 and
4.8±0.15 ‰, likely indicating a relatively stable lake level. The alternating fabric precipitated
between 2073 cal yr BP to present, and the δ
13
C values of ooids trending toward lighter values
(from ~4.5‰ to 3.7‰), suggests a largely transgressive phase in GSL. The youngest inorganic
carbon sample from north arm ooids falls off this trend as it has a significantly heavier δ
13
C of
5.6±0.15‰. Modern dissolved inorganic carbon from north arm lake water indicates aragonite
70

precipitating today would have δ
13
C values between 9.1±0.55‰ to 9.8±0.75‰, therefore the
young heavy inorganic carbon of north arm ooids likely suggests that ooid formation has
occurred after the construction of the railroad causeway in 1959, which isolated the north arm
from the riverine-influenced southern portion and thus left the reservoir to generate an enriched
δ
13
C pool, representing the first robust evidence that ooid formation has occurred in modern
times. Lastly, the entire chronology of GSL ooids from both the north and south arms of the lake
record over 2‰ variability in δ
13
C over several thousand years, which calls into question the
utility of bulk geochemical analyses from accretionary structures such as ooids in dynamic
systems such as closed-basin lakes.
INTRODUCTION
Closed-basin lakes are particularly sensitive to changes in precipitation and evaporation,
thus their records of lake level fluctuation and geochemical evolution can be utilized for climatic
reconstructions in North America (Larry Benson, Lund, Negrini, Linsley, & Zic, 2003; Fritz,
Metcalfe, & Dean, 2001; Reinemann, Porinchu, Bloom, Mark, & Box, 2009). Lacustrine
carbonates such as stromatolites and speleothems have been successfully used for reconstructing
lake level fluctuations and hydrologic variability in closed-basin lakes (Berelson, Corsetti,
Johnson, Toan, & Der, 2009; Frantz et al., 2014; Mcgee et al., 2012; Newell et al., 2017;
Petryshyn et al., 2016).
The δ
18
O composition of lake water and primary carbonates tracks evaporation and
precipitation (Li & Ku, 1997), and in closed-basin lakes, the δ
18
O and δ
13
C covary (Figure 1)
(Horton, Defliese, Tripati, & Oze, 2016; Huang et al., 2014; Talbot, 1990). Thus, we may use the
δ
13
C carbonate record in terminal lakes as a proxy for lake evaporation (regression) and filling
(transgression).
71

Great Salt Lake (Utah) is a closed-basin hypersaline lake with the propensity to record
ancient environmental change in western North America. Though paleo Lake Bonneville, the
predecessor to modern GSL, has an extensive and well-characterized hydrologic history
(Broughton et al., 2000; Currey et al., 1984; Oviatt et al., 1992, 2015), lake-level indicators are
limited over the past ~10,000 ybp of GSL’s history (Figure 2).
GSL is home to extensive shoreline microbialite deposits (Baskin, 2014; Eardley, 1938),
and their utility in understanding hydrological change in GSL has been investigated by others
(Bouton, Vennin, Mulder, et al., 2016; Newell et al., 2017; Pace et al., 2016). Though
microbialite distribution is extensive, the interpretation of isotopic data from them is not
straightforward since most microbialites are poorly lithified and often porous and clotted, with
only rare occurrences of well-defined stromatolitic laminations. Additionally, radiocarbon data
from organic material within the microbialites yields enigmatic ages suggesting either older
organic material may have been incorporated during their growth, or post-depositional
groundwater infiltration or microbial activity may deposit younger organic material in the porous
regions (Newell et al., 2017). The poorly-lithified porous structure with rare laminations
combined with the lack of finely-resolved and reliable age constraints confound our ability to
reliably interpret geochemical information acquired from microbialites in GSL.
Ooids as lake level indicators
Ooids (carbonate coated grains) may be used as an alternative archive of hydrologic
change in GSL. GSL ooids are found as shoreline deposits around the lake and have recently
been demonstrated to record over 7,000 years of lake history (Chapter 2). This study uses stable
carbon isotopes coupled with a robust radiocarbon record to present a high-resolution record of
13
C over the past 7,000 years of GSL’s history. Furthermore, we characterize the petrographic
72
fabrics of individual ooid cortices to test whether lake hydrology is a driver of ooid cortex
change.
METHODS
Ooids from the sediment-water interface were sampled from: (1) Bridger Bay on
Antelope Island in the south arm of Great Salt Lake, and (2) Spiral Jetty in the north arm of
Great Salt Lake in March 2014. Ooids were rinsed three times with deionized water and dried in
an oven at 50°C. Lake water was sampled in September 2016 for
14
C and
13
CDIC analysis. One
liter of lake water was collected with no head space and immediately treated with HgCl2 to
inhibit any microbial activity. δ
13
C values reported here were measured on splits of CO2
extracted from sequential ooid dissolution or on water-DIC directly using Gas Bench coupled
with IRMS (Finnigan Delta Plus) at the University of California at Irvine (UCI). A temporal
record of δ
13
C was created using
14
C data and age models acquired using methods outline d in
Chapter 2. Uncalibrated ages (
14
C ages) are presented herein for comparison with other
publications (Appendix B, Table 1), as well as calibrated ages using IntCal13 (Reimer et al.,
2013; Stuiver, Reimer, & Reimer, 2018) which are reported in calendar years (cal yr) BP
(Appendix B, Table 1). Ooids from each site were sieved into discrete size fractions: <355 µm,
355-500 µm, and 500 – 1000 µm. Grain mounts were made for petrographic analysis, which was
carried out using Zeiss Imager M2m microscope with Axiovision software.
Sequential stable carbon isotope record of ooids
To create a
13
C record of 355-500 µm ooids, we measured the
13
C from inorganic carbon
of fractions of CO2 collected during sequential acidification of the large size fraction of sieved
ooids (355-500 µm) from Spiral Jetty and Antelope Island. Ooids (~50g) and 150ml of deionized
water (DIW) were placed in a reaction vessel constructed from a 500ml graduated round media
73
storage bottle (VWR cat. # 89000-238) and a suspended magnetic stir rod (Figure 1 from Appendix
A). The reaction vessel was purged with N2 that was scrubbed with Ascarite-II while a stir bar
spun at 700 rpm to drive off any dissolved CO2 in the water for a total of 30 minutes. The sample
was acidified by injecting 3.3M HCl at a flow rate of 10 ml/min. Gas was shunted for the first 5
seconds of acidification to off-gas any residual N2 before collecting the sample in Tedlar bags
which had been rinsed with ultra-high purity (UHP) helium scrubbed with Ascarite-II. Gas was
collected in 3 Tedlar bags per each acidification iteration. Four discrete acidifications were
performed. The reaction vessel and its components were rinsed in 10% HCl and dried between
each acidification. The DIW in the reaction vessel was replaced, and the reaction vessel was purged
for 30 minutes with ascarite-scrubbed N2 to remove any atmospheric carbon. The DIW rinse, acid
wash, and 30-minute purge was repeated for each acidification.
RESULTS
The δ
13
C record of ooids from the south arm of GSL ranges from 3.7±0.15 to 4.7±0.15‰
(Figure 3). The south arm ooid δ
13
C record varies between 4.4±0.15 and 4.7±0.15‰ from 7498
to 2354 cal yr BP, and then trends toward lighter values over the last ~2,000 years, culminating
at 3.7±0.15‰ (513 cal yr BP). The north arm δ
13
C record varies between 4.7±0.15 and
4.4±0.15‰ from 6638 to 2073 cal yr BP; however, the outermost inorganic carbon from north
arm ooids is enriched (5.6±0.15 ‰) with respect to the rest of the carbon record (Figure 3).
The δ
13
C of dissolved inorganic carbon (DIC) from lake water sampled in September
2016 was 2.3±0.15‰ and 6.9±0.15‰ for the south arm and north arm of the lake, respectively.
For direct comparison with δ
13
C from carbonates, the δ
13
CDIC values from the north and south
arms of GSL are used to calculate the range of possible aragonite values, assuming aragonite is
precipitating in equilibrium with the lake water. Given the range of average enrichment factors
74

between aragonite and bicarbonate from precipitation experiments (Romanek, Grossman, &
Morse, 1992; Rubinson & Clayton, 1969), calculated carbon isotope ratios for aragonite
precipitating in the lake are between 9.1±0.55‰ to 9.8±0.75‰ for the northern lake water, and
between 4.5±0.55‰ to 5.2±0.75‰ for southern lake water (Figure 3, shaded bars).
Petrographic analysis
Nuclei of GSL ooids include peloids which originate from brine shrimp fecal pellets
(Eardley, 1938), detrital quartz grains, igneous minerals (augite), and aragonite, though peloids
and detrital quartz are the dominant accounting for 56.7% and 26.7% of all nuclei, respectively
(Appendix B, Table 2). Grain mounts of sieved ooids (125-250 µm, 250-355 µm, 355-500 µm
diameter) reveal that 27 out of 30 (90%) individual ooids examined display large aragonite
crystal rays as the initiating fabric next to the nucleus (Figure 4; Appendix B, Table 3). The large
aragonite rays (RAY) range from 33 to 142 µm in height, with a median ray height of 78 µm.
Subsequent fabrics alternate between banded tightly-packed radial aragonite (BR) and thin clean
radial-concentric (TCR) aragonite (Figure 4; Appendix B, Table 3).
DISCUSSION
In closed basin lakes such as GSL, δ
13
C and δ
18
O exhibit a positive correlation suggesting
the stable carbon isotope record of primary lacustrine carbonates such as ooids can be used to
reconstruct the hydrologic evolution of the lake (Figure 1). Increased evaporation or residence
time results in enriched δ
18
O values through Raleigh fractionation which is accompanied by an
enrichment in the δ
13
C of the dissolved inorganic carbon (DIC) pool. Evaporation-induced
isotopic fractionation occurs as CO2 exceeding solubility leaves the system. Evaporation leads to
an alkalinity increase, and the DIC pool becomes enriched in δ
13
C heavy bicarbonate (or
75

carbonate ions if the water pH is greater than 8.3) (Abongwa & Atekwana, 2013; Dreybrodt &
Deininger, 2014; Horton et al., 2016). Though other processes including carbon isotopic
fractionation related to respiration and photosynthesis may affect the δ
13
C of the DIC pool,
results from controlled evaporation experiments suggest that the evaporation-induced increase in
alkalinity (via equilibrium exchange with atmospheric CO2) drives the DIC enrichment (Horton
et al., 2016).
The sequentially derived stable carbon isotope record from Great Salt Lake ooids reveals
that δ
13
C varies between 4.4±0.15 and 4.8±0.15 ‰ from 7498 to 2073 cal yr BP, suggesting the
lake had a predominantly stable level during this time. Petrographic analysis reveals that the
common initial nucleating crystal fabrics amongst GSL ooids are large aragonite crystal rays.
The large aragonite rays have a median crystal height of 78.47µm, which corresponds to the first
4130-6036 years of the ooid chronology as median net growth rates of GSL ooids range from
0.013 to 0.019 um/year for south arm ooids, and 0.013 to 0.018 um/year for north arm ooids. The
absence of erosional or dissolution surfaces within the continuous aragonite rays likely indicates
that the calculated net growth rates are representative of the geochemical conditions in the lake
(i.e., high ionic strength, salinity, and seasonal temperature variability) rather than an offsetting
high precipitation and abrasion rates. The time period calculated from median aragonite crystal
ray height and net growth rates closely corresponds to the period in which there was little
variability in δ
13
C and therefore a likely stable lake level (Figure 5). The shift from large,
continuous aragonite rays to finer alternating radial aragonite textures is coincident with the shift
to lighter δ
13
C in the last ~2000 years (for south arm ooids), suggesting the cortex fabric may be
governed by geochemical changes.

76

After 2073 cal yr BP, the south arm ooid δ
13
C record becomes progressively lighter,
eventually reaching 3.7±0.15 ‰ 513 cal yr BP, suggesting an increasing lake level likely due to
increased regional precipitation. However, the youngest inorganic carbon sample from north arm
ooids (composite age of 598 cal yr BP) has a δ
13
C value of 5.6±0.15 ‰, which is considerably
heavier than the remainder of the δ
13
C record. Though the youngest sample has an age of 598 cal
yr BP, it represents a homogenization of a portion of the outermost cortex and thus, an integrated
age spanning the outermost surface of the ooid to a depth within the ooid. Any modern carbon
forming today will be homogenized within the youngest composite sample. The δ
13
C of modern
north arm lake DIC indicates that carbonates forming in the north arm of GSL today would have
a δ
13
C value between 9.1±0.55‰ to 9.8±0.75‰. This enrichment is likely the result of the
division of GSL into two arms via a rock-fill railroad causeway in 1959, isolating the north arm
from riverine inflow eventually leading to increased evaporation and hypersaline lake water on
this side. Without steady delivery of isotopically light δ
13
C from river inflow, continuous
carbonate precipitation in the north arm of GSL will leave the reservoir enriched as we see today.
The enrichment of the youngest carbonate from north arm ooids (and absence of this enrichment
in the youngest south arm ooid sample) coupled with the measured modern δ
13
CDIC suggest that
there has been ooid growth on the Northern side of GSL after the construction of the railroad
causeway in 1959. The averaging effect all the data undergo due to the nature of the experiments
made it difficult to comment on whether ooids are actively forming in GSL. The stable carbon
record of north arm ooids is the first evidence that ooids have been actively forming in the last
several decades—an untested assumption that has been perpetuated over the years. Additionally,
the stable carbon isotope record of GSL ooids records over 2‰ variability over the course of
several thousand years, which illustrates that the bulk sampling of ooids (especially in
77

climatically sensitive terminal lakes such as GSL) will likely result in geochemical data that
confounds our ability to comment on the hydrologic evolution of the system.
We can construct a stable carbon isotope mass balance (Equation 1 and 2) assuming the
chronology and stable carbon ooid data records discrete ages/values, and that the youngest
carbonate value is a mix of two end members: (1) derived “modern” north arm carbonate values
and (2) stable homogeneous lake values before the δ
13
C depletion occurring around 2000 cal yr
BP. The inorganic carbon isotope composition of the youngest north arm ooid sample (δ
13
CO) is
equal to a combination of enriched modern carbon, δ
13
CEC, and ancient isotopically light lake
carbon, δ
13
CLC (Eq. 1). The fraction of enriched modern inorganic carbon and isotopically light
homogenous lake carbon are represented by fEC and fLC, respectively (Eq. 2).

δ
13
CO = δ
13
CEC * fEC + δ
13
CLC * fLC (1)

fEC + fLC = 1 (2)

Using 5.6‰ for δ
13
CO, 4.5‰ for δ
13
CLC, and either 9.1‰ or 9.8‰ to represent the range
of possible δ
13
CEC, we can solve Eq. (1) by substituting Eq. (2) once rearranged to solve for fEC.
The results indicate that 20-24% of the youngest inorganic carbon sample from north arm ooids
came from enriched “modern” carbonate precipitation. The calibrated age of the youngest
sample (598 cal yr BP) and the calculated linear depth from Chapter 2 indicate that roughly 5
micrometers of calcium carbonate precipitation occurred over 664 years (598 cal yr BP plus 66
78

years from 1950 to 2016, when the DIC was sampled). If 20-24% of the youngest carbonate
sample originated from post-causeway enriched water, and we assume this enrichment occurred
instantaneously, we are left with a net growth rate of ~0.002 µm/yr after the construction of the
railroad causeway. These estimates agree with linear growth rates calculated from the
radiocarbon chronology of ooids, giving another confirmation of the slow net growth rates
during this time period.
The stable carbon isotope composition of modern south arm DIC (Figure 1), suggests that
aragonite precipitating from the south arm of the lake have δ
13
C values ranging from 4.5±0.55‰
to 5.2±0.75‰. If ooids have been forming in the south arm in recent years, one would expect the
youngest inorganic carbon values from south arm ooids to represent a mix of inorganic carbon
from the modern and ancient lake; however, the youngest inorganic carbon from south arm ooids
do not indicate recent precipitation. This deviation may be due to the absence of “modern”
aragonite precipitation in south arm ooids, a short interval of lake rise between 1251 and 1766
cal yr BP which resulted in a depleted inorganic carbon reservoir (resulting in the depletion of
the subsequent inorganic carbon sample), or the “modern” aragonite values may extend in the
negative direction due to seasonal fluctuation in δ
13
C as our water samples were collected in
September.
CONCLUSIONS
The stable carbon isotope record of GSL ooids suggests that Great Salt Lake had a
roughly stable lake level from 7498 to 2073 cal yr BP, with δ
13
C fluctuating from 4.4±0.15 to
4.8±0.15 ‰. Growth rates from the radiocarbon chronology of GSL ooids indicate that the large
aragonite crystal rays comprise the first 4130-6036 years of the ooid chronology as they are the
dominant initiating crystal fabric. The large aragonite crystal rays are coincident with a time
79

when GSL water remained near 4.5‰. The aragonite rays are often followed by alternating
finely crystalline radial aragonite and thin radial concentric aragonite, a shift that coincides with
a trend toward depleted carbon isotopes that began around ~2000 years ago. Additionally, the
youngest δ
13
C of inorganic carbon from north arm ooids is enriched compared to the remaining
stable carbon isotope record. Each δ
13
C value is measured on a homogenized sample over several
microns of carbonate growth, thus we interpret the heavy north arm δ
13
C of 5.6±0.15‰ to
represent a mix of carbonate from the ancient homogeneous lake and carbonate precipitating
after the construction of the railroad causeway in 1959, which left the north arm water restricted
from inflow and with a heavier pool of DIC as carbonate precipitated over time. The sequentially
derived δ
13
C record of both north and south arm ooids provides the first evidence of post-
causeway ooid formation over the last several decades and it demonstrates the utility of ooids to
record several thousand years of hydrologic change in which over 2‰ of variability is recorded
in δ
13
C.
ACKNOWLEDGMENTS
This research was supported by the Keck Carbon Observatory at the University of California,
Irvine, the SEPM (student research grant program), and Nick Rollins for help in the laboratory.
We thank Bonnie Baxter and Jaimi Butler for their support in field logistics, as well as a Carie
Frantz, Dylan Wilmeth, and Joyce Yager for their helpful assistance in the field.

80

FIGURES

Figure 1: Relationship between δ
13
C and δ
18
O in closed-basin lakes, adapted from Talbot, 1990.
δ
13
C covaries with δ
18
O in carbonates within terminal lake systems. Closed-basin lakes with a
large surface area to depth ratio (such as Great Salt Lake) tend to have a less steep slope than
lakes with a low surface area to depth ratio.

81
Figure 2: Hydrograph depicting lake level fluctuation in Lake Bonneville and Great Salt Lake
throughout the past 30,000 years (modified from Murchison, 1989; Newell et al., 2017; Oviatt,
1997, 2015; Oviatt et al., 2015). The transgressive phase (T), the overflowing phase (O), and the
regressive phase (R) are marked with gray boxes on the top of the figure. Lake Bonneville
fluctuated from ~30 cal ka until ~15 cal ka when it regressed to Great Salt Lake levels.
82

Figure 3. Stable carbon isotope (δ
13
C) record of inorganic carbonate carbon from ooids through
time. Ages are calibrated from the
14
C chronology found in Paradis et al. 2019 using the Intcal13
calibration (Table 1) (Reimer et al., 2013; Stuiver & Reimer, 1993; Stuiver et al., 2018).
Calibrated ages are given in calibrated years BP (before 1950). The grey bar shows the range of
values expected for aragonite precipitating in equilibrium with Great Salt Lake water sampled in
September 2016 from the north and south arms of GSL (Romanek et al., 1992). Data from north
arm ooids of 355-500 µm diameter are depicted by open circles, and south arm ooids of 355-500
µm diameter are closed circles.

83

Figure 4: Great Salt Lake ooid with pellet nucleus (A). Large aragonite crystal rays are often the
initiating crystal fabric against the nucleus (B). The other common crystal fabrics include banded
tightly-packed radial aragonite crystals (white arrow), and thin radial-concentric aragonite
crystals (black arrow).

84

Figure 5: A comparison of the radiocarbon chronology of 355-500 µm ooids from the southern
arm of GSL (TOP) and the δ
13
C record from inorganic carbon of ooids from the same size
fraction (BOTTOM). Radiocarbon ages are given in calibrated years BP. The shaded gray bar
indicates an approximation of the occurrence of large aragonite crystal rays throughout the
record using median crystal ray height (78 µm). The large aragonite rays are coincident with a
stable time in GSL history, when δ
13
C remained near 4.5‰, and the shift in microfacies from
large aragonite rays to finely laminated radial aragonite is coincident with a shift in δ
13
C toward
depleted values.

85

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CHAPTER 4: MICROBIAL COMMUNITIES OF LACUSTRINE OOIDS AND
CORRESPONDING NUCLEI REVEAL RELICT DNA MAY BE AN IMPORTANT SOURCE
OF DNA IN STUDIES OF BIOGENICITY
Paradis, O., Corsetti, F., Berelson, W., Stamps, B., Stevenson, B., Nunn, H., Spear, J.
ABSTRACT
Ooids are a significant component of the sedimentary rock record throughout Earth’s
history, but the mechanisms governing their formation are still debated. Traditionally viewed as
abiogenic precipitates, the role of biology in ooid formation has recently been proposed because
certain microbial metabolisms may either foster carbonate precipitation or dissolution by
manipulating dissolved inorganic carbon and alkalinity. This study evaluates the microbial
community of lacustrine ooids from the southern arm of Great Salt Lake and the brine shrimp
(Artemia) pellets upon which they commonly form. The small subunit rRNA analysis revealed
ooids and Artemia pellets have similar microbial assemblages dominated in Bacteroidetes,
Alphaproteobacteria, Deltaproteobacteria, and Gammaproteobacteria. The striking similarity
between the microbial communities of GSL ooids and brine shrimp gut microbiota suggests
relict DNA residing in oolitic nuclei may be an important source of DNA in microbial
community analysis of ooids and other accretionary structures. The calcium carbonate saturation
state of GSL is supersaturated with respect to aragonite, though it ranges from 100% to 250%
saturation seasonally. Calcification rates of GSL ooids are several orders of magnitude lower
than experimental precipitation of aragonite, suggesting aragonite may be precipitating in GSL
just above saturation, conditions that do not necessitate the geochemical manipulation of
microbiota to achieve in this system.

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INTRODUCTION
Ooids are spherical to ellipsoidal grains (usually <2mm in diameter) coated in calcium
carbonate (i.e., calcite or aragonite). Ooids are traditionally viewed as abiogenic grains, produced
through physical and chemical processes in shallow agitated water that is supersaturated with
respect to calcium carbonate (Duguid et al., 2010; Lowenstam & Epstein, 1957; Morse &
Mackenzie, 1990; Sumner & Grotzinger, 1993). Recent studies attribute modern marine ooid
formation to biogenic processes, as several microbial metabolisms are known to affect total
alkalinity (the propensity of a solution to neutralize an acid) and dissolved inorganic carbon
(DIC), which is composed of carbonic acid, bicarbonate, and carbonate ions (Diaz et al., 2013,
2014) .
Metabolisms that consume dissolved inorganic carbon (DIC), often via the consumption
of CO2, and/or generate alkalinity will promote carbonate precipitation, while metabolisms that
consume alkalinity and/or generate DIC promote calcium carbonate dissolution. This study
defines biogenic precipitation as the precipitation of calcium carbonate that occurs as a result of
the microbial metabolisms (also known as “microbially-induced precipitation” (Dupraz et al.,
2009). The likelihood that microbial processes will affect carbonate precipitation is governed by
the net effects on alkalinity and DIC by the entirety of the microbial community (not one taxon).
In addition, microbially-generated extracellular polymeric substances (EPS) may serve as a
template for carbonate precipitation when accompanied by the microbial degradation of EPS
which liberates Ca
2+
ions into the environment (Dupraz et al., 2009). The presence of unaltered
EPS alone, however, is not evidence of microbially-influenced carbonate precipitation as EPS is
known to inhibit the nucleation of calcium carbonate in the absence of EPS degradation
(Kawaguchi & Decho, 2002). Several microbial metabolisms have been identified as likely
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candidates for inducing biogenic calcium carbonate precipitation: sulfate reduction (Dupraz et
al., 2004; Visscher, Reid, & Bebout, 2000), oxygenic (Riding, 2000) and anoxygenic
photosynthesis (Bundeleva et al., 2012), and anaerobic oxidation of methane (Michaelis et al.,
2002).
Biogenicity of ooids in marine settings
Some studies on modern marine ooids have concluded microorganisms play an active
role in ooid formation based on: (1) the presence of organic matter trapped within ooid cortex
layers (Folk & Lynch, 2001; O’Reilly et al., 2017); (2) the presence of a common lipid
assemblage across modern and ancient ooids suggesting ooids are colonized by a common
distinct microbial community that potentially mediates calcification (Summons et al., 2013); (3)
high microbial diversity in marine ooids compared to the diversity found in adjacent stromatolite
systems (Diaz et al., 2013, 2014); and (4) the presence of a functional gene assemblage which is
linked to microbial metabolisms that foster calcification which surpasses the abundance of
metabolisms that foster carbonate dissolution (Diaz et al., 2014).
Studies using 16S rRNA and GeoChip4 functional gene analyses provide a picture of the
potential of the microbial community in each environment. The 16S rRNA analyses of ooids,
stromatolites, and thrombolites in the Bahamas reveal that ooids possess the highest diversity
when compared to the microbial diversity in stromatolites and thrombolites in the same setting
(Diaz et al., 2013, 2014). Functional gene analysis of ooids in the Bahamas reveals the presence
of genes responsible for metabolisms that have the potential to foster carbonate precipitation:
denitrification, oxygenic photosynthesis, sulfate reduction, and ammonification. Microbial
metabolisms that foster carbonate dissolution (sulfate oxidation, nitrification, aerobic sulfide
oxidation) have a lower relative abundance in Bahamian ooids. In addition, genes for EPS-
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degradation support the role of EPS (and the degradation thereof) in the calcification of ooids.
The presence and higher relative abundance of genes responsible for metabolisms that foster
carbonate precipitation coupled with the genetic potential for EPS degradation lead the authors to
conclude the net effect of the microbial community likely fosters carbonate precipitation (Diaz et
al., 2013, 2014), although a direct link between microbial process and carbonate precipitation
was not presented. Lipid biomarkers from ooids from Shark Bay, Western Australia, and the
Bahamas suggest that a common microbial community colonizes the ooids from both locations.
Carbon and Hydrogen isotopes from the fatty acid methyl esters indicate the lipids originate from
a diverse assemblage of microbes including sulfate-reducing bacteria, heterotrophs, and
photoautotrophs. The study suggests that the shared microbial community in marine ooids from
the Bahamas and Western Australia may be important in calcification, though the evidence for
direct involvement in calcification is circumstantial (O’Reilly et al., 2017; Summons et al., 2013)
Sulfate reducers are usually implicated in enhancing calcium carbonate precipitation by
increasing alkalinity, however depending on the local geochemistry and substrate, sulfate
reduction may actually have no net effect or even inhibit carbonate precipitation (Loyd et al.
2012, Meister 2013).
In contrast to the previous work, Duguid et al. (2010) acknowledged the presence of
microorganisms in ooids but concluded that the microbes engage in diagenetic processes which
alter the chemistry and texture of ooids after they have been formed, and thus are not playing a
role in ooid formation. The formation of micritized ooids found in stromatolites from Exuma
Cay, Bahamas has been attributed to the activities of endolithic cyanobacteria (Solentia sp.),
whereby the ooids undergo microboring and infilling of fibrous aragonite. The cycles of boring
and infilling occurs when the cyanobacteria remain at the surface of the stromatolites (Macintyre,
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Prufert-Bebout, & Reid, 2000). This study illustrates the importance of microbes in the
precipitation of carbonate in microborings and highlights how endolithic bacteria have the
potential to complicate biomarker and microbial community analysis as they are not responsible
for building the structure of the ooid.
Biogenicity of lacustrine ooids: Great Salt Lake, Utah
Though there is evidence that the microbial communities of ooids in marine settings have
the potential for fostering calcium carbonate precipitation, there is little work investigating the
potential for microbially induced or influenced carbonate precipitation on ooids in lacustrine
systems. Lacustrine ooids have occurred throughout the geologic record dating back to the
Archaean (Awramik & Buchheim, 2009). Great Salt Lake in Utah is the fourth largest terminal
lake in the world and well known for its modern occurrence of lacustrine ooids (Eardley, 1938;
Halley, 1977; Kahle, 1974; P. Sandberg, 1975). The connection between GSL’s fauna and ooid
was documented when Eardley (1938) hypothesized that brine shrimp (Artemia) fecal pellets
served as the dominant nuclei that give Great Salt Lake ooids their characteristic elongated “pill”
shape based on similarities in morphology between modern brine shrimp pellets and rod-shaped
GSL nuclei in thin-section (Figure 1). Previous studies on Great Salt Lake were concentrated in
the 1970’s and described their radial crystal fabric and morphology (Kahle, 1974; Sandberg,
1975; Halley, 1977).
More recently, it has been suggested that GSL ooids are a result of both
organomineralization and local increased alkalinity (due to strong sulfate reduction occurring in
the sediments) (Reitner, 1997). This study documents the high organic matter content of GSL
ooids and demonstrates that the most dominant sources of organic matter within ooids are insect
remains from Ephydra (brine flies) and terrestrial plant organic material, with minor components
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of microbial polysaccharides. They hypothesize the growth and fabric of GSL ooids is mediated
by organic material adhering to the surface of the ooids. The hydrocarbons found in total
sediment samples are similar to those found in purified ooids, suggesting the formation of ooids
is not connected to biological activities (biofilms), however the authors do invoke an alkalinity
pump driven by sulfate reduction (Reitner, 1997).
Recently, microbial mediation has been invoked in the formation of the oldest known
calcitic radial ooids from the 2.72 Ga Meentheena Member, Tumbiana Formation carbonate lake
system based on the presence of organic matter in close association with primary and early
diagenetic mineral phases in the ooids. The authors suggest the organic matter occurred was
incorporated during accretion and attribute the organic matter to the presence of microbial
biofilms which may be responsible for the mediation of ooid precipitation (Flannery et al., 2018).
Petrography reveals that Great Salt Lake ooids lack the extensive endolithic activity that
modern marine ooids experience which means GSL ooids are a less complex system in which to
investigate biogenicity (Figure 1, right). The goal of this study is to apply modern molecular
tools (16S rRNA sequencing) to investigate the potential of the microbial community of GSL
ooids to foster calcium carbonate precipitation via metabolic chemistry. A survey of the genetic
potential of the microbial community of ooids and the surrounding lake water will provide the
framework with which to evaluate the propensity for major and minor metabolic pathways to
either foster or inhibit calcium carbonate precipitation. In addition, the microbial community of
the brine shrimp (Artemia) pellets will be analyzed as they serve as a dominant nucleus for many
ooids in GSL (Eardley, 1938).

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METHODS
Study site and sampling
Subaqueous ooids (up to 20cm water depth) were sampled at the sediment-water interface
in Bridger Bay, Antelope Island in south arm of GSL in August 2015. Ooids were sieved into
discrete size fractions (<0.355mm, 0.355-0.500mm, and 0.500-1.00mm), pulverized using a
mortar and pestle, and technical replicates were collected from each size fraction. Unfiltered lake
water was collected in August 2015, and 960ml was centrifuged at 9,000 RPM, 4℃ for 12
minutes to pellet DNA within an hour of sampling. Unfiltered lake water from the southern arm
of GSL was collected in August 2015, with abundant brine shrimp (Artemia). The brine shrimp
fecal pellets were collected from the water using a pipette after 3 weeks had passed.
Additionally, ooid samples were collected from the north and south arms of GSL in Spiral Jetty
and Antelope Island, respectively in January 2015 for 16S rRNA analysis. The microbial
community comparison from these may be found in Appendix C. The ooid samples, DNA pellets
from lake water, and brine shrimp fecal pellets were immediately suspended in 750µl
Xpedition™ Lysis/Stabilization Solution (Zymo Research Co., Irvine, CA, United States), and
homogenized for 60 seconds using a reciprocating saw. DNA was extracted from the preserved
samples using the Zymo Research Xpedition Soil/Fecal DNA MiniPrep Extraction kit (Zymo
Research Co.) according to the manufacturer’s instructions.
Microbial community composition analysis
The extracted DNA was amplified using PCR with primers that span the V4 region of the
16S rRNA gene between positions 515 and 926 (Escherichia coli numbering), producing a
product of approximately 400 bp. The forward primer forward primer 515F-Y (GTA AAA
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CGA CGG CCA G CCG TGY CAG CMG CCG CGG TAA-3′) contains the M13 forward
primer (in bold) fused to the gene-specific forward primer (underlined). The reverse primer 926R
(5′-CCG YCA ATT YMT TTR AGT TT3′′) was unmodified from Parada, Needham, &
Fuhrman, 2016. All reactions used 5 PRIME HotMasterMix (Quanta Biosciences, Beverly, MA,
United States) to reach a final reaction volume of 50 µl. PCR thermal cycling conditions
included an initial denaturation of 95℃ for 2 min, 30 cycles of 95℃ for 30s, 50℃ for 45s, 68℃
for 45s, and a final extension of 68℃ for 10m. Positive (E.coli) and negative (no template)
controls were amplified along with the same template reactions. The reactions were purified
using Agencourt Ampure XP paramagnetic beads (Beckman Coulter Inc., Indianapolis, IN,
United States) at a 0.8x final concentration. A second cycle of PCR was used to attach a unique
12 bp barcode to each library (Stamps et al., 2016). The final barcoded products were cleaned
and concentrated using Ampure XP Paramagnetic beads, quantified using the Qubit™ dsDNA
HS assay kit (Life Technologies, Carlsbad, CA, USA), pooled in equimolar amounts, and
concentrated to a final volume of 80 µl using an Amicon Ultra-0.5 ml 30K Centrifugal Filter
(EMD Millipore Sigma, Billerica, MA, United States). The final pooled library was submitted
for sequencing on the Illumina MiSeq (Illumina Inc., San Diego, CA, United States) using
PE250 V2 chemistry.
Analyses of the small sub-unit (SSU) rRNA gene were carried out within QIIME
(Caporaso, Kuczynski, et al., 2010), wherein reads were merged and de-multiplexed, filtered at a
minimum quality score of 20, and clustered into operational taxonomic units (OTUs) using
USEARCH 6.1 (Edgar, 2010). Representative sequences for each OTU were assigned taxonomy
by mothur (Schloss et al., 2009) against the Greengenes database (DeSantis et al., 2006;
Mcdonald et al., 2012; http://greengenes.secondgenome.com; version 13_8). Representative
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sequences of each OTU were then aligned with PyNAST (Caporaso et al., 2010a) against the
Greengenes database (v13_8), and a phylogenetic tree was generated using FastTree (Price,
Dehal, & Arkin, 2010), which was used to produce a weighted UniFrac distance matrix
(Lozupone & Knight, 2005) for principal coordinate ordination.
Carbonate geochemistry
Lake water was sampled for TCO2 (H2CO3 + HCO3
-
+ CO3
2-
) analysis in January 2015,
and TCO2 data were obtained using a cavity ring-down spectrometer (CRDS; Picarro). pH was
measured using a SevenGo Duo pH meter (Mettler Toledo) in the field. Field-measured pH,
TCO2, temperature, and ionic concentration (Rupke & Mcdonald, 2012) were used to calculate
saturation indices (SI) in PHREEQ-C (Parkhurst & Appelo, 2013). Inputs for saturation index
calculations may be found in Table 1 (Appendix C). The saturation indices from PHREEQ-C
(Appendix C, Tables 2-3) were converted to saturation state values, known herein as Omega (Ω),
using the equation: Ω = 10^(SI), where SI = saturation index.
RESULTS
The water from the southern arm of GSL contained higher relative abundances of
Archaea (Euryarchaeota) compared to the pellets of the brine shrimp Artemia and ooids (Figure
2). Specifically, the water is dominated by salt-tolerant Halobacteria (Archaea), which account
for 55 – 57% of the total microbial community analyzed. After Archaea, Gammaproteobacteria
(Proteobacteria) and Bacteroidetes account for 20 – 22% and 17 – 19% of the community,
respectively. The microbial community of the Artemia pellets is dominated by Bacteroidetes (32
– 37%), Proteobacteria (36 – 40%), and Cyanobacteria (19 – 23%), most of which are
Chloroplasts from Eukaryotic organisms (likely from halophilic algae) in the water in which the
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brine shrimp were living and likely eating. The Proteobacteria in the Artemia pellets are
dominated by Gammaproteobacteria and Deltaproteobacteria.
Unsorted and sieved (<0.355mm, 0.355-0.500mm, and 0.500-1.00mm) ooids were
pulverized in triplicate, DNA extracted, and the small subunit rRNA composition was analyzed.
Unsorted ooids and ooids of <0.355mm and 0.355-0.500mm in diameter are dominated by
Bacteroidetes, Alphaproteobacteria, Deltaproteobacteria, and Gammaproteobacteria, and to a
lesser extent, Cyanobacteria and Planctomycetes. In contrast, 0.500 – 1.00mm diameter ooids
are dominated by a higher relative abundance of Alphaproteobacteria and Betaproteobacteria,
and a lower relative abundance of Cyanobacteria, Planctomycetes, and Bacteroidetes than the
remaining ooids, both unsorted and those with smaller diameters.
The microbial communities from southern arm lake water, Artemia pellets, and unsorted
and sieved ooids were compared using β-diversity analyses and visualized using principal
coordinate analysis (PCoA) (Figure 3). While the communities of the unsorted and the sieved
ooids appear broadly similar in composition (Figure 2), the coarsest size fraction (0.500 –
1.00mm diameter) ooids show a both a distinct increase in Alphaproteobacteria and
Betaproteobacteria and a decrease in Bacteroidetes and Cyanobacteria when compared to the
remaining ooid samples and therefore, the 0.500 – 1.00mm ooids were separated from the rest of
the ordination (Figure 3). The community composition of the Artemia pellets closely resembles
the communities of the unsorted, <0.355mm diameter ooids, and the 0.355 – 0.500mm diameter
ooids (Figure 2), and clusters with these samples in the ordination (Figure 3). In contrast, the
high relative abundance of Halobacteria (Archaea) in southern arm lake water separates these
samples from the ooids and Artemia pellets (Figure 3). A microbial community analysis of ooids
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from the north and south arm of GSL can be found in Appendix C. The microbial communities
of north and south arm ooids are similar, likely owing to the common core DNA in their nuclei.
The saturation state of calcium carbonate (aragonite) was calculated in PHREEQ-C
(Parkhurst & Appelo, 2013) using temperature, pH, TCO2, and ionic data found in Table 1
(Appendix C). Given an average TCO2 concentration in the southern arm of GSL of 5 mM, and
pH of 8.1, the saturation state of aragonite in the winter (~4℃) is close to saturation (Ω=1.2),
while in the summer the saturation state (Ω) is 2.5 (Figure 3).
DISCUSSION
Small subunit rRNA composition of ooids from GSL and the gut microbiota of brine
shrimp (Artemia) confirms the relationship between GSL ooid nuclei and brine shrimp pellets
that has been previously hypothesized based on morphology alone (Eardley, 1938). The
microbial community composition of brine shrimp pellets closely resembles the microbial
community of every ooid sample except the largest size fraction (0.500 – 1.00mm diameter),
which suggests that much of the DNA in GSL ooids originates from pellet nuclei. The microbial
community of the largest size fraction of ooids differs from the remaining ooids and the Artemia
pellets (Figure 2). The larger ooids are less dominated by brine shrimp fecal pellet nuclei than
smaller ooids are, and thus have a higher proportion of calcium carbonate and therefore, DNA
incorporated into the carbonate cortex. The similarity of the microbial communities of most ooid
samples and brine shrimp pellets coupled with the enrichment of those samples in fecal bacteria
(i.e., Bacteroidetes, Firmicutes) suggests the microbial community of GSL ooids may be
dominated by DNA from incipient brine shrimp pellet microbiota. Given the recent radiocarbon
chronology of Great Salt Lake ooids (Chapter 1) which gives ages of >6,000 years old for
organic carbon residing in GSL nuclei, we hypothesize a portion of the DNA recovered from
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GSL ooids is relict DNA from ancient Artemia gut microbiota. While bulk 16S rRNA analysis
gives insight into the potential of the microbial community, the presence of relict DNA may
inhibit our ability to assess biogenicity of GSL ooids as the microbiota residing on the surface of
ooids and at the carbonate-water interface is the microbial community of interest. Great Salt
Lake ooids may, however, shed light on the ancient gut microbiome of Artemia from several
thousand years ago. Artemia in GSL have a life cycle that begins with the hatching of the winter
eggs in the spring (April), which corresponds with the bloom of the halophilic algae Dunaliella.
The brine shrimp feed on algae and undergo up to four generations of turnover (hatching, feeding
on algae, and death) before the winter kill in November when temperatures fall below 6℃
(Cuellar, 1990; Post, 1977). The chloroplasts found in Artemia pellets and ooids are likely a
result of the brine shrimp consuming green algae.
Carbonate chemistry of Great Salt Lake
To investigate the biogenicity of a carbonate accretionary structure, we must determine if
the system has the propensity to precipitate calcium carbonate minerals in the absence of
microbial activity and characterize the carbonate chemistry of the environment so that we may
understand how microbial metabolism may foster carbonate mineral precipitation or dissolution.
Using any two carbonate chemistry parameters (pH, TCO2, Total Alkalinity), we can constrain
the saturation state (Ω) of the calcium carbonate mineral of question (1). Saturation states less
than 1 favor dissolution of calcium carbonate, saturation states greater than 1 have the propensity
to precipitate calcium carbonate, and a saturation state equal to 1 is at saturation.
Ω = [Ca] * [CO3] (1)
Ksp’ (aragonite/calcite)

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The saturation state of aragonite in the southern arm of GSL is 1.2 and 2.5 in the winter
(temperature minimum) and summer (temperature maximum), respectively (Figure 4). Because
the saturation state of aragonite approaches saturation (Ω = 1) in the winter due to below-
freezing water temperatures, the favorable window for appreciable calcium carbonate
precipitation is likely in the summer when lake water temperatures rise. In contrast, low latitude
subtropical marine environments where modern ooids are found (i.e., Bahamas, Turks and
Caicos) experience summer saturation state values of ~4 (Langdon et al., 2000), which may be
attributed to either the higher relative Ca concentration in marine water (~10mM Ca) compared
to GSL water (~5mM Ca) or the higher salinity in GSL, which increases the solubility of
aragonite thus reducing the saturation state. Though GSL has lower calcium carbonate saturation
than marine environments, it still has the propensity to precipitate aragonite, though likely at
lower calcification rates. Radiocarbon ages of Great Salt Lake ooids reveal net growth rates
between 0.01 – 0.1 µmol m
-2
h
-1
, several times lower than carbonate growth rates under
simulated seawater conditions (Burton & Walter, 1987; Mucci & Morse, 1983; Romanek et al.,
1992; Romanek, Morse, & Grossman, 2011; Zhong & Mucci, 1989). Biogenic carbonates such
as marine corals are sensitive to shifts in saturation state, with calcification rates covarying with
seasonal shifts in saturation state (Langdon et al., 2000). Calcification rates of colonies of the
coral S. pistillata increase exponentially as a function of aragonite saturation state above Ω=1
until they reach a plateau between Ω values of 4.2 to 5.8 (Gattuso et al., 1998). Experiments of
coral growth under reduced saturation states either by reducing calcium concentration (Gattuso,
et al., 1998) or carbonate concentration (Langdon et al., 2000), both had a significant impact on
coral calcification rates.

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Implications of relict DNA in studies of biogenicity
The role of the activity of microbes in the precipitation of ooids, stromatolites, or any
carbonate discerned from DNA analyses is not necessarily straightforward. Recently, Petryshyn
et al., (submitted) suggested that DNA within carbonates could originate from builders (microbes
that had a role in carbonate precipitation), tenants (microbes that inhabited the structure later,
e.g., endoliths or the like) and/or squatters (microbes that ended up in or on the carbonate by
accident, settling from the water column or elsewhere). The Great Salt Lake ooids provide an
opportunity to investigate the builder-tenant-squatter concept in carbonates that have enigmatic
biogenicity and form in an extreme hypersaline environment.
This study demonstrates the DNA found in GSL ooids is dominated by DNA from the
brine shrimp fecal pellet nuclei, which, in the parlance of Petryshyn et al. (submitted), would be
considered “squatter” DNA, as it was not involved in construction or later inhabiting of the
structure by microbes. However, it may be prudent to add a fourth category to the origin of
organic matter (relict) to indicate DNA that existed in the system prior to the formation of the
structure. The dominance of relict DNA in bulk ooid samples complicates the utility of 16S
rRNA analysis in assessing the biogenicity of ooids and other accretionary structures that may
harbor ancient DNA. On one hand, some might use the presence of Cyanobacteria in GSL ooids
as evidence of the potential that photosynthetic activity likely fosters carbonate precipitation.
However, the presence of chloroplasts suggests that DNA is likely from the algae the Artemia
consumed. The same may be true for ooids in modern marine systems. Ooids from Highborne
Cay, Bahamas and Carbla Beach, Australia can range from 1310±80
14
C years in age to modern
(Beaupré et al., 2015). Studies using 16S rRNA and GeoChip 4 functional gene analyses have
revealed a highly diverse population of microorganisms in Bahamian ooids that are dominated
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by metabolisms that foster carbonate precipitation (i.e., denitrification, autotrophic CO2 fixation,
sulfate reduction, and ammonification) (Diaz et al., 2013, 2014; Summons et al., 2013).
However, the presence of DNA related to those metabolisms does not necessarily mean they
would have had contact with the ooid-water interface (i.e., they may have resided in nucleus).
Because many marine ooids are typically superficial ooids (with only a few laminations), where
the thickness of the cortex is less than half of the radius of the entire ooid (Simone, 1980), the
concentration of any DNA residing in the nucleus relative to the carbonate cortex will be more
enriched due to the superficial nature of the calcium carbonate.
Raman spectroscopy reveals organic material is distributed throughout GSL ooid nuclei
and their carbonate cortices (Appendix A), confirming the results from Reitner (1997) which
demonstrated much of the organic material in GSL ooids originates from insect and terrestrial
organic matter, likely incorporated in the carbonate cortex during precipitation. These results
further highlight the caution needed when interpreting analysis of organic material bound within
ooids, as it may be unrelated to microbial biofilms. For instance, the presence of organic matter
in close association with primary and early diagenetic mineral phases in the Neoarchaean ooids
of the Tumbiana Formation is attributed to microbial biofilms which are hypothesized to mediate
the accretion of ooids (Flannery et al., 2018). However, the incorporation of organic material
from allochthonous sources is possible, which would not invoke a biogenic origin for the ooids.
That is, ooid-based microbial mats or biofilms need not be invoked to explain the presence of
organic matter within the Archean ooids. Indeed, the closest modern example would suggest it
may not be indigenous to the ooid at all.

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The microbial communities of Great Salt Lake ooids are heavily influenced by relict
DNA from brine shrimp gut microbiota which makes determining the biogenicity of GSL ooids
using 16S rRNA analysis not straightforward. The carbonate chemistry of GSL and the growth
rates of GSL ooids do not necessitate microbial intervention, therefore we put forth an abiogenic
mode of formation in the absence of clear evidence pointing toward biogenic origins.
Organomineralization is still a possibility in this system, whereby the growth of GSL ooids is
controlled by organic substances (not microbial in origin) adhered to the surface of the ooids.
CONCLUSIONS
This paper provides molecular evidence linking Great Salt Lake ooids to the Artemia
(brine shrimp) pellets that frequently serve as nuclei, raising the possibility that relict DNA from
the nucleus must be taken into account when examining the DNA contained within ooids or
other carbonate structures. Small subunit rRNA analysis demonstrates unsorted, 0.355 –
0.500mm, and <0.355mm, ooids have a similar microbial community composition enriched in
Bacteroidetes, Alphaproteobacteria, Deltaproteobacteria, and Gammaproteobacteria. The
microbial community of GSL water is a stark contrast as it is dominated by halophilic Archaea.
The similarity of the oolitic microbial community to the gut microbiota of Artemia suggests that
much of the microbial assemblage of GSL ooids may be relict DNA extracted from pellet nuclei
that are several thousand years old. Relict DNA may be an important, previously unrecognized
source of DNA in investigations of the geobiology of certain modern and ancient accretionary
structures such as ooids and stromatolites and may ultimately inhibit our ability to rely on 16S
rRNA sequencing alone to comment on biogenicity. Great Salt Lake has the propensity to
precipitate calcium carbonate year-round, though it approaches carbonate saturation in the winter
months. Slow net calcification in Great Salt Lake ooids as evidenced in the radiocarbon
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chronology in Chapter 2 may indicate aragonite is precipitating in water just above saturation
(the current conditions of GSL water) which would not necessitate the influence of microbial
metabolism.
ACKNOWLEDGMENTS
We would like to thank John Spear, Joyce Yager, Scott Perl, and Bonnie Baxter for
sampling assistance, logistics, and field support. This work was also supported by the Stevenson
lab at Oklahoma University, especially Bradley Stevenson, Blake Stamps, and Heather Nunn.

117

FIGURES

Figure 1: Brine shrimp (Artemia) fecal pellets (left) from Eardley, (1938) compared to Great Salt
Lake ooid with pellet nucleus (right). Eardley (1938) first noted the similarity in morphology and
made the connection between brine shrimp and GSL ooids.

118

Figure 2. Relative abundance of bacterial and archaeal phyla in water from the southern arm of
GSL (W1, W2, W3, W4), brine shrimp (Artemia) fecal pellets (BSP1, BSP2, BSP3), and ooids
(unsorted ooids: OUR1, OUR2, OUR3; ooids < 0.355mm diameter: OFR1, OFR2, OFR3; ooids
0.355-0.500mm diameter: OMR1, OMR2, OMR3; ooids 0.500-1.00mm diameter: OCR1, OCR2,
OCR3). The phylum Proteobacteria has been expanded (shades of blue) into the classes
Alphaproteobacteria, Betaproteobacteria, Deltaproteobacteria, and Gammaproteobacteria on
the left portion of the bars. Each individual bar represents the microbial community of a
replicate. Euryarchaeota (Archaea) are the most abundant in GSL water, while Proteobacteria
and Bacteroidetes are the most abundant phyla in both brine shrimp pellets and ooids.
119

Figure 3. Principal coordinate analysis (PCoA) of variance between microbial communities from
GSL water, brine shrimp (Artemia) pellets, and both unsorted and sieved ooids. The brine shrimp
(Artemia) pellet samples are more similar to <0.355 mm, 0.355 – 0.500 mm, and unsorted ooids
than they are to 0.500 – 1.00 mm ooids or GSL water. Great Salt Lake water from the southern
arm of the lake is enriched in halophilic archaea, thus causing them to cluster away from the rest
of the samples collected.

120

Figure 4. Saturation state of calcium carbonate (aragonite) in southern arm GSL water calculated
in the winter (A) and summer (B) using PHREEQ-C (Parkhurst and Appelo, 2013). The water
temperatures for winter and summer used were 4℃ and 26℃, respectively. The saturation state
of aragonite in the winter and summer are (black dots) are 1.2 and 2.5, respectively.
121

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136

SUPPLEMENTARY INFORMATION
APPENDIX A
Methods for bulk organic and inorganic carbon extractions on sieved ooids
To obtain bulk inorganic carbon ages on sieved and unsieved ooids, ooids were dissolved
in 3 extractions, and the results of those extractions were weighted and reported herein. The first
CO2 extraction occurred after 0.8 ml H2PO4 acid was added at 70°C and left to react for 1-2
hours. The second CO2 extraction occurred after an additional 0.8 ml H2PO4 was added and left
to react for 2 more hours at 70°C. The third CO2 extraction occurred after 2 hours at 70°C after
the second extraction, but no extra acid was added to the samples. All purified CO2 samples
were graphitized using a sealed-tube zinc reduction method (Xu et al., 2007). Graphite was
pressed into aluminum target holders and analyzed for
14
C at the Keck Carbon Cycle Accelerator
Mass Spectrometer (KCCAMS) facility at the University of California, Irvine (Southon et al.,
2004; Beverly et al., 2010). Data were normalized and background corrected using both modern
coral and radiocarbon-dead reference carbonates acidified in the same reaction vessel. Resulting
fractions of modern (FM) carbon were weighted according to yield to calculate bulk inorganic
carbon age, and
14
C data are presented according to the conventions presented in Stuiver and
Polach (1977).
To measure bulk organic carbon content and radiocarbon age from the acid insoluble
fraction, 1M HCl was added to 10g of ooids at 70°C for 24 hours until pH maintained at 1 for 2
hours. During acidification, the solution containing the sample was centrifuged and the solution
was decanted, then a new aliquot of 1M HCl was added. This process was repeated many times
until the pH maintained at 1 for 2 hours. The residuals were then rinsed with Milli-Q until the pH
became neutral.
137

Figure 1. Ooid dissolution reaction vessel. 50g of size-sieved ooids are added to the reaction vessel
with 150cc of deionized water. Ascarite-scrubbed N2 will flow through while the stir bar spins at
600 rpm for 30 min. This step ensures no atmospheric carbon remains in the reaction vessel or
water. Next, the outflow and gas inflow stopcocks are closed. For each dissolution step, 60cc of
3.3M HCl are added to the reaction vessel at a rate of 10cc/min. Tedlar bags are filled with the
resulting gas every ~3min. The gas is moved into He-rinsed and evacuated 25cc serum vials to be
submitted to UC Irvine for radiocarbon analysis.

138

Table 1. Bulk inorganic carbon extractions for sieved and unsorted ooids. The
14
C ages from each
extraction were pooled to calculate a bulk age for each sample.

Locality Grain Size (µm) Extraction Yield (mgC, Inorg) Inorg. C Fraction F
m
Bulk F
m
Bulk
14
C Age (yr BP)
Antelope Island Unsorted 1 2.29 0.6844 0.7097 ± 0.0012 0.6423 ± 0.0012 3556 ± 15
South Arm 2 0.97 0.2904 0.4952 ± 0.0010
3 0.08 0.0252 0.5086 ± 0.0021
355-500 1 2.36 0.6706 0.6676 ± 0.0012 0.6118 ± 0.0011 3947 ± 15
2 0.98 0.2792 0.4977 ± 0.0009
3 0.18 0.0501 0.5004 ± 0.0011
250-355 1 2.22 0.6584 0.6838 ±0.0012 0.6204 ± 0.0011 3834 ± 15
2 1.04 0.3092 0.5024 ± 0.0009
3 0.11 0.0324 0.4599 ± 0.0018
125-250 1 2.42 0.7559 0.7222 ± 0.0013 0.6749 ± 0.0012 3158 ± 15
2 0.72 0.2257 0.5254 ± 0.0009
3 0.06 0.0184 0.5659 ± 0.0027
Spiral Jetty Unsorted 1 3.29 0.7682 0.6482 ± 0.0011 0.6175 ± 0.0011 3872 ± 15
North Arm 2 0.84 0.1965 0.5202 ± 0.0010
3 0.15 0.0354 0.4924 ± 0.0015
355-500 1 2.99 0.7542 0.6030 ± 0.0011 0.5802 ± 0.0011 4373 ± 20
2 0.82 0.2055 0.5217 ± 0.0010
3 0.16 0.0403 0.5020 ± 0.0012
250-355 1 2.89 0.8339 0.6491 ± 0.0011 0.6263 ± 0.0011 3759 ± 15
2 0.52 0.1503 0.5133 ± 0.0011
3 0.05 0.0158 0.5010 ± 0.0031
125-250 1 3.00 0.8602 0.7359 ± 0.0013 0.7120 ± 0.0013 2728 ± 15
2 0.44 0.1253 0.5646 ± 0.0011
3 0.05 0.0145 0.5685 ± 0.0032
139

Figure 2. Scanning electron microscope images of individual ooids with close-up inset after 60ml
(A – B), 120ml (C – D), and 180ml (E – F) of 3.3M HCl was added to reaction vessel. Ooids
maintain general shape post-acidification confirming dissolution occurred fairly uniformly from
exterior to interior.

140

Table 2. Thin sections of individual ooids (A) analyzed with Raman spectroscopy (B) to map the
presence of organic matter (blue) within the ooid cortex. A Raman spectrum with a strong noise
signal suggesting the presence of organic material was selected from a GSL ooid and established
as the “organic material reference spectrum. Each point on the thin section was analyzed using
Raman spectroscopy and compared to the reference spectrum. The similarity of the measured
spectra to the reference organic matter spectrum was mapped on thin sections with a blue
overlay, with blue indicating presence of organic material.
p. Thin Section
Label
Site Grain Size (µm) Ooid Number
141 AI-oc Antelope Island – S. Arm 355 – 500 1
142 AI-oc Antelope Island – S. Arm 355 – 500 2
143 AI-oc Antelope Island – S. Arm 355 – 500 3
144 AI-oc Antelope Island – S. Arm 355 – 500 4
145 AI-oc Antelope Island – S. Arm 355 – 500 5
146 AI-om Antelope Island – S. Arm 250 – 355 1
147 AI-om Antelope Island – S. Arm 250 – 355 2
148 AI-om Antelope Island – S. Arm 250 – 355 3
149 AI-om Antelope Island – S. Arm 250 – 355 4
150 AI-om Antelope Island – S. Arm 250 – 355 5
151 AI-of Antelope Island – S. Arm 125 – 250 1
152 AI-of Antelope Island – S. Arm 125 – 250 2
153 AI-of Antelope Island – S. Arm 125 – 250 3
154 AI-of Antelope Island – S. Arm 125 – 250 4
155 AI-of Antelope Island – S. Arm 125 – 250 5
156 SJ-oc Spiral Jetty – N. Arm 355 – 500 1
157 SJ-oc Spiral Jetty – N. Arm 355 – 500 2
158 SJ-oc Spiral Jetty – N. Arm 355 – 500 3
159 SJ-oc Spiral Jetty – N. Arm 355 – 500 4
160 SJ-oc Spiral Jetty – N. Arm 355 – 500 5
161 SJ-om Spiral Jetty – N. Arm 250 – 355 1
162 SJ-om Spiral Jetty – N. Arm 250 – 355 2
163 SJ-om Spiral Jetty – N. Arm 250 – 355 3
164 SJ-om Spiral Jetty – N. Arm 250 – 355 4
165 SJ-om Spiral Jetty – N. Arm 250 – 355 5
166 SJ-of Spiral Jetty – N. Arm 125 – 250 1
167 SJ-of Spiral Jetty – N. Arm 125 – 250 2
168 SJ-of Spiral Jetty – N. Arm 125 – 250 3
169 SJ-of Spiral Jetty – N. Arm 125 – 250 4
200 µm
A
200 µm
B
141
100 µm
100 µm
A
B
142
100 µm
A
100 µm
B
143
100 µm
A
100 µm
B
144
100 µm
A
100 µm
B
145
100 µm
100 µm
A
B
146
100 µm
100 µm
A
B
147
100 µm
100 µm
A
B
148
100 µm
100 µm
A
B
149
100 µm
100 µm
A
B
150
50 µm
50 µm
A
B
151
100 µm
100 µm
A
B
152
100 µm
100 µm
A
B
153
100 µm
100 µm
A
B
154
100 µm
100 µm
A
B
155
100 µm
100 µm
A
B
156
100 µm
100 µm
A
B
157
100 µm
100 µm
A
B
158
100 µm
100 µm
A
B
159
100 µm
100 µm
A
B
160
100 µm
100 µm
A
B
161
100 µm
100 µm
A
B
162
100 µm
100 µm
A
B
163
100 µm
100 µm
A
B
164
100 µm
100 µm
A
B
165
50 µm
50 µm
A
B
166
100 µm
100 µm
A
B
167
100 µm
100 µm
A
B
168
50 µm
A
50 µm
B
169
170

APPENDIX B
Figure 1. Representative examples of fabrics most commonly found in GSL ooids as shown on
an ooids from the north arm of GSL. Fabrics include large aragonite crystal rays (A), banded
tightly-packed radial aragonite (B) and thin clean radial-concentric aragonite (C).

171

Table 1. Summary of δ
13
C and
14
C chronology for all organic and inorganic carbon samples from
sequential dissolution of ooids from north and south arms of GSL.
14
C ages have been calibrated
using Intcal 13 (Reimer et al., 2013; Stuiver & Reimer, 1993; Stuiver et al., 2018). Calibrated
ages are given in calibrated years BP (before 1950).

Relative
Area
Relative
Area
Median
AI14-10-A1-1 3.7 0.15 0.9441 0.0022 460 20 518 505 1 528 499 1 513
4.0 0.15 0.8498 0.0021 1305 25 1284 1256 0.605 1290 1226 0.703 1251
1204 1187 0.326 1211 1183 0.297
1248 1242 0.069
4.2 0.15 0.7966 0.0022 1825 25 1811 1753 0.737 1825 1699 1 1766
1744 1724 0.263
AI14-10-B1-1 4.7 0.15 0.7462 0.0023 2350 25 2362 2339 1 2438 2330 1 2354
AI14-10-B2-1 4.3 0.15 0.7130 0.0023 2720 30 2844 2782 1 2866 2760 1 2815
4.4 0.15 0.6836 0.0022 3055 30 3266 3216 0.507 3355 3205 0.935 3270
3335 3288 0.493 3203 3179 0.065
4.3 0.15 0.6042 0.0027 4050 40 4575 4499 0.596 4629 4421 0.909 4529
4486 4440 0.387 4798 4762 0.078
4777 4773 0.017 4689 4680 0.008
4642 4636 0.005
4.7 0.15 0.5732 0.0026 4470 40 5279 5164 0.638 5293 4972 1 5146
5074 5038 0.188
5133 5106 0.132
4995 4985 0.042
4.4 0.15 0.4780 0.0031 5930 60 6798 6673 0.909 6914 6636 0.994 6759
6832 6818 0.076 6927 6917 0.006
6798 6673 0.016
4.4 0.15 0.4398 0.0032 6600 60 7514 7440 0.784 7579 7424 1 7498
7560 7539 0.216
n/a 0.4617 0.0010 6210 20 7112 7068 0.458 7128 7011 0.737 7092
7057 7028 0.318 7179 7145 0.216
7168 7155 0.224 7235 7217 0.028
7209 7195 0.019
2.3 0.15 0.9638 0.0022 295 20 426 392 0.679 432 356 0.694 394
319 304 0.321 332 299 0.306
5.6 0.15 0.9213 0.0016 660 15 582 569 0.521 590 564 0.503 598
662 649 0.479 667 641 0.497
4.4 0.15 0.7700 0.0021 2100 25 2118 2040 0.95 2137 1999 1 2073
2016 2012 0.05
4.8 0.15 0.7170 0.0020 2675 25 2785 2754 1 2808 2750 0.861 2775
2844 2816 0.139
4.4 0.15 0.6159 0.0026 3895 35 4410 4294 1 4422 4234 0.992 4334
4196 4184 0.008
4.6 0.15 0.5917 0.0025 4215 35 4755 4708 0.538 4762 4627 0.632 4743
4842 4812 0.404 4852 4796 0.368
4666 4659 0.059
4.6 0.15 0.5423 0.0029 4915 45 5662 5596 0.929 5733 5588 1 5644
5699 5693 0.04
5706 5702 0.031
4.6 0.15 0.4976 0.0031 5605 50 6414 6315 0.977 6480 6299 1 6379
6431 6429 0.023
4.7 0.15 0.4838 0.0031 5830 60 6719 6561 1 6758 6490 0.979 6638
6781 6762 0.021
-18.9 0.15 0.4754 0.0008 5975 15 6847 6814 0.563 6859 6746 0.978 6814
6801 6781 0.383 6879 6871 0.022
6762 6757 0.054
6.9 0.15 0.9860 0.0023 115 20 118 64 0.537 145 55 0.573 112
259 242 0.153 267 215 0.292
139 124 0.147 47 19 0.135
232 222 0.098
38 30 0.066
Intcal13 Calibration in cal BP
68% (1 σ) 95% (2 σ)
Locality
Carbon
Sample
Sample ID
d
13
C
(‰)
±
Fraction
Modern
±
14
C Age
(BP)
±
Inorganic
(DIC)
Inorganic
(ooid)
Organic
(nuclei)
Inorganic
(DIC)
Antelope Isl.
South Arm
Spiral Jetty
North Arm
Inorganic
(ooid)
Organic
(nuclei)
AI14-10-A2-1
AI14-10-A3-1
AI14-10-B3-1
AI14-10-C2-1
AI14-10-C3-1
AI14-10-D2-1
AI14-10-D3-1
AI14-10-org
DIC-AI16-9-10
SJ14-10-A1-1
SJ14-10-D2-1
SJ14-10-D3-1
SJ14-10-org
DIC-SJ-9-9
SJ14-10-A2-1
SJ14-10-A3-1
SJ14-10-B2-1
SJ14-10-B3-1
SJ14-10-C2-1
172

Table 2. Summary of petrographic characteristics of individual ooids. Sites AI and SJ are
indicative of Antelope Island from the south arm of GSL and Spiral Jetty form the north arm of
GSL, respectively. Grain size, ooid diameters, nuclei diameters, and cortex width are measured
in µm. Major and minor ooid/nuclei diameters are a measurement of the major and minor axis of
each constituent from thin section.

Grain size No. Nucleus
Ooid diameter
major
Ooid diameter
minor
Nucleus
diameter
major
Nucleus
diameter
minor
Cortex
width
125-250 1 Peloid 372 300 156 74 111
2 Peloid 444 314 197 76 119
3 Quartz 336 293 195 95 87
4 Peloid 341 268 154 77 96
5 Quartz 357 308 104 67 125
250-355 1 Peloid 704 306 537 109 105
2 Quartz 575 455 323 202 120
3 Peloid 800 264 698 107 78
4 Quartz 388 368 175 143 110
5 Peloid 561 332 359 108 114
355-500 1 Peloid 829 377 532 68 165
2 Quartz 487 366 289 181 93
3 Peloid 396 228 250 79 73
4 Peloid 529 246 349 92 81
5 Quartz 382 304 224 134 85
125-250 1 Quartz 273 250 88 65 96
2 Quartz 324 293 90 58 115
3 Augite 290 273 119 114 78
4 Peloid 315 283 143 109 86
5 Peloid 383 321 142 80 116
250-355 1 Quartz 445 399 138 83 152
2 Augite 442 373 145 124 144
3 Peloid 453 447 144 144 151
4 Peloid 635 420 336 126 154
5 Aragonite 407 298 159 61 119
355-500 1 Peloid 908 381 668 110 135
2 Peloid 521 529 106 92 204
3 Peloid 770 527 366 116 210
4 Augite 550 491 230 171 152
5 Peloid 877 526 448 111 200

173

Table 3. Order and width (µm) of fabrics from individual ooids from Antelope Island and Spiral Jetty. The order of each fabric found
in GSL ooids from Antelope Island (south arm of GSL) and Spiral Jetty (north arm of GSL), as seen in thin section. Fabrics are
composed of large aragonite crystal rays (RAY), banded tightly-packed radial aragonite (BR) and thin clean radial-concentric
aragonite (TCR). Photos of representative examples of these textures can be found in Appendix B, Figure 1.
Fabric
Width
(µm)
Fabric
Width
(µm)
Fabric
Width
(µm)
Fabric
Width
(µm)
Fabric
Width
(µm)
Fabric
Width
(µm)
Fabric
Width
(µm)
Fabric
Width
(µm)
Fabric
Width
(µm)
Fabric
Width
(µm)
1 RAY 84 BR 9 BR 6 BR 9 TCR 6
2 RAY 62 BR 18 BR 25 TCR 5 TCR 6
3 RAY 72 TCR 9
4 RAY 87 TCR 11
5 RAY 109 BR 16
1 RAY 62 TCR 10 TCR 8 BR 13 BR 10
2 RAY 104 TCR 7 TCR 8
3 RAY 66 TCR 9 TCR 6
4 RAY 67 BR 17 TCR 13 TCR 17
5 RAY 37 RAY 29 TCR 3 RAY 32 TCR 5 BR 4
1 RAY 103 TCR 13 TCR 15 TCR 10 TCR 15
2 TCR 16 RAY 57 TCR 17
3 RAY 57 TCR 6 BR 11
4 RAY 62 BR 13
5 BR 11 TCR 10 BR 11 TCR 28 BR 22
1 RAY 96
2 RAY 103 TCR 8
3 RAY 78
4 RAY 86
5 BR 10 RAY 97 TCR 4 TCR 4
1 RAY 142 TCR 10
2 RAY 123 BR 9 BR 11
3 BR 23 BR 23 BR 60 TCR 11 BR 13 TCR 22
4 RAY 100 TCR 6 BR 9 BR 20 BR 14
5 RAY 107 BR 14
1 RAY 67 BR 30 TCR 13 TCR 15
2 RAY 74 TCR 5 BR 48 BR 18 TCR 11 TCR 17 BR 16 BR 7 TCR 6
3 RAY 33 RAY 79 TCR 4 BR 32 TCR 21 BR 20 TCR 6 TCR 6 BR 11 TCR 10
4 RAY 72 BR 65 TCR 13
5 RAY 62 BR 34 BR 51 TCR 10 TCR 8 TCR 4 TCR 5 BR 13
5th
Locality No.
Antelope Island
South Arm
Spiral Jetty
North Arm
125-250
250-355
355-500
125-250
250-355
355-500
Grain
Size (µm)
1st 2nd 3rd 4th 6th 7th 8th 9th 10th
174

APPENDIX C
Figure 1: Ooids from the northern (Spiral Jetty) and southern (Bridger Bay, Antelope Island)
arms of GSL sampled in January 2015. Ooids were sieved, pulverized using sterile mortar and
pestle, and processed according to methods outlined in Chapter 3. The microbial communities of
unsorted and sieved ooid samples resemble one another, likely owing to the core DNA residing
in the Artemia pellet nuclei.

Abstract (if available)

Abstract Ooids are laminated coated grains that are most commonly composed of calcium carbonate which precipitates around a nucleus. As accretionary structures, they have the potential to record a time series of hydrologic and geochemical change in marine and lacustrine environments. Ooids are common in carbonate environments throughout Earth’s history, but the mechanism by which they form remains unclear. In particular, the rate of ooid growth remains elusive in all but a few modern marine environments. Furthermore, modern marine ooids display tangentially-oriented aragonite crystals within their cortices, whereas many ancient ooids have radially-oriented cortices. Ooids are traditionally viewed as abiogenic precipitates, however the role of biology in ooid formation has recently been suggested because certain microbial metabolisms may either foster carbonate precipitation or dissolution by manipulating dissolved inorganic carbon and alkalinity. The following dissertation uses ooids from Great Salt Lake (GSL), Utah, a well-known site of primary radial ooids, to investigate ooid growth rate and age, the potential of ooids to resolve lacustrine hydrologic change, and the biogenicity of lacustrine ooids. ❧ We used ¹⁴C to establish a sequential chronology for ooids from GSL. Ooids from the coarse size fraction were sequentially dissolved and ¹⁴C ages were obtained for each dissolution step to create a time series of ooid growth. The results of the sequential dating indicate that the formation of coarse Great Salt Lake ooids began between 5800-6600 ± 60 ¹⁴C yr BP while their outer cortices are nearly modern. Sequentially dated ooids from Antelope Island (southern part of Lake) record a nearly linear growth history with linear growth rates of ∼0.01 – 0.015 µm/yr, whereas ooids from Spiral Jetty (NE part of Lake) record somewhat faster growth between ∼6000 and 4000 years ago (0.03 – 0.06 µm/yr) followed by a slower growth history for the remainder of their lifespan (0.003 – 0.008 µm/yr). The lifespan of Great Salt Lake radial aragonitic ooids ranges is two and six times longer than those from modern marine environments, and thus provides a unique end member for understanding the mechanisms behind radial ooid formation. The antiquity of the ooids would suggest that geochemical parameters measured from bulk ooid dissolution would integrate over ∼6000 years and thus do not represent geochemical snapshots in time as some previous studies suggest. ❧ Lake level indicators are sparse in GSL between ∼10,000 ybp and the onset of historical times, so the Holocene lake variation is somewhat poorly resolved. We evaluate the ¹⁴C age and stable carbon isotope composition of ooids that grew in the lake over the last 7,000 years. Since δ¹³C and δ¹⁸O covary in closed basin lakes, changes in the δ¹³C through time can be interpreted to represent changes in lake volume if other assumptions we make hold true. Ooids were sequentially dissolved to create a paired radiocarbon chronology and δ¹³C profile with depth in the ooid. Petrographic analysis reveals that, for the majority of ooids, the initial precipitation upon the nucleus is represented by a comparatively large aragonite crystal ray texture, followed by alternations of finely crystalline radial aragonite and thin radial-concentric layers. Net growth rates derived from radiocarbon ages indicate the large crystal ray fabric corresponds to 7498 to 2073 cal yr BP with δ¹³C between 4.4±0.15 and 4.8±0.15 ‰, likely indicating a relatively stable lake level. The alternating fabric precipitated between 2073 cal yr BP to present, and the δ¹³C values of ooids trending toward lighter values (from ∼4.5‰ to 3.7‰), suggests a largely transgressive phase in GSL. The youngest inorganic carbon sample from north arm ooids falls off this trend as it has a significantly heavier δ¹³C of 5.6±0.15‰. Modern dissolved inorganic carbon from north arm lake water indicates aragonite precipitating today would have δ¹³C values between 9.1±0.55‰ to 9.8±0.75‰, therefore the young heavy inorganic carbon of north arm ooids likely suggests that ooid formation has occurred after the construction of the railroad causeway in 1959, which isolated the north arm from the riverine-influenced southern portion and thus left the reservoir to generate an enriched δ¹³C pool, representing the first robust evidence that ooid formation has occurred in modern times. Furthermore, the entire chronology of GSL ooids from both sample sites record over 2‰ variability in δ¹³C over several thousand years, which calls into question the utility of bulk geochemical analyses from accretionary structures such as ooids in dynamic systems such as closed-basin lakes. ❧ To investigate the biogenicity of ooids in GSL, we evaluated the microbial community of ooids from the southern arm of GSL and the brine shrimp (Artemia) pellets upon which they commonly form. The small subunit rRNA analysis revealed ooids and Artemia pellets have similar microbial assemblages dominated in Bacteroidetes, Alphaproteobacteria, Deltaproteobacteria, and Gammaproteobacteria. The striking similarity between the microbial communities of GSL ooids and brine shrimp gut microbiota suggests relict DNA residing in oolitic nuclei may be an important source of DNA in microbial community analysis of ooids and other accretionary structures. The calcium carbonate saturation state of GSL is supersaturated with respect to aragonite, though it ranges from 100% to 250% saturation seasonally. Calcification rates of GSL ooids are several orders of magnitude lower than experimental precipitation of aragonite, suggesting aragonite may be precipitating in GSL just above saturation, conditions that do not necessitate the geochemical manipulation of microbiota to achieve in this system.

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University of Southern California Dissertations and Theses

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Creator Paradis, Olivia Piazza (author)

Core Title Great Salt Lake ooids: insights into rate of formation, potential as paleoenvironmental archives, and biogenicity

School College of Letters, Arts and Sciences

Degree Doctor of Philosophy

Degree Program Geological Sciences

Publication Date 05/10/2019

Defense Date 03/20/2019

Publisher University of Southern California (original), University of Southern California. Libraries (digital)

Tag Carbon,carbonate,Great Salt Lake,OAI-PMH Harvest,ooid,radiocarbon,stable isotopes

Format application/pdf (imt)

Language English

Contributor Electronically uploaded by the author (provenance)

Advisor Corsetti, Frank (committee chair), Berelson, William (committee member), Heidelberg, John (committee member)

Creator Email oparadis8@gmail.com,opiazza@usc.edu

Permanent Link (DOI) https://doi.org/10.25549/usctheses-c89-156015

Unique identifier UC11660535

Identifier etd-ParadisOli-7415.pdf (filename),usctheses-c89-156015 (legacy record id)

Legacy Identifier etd-ParadisOli-7415.pdf

Dmrecord 156015

Document Type Dissertation

Format application/pdf (imt)

Rights Paradis, Olivia Piazza

Type texts

Source University of Southern California (contributing entity), University of Southern California Dissertations and Theses (collection)

Access Conditions The author retains rights to his/her dissertation, thesis or other graduate work according to U.S. copyright law. Electronic access is being provided by the USC Libraries in agreement with the a...

Repository Name University of Southern California Digital Library

Repository Location USC Digital Library, University of Southern California, University Park Campus MC 2810, 3434 South Grand Avenue, 2nd Floor, Los Angeles, California 90089-2810, USA