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BIOMINERALIZATION OF GIANT CLAM SHELLS (TRIDACNA GIGAS): IMPLICATIONS FOR PALEOCLIMATE APPLICATIONS by MICHELLE ELIZABETH GANNON ALBERTO PÉREZ-HUERTA, COMMITTEE CO-CHAIR PAUL AHARON, COMMITTEE CO-CHAIR C. FRED ANDRUS NATASHA T. DIMOVA JULIE OLSON A THESIS Submitted in partial fulfillment of the requirements for the degree of Master of Science in the Department of Geological Sciences in the Graduate School of The University of Alabama TUSCALOOSA, ALABAMA 2016

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Page 1: TRIDACNA GIGAS - Acumenacumen.lib.ua.edu/content/u0015/0000001/0002339/u0015...The giant clam, Tridacna gigas, is the largest known bivalve and one of the largest members of the Phylum

BIOMINERALIZATION OF GIANT CLAM SHELLS

(TRIDACNA GIGAS): IMPLICATIONS FOR

PALEOCLIMATE APPLICATIONS

by

MICHELLE ELIZABETH GANNON

ALBERTO PÉREZ-HUERTA, COMMITTEE CO-CHAIR

PAUL AHARON, COMMITTEE CO-CHAIR

C. FRED ANDRUS

NATASHA T. DIMOVA

JULIE OLSON

A THESIS

Submitted in partial fulfillment of the requirements

for the degree of Master of Science

in the Department of Geological Sciences

in the Graduate School of

The University of Alabama

TUSCALOOSA, ALABAMA

2016

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Copyright Michelle Elizabeth Gannon 2016

ALL RIGHTS RESERVED

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ABSTRACT

The giant clam, Tridacna gigas, is an important faunal component of Indo-Pacific reef

ecosystems, for which its shell is often used as an environmental archive for modern and past

climates. This thesis is a study of the shell microstructure of modern specimens from Palm

Island, Great Barrier Reef (GBR), Australia and Huon Peninsula, Papua-New Guinea (PNG),

using a combination of petrography, scanning electron microscopy (SEM), electron backscatter

diffraction (EBSD) and Raman spectroscopy, as well as a microstructural comparison of fossil T.

gigas through 200 ka from PNG.

Daily growth increments are recognizable in all specimens through ontogeny within the

internal layer. For modern T. gigas from PNG, increments are composed of pairs of organized

aragonitic needles and compact, oblong crystals, whereas modern specimens from GBR are

composed of shield-like crystals. The combination of nutrient availability and rainfall are likely

the most significant factors controlling shell growth and it may explain the observed differences

in microstructure. The external layers are composed of a dendritic microfabric, significantly

enriched in 13

C compared to the internal layer, suggesting a different metabolic control on layer

secretion. The internal and external layers are likely mineralized independent from each other,

associated with the activity of a specific mantle organ.

Furthermore, needles similar to those of modern T. gigas from PNG, are observed and the

widths are measured in the set of fossil T. gigas. An exception includes two mid-Holocene-aged

individuals, composed of elongated crystals, oblique to the outside of the shell. The results show

that widths follows a cyclic pattern, similar to those of solar radiation variability, suggesting

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there is a relationship between solar activity and the width of aragonitic needles. Differences

between modern and mid-Holocene T. gigas, are likely associated with fundamental

environmental differences.

The results of this study, pointing to locality and environmental dependence, layer

specific mantle biomineralization, and co-variation between needle width and solar modulation,

advance the potential of giant clam shells to assist in the reconstruction of many climate

parameters that were previously limited to chemical analyses. Microstructural results are

additionally applicable in engineering and medical research fields.

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DEDICATION

This thesis is dedicated to everyone who has supported me through the process of

pursuing the work required to complete my master’s research, especially my family and friends.

This could not have been done without the help of my understanding parents, Roses and Jerry.

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LIST OF ABBREVIATIONS AND SYMBOLS

ANW Aragonitic Needle Width

ASIL Alabama Stable Isotope Laboratory

Ba Barium

10

Be Beryllium-10

C Celsius

13

C Carbon-13

Ca Calcium

CAF Central Analytical Facility

cm Centimeter

cm-1

Per Centimeter

D/L D-alloisoleucine/L-isoleucine

E East

EBSD Electron Backscatter Diffraction

e.g. Example

ENSO El Niño Southern Oscillation

FE-SEM Field Emission Scanning Electron Microscope

Fig. Figure

GBR Great Barrier Reef

gcm-1

Grams per Centimeter

HCl Hydrochloric Acid

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IRMS Isotope Ratio Mass Spectrometer

K Kanzarua

ka Thousands of Years

km Kilometer

kV Kilovolts

m Meter

Mg Magnesium

µm Micrometer

µM Micromole

mm Millimeter

n Population size

nA NanoAmpere

NBS-19 National Bureau of Standards-19

nm Nanometer

18

O Oxygen-18

231

Pa Protactiunium-231

PT Palm Tridacna

PNG Papua-New Guinea

S South

SEM Scanning Electron Microscopy

se Standard Error

Sr Strontium

SST Sea Surface Temperature

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230Th Thorium-230

T. gigas Tridacna gigas

238

U Uranium-238

235

U Uranium-235

U Uranium

UV Ultraviolet

V-PBD Vienna Pee Dee Belemnite

° Degree

δ Delta

= Equal to

> Greater than

< Less than

% Percent

‰ Permil

± Plus or Minus

σ Standard Deviation

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ACKNOWLEDGMENTS

The graduate study of the senior author was supported by a W. Gary Hooks Endowed Geology

Fund and a University of Alabama Research Graduate Council (RGC) Award. Thanks are

extended to Johnny Goodwin and the Central Analytical Facility (CAF) for assistance and

training on the SEM, Dr. Joe Lambert and the Alabama Stable Isotope Laboratory (ASIL) for

facilitating isotope analyses, and Patrick Sipe and Gregory Dye for assistance in obtaining

Raman spectra. Finally, thanks to Sara Kozmor for collaborative ideas.

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CONTENTS

ABSTRACT .................................................................................................................................... ii

DEDICATION ............................................................................................................................... iv

LIST OF ABBREVIATIONS AND SYMBOLS ............................................................................v

ACKNOWLEDGMENTS ........................................................................................................... viii

LIST OF TABLES ........................................................................................................................ xi

LIST OF FIGURES ..................................................................................................................... xii

1.INTRODUCTION .......................................................................................................................1

2. A BIOMINERALIZATION STUDY OF GIANT CLAM (TRIDACNA GIGAS) SHELLS ......3

3. MICROSTRUCTURE VARIABILITY IN FOSSIL GIANT CLAMS TRIDACNA GIGAS)

FROM THE HUON PENINSULA, PAPUA NEW GUINEA: AN ARCHIVE OF SOLAR

MODULATION? .......................................................................................................................32

REFERENCES .............................................................................................................................53

APPENDIX I: BIOLOGY ............................................................................................................61

APPENDIX II: SEASONAL GROWTH.......................................................................................64

APPENDIX III: NEEDLE MEASUREMENTS ...........................................................................67

APPENDIX IV: DAILY GROWTH .............................................................................................77

APPENDIX V: DAYTIME AND NIGHTTIME GROWTH ........................................................78

APPENDIX VI: DAILY HIGH TIDES .........................................................................................79

APPENDIX VII: SOLAR IRRADIANCE CALCULATIONS .....................................................86

APPENDIX VIII: PRECIPITATION AND GROWTH ................................................................90

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APPENDIX VII: RAMAN SPECTROSCOPY .............................................................................93

APPENDIX VIII: STABLE ISOTOPIC ANALYSIS ................................................................119

APPENDIX IX: TERRACE CORRELATIONS .........................................................................123

APPENDIX X: TERRACE AGES ..............................................................................................125

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LIST OF TABLES

1.1. Solar irradiance calculations ..................................................................................................26

2.1. Geologic ages of coral reef terraces .......................................................................................37

2.2. Fossil specimens ....................................................................................................................38

2.3. Microstructure of fossil specimens ........................................................................................47

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LIST OF FIGURES

1.1. Schematic representation of a shell section ..............................................................................5

1.2. Shell sections of T. gigas specimens and collection sites .........................................................7

1.3. Shell sections at the umbo region .............................................................................................8

1.4. Petrographic and SEM observations of daily growth lines .....................................................13

1.5. Microstructure of the internal layer of Palm Island ................................................................14

1.6. Schematic representation of microstructure measurements (modern) ....................................15

1.7 Microstructure of the internal layer of Huon Peninsula ...........................................................16

1.8. Raman spectroscopy ...............................................................................................................19

1.9. Seasonal shell growth ............................................................................................................21

1.10. Shell growth and tidal influences ..........................................................................................22

1.11. Shell growth, precipitation and insolation ............................................................................24

1.12. External microstructure and stable carbon isotopes .............................................................28

1.13. Schematic representation of internal organs ........................................................................29

1.14. Schematic representation of mantles and shell layers ..........................................................30

2.1. Fossil sample collection locations (Huon Peninsula ) ............................................................36

2.2. Shell sections of fossil specimens ...........................................................................................40

2.3. Schematic representation of microstructure measurements (fossil) .......................................41

2.4. Shell section, petrographic and SEM representation of modern T. gigas ...............................42

2.5. Petrographic images of fossil specimens ................................................................................43

2.6. High resolution SEM images of fossil specimens .................................................................44

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2.7. T. gigas microstructure types ..................................................................................................45

2.8. EBSD comparison of Holocene and modern T. gigas ...........................................................46

2.9. Aragonitic needle measurements and correlations .................................................................49

A.1. Seasonal growth increments: modern GBR ..........................................................................65

A.2 Seasonal growth increments: modern PNG ............................................................................66

A.3 Stable isotope sampling ........................................................................................................119

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1. INTRODUCTION

Giant clam (Tridacna gigas) shells are considered excellent bioarchives of their

surrounding environment and have been used in paleoclimate studies (Aharon, 1983; Aharon and

Chappell, 1986; Watanabe et al., 2004; Yan et al., 2014; Ayling et al., 2015; Warter and Müller,

2016). Their restricted range in Indo-Pacific coral reefs allows for their shell chemistry and

morphology to represent the environments that drive many global climate factors such as El

Niño-Southern Oscillation, monsoons, and oscillation of the Intertropical Convergence Zone

(Chiang, 2009), making them ideal specimens. However, studies of these shells often contain

many common assumptions: i.) presence of regular growth increments at daily, monthly and

seasonal scales using imaging techniques with varying resolutions; ii.) shells grow continuously;

and iii.) diagenetic alteration is present in fossil specimens. The objectives of this master’s thesis

research are to characterize the microstructure of T. gigas through time, ontogeny and locality,

simultaneously addressing the common assumptions. In order to thoroughly explore each

objective, several methodologies were employed: petrographic analysis, scanning electron

microscopy (SEM), electron backscatter diffraction (EBSD), stable isotope mass spectrometry

and Raman spectroscopy.

This thesis is composed of two manuscripts. The first, “A Biomineralization Study of the

Giant Clam (Tridacna gigas) Shells,” addresses modern T. gigas from Palm Island, Great Barrier

Reef, Australia and Huon Peninsula, Papua New Guinea. This manuscript describes the shells’

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microstructural differences between these localities and discusses several environmental

parameters that might affect shell growth. Additionally suggested is a mechanism through which

the internal and external layers of T. gigas might be secreted, with respect to the lateral and

siphonal mantles, likely responsible for their complexities.

The second paper entitled: “Microstructure Variability in Fossil Giant Clams (Tridacna

gigas) from the Huon Peninsula, Papua New Guinea: An archive of solar modulation?” addresses

T. gigas microstructure through time (~134.1 ka), utilizing specimens from the raised coral reef

terraces of the Huon Peninsula, Papua New Guinea. Presented is a co-variance between the

oscillation of the widths of aragonitic needles, the major component of T. gigas microstructure,

and solar modulation.

Finally, several appendices are included that provide additional insights that have not

been incorporated into the manuscript chapters for publication. These include data tables as well

as a description of seasonal growth increments, noted in modern specimens.

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2. A BIOMINERALIZATION STUDY OF THE GIANT CLAM (TRIDACNA GIGAS) SHELLS

Gannon, M. E.1, A. Pérez-Huerta

1, P. Aharon

1, and S. C. Street

2

1Department of Geological Sciences, The University of Alabama, Tuscaloosa

2Department of Chemistry, The University of Alabama, Tuscaloosa

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Abstract

The giant clam, Tridacna gigas, is an important faunal component of reef ecosystems within the

Indo-Pacific region. In addition to its ecological role, shells of this bivalve species are useful

bioarchives for past climate and environmental reconstructions. However, the biomineralization

processes involved in the shell aragonite deposition are insufficiently understood in order to

confidently make chemical and other analytical measurements. Here, we present a study of the

shell microstructure of modern specimens from Palm Island, Great Barrier Reef (GBR) of

Australia and Huon Peninsula, Papua New Guinea (PNG), using a combination of petrography,

scanning electron microscopy (SEM), electron backscatter diffraction (EBSD) and Raman

spectroscopy. Daily growth increments are recognizable in all specimens through ontogeny, and

counting these growth lines provides a robust specimen age estimate. For the internal layers,

paired increments of organized aragonitic needles and compact, oblong crystals are recognized

for a specimen from PNG, whereas specimens from GBR are composed of shield-like crystals.

The combination of nutrient availability, rainfall and solar irradiance are likely to be the most

significant factors controlling shell growth and it may explain the observed differences in

microstructure. The external layer, identical in all specimens, is composed of a dendritic

microfabric and it is significantly enriched in 13

C compared to the internal layer suggesting a

different metabolic control on layer secretion. This study proposes that the mineralization of the

internal and external layers is independent from each other and associated with the activity of a

specific mantle. Future studies using T. gigas shells as bioarchives need to consider the

microstructure as it reflects the environment in which the individual lived as well as the

differences in mineralization pathways of internal and external layers.

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Figure 1.1, Schematic representation of a shell

section, from the umbo to the posterior region,

indicating the location of the external and

internal layers, and the separation of the first

(FG) and last (LG) growth stages through

ontogeny as recorded within the internal layer.

Introduction

The giant clam, Tridacna gigas, is the largest known bivalve and one of the largest members of

the Phylum Mollusca (Rosewater, 1965). T. gigas is a characteristic invertebrate in coral reef

environments throughout the Indo-Pacific ocean region, mainly between 100° to 180° of

longitude in the Southern Hemisphere (Rosewater, 1965), with a limited distribution due to

complicated reproduction (Braley, 1984). This species is found in shallow waters of fringing,

barrier, and lagoonal reefs typically no deeper than 10 m (Aharon and Chappell, 1986). Besides

its ecological importance in coral reefs, shells of T. gigas have been used to extract paleoclimate

information (e.g., Aharon et al., 1980; Aharon, 1983; Aharon, 1985; Aharon and Chappell, 1986;

Watanabe et al., 2004; Elliot et al., 2009; Welsh et al., 2011; Batenburg et al., 2011; Sano et al.,

2012; Yan et al., 2013), especially for the low latitude tropics in which well documented records

are rare (Sano et al., 2012). These clams are considered valuable bioarchives because their shells

are very dense with a high growth rate,

resistant to extreme diagenesis in fossil

specimens relative to coral skeletons (Veeh

and Chappell, 1970), and may have long life

spans (Watanabe et al., 2004). Internal and

external shell morphologies have previously

been analyzed to distinguish Tridacna

species (Rosewater, 1965) and provide a

temporal context for geochemical analyses

(Fig. 1.1). However, shell biomineralization

and growth of these clams are insufficiently

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understood in order to constructively plan chemical analyses, besides some basic descriptions of

shell microstructures (e.g., Taylor, 1973; Watanabe et al., 2004) and histology of mantle tissues

(Norton and Jones, 1992).

The aim of this study is to provide a detailed description of the shell microstructure and

growth features, focusing primarily on the internal layers, of T. gigas shells collected from two

different geographical regions and reef settings. An additional objective is to evaluate common

assumptions about shell biomineralization involving continuous deposition of aragonite

throughout the ontogeny (Watanabe et al., 2004) and the interpretation of daily growth

increments in reference to shell microstructures (Aharon and Chappell, 1986; Watanabe et al.,

2004; Sano et al., 2012). Finally, a model of shell biomineralization, based on the activity of

siphonal and lateral mantles, is proposed.

Materials and Methods

Samples

Two live specimens (PT-1 and PT-3) were collected from Palm Island, the Great Barrier Reef

(GBR) of Australia, on May 27, 1980 (Fig. 1.2). From the umbo to the posterior region, PT-1

and PT-3 are 31 cm long and 6.5 cm thick and 55 cm long and 16 cm thick at the umbo,

respectively. At this site, coral reefs reside in the GBR lagoon where sea surface temperatures

(SST) vary between 23°C and 28°C, with a mean seasonal temperature amplitude of about 3°C

(Aharon, 1991). These reefs receive a daily influx of fresh water from river channel inputs

(Crossland and Barnes, 1983) and therefore, the nutrient supply to the lagoonal reef is mostly

terrestrial based, composed of phosphate, ammonium and nitrate (Furnas and Mitchell, 1983).

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Rainfall also controls the large inputs of freshwater as storm events causing flash floods in this

region are common during the Austral summer (King et al., 2001).

For an inter-specimen comparison, an additional T. gigas sample (K-133) was added to

the study. It was collected alive on September 15, 1977 from an active fringing reef on the Huon

Peninsula, Papua New Guinea (PNG; Fig. 1.2). From the umbo to posterior region, K-133 is 28.6

cm tall and 7.3 cm thick at the umbo. The average SST at this location is 27.9 ± 0.9°C with a

seasonal amplitude of about 2.5°C (Aharon and Chappell, 1986). This specimen is a modern

representative of T. gigas fossils from uplifted coral reef terraces at the same location that were

Figure 1.2, Shell sections of

T. gigas specimens and

collec-tion sites. Longitudinal

shell sections, from umbo to

post-erior regions, of the

specimens (K-133 (a) and PT-

1 (b)) collected in Papua-New

Guinea and Australia. (c-e),

Details of collection sites at

the Huon Peninsula, Papua-

New Guinea (c) and Palm

Island, Great Barrier Reef,

Australia (d), with white

circles showing the location

of specimen collection and

white triangles showing the

location of rainfall stations

whose data are used in this

study.

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previously used in paleoclimate research (Aharon et al., 1980; Aharon, 1983; Aharon, 1985;

Aharon and Chappell, 1986). Elongated coral reefs of PNG occur along the steep Vitiaz Strait

shorelines, and form either narrow fringing reefs that follow the coastal contours or barrier reefs

enclosing shallow, narrow and elongated lagoons (Chappell, 1974b). Fringing, open ocean reefs

Figure 1.3, Shell sections of T. gigas at

the umbo region used for analyses

during the first and last seasons of

growth (a: K-133; b: PT-1; c: PT-3).

Boxes on each individual represent the

locations of cuts made to produce shell

block samples for SEM, EBSD and

Raman spectroscopy analyses, and

circles designate the first or last season

of growth. The pallial line on each

individual, separating external (EL) and

internal (IL) layers is represented by a

dashed line. First season of growth is

closest to the pallial line while the last

growth is farthest from the pallial line.

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have water flowing through them constantly (Aharon and Chappell, 1986), bringing offshore

nutrients into the reef environment (Belda et al., 1993). The coral reef setting along the coast of

PNG differs from that of the GBR where water enters the system nearly exclusively through

surface water channel inputs. Thus, the flux of nutrient supply to the fringing reefs of PNG is

likely higher than that of the GBR lagoon reefs (Belda et al., 1993) and therefore PNG is less

dependent on local rainfall. La Niña events associated with El Niño-Southern Oscillation affect

PNG, bringing periods of increased rainfall to the region every 2.5 to 7 years (Tudhope et al.,

2001).

Specimens from GBR and PNG were selected for this study as they were generously

made available from Dr. Paul Aharon’s personal collection. Tridacna from these locations have

been used in several other studies including: Chappell and Polach, 1972; Aharon, 1980; Aharon

and Chappell, 1986; Aharon, 1991; Hearty and Aharon, 1993; Elliot et al., 2009; Welsh et al.,

2011; Ayling et al., 2015; among others. Due to the popularity of Tridacna studies from these

locations and the availability of these specimens, it seemed wise to incorporate all modern

specimens available for this work.

Methods

A comparison of the first season of growth, internal from the pallial line, and the last season of

growth before capture, was made among analyzed specimens using SEM and EBSD techniques

(Fig. 1.3). However, petrographic analyses were made on the entire inner layer of K-133 (PNG)

and PT-1 (GBR) to estimate the total growth per lunar month.

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Petrography

Thin sections of the entire internal layers for K-133 and PT-1 were analyzed using a Nikon

stereoscopic microscope and SPOT Advanced imaging software at the Alabama Stable Isotope

Laboratory (ASIL) in the Department of Geological Sciences of The University of Alabama. Due

to the size of PT-3, a thin section representing the entire shell was not available.

Scanning Electron Microscopy (SEM)

Shell sections of T. gigas from the first and last stages of ontogenetic growth (Fig. 1.1) within the

internal layer were cut, first using a Hillquist Trim Saw and subsequently a Buehler Isomet 1000

for high precision, embedded in Buehler EpoxiCure 2 resin and hardener, and ground using sand

paper from coarse to fine grit size. Subsequently, each sample was polished using alumina oxide

of 1.0 µm and 0.3 µm and etched for 30 seconds using 2% HCl. Samples were coated with

approximately 20 nm of gold. SEM analyses were performed using a field emission scanning

electron microscope (FE-SEM) JEOL 7000 located in the Central Analytical Facility (CAF) of

The University of Alabama. Imaging was obtained at high vacuum, using a medium probe

current of 8 nA, and an accelerating voltage of 8 kV.

Electron Backscatter Diffraction (EBSD)

Samples used for SEM were re-polished and coated with 2.5 nm of carbon for EBSD analysis

(Pérez-Huerta and Cusack, 2009). The EBSD study was carried out with an Oxford Nordlys

camera mounted on a field emission scanning electron microscope (FE-SEM) JEOL 7000

located in the CAF of The University of Alabama. EBSD data were collected with Oxford Aztec

2.0 software at high vacuum, 20 kV, a large probe current of 15 nA, working distance of 10 mm

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and a resolution of 1.15 μm step size for crystallographic maps. Finally, data were analyzed

using OIM 5.3 from EDAX-TSL. In this study, EBSD data are represented by crystallographic

maps and pole figures in reference to the {001} plane of aragonite (see further details in Pérez-

Huerta et al., 2011).

Raman Spectroscopy

Ultrapolished samples were analyzed with a Jobin-Yvon HR800 UV Raman Spectrometer using

a wavelength of 100 nm and spot size of 5 µm in the Department of Chemistry at the University

of Alabama.

Carbon Stable Isotopes

Powder samples of T. gigas shells K-133 and PT-1 were acquired using a computer-assisted,

New Wave Research Leica GZ6 micromill and analyzed for δ13

C across the pallial line, sampling

both the external and internal layers of the shells, thought to be contemporaneous based on the

predicted direction of growth. The internal layer was sampled parallel to the pallial line at a high

resolution interval of 50 µm for approximately the first season of life [K-133 = 3 mm, and PT-1

= 1.6 mm]. In the external layer, parallel, non-continuous trenches were drilled every 120 µm at

approximately 45° from the pallial line. δ13

C is reported relative to the conventional Vienna Pee

Dee Belemnite standard (VPDB) calibrated against the NBS-19 standard. Analyses were

conducted in the ASIL using a ThermoFinnigan Delta Plus Isotope Ratio Mass Spectrometer

(IRMS) modified with a Gasbench for orthophosphoric acid digestion and online gas extraction.

The overall precision and reproducibility of the isotope measurements during the study was

0.06‰ on the basis of NBS-19 standard repeats (n = 27).

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Results

Shell Growth Features

Giant clams from both GBR and PNG exhibit regular growth lines that have been previously

interpreted as daily growth increments (Fig. 1.4; see also Aharon and Chappell, 1986; Watanabe

et al., 2004). A comparison of petrographic and SEM observations was conducted for samples K-

133 and PT-1 (Fig. 1.4) in order to correlate the daily growth lines with the microstructure.

Under SEM, the presence of these growth lines is more marked in the specimen from PNG (Fig.

1.4c), having an average thickness of 32.7 ± 2.5 µm (1σ, n = 10) during the first stage of growth

(Fig. 1.3a). Throughout the individual’s ontogeny, these growth increments become thinner, with

an average thickness of 21.2 ±5.8 µm (1σ, n = 10) at the last stage of growth (Fig. 1.3a).

Microscopically visible growth increments of specimens from the first stage of growth for the

GBR specimen (PT-1; Fig. 1.3b) have an average thickness of 26.1 ± 3.3 µm (1σ, n = 10) (Fig.

1.4d). At the last stage of growth (Fig. 1.3b), SEM reveals smaller increment thickness with an

average of 20.6 ± 3.2 µm (1σ, n = 10). These observations show a trend in the vertical reduction

of the daily growth line thickness with increasing age between first and last growth (Fig. 1.3)

specimens from both localities. The reduction in vertical daily growth thickness is likely caused

by shell extension with age that could be governed by mass conservation.

Shell Microstructure and Microtexture

The microstructure of the internal layers of giant clams collected from Palm Island, GBR, is

consistent between specimens and composed of shield-like aragonite crystals that do not vary in

morphology along a growth line (Fig. 1.5). These crystals tend to widen through the ontogeny at

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an independent rate for each one of the two specimens (Fig. 1.6 for measuring methodology).

The average crystal width in the first season for PT-3 is 2.7 ± 0.6 µm, (1σ, n = 20) (Figs. 1.3c

and 1.5a) and PT-1 is 0.9 ± 0.2 µm (1σ, n = 20) (Figs. 1.3b and 1.5b). The last season of growth

has average crystal thicknesses for PT-3 of 2.9 ± 1.2 µm (1σ, n = 20) (Figs. 1.3c and 1.5c) and

for PT-1 of 3.6 ± 1.5 µm (1σ, n = 20) (Figs. 1.3b and 1.5d). The microtextural characterization of

these aragonite crystals by EBSD was unsuccessful because diffraction was not recorded in

either specimen and no crystallographic information was obtained.

Unlike the GBR samples, the specimen from PNG exhibits couplets of two layers for

each daily growth increment (Fig.1.7): (i) a layer with a mean thickness of 33.5 ± 6.2 µm (1σ, n

= 12), consisting of well-organized, complex prismatic, orthogonal, aragonitic needles, and a

Figure 1.4, Petrographic and SEM observations of daily growth increments. Petrographic

analysis allows for the visual observation of wide-view daily growth increments: (a) K-133

and (b) PT-1; SEM images show distinct increments and suggest microstructural changes at a

low resolution: (c) K-133 and (d) PT-1.

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thinner increment with a mean thickness of 9.7 ± 3.3 µm (1σ, n = 12) composed of small,

clinogonal, oblique crystals propagating at an angle from the needles. The occurrence of these

couplets can be traced throughout the ontogenetic growth recorded within the internal layers, but

the aragonitic needles within the thicker bands tend to widen through ontogeny (Figs. 1.3a and

1.7; Fig. 1.6 for needle measuring methodology). Needles present in the first season of growth

average 3.0 ± 0.7 µm in width (1σ, n = 20) (Fig. 1.7a) and retain their morphology during later

growth, averaging 4.6 ± 1.3 µm in width (1σ, n= 20) (Fig. 1.7b). Unlike the specimens from

Palm Island, EBSD for the first and last seasons of growth successfully diffracted, producing in

situ crystallographic data to understand the mineralization of these crystal structure (Fig. 1.7).

The crystallographic map and pole figure indicate that there is microtextural continuity between

Figure 1.5, Microstructure of the internal layer of T. gigas specimens from Palm Island

showing wide, shield-like aragonite crystals for the first stages of growth in PT-3 (a) and PT-

1 (b) and those regions of the shell deposited at the end of the individual’s life [PT-3 (c) and

PT-1 (d)]. Arrow denotes the direction of growth.

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Figure 1.6, Width measurements were acquired during SEM imaging of the internal

layers (IL) overlaying the first season of growth (circle). Black box: section of shell

prepared for analysis; EL: External Layer. This example is K-133 (PNG), last

season of life.

the orthogonal needles and small, clinogonal, oblique crystals, representing single crystals with

the c-axis parallel to needle elongation and perpendicular to the visible growth lines.

EBSD of modern T. gigas allows for the in situ characterization of the microtexture of the

shell in original form. This will provide an understanding of the consistency regarding the

expected crystallographic context for future studies. T. gigas specimens of unknown age may

have been subjected to diagenetic processes which will likely show disorganization in which

case the shell might not be best fit for further analyses (Chappell and Polach, 1972).

Discussion

Shell Growth Features

Internal layers in hand-held, sectioned samples are characterized by macroscopically visible

alternation of light and darks bands, with each pair likely representing about one year of growth

(Aharon and Chappell, 1986). Counting these bands is the primary tool to obtain ages for modern

and fossil T. gigas shells (e.g., Aharon, 1991; Patzold et al., 1991; Watanabe et al., 2004; Elliot

et al., 2009; Welsh et al., 2011; Batenburg et al., 2011; Yan et al., 2013). Using this

methodology, for example, the age of the PT-3 specimen used in this study was estimated to be

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16 years (Aharon, 1991). Another approach for specimen-age determination is to use the

thickness of each individual, which is estimated to be 2 cm per year (Aharon and Chappell,

1986). Watanabe et al. (2004) reported the finding of a 60 year old T. gigas specimen that is 36

cm thick at its maximum growth (umbo regions). The specimen PT-3 is 16 cm thick at the umbo,

but only 16 years old based on band counting creating a large discrepancy between utilizing the

thickness of the shell versus the counting of annual bands. Besides morphological features,

chemical information is also used to determine the age of an individual. Oxygen isotopes

oscillate seasonally (Aharon and Chappell, 1986), being more enriched in 18

O during Austral

Figure 1.7, Elongated aragonite needle and small crystal packages compose the

microstructure of K-133 from PNG likely representing day time (DT) and night time (NT)

growth. The microstructure is consistent through the first stage of growth (a) and the last

season of the individual’s life (b). EBSD maps show the microtexture of this individual in

relation to the microstructure; the crystallographic orientation map (c) and pole figure (d)

show that the c-axis of aragonite is parallel to the elongation of needles and small crystals and

perpendicular to the daily growth increments (dashed lines), further displayed by the pole

figure (d). Arrow denotes the direction of growth.

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winter months and depleted in 18

O during the Austral summer months (Watanabe et al., 2004;

Elliot et al., 2009; Welsh et al., 2011; Batenburg et al., 2011; Yan et al., 2013). Utilizing δ18

O is

effective, although it requires accounting for vital effects and climate anomalies, which is hard to

do for fossil specimens.

An additional method of shell dating is proposed by counting the thinnest growth lines,

observed in all specimens under a petrographic microscope and SEM observations (Fig. 1.4).

These growth lines are accepted to be daily growth increments (Aharon and Chappell, 1986;

Sano et al., 2012) and can be counted in order to determine ontogenic age, which has also been

used in Tridacna maxima (Duprey et al., 2015). Using petrographic and SEM analysis, PT-1 age

before capture is approximately 46 months old and K-133 is approximately 44 months old at the

time of capture, and both have similar thicknesses of 6.5 cm and 7.3 cm, respectively. The large

age discrepancy between both GBR specimens (PT-3: 16 years and PT-1: 46 months) is likely

caused by the confusion of macroscopically visible light and dark banding, because there are

often subsets of smaller light and dark bands within the annual band that do not seem to correlate

to regular growth increments, allowing for erroneous age dating.

Daily band counting is thought to be the best possible method to estimate ontogenic ages

because there is less uncertainty, unless there is a significant break in growth (e.g., anomalous

seasonal growth interruption). However, the method is time-consuming and relies on good

sample preservation and thus, it could be a useful complement with annual band counting and

chemical data for fossils.

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Differences in Shell Microstructure and Microtexture

Although all studied specimens are assigned to the species Tridacna gigas and have an identical

external morphology, there are substantial differences in the microstructure and microtexture of

their internal shell layers. Specimens from GBR have an identical microstructure, composed of

shield-like aragonite crystals, while the specimen from PNG is composed of paired growth

increments of wide bands of orthogonal needles and narrow bands of small, clinogonal, aragonite

crystals that propagate at an angle from the needles (Figs. 1.5 and 1.6). These paired increments

are likely to represent daytime and nighttime deposition as suggested by Sano et al. (2012) on the

basis of solar insolation and measured Sr/Ca variability through the daily cycle. The

microstructural changes documented here for the PNG specimen support the Sano et al., (2012)

contention and are likely driven by the dissimilar calcification rates controlled by the

photosynthetic activity of zooxanthellae that are active during daytime and shut-off at night.

Although daily growth lines are recognizable at low resolution in the two GBR specimens the

paired day/night increments are apparently absent. Also, the more irregular morphology of

aragonite crystals and less defined crystal boundaries may suggest faster secretion of

microstructural components.

Besides a dissimilar microstructure, microtextural differences are also present between

specimens from the two locations. EBSD analysis of the specimen K-133 (PNG) produced

consistent aragonite diffraction patterns, allowing the comparison of microstructure to preferred

crystallographic orientations (Fig. 1.6), but there was no diffraction for the specimens from GBR

(PT-1 and PT-3). The absence of EBSD diffraction in biominerals has been attributed to the (i)

the presence of amorphous mineral phases; (ii) organics (e.g., Dalbeck et al., 2006); (iii) size of

crystals (Humphreys, 2004); and (iv) weak crystallization (Carlson, 1980; Kats et al., 1993).

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Figure 1.8, Raman spectra show the aragonite defining peaks at

wavelengths of 706 and 1085, and the absence of peaks associated with

organic components. Each spectrum is the average of four data points: (1)

PT-1 internal layer early ontogeny, (2) PT-1 internal layer late ontogeny,

(3) K-133 external, (4) PT-3 external, (5) K-133 internal layer late

ontogeny, (6) K-133 internal layer early ontogeny.

Raman spectroscopy was performed in order to resolve the absence of diffraction in specimens

from GBR and to determine any possible differences in the mineralization of internal and

external layers in the studied specimens. Data were collected for PT-1 and K-133 at four points

within the internal layer in both the first and last growth (Fig. 1.1). Results show the presence of

identical aragonite defining peaks at 706 and 1085 cm-1

(Fig. 1.7; Urmos et al., 1991) and the

absence of peaks attributed to organic components. These findings rule out the presence of

amorphous and organic phases in GBR specimens and favor the weak crystallization hypothesis

for the absence of EBSD diffraction at the activation volume (< 50 nm).

The discrepancies in microstructure and microtexture observed between specimens from

GBR and PNG could be attributed to either speciation or environmental factors. External

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morphologies of different giant clam species have been well assessed (Rosewater, 1965) and the

individuals in this study were confidently assigned to the species T. gigas. Analysis of genetic

differentiation of T. gigas in the Great Barrier Reef spanning over 1000 km showed that there

was genetic variability, although no significant differences were reported (Benzie and Williams,

1992). Due to the combination of the lack of speciation in distal reefs, as well as panmictic

reproduction, any genetic disturbance could be devastating for the species because the genes

would spread through the population rapidly (Benzie and Williams, 1992). In the current study,

the distance between specimen collection locations is approximately 900 km, which is shorter

than the variation between the farthest sampling sites utilized by Benzie and Williams (1992),

reporting no significant genetic deviations. Thus, it is assumed that the GBR and PNG localities

are sufficiently proximal to be considered members of the same reproductive population,

containing similar genetic components. Therefore, environmental differences between locations

are the likely explanation for observed differences in the internal layers of studied T. gigas

shells.

Environmental Factors Influencing Shell Growth

Biomineralization of the T. gigas shell is tightly controlled biologically, but also highly

influenced by site-specific environmental parameters (Sano et al., 2012). The influence of

external factors on shell growth is dependent on the scale of observation, with the smallest being

diurnal, while other factors are recognizable at the seasonal level.

On the diurnal scale, a symbiotic relationship maintained with photosynthetic

zooxanthellae, which live within the tissues of the siphonal mantle (Norton and James, 1992),

cause T. gigas to display brilliant blue and green colors when their valves are opened

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Figure 1.9, Growth tends to increase during the Austral summer and decrease in the Austral

winter for both PT-1 (GBR) and K-133 (PNG). A selection of seasonal data is shown.

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(Rosewater, 1965). Zooxanthellae depend on sunlight for their production of glucose, which is

essential in giant clam nutrition (Rees et al., 1993). According to Sano et al. (2012), giant clams

are supplied with nutrients at a higher rate during the daytime and thus calcify a thicker daytime

growth increment during sunlight hours.

Careful petrographic daily band counting yields total growth per month based on sets of

29 days (typical lunar month is 29.53 days). Therefore, growth along the same transect within the

shell can be assumed to represent the overall activity of the individual. Growth seems to be

partially influenced by general seasonality (Fig. 1.9). The rainy season, Austral summer in the

GBR and PNG, tends to be positively correlated with growth, while the dry season, Austral

winter, tends to have a negative correlation with growth. This effectively means that as the

season progresses, the amount of growth per lunar month is increasing. The association is more

pronounced in PT-1, (GBR) and subtle in K-133, (PNG); this might be due to anomalous

weather patterns, such as El Niño Southern Oscillation or influenced by differing environmental

fluctuations seen between the two locations, discussed below.

Figure 1.10, The quantified growth of approximately 29 day intervals for (a) K-133 (PNG) and

(b) PT-1 (GBR) are plotted in a grey and white dashed line and the time of day of the highest

tide, every seven days for (a) Dreger Harbour, PNG and (b) Lucinda, Australia, is represented

by the black line. Every seventh day was plotted for a clearer visibility and is thought to be

representative of the days surrounding. There is no apparent correlation between total growth

and the time of day of the highest tide. Tidal data from http://tides.mobilegeographics.com.

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At the seasonal level, there are several factors that may influence shell growth dynamics:

seawater temperature, tides, rainfall, solar irradiance and nutrient availability. Seawater

temperature variability is not considered to be important in the growth of these specimens

because seasonal differences in temperature are minimal at collection sites (Aharon, 1991). Tidal

cycles have been shown to correspond to growth at a 14-day periodicity, represented by the

ability to determine the breaks between or general width of daily growth increments (Pannella

and MacClintock, 1986). This noted periodicity is likely due to the prominence of either daytime

or nighttime increments, limited by the time of day of the highest tide. For instance, if the highest

tide occurs during the daytime, when zooxanthellae are active, it is thought that this would

disrupt maximum potential growth. If the highest tide is during the nighttime, the growth would

likely not be affected. To investigate this hypothesis, tidal records for Dreger Harbour, Papua

New Guinea (6.6500° S, 147.8667° E) and Lucinda, Australia (18.5167° S, 146.3333° E)

obtained from http://tides.mobilegeographics.com) and times of highest tide and total growth

were plotted (Fig. 1.10). The times of highest tides tend to oscillate, although maintain a seasonal

high or low, and although the total growth is only measured in terms of lunar months, plotting

these on a time series should show a correlation, where present. The growth of both specimens,

K-133 (PNG) and PT-1 (GBR), show no correspondence with daily tidal cycles as are observed

in other bivalve species (e.g., Pannella and MacClintock, 1968; Tran et al., 2011; Schöne and

Surge, 2012).

On the other hand, seasonal rainfall variability is greatly enhanced during the monsoons

that move through the region during the Austral summer (November through April) and the

relative drought during the Austral winter (May through October; Williamson and Hancock,

2005). Growth is plotted against rainfall records in Figure 1.11 from nearby stations used as a

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proxy for collection sites (Madang, Papua New Guinea, IAEA/WMO, 2014; Orpheus Island,

Great Barrier Reef, Australia, Bureau of Meteorology, Government of Australia, 2014). In

general, rainfall amount and total growth in both K-133 and PT-1 exhibit an inverse relationship

suggesting the existence of a linkage between the two parameters (Fig. 1.11), although the use of

data from proxy locations might introduce some error resulting less significant correspondence.

Solar irradiance has previously been correlated with enhanced daily shell growth, through

the analysis of Sr/Ca (Sano et al., 2012; Hori et al., 2015). This is thought to be due to the

influence from photosymbiosis. Therefore, in times that there are fewer cloudy days, there

should be more input from zooxanthellae, allowing the shell to have enhanced growth. Because

Figure 1.11, Growth lines of approximately 29 daily increments are marked on the whole

shell petrographic images with blue lines and shown plotted in red (K-133: a and c; PT-1 b

and d). Rainfall data, plotted in blue, is from Madang, Papua New Guinea (IAEA/WMO

(2014) and Orpheus Island, Great Barrier Reef, Australia (Bureau of Meteorology,

Government of Australia (2014). Calculated solar irradiance is in yellow (Nix and Kalma,

1972; Aharon (unpublished dissertation); NASA Langley Research Center, Atmospheric

Science Data Center).

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this data was not directly collected for regions of interest during the lifespan of the T. gigas, an

estimate of solar irradiance is required. An equation approximating the relationship of rainfall

and level of cloudiness was developed by Nix and Kalma (1972) specifically for the latitude of

6˚S, where the Huon Peninsula is located and further employed by Aharon (unpublished doctoral

dissertation):

Q = QA [(-0.5 • (p/Σp)) + (ΣQ/ΣQA) + 0.054]

Data for each variable from both locations were obtained from the NASA Langley Research

Center, Atmospheric Science Data Center for PNG (6˚S, 147.5˚E) and GBR (18.517˚S, 146.3˚E),

including insolation at the top of the atmosphere at the monthly (QA) and annual (ΣQA) scales

and annual effective solar radiation (ΣQ; Table 1.1). The independent variable influencing

changes in solar irradiance is monthly rainfall precipitation (p). Annual precipitation (Σp) is a

based on the sum of the average monthly precipitation (Table 1.1).

When plotted with growth, no direct relationship can be derived (Fig. 1.11). Rainfall data

for both sites during the lifespan of each T. gigas is a proxy because it was not collected prior to

the field work for initial studies (Madang, PNG and Orpheus Island, GBR, Australia) and

therefore is an approximation, likely adding error to the calculations for solar irradiance.

Additionally, because the equation was developed specifically for use in the Huon Peninsula, the

application to the GBR is an over-simplification of a more complicated system, which requires

further empirical testing to prove usefulness. With more representative rainfall and solar

irradiance data, a better correlation between these parameters and growth is predicted.

Nutrient availability in the environment in which T. gigas specimens live may impact

their ability to mineralize their shell. Major nutrients, including nitrogen and phosphorus, tend to

differ depending on the coral reef type. Typically, coral reefs are nutrient depleted and thought to

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be sinks of phosphorus and producers of carbon and nitrogen (Crossland and Barnes, 1983).

Open ocean environments, such as the fringing coral reef along the Huon Peninsula (PNG), have

between 0 and 4 µM of nitrogen and between 0 and 0.6 µM of phosphorous (Belda and

Yellowlees, 1993) and are relatively nutrient depleted. In contrast, the inshore reefs along the

GBR contain >1 µM of nitrogen and >2 µM of phosphorus derived through overland channel

inputs that introduce nitrogen and phosphorus to the system (Crossland and Barnes, 1983; Belda

and Yellowlees, 1993). A study from the Palm Passage, GBR, in 1983, only three years after the

collection of the GBR T. gigas, shows that nearby water quality samples contained between 0 –

0.045 µM nitrogen of predominately ammonium but also included nitrate and nitrite; ammonium

peaks between December and February while nitrate peaks between March and June.

Table 1.1, Values of data used in solar irradiance calculations for the purpose of estimating

monthly cloudiness. Data were obtained from the NASA Langley Research Center

Atmospheric Science Data Center.

Huon Peninsula Great Barrier Reef

(6˚S, 147.5˚E) (18.517˚S, 146.3˚E)

Month

1 912.0 989.5

2 920.7 946.5

3 903.4 869.0

4 851.0 757.2

5 783.8 651.3

6 746.8 597.1

7 761.5 620.4

8 816.5 708.1

9 869.0 819.1

10 903.4 912.0

11 903.4 972.3

12 903.4 998.1

Parameter Huon Peninsula Great Barrier Reef

Σp (mm) 2602.69 1147.5

ΣQ (cal/cm2/day) 173.2 ± 42.5 161.4 ± 85.8

ΣQA (cal/cm2/day) 312.7 ± 63.3 299.5 ± 149.2

Monthly Insolation (cal/cm2/day)

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Phosphorous ranged between 0.004 – 0.009 µM (Phosphate; Furnas and Mitchell, 1983). If these

results are representative of the very proximal location where PT-1 lived, increased nitrogen

might be responsible for some of the growth seen during the dry seasons as the rainy season

likely incorporated ammonium and nitrate into the lagoon. Although zooxanthellae can uptake

ammonium (Fitt et al., 1993), over a brief period of time, the nitrogen was likely fixed to become

nitrate and consumed by phytoplankton, which subsequently were filtered by the clams. The

large difference in nutrient availability between collection sites could be an important factor to

explain differences in growth as well as microstructure in the studied T. gigas specimens.

The combined processes surrounding these environmental parameters lead to the

conclusion that higher concentrations of nutrients, influenced by rainfall, and phytoplankton,

combined with an increase in solar irradiance, as suggested by Sano et al., (2012) and calculated

utilizing rainfall, would favor a faster growth rate. Therefore, the assumed seasonal order of

events begins with a period of increased rainfall, during which T. gigas should have a slower

growth rate due to an inferred increase in cloudiness preventing optimal photosynthesis (Sano et

al., 2012), followed by a period of enhanced solar irradiance and likely a spike in nutrient

availability, yielding increased shell growth (Fig. 1.11).

Biomineralization Model

Any model attempting to explain the biomineralization of T. gigas shells has to account for the

secretion of both internal and external layers (Fig. 1.12). Although the geochemistry of the

external layer has been studied (Elliot et al., 2009), its microstructure has not been fully

described and it is assumed to be composed of crossed lamellar aragonite (Patzold et al., 1991;

Lin et al., 2006). However, the external layer of another giant clam species, Hippopus hippopus,

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has been described to have a dendritic nature (Taylor, 1973), which is a precursor of the crossed

lamellar structure, although it does not have a first order lamellae (Kouchinsky, 2000). Dendritic

growth patterns are consistent with the current observations (Fig. 1.12). The microstructure of

the external layers is identical for all studied individuals, confirmed by Raman spectroscopy data

which show nearly identical intensities of Raman scattering (Fig. 1.8). This supports the idea that

external layers are secreted in the same manner, and possibly more highly controlled biologically

than the internal layers because the internal layers tend to differ while the external are identical.

Figure 1.12, The dendritic microstructure of the external layer of K-133 (PNG) compared

with the prismatic internal microstructure displays that the layers are thought to propagate

bi-directionally, as shown by arrows, the white box in (a) shows a representative area from

where (b) was imaged; the external layer is composed of two crystal orientations. Stable

carbon isotope profiles of the external and internal layers across the pallial line: (c) K-133,

from PNG and (d) PT-1, Palm Island, GBR.

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To determine possible differences in the metabolic pathways of shell deposition for both

internal and external layers, δ13

C was measured in the external layer and in the first season of

growth in the internal layer of both PT-1 and K-133 (Fig. 1.12). In K-133 there is a large positive

δ13

C shift of between 0.29 and 0.67 (‰VPDB) across the pallial line that separates external and

internal layers within a 95% confidence interval. The external δ13

C in the GBR specimen PT-1 is

also substantially higher than that of the internal by between 1.40 and 1.77 (‰VPDB) in a 95%

confidence interval, consistent with previous observations by Elliot et al. (2009) who found a

difference of about 2 (‰VPDB). As δ13

C appears to be consistently higher within the external

layer than internal, it further suggests that there are likely two metabolic pathways responsible

for shell secretion.

T. gigas has two mantle organs, which is uncommon in mollusks, and it may cause the

observed discrepancies between the internal and external shell layers. The siphonal mantle is in

Figure 1.13, Schematic representation of the

mantles and water currents in Tridacna gigas

(modified after Norton and James, 1992):

Siphonal mantle (SM), ctenidia (CT), incurrent

water (IWC), excurrent water chamber (ECW),

digestive mass (DM), and lateral mantle (LM).

Zooxanthellae tubular system is denoted by the

red pathway. The mechanism by which water is

filtered is shown by the blue pathway: incurrent

water moves through ctenidia and food is

removed within the ctenidia gills, processed,

and ultimately ending up in the digestive mass.

The digestive mass shares a boundary with the

hinge gland, which is partially bounded by the

lateral mantle and extra pallial space. The

lateral mantle is a membranous lining of the

inside of the valves and is confined by the

excurrent water chamber, which can hold and

squirt water as a method of deterring predators

at rates dependent on the individuals size (Neo

and Todd, 2011).

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Figure 1.14, Schematic representation of external (EL) and internal (IL) layers

of T. gigas shell in relation to the position of two mantles (blue: siphonal

mantle, green: lateral mantle; a). (b) Three folds of the siphonal mantle: outer

fold (OF), middle fold (MF), and inner fold (IF), responsible for the secretion of

the dendritic microstructure observed in the external layer of the shell (c). (d)

Cuboidal epithelial cells line of the lateral mantle, responsible for the secretion

of shield-like crystals in the internal layers of specimens from the Palm Island

(e) and the orthogonal needles and clinogonal crystals of internal layers in the

specimen from the PNG (f).

direct contact with the external layer while the lateral mantle lines the inside of the shell (Fig.

1.13). These results suggest that the siphonal mantle is responsible for the secretion of the

external shell layer. Histological studies show that there are three folds along the margin of this

mantle: inner fold, middle fold and, outer fold (Norton and James, 1992; Fig. 1.14). Each fold

displays tightly packed columnar epithelial cells on the outside, which are not exposed (Norton

and James, 1992; Fig. 1.14). The loosely folded inside of the inner fold is the colorful, visible

part of the mantle that holds the zooxanthellae (Norton and James, 1992). The size of the

dendritic structures produced is likely a direct representation of the space the epithelial cell had

to fill. Thus, it is proposed that tight folds are likely responsible for the third order lamellae while

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loose folds are likely responsible for the second order lamellae, defining the dendritic

microstructure

The lateral mantle is connected to the siphonal mantle at the pallial line, below which the

internal layer of the shell is covered with a very thin layer of membranous material (Norton and

James, 1992; Fig. 1.13). Simple cuboidal epithelial cells cover the lateral mantle along the shell

and are likely responsible for secreting the aragonitic needles or shield-like crystals that compose

internal microstructures of both GBR and PNG specimens (Fig. 1.13). The cuboidal epithelia are

not folded which explains the simplicity of the aragonite microstructure they produce. It is

possible that the size of the epithelial cells depends on the environment in which the clam lives,

providing an explanation for different microstructures produced in each unique locality, besides

nutrient availability.

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3. MICROSTRUCTURE VARIABILITY IN FOSSIL GIANT CLAMS (TRIDACNA GIGAS)

FROM THE HUON PENINSULA, PAPUA NEW GUINEA: AN ARCHIVE OF SOLAR

MODULATION?

M. E. Gannon1, P. Aharon1 and A. Pérez-Huerta1

1Department of Geological Sciences, The University of Alabama, Tuscaloosa

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Abstract

Massive, aragonitic shells of fossil giant clams, Tridacna gigas, are often well preserved and

provide robust bioarchives for paleoclimate studies because they are long-lived and exhibit fast

growth rates. Modern and fossil specimens from the uplifted coral reef terraces of the Huon

Peninsula, Papua New Guinea, display daily growth increments at the microscale that are

composed of pairs of complex prismatic aragonitic needles (daytime) and small crystals, oblique

to needles (nighttime). An exception includes two mid-Holocene-aged individuals, composed of

elongated, clinogonal aragonite crystals, oblique to the outside of the shell. Daily growth of

modern T. gigas has been documented to correlate with solar irradiance, and it is likely that fossil

T. gigas do also because similar growth increments are displayed. This study assesses fossil T.

gigas spanning the last 200 ka for microstructural differences as well as monitoring aragonitic

needle width (ANW) through time. The results show that ANW follows a cyclic pattern, similar

to those of solar radiation variability and δ18O, suggesting a relationship between solar irradiance

and the width of aragonitic needles over the past 200 ka. Due to microstructural and

microtextural differences between modern and mid-Holocene T. gigas, it is likely that there are

fundamental environmental differences between these times. The results of this study, suggesting

co-variation between ANW of fossil T. gigas and solar modulation, advance the potential of

giant clams to assist in the reconstruction of past solar modulation at a resolution higher than so

far achieved.

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Introduction

The study of past seasonal climate variability is important in order to understand current and

future climate trends. Ice cores, sediments, and fossils serve as useful archives of paleoclimate

data based on the analysis of climate proxies. However, well documented records are rare for the

tropics (Sano et al., 2012; Duprey et al., 2015; NOAA Index of Public Paleoclimatology

Datasets) although low latitudes predominantly control global climate, including El Niño

Southern Oscillation (ENSO) as well as Asian and African monsoons (Chiang, 2009).

Reef building organisms (e.g., corals, bivalves, algae) are important high resolution

bioarchives of the ambient environment and are extremely sensitive to climate shifts that are

embedded in the chemistry of their carbonate exoskeletons (McCulloch et al., 1999; Watanabe et

al., 2004; McGregor and Gagan, 2004; Elliot et al., 2009; Batenburg et al., 2011; Welsh et al.,

2011; Shöne and Surge, 2012). Giant clams, Tridacna species, are particularly useful bioarchives

of paleoclimate and paleoenvironmental change because of the following reasons: (i) their

massive aragonitic shells are typically well preserved (Hearty and Aharon, 1988; Romanek and

Grossman, 1989; Welsh et al., 2011; Warter et al., 2015); (ii) high growth rates afford daily to

seasonal resolution (Aharon and Chappell, 1986; Watanabe et al., 2004; Elliot et al., 2009;

Batenburg et al., 2011; Welsh et al., 2011; Yan et al., 2013; Duprey et al., 2015; Warter et al.,

2015); (iii) massive aragonite is deposited in oxygen isotope equilibrium and carbon isotope

equivalency with the ambient seawater (Aharon and Chappell, 1986; Jones et al., 1986; Aharon,

1991; Watanabe et al., 2004; Ayling, 2006; Batenburg et al., 2011; Welsh et al., 2011) and (iv)

major and trace chemical elements, (e.g., Sr, Mg, Ba) normalized to Ca, are useful proxies of the

ambient seawater environment (Elliot et al., 2009; Sano et al., 2012; Yan et al., 2013; Warter et

al., 2015; Hori et al., 2015; Yan et al., 2015). Yet, in spite of the recent flurry of geochemical

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proxy studies listed above, the variability of Tridacna shell microstructure in modern and fossil

specimens that may impact the records is rarely considered because it is thought that

macroscopic shell growth is relatively understood. For example, it has been documented that

bundles (botryoids) consisting of fibrous aragonitic needles in the internal layers of T. gigas tend

to widen in fossil samples while still preserving the original aragonite mineralogy (Chappell and

Polach, 1972; Aharon and Chappell, 1986). The question of whether the changing aragonite

needle width (ANW), previously observed in the fossil samples, shows an geologic age-

dependency or is variable and controlled by changes in ambient factors has not yet been

explored. The aim of this study is to quantitatively describe the microstructure of the internal

layers of fossil T. gigas shells of Late Pleistocene age in order to establish their variability and

explore whether ANW is governed by age or ambient changes.

Geological Context

The Huon Peninsula on the eastern borderland of Papua New Guinea (PNG) (6°S and 148°E)

(Fig. 2.1) is considered one of the best examples of raised coral reef terraces worldwide

(Pirazzoli et al., 1991). This region is subjected to an uplift rate of up to 2.5 m per 1000 years

due to massive compression of the Australian and Pacific Plates (Chappell, 1974a; Cutler et al.,

2003). The terraces have been the subject of extensive studies over several decades (Veeh and

Chappell, 1970; Chappell and Polach, 1972; Bloom et al., 1974; Aharon and Chappell, 1986;

Stein et al., 1993; Cutler et al., 2003; Ayling, 2006, among others) and yielded one of the most

detailed Late Pleistocene sea level change histories that confirmed the Milankovitch theory of ice

ages (Veeh and Chappell, 1970; Bloom et al., 1974; Aharon and Chappell, 1986; Chappell et al.,

1996; Lambeck and Chappell, 2001; Yokoyama et al., 2001; Cutler et al., 2003). Fringing,

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Figure 2.1, Plan view map of uplifted coral reef terraces along the

Sialum section of the Huon Peninsula, Papua-New Guinea (From

Aharon and Chappell, 1986). Terrace numbers are marked on

appropriate terrace locations (Table 2.1). RMF: Ramu-Markham

Fault.

barrier and lagoonal reefs are distinguished in the field on the basis of their geomorphology

(Chappell, 1974b; Aharon, 1983; Aharon and Chappell, 1986). The terrace units contain a

variety of coral species of Indo-Pacific affinity (Veeh and Chappell, 1970; Bloom et al., 1974;

Aharon and Chappell, 1986; Stein et al., 1993; Yokoyama et al., 2001), bivalves, coralline algae

and fine to coarse-grained reef sediments that are lithified into hard limestone (Chappell, 1974b;

Aharon and Chappell, 1986). Tridacna clams are commonly well preserved and often found in

growth position on the raised coral reef terraces.

Terrace Ages

The age assignment of the terraces, important to our evaluation of ANW variability in time, is

based on precise 238U/230Th and 235U/231Pa dating of relatively [U]-rich aragonitic corals (Veeh

and Chappell, 1970; Bloom et al., 1974; Aharon and Chappell, 1986; Stein et al., 1993; Cutler et

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al., 2003). Because of the rarity of diagenetically unaltered corals on the terraces on the one hand

and the impracticality of using well preserved but [U]-poor T. gigas on the other, the radiocarbon

method was used to date pristine T. gigas from terraces younger than 40 ka (1 ka = 1000 years)

(Veeh and Chappell, 1970; Bloom et al., 1974). Additionally, D-alloisoleucine/L-isoleucine

(D/L) ratios were measured in T. gigas whose ages were calibrated against radiometrically dated

coral reef terraces from Huon Peninsula. The relationship between the D/L ratios and the

radiometric ages offers an alternative method for estimating undated or insufficiently dated

terraces (Hearty and Aharon, 1988).

With an exception below, Table 2.1 offers a summary of the terrace ages I to VIII (Fig.

2.1) using U/Th and U/Pa age data from corals published so far (Veeh and Chappell, 1970;

Bloom et al., 1974; Aharon and Chappell, 1986; Stein et al., 1993; Cutler et al., 2003). The

exception includes two mid-Holocene T. gigas that were dated recently using the radiocarbon

method (Table 2.1). Careful consideration was given to each reported age and, in order to be

accepted for use in the current study, samples must have been in pristine condition, free of

Table 2.1, Weighted means and errors of accepted radiometric dates for each terrace based on

recent and previous studies (see text). Terrace numbers according to Figure 2.1. Terrace Dating Method Age (ka)

1Error (ka)

2Range (ka) n Reef Material

I Δ14

C 5.4 3

0.07 5.3-5.5 2 T. gigas

II U-Th & Pa-U 36.7 6, 8

0.01 31-37 5 coral

IIIb U-Th 41.7 5, 6

0.6 40-42 4 coral

IIIa U-Th 51.0 4, 6

0.6 49-53 3 coral

IV U-Th 62.5 4, 5, 6

0.6 57-74 7 coral

V U-Th & Pa-U 92.1 5, 6, 8

3.1 85-95 7 coral

VI U-Th 107.0 5, 6

0.2 106-108 3 coral

VIIb U-Th 119.1 4, 6, 7

1.9 116-124 8 coral

VIIa U-Th 134.1 4, 5, 6, 7

1.4 132-142 8 coral

VIII U-Th 198.7 7

4.6 180-190 1 coral

1Weighted Mean;

2Weighted Mean Error;

3Aharon, personal communication;

4Veeh and Chappell, 1970;

5Bloom et al., 1974;

6Aharon and Chappell, 1986;

7Stein et al., 1992;

8Cutler et al., 2003.

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diagenetic effects. If a particular coral age was significantly different from others on the same

terrace unit it was considered suspect and discarded (Table 2.1).

Methods

Materials Studied

Fossil specimens used in this study were collected by P. Aharon from the Huon Peninsula, PNG,

during three months of field work in the fall of 1977 (Figs. 2.1 and 2.2; Table 2.2). During this

collection, specimens were acquired only if both valves were found together and attached to the

limestone substrate (Aharon and Chappell, 1986). Petrographic and SEM observations were

made on the internal layer of the first season of growth within 1 to 2 millimeters of the umbo

(Fig. 2.3). In specimens where the umbo was not preserved, the measurements were made in the

area that was thought to be closest to that region. Resulting observations of the microstructure of

Table 2.2, Age assignment (Table 2.1) and specific reef

morphology of the sampling sites for each specimen

investigated in this study (see Fig. 2.2).

Sample Terrace Terrace Age (ka) Reef Facies

K-133 Modern 0 fringing reef crest

K-134 I 5.35 ± 0.02 1

fringing reef crest

K-135 I 5.52 ± 0.02 1

fringing reef crest

K-9 IIIb 41.7 ± 0.6 regression slope-fringing

K-8 IV 62.5 ± 0.6 fringing reef crest

K-14 V 92.1 ± 3.1 fringing reef crest

K-15 V 92.1 ± 3.1 fringing reef crest

K-17 VI 107 ± 0.2 fringing reef crest

K-46 VI 107 ± 0.2 fringing reef crest

K-131 VIIb 119.1 ± 1.9 regression slope-fringing

K-126 VIIa 134.1 ± 1.4 barrier reef

K-24 VIIa 134.1 ± 1.4 lagoonal

K-38 VIIa 134.1 ± 1.4 fringing reef crest1Aharon, peronal communication.

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internal layers were then compared to those from a modern specimen (K-133) collected from the

same locality (Fig. 2.4).

Optical Microscopy

Thin sections of the internal layer were analyzed using a Nikon stereoscopic microscope and

SPOT Advanced imaging software at the Alabama Stable Isotope Laboratory (ASIL) in the

Department of Geological Sciences of The University of Alabama.

Scanning Electron Microscopy (SEM)

Shell sections of the internal layer from first stage of growth of each T. gigas (Fig. 2.3) were cut,

embedded in resin, and ground using sand paper from coarse to fine grit size. Subsequently, each

sample was polished using alumina oxide of 1.0 µm and 0.3 µm and etched for 30 seconds using

2% HCl. Samples were coated with ~ 20 nm of gold. SEM analyses were performed using a

Field Emission Scanning Electron Microscope (FE-SEM) JEOL 7000 located in the Central

Analytical Facility (CAF) of The University of Alabama. Imaging was obtained at high vacuum,

using a medium probe current of 8 nA, and an accelerating voltage of 8 kV.

Electron Backscatter Diffraction (EBSD)

Samples used for SEM were re-polished and coated with 2.5 nm of carbon for EBSD analysis

(Pérez-Huerta and Cusack, 2009). The EBSD study of modern T. gigas (K-133) was carried out

with an Oxford Nordlys camera mounted on a Field Emission Scanning Electron Microscope

(FE-SEM) JEOL 7000 located in the Central Analytical Facility (CAF) of The University of

Alabama. EBSD data were collected with Oxford Aztec 2.0 software at high vacuum, 20 kV, a

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Figure 2.2, Shell internal layer sections of fossil T. gigas specimens.

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Figure 2.3, Aragonitic needle width (ANW) measurements were

acquired during SEM imaging of the internal layers (IL) overlaying

the first season of growth (black circle). Black box: section of shell

prepared for analysis; EL: External Layer. This example is K-15 from

Terrace V (see Fig. 2.2).

large probe current of 15 nA, working distance of 10 mm and a resolution of 1.15 μm step size

for crystallographic maps. Data for a mid-Holocene-aged specimen (K-134) were acquired using

a TESCAN LYRA FESEM also located in the Central Analytical Facility (CAF) at the

University of Alabama. EBSD data of K-134 were collected at high vacuum, 30 kV, a large

probe current (beam size) of 20 nA, working distance of 12.34 mm and a resolution of 1.0 μm

step size. Finally, data were analyzed using OIM 5.3 from EDAX-TSL. EBSD data are

represented by crystallographic maps and pole figures in reference to the {001} plane of

aragonite (see further details in Pérez-Huerta et al., 2011).

Results

Shell Growth Features

Hand specimens show macroscopic light and dark banding (Fig. 2.2), with coupled increments

thought to represent yearly couplets of seasonal growth (Aharon and Chappell, 1986). Based on

our petrographic observations, most fossil specimens exhibit regular growth lines, composed of

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paired light and dark increments that have been previously interpreted as daily growth lines

(Aharon and Chappell, 1986; Watanabe et al., 2004) (Fig. 2.5) consistent with the modern

specimen (Fig. 2.4b). The mid-Holocene-age specimen (K134) observed petrographically, does

not have clear daily growth increments regularly through ontogeny, but increments can be

identified in several locations (Fig. 2.5). Although the dissimilar structure is consistent in the

entire section, complicated sample preparation could cause this effect.

Figure 2.4, Modern T. gigas (K-133) is

the benchmark specimen to which fossils

are compared. (a) Section of K-133; (b)

Photomicrograph showing regular paired

light and dark bands that represent a daily

growth increment; plane view; (c) SEM

image of a single daily growth increment

showing aragonitic needles with small,

clinogonal, oblique crystals, representa-

tive of the overall microstructure found in

this individual. Arrow denotes direction

of growth.

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Shell Microstructure

Observations under high-resolution SEM imaging show that the entire ontogeny of modern T.

gigas (K-133) is composed of pairs of complex prismatic aragonitic needles, orthogonal to the

Figure 2.5, Photomicrographs of fossil T. gigas from thin sections; plane view. Light and dark

pairs represent daily growth increments. Arrow denotes direction of growth.

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external layer of the shell (Majewske, 1974; Flügel, 2013) and small, clinogonal aragonite

crystals, oblique to the aragonitic needles (Fig. 2.4c). Imaging of fossil T. gigas indicate that

daily growth increments are visible in most specimens and are also composed of complex

Figure 2.6, High resolution SEM images of fossil specimens exhibiting two morphological

types (Table 2.3). Mid-Holocene-aged T. gigas have elongated, clinogonal aragonite crystals,

oblique to the outside of the shell, an example is outlined a dashed, black line for K-134 and

K-135. Other T. gigas display orthogonal, complex prismatic aragonite needles. Arrow

denotes direction of growth.

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prismatic aragonite needles with an additional increment of small, clinogonal aragonite crystals,

oblique to the aragonitic needles (K-8, K-14, K-15, K-17, K-24, K-131 and K-38; Figs. 2.6 and

2.7b), similar to those present in the modern specimen (K-133; Fig. 2.4c). Several specimens

have poorly defined small, clinogonal crystals making the aragonitic needles appear to end

abruptly (K-9 and K-126; Figs. 2.6 and 2.7a) although their overall microstructure is comparable

with most of the other specimens. A T. gigas from Terrace VI, K-46, contains increments of

small, clinogonal, oblique aragonite crystals and poorly defined aragonitic needles although the

measured ANW is nearly the same as that of the other T. gigas (K-17) from Terrace VI, (see

Discussion). An exception includes mid-Holocene-age samples, K-134 and K-135, which are

composed of elongated, clinogonal crystals, oblique to the external shell layer (Figs. 2.6 and

2.7c).

Microtextural analyses of modern T. gigas show there is tight control over the

crystallographic domain of the aragonitic needles and small crystals, which are also well defined

in the crystallography map and pole figure (Fig. 2.8c and 2.8d). The c-axis is parallel to the

elongation of the needles, shown by the a- and b-axes alternating between needles.

Crystallographic preference extends through the small, clinogonal crystal increment, denoted by

the white dashed line (Fig. 2.8c). A mid-Holocene-aged sample, K-134, differs from the modern

in terms of microstructure as well as microtexture, displayed by the crystallography map and

Figure 2.7, Two needle morphological types documented in T.

gigas (a) orthogonal, aragonitic needles (daytime deposition)

with increments of small, clinogonal crystals, oblique to needles

(nighttime deposition) (K-8, K-14, K-15, K-17, K-24, K-131, K-

133 (Fig. 2.4a) and K-38; see Figs. 2.2 and 2.6), although several

samples have poorly defined small, clinogonal, oblique crystals

that are not easily visible (K-9 and K-126; see Figs. 2.2 and 2.6);

(b) elongated, clinogonal, oblique crystals that are primarily

observed in specimens of mid-Holocene-age (K-134 and K-135).

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pole figure obtained by EBSD (Fig. 2.8a and 2.8b). Elongated, clinogonal crystals, oblique to the

outside of the shell, are well-defined and present single crystallographic domains. Although the

c-axis of aragonite is overall parallel to the elongation of the crystals in each specimen, K-134

has less organization of the orientation of the crystallographic domains, represented by the

variability in the pole figure (Fig. 2.8b).

Figure 2.8, EBSD crystallographic maps and pole figures for a mid-Holocene-aged specimen

(K-134: a and b) and modern specimen (K-133: c and d) show that the c-axis is elongated in

the direction of growth, however the pole figures show that the modern specimen (d) had

more control over growth than the Holocene-age specimen (b) because the poles are more

constricted along the center of the figure (c-axis). White dashed lines constrict the nighttime

(NT) interval from the daytime (DT), displaying the retained crystallographic domain through

each interval (c). Red spots (a) and black spots (c) are areas of non-diffraction, which are

different colors due to the different equipment used to obtain EBSD data.

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Discussion

Aragonite Needle Width (ANW) Variability

Chappell and Polach (1972) proposed that botryoids of aragonite needles increase in width

through time due to diagenetic alteration from aragonite to calcite although needles retain their

morphology (Chappell and Polach, 1972; Aharon and Chappell, 1986; Ayling, 2006).

Differences in aragonite and calcite crystal densities (2.930 gcm-3

and 2.710 gcm-3

, respectively)

may explain the authors’ observations. It has also been suggested that neomorphism from

aragonite to a secondary aragonite is possible (Moir, 1990; Ayling, 2006). EBSD maps show that

there is no calcite in either modern or mid-Holocene specimens (Fig. 2.8). Additionally,

orthogonal needles (K-133) and clinogonal, oblique crystals (K-134 and K-135) are well defined

crystallographically ruling out a secondary phase of aragonite. If aragonite dissolution caused by

diagenesis were present (other than slight effects from etching with dilute HCl, required in SEM

Sample Terrace Age (ka) Microstructure

Average

ANW (µm)

(n = 20) σ se

K-133 Modern 0 Needles 3.04 0.69 0.15

K-134 I 5.4 Oblique Crystals 1.56 1.32 0.30

K-135 I 5.4 Oblique Crystals 1.13 0.40 0.09

K-9 IIIb 41.7 Needles 1.07 0.43 0.10

K-8 IV 62.5 Needles 2.43 1.12 0.25

K-14 V 92.1 Needles 1.32 0.48 0.11

K-15 V 92.1 Needles 1.65 0.59 0.13

K-17 VI 107 Needles 4.54 1.02 0.23

K-46 VI 107 Needles 4.52 1.74 0.39

K-131 VIIb 119.1 Needles 2.62 0.81 0.18

K-126 VIIa 134.1 Needles 4.50 1.10 0.25

K-24 VIIa 134.1 Needles 2.21 0.59 0.13

K-38 VIIa 134.1 Needles 3.08 0.71 0.14

σ: standard deviation; se: standard error

Table 2.3, Microstructure type, average needle width, 1σ standard deviation and standard

error of the mean for each T. gigas individual.

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sample preparation), then the EBSD analyses paired with SEM imaging would have recognized

it. Hence the aragonitic needles of each T. gigas are likely representatives of the geologic time in

which it lived. Due to the discrepancies in microstructure and microtexture between the mid-

Holocene specimens and other fossil T. gigas, measurements made on these crystals might not be

comparable with those of the aragonitic needles, however they are included in the ANW

comparison in this study. While it is found that ANW does increase through time, the upward

trend is not uniform but rather cyclical (Fig. 2.9a; Table 2.3). The relationship between daily

growth and solar irradiance, previously established in modern T. gigas by Sano et al. (2012) and

Hori et al. (2015), opens the opportunity for reconstructing past solar modulation (total

irradiance) variability using the ANW cyclicity in fossil T. gigas. According to van Geel et al.

(1999) and Sharma (2002), solar modulation is caused by complex interactions between galactic

cosmic rays, solar wind (solar magnetic field intensity) and the geomagnetic field intensity of the

Earth. Radiocarbon and 10

Be-based solar modulation reconstructed for the last 1000 years

exhibits an excellent match with the Schwabe (11-yr) irradiance cycle suggesting that solar

forcing may have contributed to a proportion of the warming since 1860 AD (Lean et al., 1995;

Muscheler et al., 2007).

Solar modulation variability has been estimated for the past 200 ka by Sharma (2002) on

the basis of 10

Be variability in deep-sea cores. According to the Sharma (2002) model, the

strongest solar modulation occurred between 111-125 ka followed by a general weakened trend

to the Holocene followed by modern high values (Fig. 2.9b). The solar modulation model of

Sharma (2002) matches well with the contemporaneous benthic marine δ18

O time-series (Grootes

et al., 1993; Johnson et al., 1997; Herbert et al., 2001; Lisiecki and Lisiecki, 2002) suggesting

that in general, warm periods during the interglacials (190 ka, 125 ka and modern) are typified

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Figure 2.9, Mean and standard error of ANW for each individual

giant clam plotted against age (a; Tables 2.2 & 2.3). Open boxes

represent the mid-Holocene-aged specimens with microstructure

varying from of other specimens. The black dashed line is a cubic

spline, showing the interpolation between ANW as it changes

through time. (b) Normalized solar modulation for the past 200 ka

(black dots) is plotted with δ18

O values (red line) (Lisiecki and

Lisiecki, 2002). Blue shaded and white boxes denote phase changes

and transitions between interglacial (blue) and glacial (white)

intervals. A rough correspondence is apparent between solar

modulation, δ18

O excursions and ANW variability [(b) modified

after Sharma, 2002].

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by high solar modulation and lower δ18

O values whereas colder intervals during interstadials and

glaciations are associated with weaker solar modulation and higher δ18

O values.

Aragonite secretion at the microstructural level is controlled diurnally by photosynthetic

processes due to T. gigas’ symbiotic relationship with zooxanthellae (Watanabe et al., 2004;

Sano et al., 2012; Yan et al., 2013; Hori et al., 2015). During daylight, photosynthesis of the

zooxanthellae allows the shell to calcify at a higher rate, thus producing the organized aragonitic

needles (Fig. 2.7). It follows that growth of fossil T. gigas also occurred under the diurnal

influence due to the presence of daytime and nighttime increments, similar to the modern K-133

(Fig. 2.4). Given the documented relationship between solar irradiance and growth attributed to

photosynthesis for modern T. gigas (Sano et al., 2013), we test below the hypothesis of a

correspondence between solar modulation change through time and giant clam growth pattern

expressed in terms of aragonite needle width (ANW) variability.

Notwithstanding that the number of T. gigas fossils in the pool of samples is scant, the

matching of the phase mode changes in the ANW record with those of the solar modulation and

δ18

O records is intriguing (Fig. 2.9). For example, the rise in solar modulation during the

penultimate interglacial, the last interglacial and the modern, accompanied by δ18

O negative

excursions, matches reasonably well with rises in ANW values of fossil T. gigas associated with

the coral reef terraces VIII at 198.7 ± 4.6 ka, VIIa at 134.1 ± 1.4 ka and VIIb at 119.1 ± 1.9 ka

and the modern reef. Substantial drops in solar modulation associated with positive δ18

O

excursions correspond to drops in ANW values of coral reef terraces formed during interstadials

(Terrace V at 92.1 ± 3.1 ka, Terrace IV at 62.5 ± 0.6 ka and Terrace IIIb, at 41.7 ± 0.6 ka).

Discrepancies between the exact timing of phase changes are discerned between ANW record in

Figure 2.9a and the records in Figure 2.9b. For example, whereas the maximum solar modulation

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values occur at around 124 ka, the ANW values peak 17 ka later at 107 ±0.2 ka. Additionally,

two fossil T. gigas samples from coral reef terrace VIIa at 134 ka yield significantly different

ANW values (Table 2.3 and Fig. 2.9a) for reasons that are not understood. These observed

discrepancies suggest either the presence of unidentifiable artifacts in the databases or some

leads and lags in the systems.

The discrepancies in microstructure between modern, K-133 and mid-Holocene-age T.

gigas, K-134 and K-135, as well as the microtextural differences between K-133 and K-134,

suggest that there are environmental differences between these times. Variations that could affect

shell microstructure include solar modulation, rainfall and nutrient availability. Based on these

circumstances, it is impossible to determine the reason for differences in biomineralization

mechanisms between modern and Holocene times though it is thought that they are likely

environmentally influenced.

Conclusions

Modern and fossil T. gigas from the raised coral reef terraces of the Huon Peninsula are

composed of orthogonal, complex, prismatic, aragonitic needles representing daylight deposition

and small, crystals, oblique to the needles, representing nighttime deposition. Mid-Holocene-

aged specimens are exceptional in their microstructure being composed of elongated aragonite

crystals, oblique to the outside of the shell. T. gigas depend on the solar irradiance for enhanced

daytime growth, which is a significant part of their shell deposition strategy; when solar

irradiance is high, the individual clams produce wider needles. The ANW in fossil specimens

shows phase-change modes that roughly match solar modulation and δ18

O variability over the

past 200 ka. Discrepancies between modern and mid-Holocene T. gigas are most likely due to

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yet unidentified environmental changes that cause a biological response. These initial results

suggest that shell microstructure (ANW) is likely affected by variations in solar activity and can

be quantified for comparisons through geologic time. Future studies will need a much larger

sample pool of wider age distribution to confirm the results reported here.

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REFERENCES

Aharon, P., personal communication.

Aharon, P. Stable isotope geochemistry of a late Quaternary coral reef sequence, New

Guinea: application of high resolution data to palaeoclimatology. (Unpublished doctoral

dissertation).

Aharon, P. (1983). 140,000-yr isotope climatic record from raised coral reefs in New Guinea.

Nature 304, 720-723.

Aharon, P. (1985) Carbon Isotope Record of Late Quaternary Coral Reefs: Possible Index of

Sea Surface Paleoproductivity. Geophysical Monograph 32, American Geophysical

Union, Washington, D.C., p. 343-355.

Aharon, P. (1991). Recorders of reef environment histories: stable isotopes in corals, giant clams,

and calcareous algae. Coral Reefs 10, 71-90.

Aharon, P., J. Chappell. (1986). Oxygen Isotopes, Sea Level Changes and the Temperature

History of a Reef Environment in New Guinea over the Last 105 Years.

Palaeogeography, Palaeoclimatology, Palaeoecology 56, 337-379.

Aharon, P., J. Chappell, and Compston, W. (1980). Stable isotopes and sea level data from New

Guinea support Antarctic ice-surge theory of ice ages: Nature 283, 649-651.

Åse, L-E. (1981). Studies of Shores and Shore Displacement on the Southern Coast of Kenya.

Especially in Kilifi District. Geografiska Annaler. Series A, Physical Geography 63, 303-

310.

Ayling, B. F. (2006). Seasonal Paleoclimates of the MIS 5E, 9 and 11 Interglacials, Using

Geochemical Proxies in Porites and Tridacna. (Unpublished doctoral dissertation).

Australian National University. Canberra, Australia.

Batenburg, S. J., G. Reichart, T. Jilbert, M. Janse, F. P. Wesselingh, W. Renema. (2011).

Interannual climate variability in the Miocene: High resolution trace element and stable

isotope ratios in giant clams. Palaeogeography, Palaeoclimatology, Palaeoecology 306,

75-81.

Belda, C. A., C. Cuff, and D. Yellowlees. (1993). Modification of Shell Formation in the Giant

Clam Tridacna gigas at Elevated Nutrient Levels in Sea Water. Marine Biology 117,

251-257.

Page 68: TRIDACNA GIGAS - Acumenacumen.lib.ua.edu/content/u0015/0000001/0002339/u0015...The giant clam, Tridacna gigas, is the largest known bivalve and one of the largest members of the Phylum

54

Benzie, J. A. H., S. T. Williams. (1992). No genetic differentiation of giant clam (Tridacna

gigas) populations in the Great Barrier Reef, Australia. Marine Biology 113, 373-377.

Bloom, A. L., W.S. Broecker, J. M. A. Chappell, R. K. Matthews, K. J. Mesolella. (1974).

Quaternary Sea Level Fluctuations on a Tectonic Coast: New 230Th

/234

U Dates from the

Huon Peninsula, New Guinea. Quaternary Research 4, 185-205.

Braley, R. D. (1984). Reproduction in the Giant Clams Tridacna gigas and T. deresa In situ on

the North-Central Great Barrier Reef, Australia, and Papua New Guinea. Coral Reefs 3,

221-227.

Bureau of Meteorology, Government of Australia. (2014) Climate and past weather. Station:

32083, Orpheus Island. http://www.bom.gov.au/climate/data.

Carlson, W. D. (1980). The calcite-aragonite equilibrium: effects of Sr substitution and anion

orientational disorder. American Mineralogist 65, 1252-1262.

Chappell, J., H. A. Polach. (1972). Some Effects of Partial Recrystallisation on 14

C Dating Late

Corals and Molluscs. Quaternary Research 2, 244-252.

Chappell, J. (1974a). Upper Mantle Rheology in a Tectonic Region: Evidence From New

Guinea. Journal of Geophysical Research 79, 390-398.

Chappell, J. (1974b). Geology of Coral Terraces, Huon Peninsula, New Guinea: A Study of

Quaternary Tectonic Movements and Sea-Level Changes. Geological Society of America

Bulletin 85, 553-570.

Chappell, J., A. Omura, T. Esat, M. McCulloch, J. Pandolfi, Y. Ota, B. Pillans. (1996).

Reconsiliation of late Quaternary sea levels derived from coral terraces at Huon

Peninsula with deep sea oxygen isotope records. Earth and Planetary Science Letters

141, 227-236.

Chiang, J. C. H. (2009). The Tropics in Paleoclimate. Annual Review of Earth Planet Science 37,

263-297.

Clement, A. C., R. Seager, M. A. Cane. (2000). Suppression of El Niño during the mid-Holocene

by changes in the Earth’s orbit. Paleoceanography 15, 731-737.

COHAP Members. (1988). Climatic Changes of the Last 18,000 Years: Observations and Model

Simulations. Science 241, 1043-1052.

Crossland, C. J., D. J. Barnes. (1983). Dissolved Nutrients and Organic Particulates in Water

Flowing over Coral Reefs at Lizard Island. Australian Journal of Marine Freshwater

Research 34, 853-844.

Cutler, K. B., R. L. Edwards, F. W. Taylor, H. Cheng, J. Adkins, C. D. Gallup, P. M. Cutler, G.

Page 69: TRIDACNA GIGAS - Acumenacumen.lib.ua.edu/content/u0015/0000001/0002339/u0015...The giant clam, Tridacna gigas, is the largest known bivalve and one of the largest members of the Phylum

55

S. Burr, A. L. Bloom. (2003). Rapid sea-level fall and deep-ocean temperature change

since the last interglacial period. Earth and Planetary Science Letters 206, 253-271.

Dalbeck, P., J. England, M. Cusack, M. R. Lee, A. E. Fallick. (2006). Crystallography and

chemistry of the calcium carbonate polymorph switch in M. edulis shells. European

Journal of Mineralogy 18, 601-609.

Deane, E. M., R. W. O’Brien. (1980). Composition of the haemolymph of Tridacna maxima

(Mollusca: Bivalvia). Comparative Biochemistry and Physiology 66A, 339-341.

Dodge, R. E., R. G. Fairbanks, L. K. Benninger, F. Maurrasse. (1983). Pliestocene Sea Levels for

Raised Coral Reefs of Haiti. Science 219, 1423-1425

Duprey, N., C. E. Lazareth, C. Dupouy, J. Butscher, R. Farman, C. Maes, G. Cabioch. (2015).

Calibration of seawater temperature and δ18

Oseawater signals in Tridanca maxima’s δ18

Oshell

record based on in situ data. Coral Reefs 34, 437-450.

Elliot, M., K. Welsh, C. Chilcott, M. McCulloch, J. Chappell, B. Ayling. (2009). Profiles of trace

elements and stable isotopes derived from giant long-lived Tridacna gigas bivalves:

Potential applications in paleoclimate studies. Palaeogeography, Palaeoclimatology,

Palaeoecology 280, 132-142.

Fitt, K., T. Rees, R. Braley, J. Lucas, D. Yellowlees. (1993). Nitrogen flux in giant clams: size-

dependency and relationship of zooxanthellae density and clam biomass in the uptake of

dissolved inorganic nitrogen. Marine Biology 117, 381-386.

Flater, David. Tides. Mobile Geographics. http://tides.mobilegeographics.com.

Flügel, E. (2013). Microfacies of Carbonate Rocks: Analysis, Interpretation and Application.

Springer Science & Business Media.

Furnas, M., A. Mitchell. (1983). A Hydrologic, Nutrient and Phytoplankton Biomass Survey in

Palm Passage (Central Great Barrier Reef) during 1983. Australian Institute of Marine

Science, Data Report: Oceanographic Series, AIMS-OS-84-1.

Gagan, M. K., L. K. Ayliffe, D. Hopley, J. A. Cali, G. E. Mortimer, J. Chappell, M. T.

McCulloch, M. J. Head. (1998). Temperature and Surface-Ocean Water Balance of the

Mid-Holocene Tropical Western Pacific. Science 279, 1014-1018.

Gannon, M., A. Pérez-Huerta, P. Aharon. (In submission). A Biomineralization Study of the

Giant Clam (Tridacna gigas) Shells. In submission to Coral Reefs.

Grootes, P. M., M. Stuiver, J. W. C. White, S. Johnsen, J. Jouzel. (1993). Comparison of oxygen

isotope records from the GISP2 and GRIP Greenland ice cores. Nature 366, 552-554.

Hearty, P. J., P. Aharon. (1988). Amino acid chronostratigraphy of late Quaternary coral reefs:

Page 70: TRIDACNA GIGAS - Acumenacumen.lib.ua.edu/content/u0015/0000001/0002339/u0015...The giant clam, Tridacna gigas, is the largest known bivalve and one of the largest members of the Phylum

56

Huon Peninsula, New Guinea, and the Great Barrier Reef, Australia. Geology 16, 579-

583.

Herbert, T. D., J. D. Schuffert, D. Andreasen, L. Heusser, M. Lyle, A. Mix, A. C. Ravelo, L. D.

Stott, J. C. Herguera, 2001, Collapse of the California current during glacial maxima

linked to climate change on land, Science v. 293: p71-76.

Hori, M., Y. Sano, A. Ishida, N. Takahata, K. Shirai, T. Watanabe. (2015). Middle Holocene

daily light cycle reconstructed from the strontium/calcium ratios of a fossil giant clam

shell. Nature Scientific Reports 5, 1-5.

Humphreys, F. J. (2004). Characterisation of fine-scale microstructures by electron backscatter

diffraction (EBSD). Scripta Materialia 51, 771-776.

IAEA/WMO. (2014). Global Network of Isotopes in Precipitation. http://www.iaea.org/water.

Jones, D. S., D. F. Williams, C. S. Romanek. (1986). Life Histories of Symbiont-Bearing Giant

Clams from Stable Isotope Profiles. Science 231, 46-48.

Johnson S. J., H. B. Clausen, W. Dansgaard, N. S. Gunderstrup, C. U. Hammer, U. Andersen, K.

K. Andersen, C. S. Hvidberg, D. Dahl-Jensen, J. P. Stffensen, H. Shoji, A. E.

SveinbjöRnsdóttir, J. White, J. Jouzel, D. Fisher. (1997). The δ18

O record along the

Greenland Ice Core Project deep ice core and the problem of possible Eemian climatic

instability. Journal of Geophysical Research: Oceans 102, 26,397-26,410.

Kats, E. I., V. V. Lebedev, A. R. Muratov. (1993). Weak crystallization theory. Physics Reports

228, 1-91.

King, B., F. McAllister, E. Wolanski, T. Done, S. Spagnol. (2001). Chapter 10: River Plume

Dynamics in the Central Great Barrier Reef. Oceanic Processes of Coral Reefs: Physical

and Biological links in the Great Barrier Reef (ed. E. Wolanski.) 145-159.

Kobayashi, I. (1969). Internal Microstructure of the Shell of Bivalve Molluscs. American

Zoologist 9, 663-672.

Kouchinsky, A. (2000). Shell microstructures in Early Cambrian molluscs. Acta

Palaeontological Polonica 45, 119-150.

Lambeck, K., J. Chappell. (2001). Sea Level Change Through the Last Glacial Cycle. Science

292, 679-686.

Lean, J., J. Beer, R. Bradley. (1995). Reconstruction of solar irradiance since 1610: Implications

for climate change. Geophysical Research Letters 22, 3195-3198.

Lin, A. Y. M, Meyers, M. A., Vecchio, K. S. (2006) Mechanical properties and structure of

Strombus gigas,Tridacna gigas, and Haliotis rufescens sea shells: A comparative study.

Page 71: TRIDACNA GIGAS - Acumenacumen.lib.ua.edu/content/u0015/0000001/0002339/u0015...The giant clam, Tridacna gigas, is the largest known bivalve and one of the largest members of the Phylum

57

Materials Science and Engineering 26, 1380-1389.

Lisiecki, L. E., P. A. Lisiecki. (2002). Application of dynamic programming to the correlation of

paleoclimate records. Paleoceanography 17, 1-12.

Majewske, O. P. (1974). Recognition of Invertebrate Fossil Fragments in Rock and Thin

Sections. Brill Achive 13.

McCulloch, M. T., A. W. Tudhope, T. M. Esat, G. E. Mortimer, J. Chappell, B. Pillans, A.

Chivas, A. Omura. (1999). Coral Record of Equatorial Sea-Surface Temperaures During

the Penultimate Deglaciation at Huon Peninsula. Science 283, 202-204.

McGregor, H. V., M. K. Gagan. (2004). Western Pacific coral δ18

O records of anomalous

Holocene variability in the El Niño-Southern Oscillation. Geophysical Research Letters

31, 1-4.

Moir, B. G. (1990). Comparative Studies of “Fresh” and “Aged” Tridacna gigas Shell:

Preliminary Investigations of a Reported Technique for Pretreatment of Tool Material.

Journal of Archaeological Science 17, 329-345.

Muscheler, R., F. Joos, J. Beer, S. A. Müller, M. Vonmoos, I. Snowball. (2007). Solar activity

during the last 1000 yr inferred from radionuclide records. Quaternary Science Reviews

26, 82-97.

NASA Langley Research Center, Atmospheric Science Data Center. eosweb.larc.nasa.gov/

Neumann, D. (1981). Tidal and Lunar Rhythms. Biological Rhythms, 351-380.

Neo, M. L., P. A. Todd. (2011). Quantification of water squirting by juvenile fluted giant clams

(Tridacna squamosa L.). Japan Ethological Society 29, 85-91.

Nix, H., J. Kalma. (1972). Climate as a dominant control in the biogeography of northern

Australia and New Guinea. Bridge and barrier: a natural and cultural history of Torres

Strait (ed. D. Walker). 61-91.

NOAA Index of Public Paleoclimatology Datasets. (ftp://ftp.ncdc.noaa.gov/pub/data/paleo/)

NOAA. (2009). National Weather Service Glossary. (weather.gov/glossary/)

Norton, J. H., G. W. Jones. (1992). The Giant Clam: An Anatomical and Histological Atlas.

Queensland Department of Primary Industries. Australian Centre for International

Agricultural Research.

Norton, J. H, M. A. Shepard, H. M. Long, W. K. Fitt. (1992). The Zooxanthellal Tubular System

in the Giant Clam. Biological Bulletin 183, 503-506.

Page 72: TRIDACNA GIGAS - Acumenacumen.lib.ua.edu/content/u0015/0000001/0002339/u0015...The giant clam, Tridacna gigas, is the largest known bivalve and one of the largest members of the Phylum

58

Pannella, G., C. MacClintock. (1968). Biological and Environmental Rhythms Reflected in

Molluscan Shell Growth. Memoir (Journal of Paleontology) 2, 64-80.

Pérez-Huerta, A., M. Cusack. (2009). Optimizing Electron Backscatter Diffraction of Carbonate

Biominerals—Resin Type and Carbon Coating. Microscopy and Microanalysis 15, 197-

203.

Pérez-Huerta, A., M. Cusack, P. Dalbeck. (2011). Crystallographic contribution to the vital effect

in biogenic carbonates Mg/Ca thermometry. Earth and Environmental Science

Transactions of the Royal Society of Edinburgh 102, 35-41.

Pirazzoli, P. A., U. Radtke, W. S. Hantoro, C. Jounnic, C. T. Hoant, C. Causse, M. Borel Best.

(1991). Quaternary Raised Coral-Reef Terraces on Sumba Island, Indonesia. Science

252, 1834-1836.

Radtke, U., G. Schellmann. (2005). Timing and Magnitude of Sea Level Change During MIS 5

Derived from Barbados Coral Reef Terraces: A Critical Literature Review and New Data.

Journal of Coastal Research SPECIAL ISSUE 42, 52-62.

Radtke, U., G. Schellmann. (2006). Uplift History along the Clermont Nose Traverse on the

West Coast of Barbados during the Last 500,000 Years-Implications for Paleo-Sea Level

Reconstructions. Journal of Coastal Research 22, 350-355.

Rees, T. A. V., W. K. Fitt, B. Baillie, D. Yellowlees. (1993). A Method for Temporal

Measurement of Hemolymph Composition in the Giant Clam Symbiosis and its

Application to Glucose and Glycerol Levels During a Diel Cycle. Limnology and

Oceanography 38, 213-217.

Rodríguez-Tovar, F. J. (2014). Orbital Climate Cycles in the Fossil Record: From Semidiurnal to

Million-Year Biotic Responses. Annual Review of Earth Planetary Science 42, 69-102.

Romanek, C. S., E. L. Grossman. (1989). Stable Isotope Profiles of Tridacna maxima as

Environmental Indicators. The Society of Economic Paleontologists and Mineralogists 4,

402-413.

Rosewater, J. (1965). The Family Tridacnidae in the Indo-Pacific. Indo-Pacific Mollusca 1: 347-

396.

Sano, Y., Kobayashi, S., Shirai, K., Takahata, N., Matsumoto, K., Watanabe, T., Sowa, K., Iwai,

K. (2012). Past daily light cycle recorded in the strontium/calcium ratios of giant clam

shells. Nature Communications 3, Article: 761.

Shöne, B. R., D. M. Surge. (2012). Bivalve Sclerochronology and Geochemistry. Treatise on

Invertebrate Paleontology, Part N, Revised 1, Chapter 14.

Sharma, M. (2002). Variations in solar magnetic activity during the last 200 000 years: is there a

Page 73: TRIDACNA GIGAS - Acumenacumen.lib.ua.edu/content/u0015/0000001/0002339/u0015...The giant clam, Tridacna gigas, is the largest known bivalve and one of the largest members of the Phylum

59

Sun-climate connection? Earth and Planetary Science Letters 199, 459-472.

Stein, M., G. J. Wasserburg, P. Aharon, J. H. Chen, Z. R. Zhu, A. Bloom, J. Chappell. (1993).

TIMS U-series dating and stable isotopes of last interglacial event in Papua New Guinea.

Geochimica et Comsmochimica Acta 57, 2541-2554.

Steinhilber, F., J. A. Abreu, J. Beer. (2008). Solar modulation during the Holocene. Astrophysics

and Space Sciences Transactions 4, 1-6.

Taylor, J. D. (1973). The Structural Evolution of the Bivalve Shell. Palaeontology 16, 519-534.

Tran, D., A. Nadau, G. Durrieu, P. Ciret, J. Parisot, J. Massabuau. (2011). Field Chronobiology

of a Molluscan Bivalve: How the Moon and Sun Cycles Interact to Drive Oyster Activity

Rhythms. Chronobiology International 28, 307-317.

Tudhope, A. W., C. P. Chilcott, M. T. McCulloch, E. R. Cook, J. Chappell, R. M. Ellam, D. W.

Lea, J. M. Lough, G. B. Shimmield. (2001). Variability in the El Niño-Southern

Oscillation through a Glacial-Interglacial Cycle. Science 291, 1511-1517.

Urmos, J., S. K., Sharma, F. T. Mackenzie. (1991). Characterization of some biogenic carbonates

with Raman spectroscopy. American Mineralogist 76, 641-646.

van Geel, B., O. M. Raspopov, H. Renssen, J. van der Plicht, V. A. Dergachev, H. A. J. Meijer.

(1999). The role of solar forcing upon climate change. Quaternary Science Reviews 18,

331-338.

Veeh, H. H., J. Chappell. (1970). Astronomical Theory of Climatic Change: Support from New

Guinea. Science 167, 862-865.

Warter, V., W. Müller, F. P. Wesselingh, J. A. Todd, W. Renema. (2015). Late Miocene

Seasonal to Subdecadal Climate Variability in the Indo-West Pacific (East Kalimantan,

Indonesia) Preserved in Giant Clams. Palaios 30, 66-82.

Warter, V. W. Müller. (2016). Daily growth and tidal rhythms in Miocene and modern giant

clams revealed via ultra-high resolution LA-ICPMS – A novel methodological approach

towards improved sclereochemistry. Palaeogeography, Palaeoclimatology,

Palaeoecology. In press.

Watanabe, T., A. Suzuki, H. Kawahata, H. Kan, and S. Ogawa. (2004). A 60-year Isotopic

Record from a Mid-Holocene Fossil Giant Clam (Tridacna Gigas) in the Ryukyu Islands:

Physiological and Paleoclimatic Implications. Palaeogeography, Palaeoclimatology,

Palaeoecology 212, 343-354.

Weiner, S., P. M. Dove. (2003). An Overview of Biominrealization Processes and the Problem

of the Vital Effect. [Eds. P. M. Dove, J. J. DeYoreo, and S. Weiner]. Reviews in

Mineralogy and Geochemistry: Biomineralization 54, 1-30.

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60

Welsh, K., M. Elliot, A. Tudhope, B. Ayling, J. Chappell. (2011). Giant bivalves (Tridacna

gigas) as recorders of ENSO variability. Earth and Planetary Science Letters 307, 266-

270.

Williamson, A., G. Hancock. (2005). The Geology and Mineral Potential of Papua New Guinea.

Papua New Guinea Department of Mining, 30-41.

Yan, H., D. Shao, Y. Wang, L. Sun. (2013) Sr/Ca profile of long-lived Tridacna gigas bivalves

from South China Sea: A new high-resolution SST proxy. Geochimica et Cosmochimica

Acta 112, 52-65.

Yan, H., L. Sun, D. Shao, Y. Wang. (2015). Seawater Temperature seasonality in the South

China Sea during the late Holocene derived from high-resolution Sr/Ca ratios of Tridacna

gigas. Quaternary Research 83, 298-306.

Yan, H., Y. Wang, L. Sun. (2014). High resolution oxygen isotope and grayscale records of a

medieval fossil giant clam (Tridacna gigas) in the South China Sea: physiological and

paleoclimatic implications. Acta Oceanologica Sinica 33, 18-25.

Yokoyama, Y., T. M. Esat, K. Lambeck. (2001). Coupled climate and sea-level changes deduced

from Huon Peninsula coral terraces of the last ice age. Earth and Planetary Science

Letters 193, 579-587.

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APPENDIX I: BIOLOGY

Relevant Anatomy

Organs responsible for shell production are those involved with the organism’s respiration and

digestion (Fig. 1.12). The siphonal mantle is visible when the valves are opened. Iridophores of

brilliant shades of blue and green are characteristic of T. gigas (Rosewater, 1965), produced by

photosynthetic zooxanthellae making this species unique from most other bivalves (Norton, et

al., 1992). Zooxanthellae live within a tubular system that extends from the siphonal mantle to

the digestive mass (Norton et al., 1992). The siphonal mantle also holds the internal and external

siphons that are responsible for filtering water in to and out of the organism. Surrounding these

orifices are sensory hyaline organs that allow the animal to detect predators, increasing their

awareness and longevity (Norton et al., 1992). Incurrent water moves through ctenidia and food

is filtered out within the ctenidial gills, processed, and ultimately ending up in the digestive mass

(Norton and Jones, 1992). The digestive mass shares a boundary with the hinge gland, which is

partially bounded by the lateral mantle and extra pallial space (Norton and Jones, 1992). The

lateral mantle is a membranous lining of the inside of the valves and is confined by the excurrent

water chamber, which can hold and squirt water as a method of deterring predators at rates

dependent on the individuals size (Neo and Todd, 2011). The pallial line is the location on the

shell where the siphonal and lateral mantles attach to the valves. This is also the interface of the

internal and external layers of shell (Norton et al., 1992).

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Feeding Mechanisms

In order to maintain rapid shell growth, a large source of nutrients is required. T. gigas fulfills

this need through two feeding mechanisms: traditional filter feeding, and symbiotic

dinoflagellate zooxanthellae. In the juvenile stage, the clam ingests and retains photosynthetic

zooxanthellae within their siphonal mantle (Fitt and Trench, 1981) by the means of sequestering

through a tubular system from their digestive mass (Norton et al., 1992). Zooxanthellae provide

the individual with carbon for calcification in the forms of glycerol and glucose (Rees et al,

1993). They can also be harvested if there is a lack of phytoplankton in the water, obtained

during filter feeding. During early ontogeny, T. gigas depends on both sources of nutrients; as it

grows, it becomes increasingly dependent on the symbiotic relationship: particulate carbon from

filter feeding decreases in absorbance from 65% to 34% (Klumpp et al., 1992).

Reproduction

T. gigas are sequential simultaneous hermaphrodites; individuals produce both egg and sperm in

a sequential pattern, although they are not spawned at the same time (Braley, 1984). Upon

reaching sexual maturity between 10 years (Jones, et al., 1986) and 12 years of age (Watanabe et

al., 2004), the gonads develop initially as male components and later add female ovaries (Norton

and Jones, 1992). The gonads are located along the side of the wall of the digestive mass (Norton

and Jones, 1992), likely allowing for easy reallocation of nutrients from shell secretion during

spawning, which is associated with decreased summer growth at the onset of sexual maturity

(Romanek and Grossman, 1989). Lunar periodicity affects when spawning occurs, however it

differs with locality ranging from on the day of the new moon to 13 days after the new moon

(Braley, 1984). The lunar month is often recorded within the shell in patterns of simple and

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complex (light and dark) daily growth increments that remain fairly consistent in thickness

through the duration of the month (Pannella and MacClintock, 1968). Due to the energy and

nutrients spawning requires, there could potentially be microstructural influences visible in the

shell.

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APPENDIX II: SEASONAL GROWTH INCREMENTS

SEM band counting and assuming daily increments for the growth lines, light and dark

microscopically visible increments at low resolution, displays microstructural changes at a

regular interval every two to three lunar months (29.53 days per lunar month) for all modern

specimens analyzed. GBR specimens showed a microstructural modification at growth breaks

represented by a band of organized needles bounded on either side by the normal growth of

shield-like aragonite crystals (Fig. A.1). Growth breaks at these intervals are also apparent in the

T. gigas from PNG (Fig. A.2). The microstructure in K-133 is composed of small disorganized

crystals in bands that span the height of several days of growth (Figs. 1.7a and 1.7b). The EBSD

crystallography map and pole figure of a K-133 seasonal break shows a continuity of the c-axis

of aragonite following the elongation of needles in the direction of growth, although the

diffraction is reduced and corresponds to the small crystal size present at these seasonal break

increments (Fig. A.2c and A.2d).

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Figure A.1, Seasonal breaks are seen at regular intervals of approximately three months in

GBR specimens by semi-organized needles with a noticeable break in growth for early

ontogenies [PT-3 (a) and PT-1 (b)] as well as during the last stage of the individual’s life [PT-

3 (c) and PT-1 (d)].

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Figure A.2, Seasonal increments in the T. gigas from PNG are composed of tiny

disorganized crystals in early ontogeny (a) and later ontogeny (b). Reduced size of the

crystals at seasonal breaks is seen in microtexture EBSD mapping (c) which retains c-axis in

the same position as non-seasonal increments (d).

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APPENDIX III: NEEDLE MEASUREMENTS

(See Figs. 1.6 and 2.3)

Age (Table 2.1) Specimen n Layer (Fig. 1.1) Needle Width (µm)

Modern K-133 1 FG 3.69

Modern K-133 2 FG 2.04

Modern K-133 3 FG 2.62

Modern K-133 4 FG 1.57

Modern K-133 5 FG 4.71

Modern K-133 6 FG 2.35

Modern K-133 7 FG 3.45

Modern K-133 8 FG 2.66

Modern K-133 9 FG 2.74

Modern K-133 10 FG 3.46

Modern K-133 11 FG 2.97

Modern K-133 12 FG 2.93

Modern K-133 13 FG 3.17

Modern K-133 14 FG 3.08

Modern K-133 15 FG 2.75

Modern K-133 16 FG 2.93

Modern K-133 17 FG 2.58

Modern K-133 18 FG 3.98

Modern K-133 19 FG 3.82

Modern K-133 20 FG 3.36

Modern K-133 1 LG 8.01

Modern K-133 2 LG 3.41

Modern K-133 3 LG 3.32

Modern K-133 4 LG 3.75

Modern K-133 5 LG 3.99

Modern K-133 6 LG 3.41

Modern K-133 7 LG 5.79

Modern K-133 8 LG 6.21

Modern K-133 9 LG 6.78

Modern K-133 10 LG 4.78

Modern K-133 11 LG 4.48

Modern K-133 12 LG 3.28

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Age (Table 2.1) Specimen n Layer (Fig. 1.1) Needle Width (µm)

Modern K-133 13 LG 3.50

Modern K-133 14 LG 4.05

Modern K-133 15 LG 4.02

Modern K-133 16 LG 5.78

Modern K-133 17 LG 6.16

Modern K-133 18 LG 3.88

Modern K-133 19 LG 3.23

Modern K-133 20 LG 5.02

Modern PT-1 1 FG 1.03

Modern PT-1 2 FG 0.72

Modern PT-1 3 FG 1.15

Modern PT-1 4 FG 1.00

Modern PT-1 5 FG 0.75

Modern PT-1 6 FG 0.72

Modern PT-1 7 FG 0.62

Modern PT-1 8 FG 0.75

Modern PT-1 9 FG 1.12

Modern PT-1 10 FG 1.12

Modern PT-1 11 FG 0.87

Modern PT-1 12 FG 1.25

Modern PT-1 13 FG 0.87

Modern PT-1 14 FG 0.84

Modern PT-1 15 FG 0.59

Modern PT-1 16 FG 0.93

Modern PT-1 17 FG 0.84

Modern PT-1 18 FG 0.69

Modern PT-1 19 FG 0.75

Modern PT-1 20 FG 0.90

Modern PT-1 1 LG 5.10

Modern PT-1 2 LG 3.02

Modern PT-1 3 LG 4.01

Modern PT-1 4 LG 4.10

Modern PT-1 5 LG 1.59

Modern PT-1 6 LG 2.33

Modern PT-1 7 LG 3.01

Modern PT-1 8 LG 2.31

Modern PT-1 9 LG 4.34

Modern PT-1 10 LG 2.92

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Age (Table 2.1) Specimen n Layer (Fig. 1.1) Needle Width (µm)

Modern PT-1 11 LG 2.86

Modern PT-1 12 LG 3.44

Modern PT-1 13 LG 2.33

Modern PT-1 14 LG 8.76

Modern PT-1 15 LG 3.81

Modern PT-1 16 LG 3.75

Modern PT-1 17 LG 3.54

Modern PT-1 18 LG 4.49

Modern PT-1 19 LG 1.32

Modern PT-1 20 LG 4.33

Modern PT-3 1 FG 3.50

Modern PT-3 2 FG 2.34

Modern PT-3 3 FG 3.63

Modern PT-3 4 FG 2.95

Modern PT-3 5 FG 3.77

Modern PT-3 6 FG 3.63

Modern PT-3 7 FG 3.26

Modern PT-3 8 FG 1.19

Modern PT-3 9 FG 1.81

Modern PT-3 10 FG 2.94

Modern PT-3 11 FG 2.57

Modern PT-3 12 FG 2.27

Modern PT-3 13 FG 3.03

Modern PT-3 14 FG 2.32

Modern PT-3 15 FG 2.88

Modern PT-3 16 FG 2.82

Modern PT-3 17 FG 2.78

Modern PT-3 18 FG 2.08

Modern PT-3 19 FG 3.07

Modern PT-3 20 FG 2.02

Modern PT-3 1 LG 3.36

Modern PT-3 2 LG 1.50

Modern PT-3 3 LG 2.15

Modern PT-3 4 LG 2.47

Modern PT-3 5 LG 3.06

Modern PT-3 6 LG 2.72

Modern PT-3 7 LG 2.90

Modern PT-3 8 LG 1.92

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Age (Table 2.1) Specimen n Layer (Fig. 1.1) Needle Width (µm)

Modern PT-3 9 LG 2.95

Modern PT-3 10 LG 4.65

Modern PT-3 11 LG 3.73

Modern PT-3 12 LG 1.37

Modern PT-3 13 LG 4.27

Modern PT-3 14 LG 2.72

Modern PT-3 15 LG 3.13

Modern PT-3 16 LG 3.00

Modern PT-3 17 LG 2.99

Modern PT-3 18 LG 2.86

Modern PT-3 19 LG 3.76

Modern PT-3 20 LG 2.06

5.35 ± 0.02 K-134 1 FG 1.19

5.35 ± 0.02 K-134 2 FG 1.54

5.35 ± 0.02 K-134 3 FG 1.69

5.35 ± 0.02 K-134 4 FG 2.70

5.35 ± 0.02 K-134 5 FG 0.75

5.35 ± 0.02 K-134 6 FG 0.77

5.35 ± 0.02 K-134 7 FG 1.63

5.35 ± 0.02 K-134 8 FG 0.88

5.35 ± 0.02 K-134 9 FG 1.13

5.35 ± 0.02 K-134 10 FG 1.89

5.35 ± 0.02 K-134 11 FG 1.88

5.35 ± 0.02 K-134 12 FG 2.26

5.35 ± 0.02 K-134 13 FG 0.78

5.35 ± 0.02 K-134 14 FG 4.01

5.35 ± 0.02 K-134 15 FG 2.00

5.35 ± 0.02 K-134 16 FG 2.33

5.35 ± 0.02 K-134 17 FG 0.75

5.35 ± 0.02 K-134 18 FG 1.02

5.35 ± 0.02 K-134 19 FG 0.84

5.35 ± 0.02 K-134 20 FG 1.14

5.52 ± 0.02 K-135 1 FG 0.19

5.52 ± 0.02 K-135 2 FG 0.09

5.52 ± 0.02 K-135 3 FG 0.53

5.52 ± 0.02 K-135 4 FG 0.13

5.52 ± 0.02 K-135 5 FG 0.09

5.52 ± 0.02 K-135 6 FG 0.28

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Age (Table 2.1) Specimen n Layer (Fig. 1.1) Needle Width (µm)

5.52 ± 0.02 K-135 7 FG 0.00

5.52 ± 0.02 K-135 8 FG 0.09

5.52 ± 0.02 K-135 9 FG 0.07

5.52 ± 0.02 K-135 10 FG 0.31

5.52 ± 0.02 K-135 11 FG 0.15

5.52 ± 0.02 K-135 12 FG 0.06

5.52 ± 0.02 K-135 13 FG 0.28

5.52 ± 0.02 K-135 14 FG 0.30

5.52 ± 0.02 K-135 15 FG 0.36

5.52 ± 0.02 K-135 16 FG 0.00

5.52 ± 0.02 K-135 17 FG 0.07

5.52 ± 0.02 K-135 18 FG 0.01

5.52 ± 0.02 K-135 19 FG 0.00

5.52 ± 0.02 K-135 20 FG 0.21

41.7 ± 0.6 K-9 1 FG 0.85

41.7 ± 0.6 K-9 2 FG 1.27

41.7 ± 0.6 K-9 3 FG 0.95

41.7 ± 0.6 K-9 4 FG 1.60

41.7 ± 0.6 K-9 5 FG 1.05

41.7 ± 0.6 K-9 6 FG 0.72

41.7 ± 0.6 K-9 7 FG 0.87

41.7 ± 0.6 K-9 8 FG 1.25

41.7 ± 0.6 K-9 9 FG 0.87

41.7 ± 0.6 K-9 10 FG 0.52

41.7 ± 0.6 K-9 11 FG 1.12

41.7 ± 0.6 K-9 12 FG 1.25

41.7 ± 0.6 K-9 13 FG 0.55

41.7 ± 0.6 K-9 14 FG 1.25

41.7 ± 0.6 K-9 15 FG 0.72

41.7 ± 0.6 K-9 16 FG 1.37

41.7 ± 0.6 K-9 17 FG 2.05

41.7 ± 0.6 K-9 18 FG 1.15

41.7 ± 0.6 K-9 19 FG 0.75

41.7 ± 0.6 K-9 20 FG 1.12

62.5 ± 0.6 K-8 1 FG 3.54

62.5 ± 0.6 K-8 2 FG 3.70

62.5 ± 0.6 K-8 3 FG 3.75

62.5 ± 0.6 K-8 4 FG 3.29

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Age (Table 2.1) Specimen n Layer (Fig. 1.1) Needle Width (µm)

62.5 ± 0.6 K-8 5 FG 0.83

62.5 ± 0.6 K-8 6 FG 5.07

62.5 ± 0.6 K-8 7 FG 2.75

62.5 ± 0.6 K-8 8 FG 1.92

62.5 ± 0.6 K-8 9 FG 2.27

62.5 ± 0.6 K-8 10 FG 2.55

62.5 ± 0.6 K-8 11 FG 0.94

62.5 ± 0.6 K-8 12 FG 2.71

62.5 ± 0.6 K-8 13 FG 1.62

62.5 ± 0.6 K-8 14 FG 1.28

62.5 ± 0.6 K-8 15 FG 2.00

62.5 ± 0.6 K-8 16 FG 0.82

62.5 ± 0.6 K-8 17 FG 2.09

62.5 ± 0.6 K-8 18 FG 1.33

62.5 ± 0.6 K-8 19 FG 3.70

62.5 ± 0.6 K-8 20 FG 2.48

92.1 ± 3.1 K-14 1 FG 1.82

92.1 ± 3.1 K-14 2 FG 0.97

92.1 ± 3.1 K-14 3 FG 1.13

92.1 ± 3.1 K-14 4 FG 0.79

92.1 ± 3.1 K-14 5 FG 2.94

92.1 ± 3.1 K-14 6 FG 1.08

92.1 ± 3.1 K-14 7 FG 1.33

92.1 ± 3.1 K-14 8 FG 1.01

92.1 ± 3.1 K-14 9 FG 2.08

92.1 ± 3.1 K-14 10 FG 1.25

92.1 ± 3.1 K-14 11 FG 0.96

92.1 ± 3.1 K-14 12 FG 1.58

92.1 ± 3.1 K-14 13 FG 1.34

92.1 ± 3.1 K-14 14 FG 1.20

92.1 ± 3.1 K-14 15 FG 0.95

92.1 ± 3.1 K-14 16 FG 1.08

92.1 ± 3.1 K-14 17 FG 1.41

92.1 ± 3.1 K-14 18 FG 1.25

92.1 ± 3.1 K-14 19 FG 1.40

92.1 ± 3.1 K-14 20 FG 0.89

92.1 ± 3.1 K-15 1 FG 2.54

92.1 ± 3.1 K-15 2 FG 0.91

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Age (Table 2.1) Specimen n Layer (Fig. 1.1) Needle Width (µm)

92.1 ± 3.1 K-15 3 FG 2.59

92.1 ± 3.1 K-15 4 FG 1.16

92.1 ± 3.1 K-15 5 FG 2.39

92.1 ± 3.1 K-15 6 FG 2.83

92.1 ± 3.1 K-15 7 FG 2.11

92.1 ± 3.1 K-15 8 FG 1.66

92.1 ± 3.1 K-15 9 FG 1.88

92.1 ± 3.1 K-15 10 FG 1.77

92.1 ± 3.1 K-15 11 FG 1.67

92.1 ± 3.1 K-15 12 FG 1.21

92.1 ± 3.1 K-15 13 FG 1.49

92.1 ± 3.1 K-15 14 FG 0.66

92.1 ± 3.1 K-15 15 FG 1.35

92.1 ± 3.1 K-15 16 FG 1.21

92.1 ± 3.1 K-15 17 FG 1.71

92.1 ± 3.1 K-15 18 FG 0.86

92.1 ± 3.1 K-15 19 FG 1.71

92.1 ± 3.1 K-15 20 FG 1.32

107.0 ± 0.2 K-17 1 FG 4.23

107.0 ± 0.2 K-17 2 FG 4.27

107.0 ± 0.2 K-17 3 FG 3.01

107.0 ± 0.2 K-17 4 FG 4.58

107.0 ± 0.2 K-17 5 FG 5.64

107.0 ± 0.2 K-17 6 FG 3.68

107.0 ± 0.2 K-17 7 FG 4.93

107.0 ± 0.2 K-17 8 FG 5.31

107.0 ± 0.2 K-17 9 FG 4.70

107.0 ± 0.2 K-17 10 FG 5.41

107.0 ± 0.2 K-17 11 FG 4.36

107.0 ± 0.2 K-17 12 FG 5.40

107.0 ± 0.2 K-17 13 FG 3.63

107.0 ± 0.2 K-17 14 FG 5.86

107.0 ± 0.2 K-17 15 FG 5.40

107.0 ± 0.2 K-17 16 FG 3.88

107.0 ± 0.2 K-17 17 FG 2.62

107.0 ± 0.2 K-17 18 FG 4.48

107.0 ± 0.2 K-17 19 FG 6.45

107.0 ± 0.2 K-17 20 FG 2.87

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Age (Table 2.1) Specimen n Layer (Fig. 1.1) Needle Width (µm)

107.0 ± 0.2 K-46 1 FG 4.77

107.0 ± 0.2 K-46 2 FG 1.68

107.0 ± 0.2 K-46 3 FG 3.43

107.0 ± 0.2 K-46 4 FG 1.58

107.0 ± 0.2 K-46 5 FG 4.70

107.0 ± 0.2 K-46 6 FG 1.99

107.0 ± 0.2 K-46 7 FG 6.43

107.0 ± 0.2 K-46 8 FG 4.27

107.0 ± 0.2 K-46 9 FG 8.14

107.0 ± 0.2 K-46 10 FG 4.69

107.0 ± 0.2 K-46 11 FG 5.52

107.0 ± 0.2 K-46 12 FG 2.19

107.0 ± 0.2 K-46 13 FG 7.37

107.0 ± 0.2 K-46 14 FG 5.37

107.0 ± 0.2 K-46 15 FG 3.01

107.0 ± 0.2 K-46 16 FG 5.23

107.0 ± 0.2 K-46 17 FG 5.12

107.0 ± 0.2 K-46 18 FG 5.16

107.0 ± 0.2 K-46 19 FG 4.56

107.0 ± 0.2 K-46 20 FG 5.17

119.1 ± 1.9 K-131 1 FG 1.98

119.1 ± 1.9 K-131 2 FG 4.19

119.1 ± 1.9 K-131 3 FG 2.92

119.1 ± 1.9 K-131 4 FG 2.21

119.1 ± 1.9 K-131 5 FG 2.37

119.1 ± 1.9 K-131 6 FG 2.29

119.1 ± 1.9 K-131 7 FG 1.98

119.1 ± 1.9 K-131 8 FG 2.79

119.1 ± 1.9 K-131 9 FG 2.19

119.1 ± 1.9 K-131 10 FG 1.85

119.1 ± 1.9 K-131 11 FG 3.39

119.1 ± 1.9 K-131 12 FG 3.05

119.1 ± 1.9 K-131 13 FG 3.62

119.1 ± 1.9 K-131 14 FG 1.93

119.1 ± 1.9 K-131 15 FG 3.72

119.1 ± 1.9 K-131 16 FG 2.71

119.1 ± 1.9 K-131 17 FG 1.77

119.1 ± 1.9 K-131 18 FG 1.87

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Age (Table 2.1) Specimen n Layer (Fig. 1.1) Needle Width (µm)

119.1 ± 1.9 K-131 19 FG 2.06

119.1 ± 1.9 K-131 20 FG 3.57

134.1 ± 1.4 K-126 1 FG 4.15

134.1 ± 1.4 K-126 2 FG 3.79

134.1 ± 1.4 K-126 3 FG 3.77

134.1 ± 1.4 K-126 4 FG 5.89

134.1 ± 1.4 K-126 5 FG 4.30

134.1 ± 1.4 K-126 6 FG 3.90

134.1 ± 1.4 K-126 7 FG 3.09

134.1 ± 1.4 K-126 8 FG 4.57

134.1 ± 1.4 K-126 9 FG 2.95

134.1 ± 1.4 K-126 10 FG 3.62

134.1 ± 1.4 K-126 11 FG 5.81

134.1 ± 1.4 K-126 12 FG 4.02

134.1 ± 1.4 K-126 13 FG 5.62

134.1 ± 1.4 K-126 14 FG 5.11

134.1 ± 1.4 K-126 15 FG 6.88

134.1 ± 1.4 K-126 16 FG 6.43

134.1 ± 1.4 K-126 17 FG 3.92

134.1 ± 1.4 K-126 18 FG 4.70

134.1 ± 1.4 K-126 19 FG 4.57

134.1 ± 1.4 K-126 20 FG 2.95

134.1 ± 1.4 K-24 1 FG 3.20

134.1 ± 1.4 K-24 2 FG 1.95

134.1 ± 1.4 K-24 3 FG 1.79

134.1 ± 1.4 K-24 4 FG 2.46

134.1 ± 1.4 K-24 5 FG 1.85

134.1 ± 1.4 K-24 6 FG 2.08

134.1 ± 1.4 K-24 7 FG 1.34

134.1 ± 1.4 K-24 8 FG 1.57

134.1 ± 1.4 K-24 9 FG 3.02

134.1 ± 1.4 K-24 10 FG 2.22

134.1 ± 1.4 K-24 11 FG 1.76

134.1 ± 1.4 K-24 12 FG 3.01

134.1 ± 1.4 K-24 13 FG 1.43

134.1 ± 1.4 K-24 14 FG 3.40

134.1 ± 1.4 K-24 15 FG 1.68

134.1 ± 1.4 K-24 16 FG 2.55

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Age (Table 2.1) Specimen n Layer (Fig. 1.1) Needle Width (µm)

134.1 ± 1.4 K-24 17 FG 1.99

134.1 ± 1.4 K-24 18 FG 2.47

134.1 ± 1.4 K-24 19 FG 1.77

134.1 ± 1.4 K-24 20 FG 2.57

198.7 ± 4.6 K-38 1 FG 2.66

198.7 ± 4.6 K-38 2 FG 2.21

198.7 ± 4.6 K-38 3 FG 3.59

198.7 ± 4.6 K-38 4 FG 3.48

198.7 ± 4.6 K-38 5 FG 3.05

198.7 ± 4.6 K-38 6 FG 3.45

198.7 ± 4.6 K-38 7 FG 3.21

198.7 ± 4.6 K-38 8 FG 2.83

198.7 ± 4.6 K-38 9 FG 3.60

198.7 ± 4.6 K-38 10 FG 2.42

198.7 ± 4.6 K-38 11 FG 3.93

198.7 ± 4.6 K-38 12 FG 5.32

198.7 ± 4.6 K-38 13 FG 2.03

198.7 ± 4.6 K-38 14 FG 3.44

198.7 ± 4.6 K-38 15 FG 2.50

198.7 ± 4.6 K-38 16 FG 3.13

198.7 ± 4.6 K-38 17 FG 2.44

198.7 ± 4.6 K-38 18 FG 1.81

198.7 ± 4.6 K-38 19 FG 2.58

198.7 ± 4.6 K-38 20 FG 3.91

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APPENDIX IV: DAILY GROWTH

Low Resolution Daily Band Thickness (µm)

Sample K-133 PT-1

Shell Location FG LG FG LG

1 31.90 17.57 19.99 17.47

2 34.04 17.80 27.43 18.08

3 31.90 20.08 22.28 22.29

4 31.90 25.60 25.71 18.28

5 37.20 12.30 22.86 17.88

6 31.40 14.80 28.57 23.08

7 35.10 22.09 28.57 27.92

8 31.40 24.50 31.43 22.01

9 34.04 23.50 26.86 20.94

10 27.66 33.45 27.43 18.26

Average 32.65 21.17 26.11 20.62

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APPENDIX V: DAYTIME AND NIGHTTIME GROWTH

High Resolution Band Thickness (µm)

Sample K-133

Needle type Daytime Nighttime

1 30.30 10.65

2 46.80 5.15

3 27.50 10.40

4 29.26 13.30

5 33.84 7.69

6 36.97 9.95

7 32.80 13.30

8 30.72 5.68

9 31.60 11.60

10 42.99 15.14

11 35.10 9.10

12 23.60 4.43

Average 33.46 9.70

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APPENDIX VI: HIGH TIDE DATA

Dreger Harbor, Papua New Guinea

(6.6500° S, 147.8667° E)

Lucinda, Australia

(18.5167° S, 146.3333° E)

Date

Time of day of the highest

tide

Date

Time of day of the highest

tide

7-Apr-74 3:35:00 AM

8/7/76 8:00 PM

14-Apr-74 5:02:00 AM

8/14/76 11:25 PM

21-Apr-74 2:54:00 AM

8/21/76 7:14 PM

28-Apr-74 4:34:00 AM

8/28/76 10:34 AM

5-May-74 2:38:00 AM

9/4/76 7:06 PM

12-May-74 4:15:00 AM

9/11/76 10:15 AM

19-May-74 1:51:00 AM

9/18/76 6:18 PM

26-May-74 3:56:00 AM

9/25/76 9:34 AM

2-Jun-74 1:36:00 AM

10/2/76 6:06 PM

9-Jun-74 3:39:00 AM

10/9/76 9:22 AM

16-Jun-74 12:15:00 AM

10/16/76 5:07 PM

23-Jun-74 3:29:00 AM

10/23/76 8:42 AM

30-Jun-74 12:00:00 AM

10/30/76 4:53 PM

7-Jul-74 3:10:00 AM

11/6/76 8:39 AM

14-Jul-74 10:24:00 PM

11/13/76 2:35 PM

21-Jul-74 3:07:00 AM

11/20/76 7:56 AM

28-Jul-74 8:39:00 PM

11/27/76 3:03 PM

4-Aug-74 2:42:00 AM

12/4/76 8:04 AM

11-Aug-74 7:40:00 PM

12/11/76 11:59 AM

18-Aug-74 2:42:00 AM

12/18/76 7:12 AM

25-Aug-74 6:48:00 PM

12/25/76 12:32 PM

1-Sep-74 2:04:00 AM

1/1/77 7:35 AM

8-Sep-74 6:11:00 PM

1/8/77 10:43 AM

15-Sep-74 2:06:00 AM

1/15/77 6:27 AM

22-Sep-74 5:35:00 PM

1/22/77 11:02 AM

29-Sep-74 4:03:00 PM

1/29/77 7:04 AM

6-Oct-74 5:06:00 PM

2/5/77 9:41 AM

13-Oct-74 3:55:00 PM

2/12/77 5:29 AM

20-Oct-74 4:39:00 PM

2/19/77 9:53 AM

27-Oct-74 3:07:00 PM

2/26/77 6:24 AM

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Dreger Harbor, Papua New Guinea

(6.6500° S, 147.8667° E)

Lucinda, Australia

(18.5167° S, 146.3333° E)

Date

Time of day of the highest

tide Date

Time of day of the highest

tide

3-Nov-74 4:14:00 PM

3/5/77 8:42 AM

10-Nov-74 3:03:00 PM

3/12/77 3:53 AM

17-Nov-74 3:53:00 PM

3/19/77 8:53 AM

24-Nov-74 2:18:00 PM

3/26/77 5:01 AM

1-Dec-74 3:33:00 PM

4/2/77 7:43 AM

8-Dec-74 2:28:00 PM

4/9/77 1:25 AM

15-Dec-74 3:18:00 PM

4/16/77 7:57 AM

22-Dec-74 3:22:00 PM

4/23/77 9:22 AM

29-Dec-74 2:27:00 PM

4/30/77 6:40 AM

5-Jan-75 3:29:00 PM

5/7/77 11:48 AM

12-Jan-75 2:23:00 PM

5/14/77 7:41 PM

19-Jan-75 7:31:00 AM

5/21/77 11:32 PM

26-Jan-75 1:56:00 PM

5/28/77 6:21 PM

2-Feb-75 6:08:00 AM

6/4/77 11:39 PM

9-Feb-75 1:54:00 PM

6/11/77 7:02 PM

16-Feb-75 5:39:00 AM

6/18/77 10:28 PM

23-Feb-75 1:04:00 PM

6/25/77 3:53 AM

2-Mar-75 4:55:00 AM

7/2/77 10:29 PM

9-Mar-75 1:48:00 PM

7/9/77 6:21 PM

16-Mar-75 4:30:00 AM

7/16/77 9:29 PM

23-Mar-75 6:08:00 AM

7/23/77 1:42 AM

30-Mar-75 3:55:00 AM

7/30/77 9:26 PM

6-Apr-75 5:35:00 AM

8/6/77 5:27 PM

13-Apr-75 3:30:00 AM

8/13/77 8:32 PM

20-Apr-75 5:18:00 AM

8/20/77 1:03 PM

27-Apr-75 3:01:00 AM

8/27/77 8:26 PM

4-May-75 4:56:00 AM

9/3/77 3:16 PM

11-May-75 2:33:00 AM

9/10/77 7:36 PM

18-May-75 4:51:00 AM

9/17/77 11:28 AM

25-May-75 2:10:00 AM

9/24/77 7:28 PM

1-Jun-75 4:31:00 AM

10/1/77 11:22 AM

8-Jun-75 1:33:00 AM

10/8/77 6:38 PM

15-Jun-75 4:32:00 AM

10/15/77 10:22 AM

22-Jun-75 1:12:00 AM

10/22/77 6:30 PM

29-Jun-75 4:11:00 AM

10/29/77 10:17 AM

6-Jul-75 12:04:00 AM

11/5/77 5:35 PM

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Dreger Harbor, Papua New Guinea

(6.6500° S, 147.8667° E)

Lucinda, Australia

(18.5167° S, 146.3333° E)

Date

Time of day of the highest

tide Date

Time of day of the highest

tide

13-Jul-75 4:13:00 AM

11/12/77 9:28 AM

20-Jul-75 12:00:00 AM

11/19/77 5:26 PM

27-Jul-75 3:48:00 AM

11/26/77 9:28 AM

3-Aug-75 11:52:00 PM

12/3/77 3:55 PM

10-Aug-75 3:50:00 AM

12/10/77 8:40 AM

17-Aug-75 8:40:00 PM

12/17/77 4:01 PM

24-Aug-75 3:20:00 AM

12/24/77 8:47 AM

31-Aug-75 7:45:00 PM

12/31/77 12:43 PM

7-Sep-75 3:24:00 AM

1/7/78 7:54 AM

14-Sep-75 6:36:00 PM

1/14/78 1:40 PM

21-Sep-75 4:20:00 PM

1/21/78 8:08 AM

28-Sep-75 6:15:00 PM

1/28/78 11:12 AM

5-Oct-75 3:57:00 PM

2/4/78 7:07 AM

12-Oct-75 5:34:00 PM

2/11/78 11:40 AM

19-Oct-75 3:21:00 PM

2/18/78 7:28 AM

26-Oct-75 5:22:00 PM

2/25/78 10:06 AM

2-Nov-75 3:02:00 PM

3/4/78 6:12 AM

9-Nov-75 4:51:00 PM

3/11/78 10:23 AM

16-Nov-75 2:25:00 PM

3/18/78 6:39 AM

23-Nov-75 4:44:00 PM

3/25/78 9:06 AM

30-Nov-75 2:57:00 AM

4/1/78 4:57 AM

7-Dec-75 4:20:00 PM

4/8/78 9:21 AM

14-Dec-75 1:23:00 PM

4/15/78 5:27 AM

21-Dec-75 4:14:00 PM

4/22/78 8:40 PM

28-Dec-75 12:00:00 AM

4/29/78 3:02 AM

4-Jan-76 3:55:00 PM

5/6/78 9:04 PM

11-Jan-76 11:23:00 AM

5/13/78 2:46 AM

18-Jan-76 3:17:00 PM

5/20/78 7:53 PM

25-Jan-76 7:28:00 AM

5/27/78 12:59 AM

1-Feb-76 3:34:00 PM

6/3/78 8:22 PM

8-Feb-76 6:56:00 AM

6/10/78 11:14 AM

15-Feb-76 3:22:00 PM

6/17/78 7:07 PM

22-Feb-76 5:50:00 AM

6/24/78 11:36 AM

29-Feb-76 3:11:00 PM

7/1/78 7:42 PM

7-Mar-76 5:32:00 AM

7/8/78 11:11 PM

14-Mar-76 2:53:00 PM

7/15/78 6:19 PM

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Dreger Harbor, Papua New Guinea

(6.6500° S, 147.8667° E)

Lucinda, Australia

(18.5167° S, 146.3333° E)

Date

Time of day of the highest

tide Date

Time of day of the highest

tide

21-Mar-76 4:44:00 AM

7/22/78 11:08 PM

28-Mar-76 4:46:00 AM

7/29/78 7:00 PM

4-Apr-76 4:27:00 AM

8/5/78 10:00 PM

11-Apr-76 4:16:00 AM

8/12/78 5:20 PM

18-Apr-76 3:49:00 AM

8/19/78 9:56 PM

25-Apr-76 4:07:00 AM

8/26/78 6:10 PM

2-May-76 3:32:00 AM

9/2/78 8:56 PM

9-May-76 4:05:00 AM

9/9/78 3:34 PM

16-May-76 3:03:00 AM

9/16/78 8:52 PM

23-May-76 4:04:00 AM

9/23/78 5:01 PM

30-May-76 2:43:00 AM

9/30/78 7:56 PM

6-Jun-76 4:45:00 AM

10/7/78 12:51 PM

13-Jun-76 2:23:00 AM

10/14/78 7:52 PM

20-Jun-76 11:51:00 PM

10/21/78 2:21 PM

27-Jun-76 1:56:00 AM

10/28/78 6:59 PM

4-Jul-76 4:57:00 AM

11/4/78 11:22 AM

11-Jul-76 1:48:00 AM

11/11/78 6:56 PM

18-Jul-76 7:56:00 PM

11/18/78 11:29 AM

25-Jul-76 12:58:00 AM

11/25/78 6:03 PM

1-Aug-76 7:00:00 PM

12/2/78 10:18 AM

8-Aug-76 1:01:00 AM

12/9/78 6:01 PM

15-Aug-76 6:21:00 PM

12/16/78 10:20 AM

22-Aug-76 12:00:00 AM

12/23/78 4:56 PM

29-Aug-76 5:40:00 PM

12/30/78 9:24 AM

5-Sep-76 12:00:00 AM

1/6/79 4:59 PM

12-Sep-76 5:10:00 PM

1/13/79 9:28 AM

19-Sep-76 6:42:00 PM

1/20/79 1:48 PM

26-Sep-76 4:36:00 PM

1/27/79 8:33 AM

3-Oct-76 5:46:00 PM

2/3/79 3:16 PM

10-Oct-76 4:09:00 PM

2/10/79 8:39 AM

17-Oct-76 5:51:00 PM

2/17/79 11:30 AM

24-Oct-76 3:40:00 PM

2/24/79 7:41 AM

31-Oct-76 5:09:00 PM

3/3/79 12:18 AM

7-Nov-76 3:16:00 PM

3/10/79 7:50 AM

14-Nov-76 5:32:00 PM

3/17/79 10:24 AM

21-Nov-76 4:29:00 AM

3/24/79 6:43 AM

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Dreger Harbor, Papua New Guinea

(6.6500° S, 147.8667° E)

Lucinda, Australia

(18.5167° S, 146.3333° E)

Date

Time of day of the highest

tide Date

Time of day of the highest

tide

28-Nov-76 4:53:00 PM

3/31/79 11:57 PM

5-Dec-76 2:09:00 PM

4/7/79 6:56 AM

12-Dec-76 5:14:00 PM

4/14/79 10:12 PM

19-Dec-76 1:47:00 PM

4/21/79 5:35 AM

26-Dec-76 4:41:00 PM

4/28/79 10:47 PM

2-Jan-77 1:18:00 PM

5/5/79 5:50 AM

9-Jan-77 4:51:00 PM

5/12/79 9:26 PM

16-Jan-77 12:55:00 PM

5/19/79 4:02 AM

23-Jan-77 4:25:00 PM

5/26/79 9:54 PM

30-Jan-77 10:15:00 AM

6/2/79 4:11 AM

6-Feb-77 4:25:00 PM

6/9/79 8:40 PM

13-Feb-77 7:16:00 AM

6/16/79 2:02 AM

20-Feb-77 4:05:00 PM

6/23/79 9:06 PM

27-Feb-77 6:52:00 AM

6/30/79 1:00 AM

6-Mar-77 3:58:00 PM

7/7/79 7:53 PM

13-Mar-77 5:45:00 AM

7/14/79 12:18 AM

20-Mar-77 4:19:00 AM

7/21/79 8:19 PM

27-Mar-77 5:32:00 AM

7/28/79 11:45 PM

3-Apr-77 3:42:00 AM

8/4/79 7:05 PM

10-Apr-77 4:47:00 AM

8/11/79 11:46 PM

17-Apr-77 3:25:00 AM

8/18/79 7:31 PM

24-Apr-77 4:35:00 AM

8/25/79 10:24 PM

1-May-77 2:49:00 AM

9/1/79 6:12 PM

8-May-77 4:01:00 AM

9/8/79 10:26 PM

15-May-77 2:36:00 AM

9/15/79 6:38 PM

22-May-77 3:49:00 AM

9/22/79 9:17 PM

29-May-77 1:56:00 AM

9/29/79 5:03 PM

5-Jun-77 3:25:00 AM

10/6/79 9:04 AM

12-Jun-77 1:53:00 AM

10/13/79 5:35 PM

19-Jun-77 3:11:00 AM

10/20/79 8:24 AM

26-Jun-77 12:00:00 AM

10/27/79 2:58 PM

3-Jul-77 2:59:00 AM

11/3/79 8:15 AM

10-Jul-77 11:05:00 PM

11/10/79 4:04 PM

17-Jul-77 2:37:00 AM

11/17/79 7:44 AM

24-Jul-77 9:01:00 PM

11/24/79 12:32 PM

31-Jul-77 2:40:00 AM

12/1/79 7:31 AM

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84

Dreger Harbor, Papua New Guinea

(6.6500° S, 147.8667° E)

Lucinda, Australia

(18.5167° S, 146.3333° E)

Date

Time of day of the highest

tide Date

Time of day of the highest

tide

7-Aug-77 7:53:00 PM

12/8/79 12:58 PM

14-Aug-77 1:59:00 AM

12/15/79 7:12 AM

21-Aug-77 6:53:00 PM

12/22/79 11:08 AM

28-Aug-77 2:15:00 AM

12/29/79 6:48 AM

4-Sep-77 6:19:00 PM

1/5/80 11:12 AM

11-Sep-77 12:56:00 AM

1/12/80 6:45 AM

18-Sep-77 5:37:00 PM

1/19/80 10:05 AM

25-Sep-77 1:29:00 AM

1/26/80 6:00 AM

2-Oct-77 5:12:00 PM

2/2/80 10:03 AM

9-Oct-77 4:16:00 PM

2/9/80 6:19 AM

16-Oct-77 4:38:00 PM

2/16/80 9:07 AM

23-Oct-77 4:18:00 PM

2/23/80 4:50 AM

30-Oct-77 4:17:00 PM

3/1/80 9:05 AM

3/8/80 5:29 AM

3/15/80 8:09 AM

3/22/80 2:29 AM

3/29/80 8:09 AM

4/5/80 10:22 AM

4/12/80 7:09 AM

4/19/80 12:15 AM

4/26/80 7:13 AM

5/3/80 11:13 PM

5/10/80 6:03 AM

5/17/80 10:46 AM

5/24/80 6:13 AM

5/31/80 10:20 PM

6/7/80 4:44 AM

6/14/80 10:50 PM

6/21/80 4:59 AM

6/28/80 9:28 PM

7/5/80 3:00 AM

7/12/80 9:48 PM

7/19/80 2:26 AM

7/26/80 8:35 PM

8/2/80 1:03 AM

8/9/80 8:51 PM

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85

Lucinda, Australia

(18.5167° S, 146.3333° E)

Date

Time of day of the highest

tide

8/16/80 12:29 PM

8/23/80 7:41 PM

8/30/80 12:06 PM

9/6/80 7:56 PM

9/13/80 10:33 PM

9/20/80 6:44 PM

9/27/80 10:48 AM

10/4/80 7:01 PM

10/11/80 9:52 AM

10/18/80 5:41 PM

10/25/80 9:48 AM

11/1/80 6:02 PM

11/8/80 9:07 AM

11/15/80 4:13 PM

11/22/80 8:58 AM

11/29/80 4:48 PM

12/6/80 8:28 AM

12/13/80 1:43 PM

12/20/80 8:12 AM

12/27/80 2:32 PM

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APPENDIX VII: SOLAR IRRADIANCE CALCULATIONS

Palm Island, Great Barrier Reef, Australia

(18.517˚S, 146.3˚E)

Q = QA [(-0.5 • (p/Σp)) + (ΣQ/ΣQA) + 0.054]

Σp: 1147.5 mm

ΣQ: 161.4 ± 85.8 (cal/cm2/day)

ΣQA: 299.5 ± 149.2 (cal/cm2/day)

Monthly

Insolation

(cal/cm2/day)

Monthly

Rainfall

(mm)

Total Solar

Irradiance

(cal/cm2/day)

Month QA p Q

30-Aug-76 708.1 0.9 419.7

30-Sep-76 819.1 3.2 484.6

30-Oct-76 912.0 22.9 531.8

30-Nov-76 972.3 70.6 546.7

30-Dec-76 998.1 317.8 453.7

30-Jan-77 989.5 97.6 544.7

28-Feb-77 946.5 478.4 364.0

30-Mar-77 869.0 271.8 412.5

30-Apr-77 757.2 84.6 421.1

30-May-77 651.3 180.8 335.0

30-Jun-77 597.1 0.2 354.1

30-Jul-77 620.4 0.4 367.8

30-Aug-77 708.1 22.2 413.1

30-Sep-77 819.1 20.4 478.5

30-Oct-77 912.0 2.4 539.9

30-Nov-77 972.3 33.4 562.4

30-Dec-77 998.1 127.6 536.4

30-Jan-78 989.5 436.6 398.6

28-Feb-78 946.5 210.2 474.6

30-Mar-78 869.0 76.4 486.4

30-Apr-78 757.2 115.2 411.0

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Monthly

Insolation

(cal/cm2/day)

Monthly

Rainfall

(mm)

Total Solar

Irradiance

(cal/cm2/day)

Month QA p Q

30-May-78 651.3 37 375.8

30-Jun-78 597.1 2.6 353.4

30-Jul-78 620.4 21.8 362.0

30-Aug-78 708.1 29 411.0

30-Sep-78 819.1 18.4 479.2

30-Oct-78 912.0 13.4 535.6

30-Nov-78 972.3 49 555.8

30-Dec-78 998.1 60.8 565.5

30-Jan-79 989.5 183.6 507.6

28-Feb-79 946.5 217.4 471.6

30-Mar-79 869.0 309.6 398.1

30-Apr-79 757.2 53.2 431.5

30-May-79 651.3 5 384.9

30-Jun-79 597.1 34.4 345.2

30-Jul-79 620.4 9.4 365.4

30-Aug-79 708.1 0.4 419.8

30-Sep-79 819.1 10.8 481.9

30-Oct-79 912.0 15.8 534.6

30-Nov-79 972.3 0.4 576.4

30-Dec-79 998.1 185 511.5

30-Jan-80 989.5 257.4 475.8

29-Feb-80 946.5 94.8 522.2

30-Mar-80 869.0 153.6 457.2

30-Apr-80 757.2 21.4 442.0

15-May-80 651.3 41.1 374.6

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Huon Peninsula, Papua New Guinea

(6˚S, 147.5˚E)

Q = QA [(-0.5 • (p/Σp)) + (ΣQ/ΣQA) + 0.054]

Σp: 2602.69 mm

ΣQ: 173.2 ± 42.5 (cal/cm2/day)

ΣQA: 312.7 ± 63.3 (cal/cm2/day)

Monthly

Insolation

(cal/cm2/day)

Monthly

Rainfall

(mm)

Total Solar

Irradiance

(cal/cm2/day)

Month QA p Q

15-Apr-74 851.0 434.0 446.2

15-May-74 783.8 305.0 430.5

15-Jun-74 746.8 338.0 405.4

15-Jul-74 761.5 84.0 450.5

15-Aug-74 816.5 133.0 475.4

15-Sep-74 869.0 96.0 512.1

15-Oct-74 903.4 265.0 503.1

15-Nov-74 903.4 279.0 500.7

15-Dec-74 903.4 488.0 464.4

15-Jan-75 912.0 383.0 487.2

15-Feb-75 920.7 449.0 480.1

15-Mar-75 903.4 352.0 488.0

15-Apr-75 851.0 308.0 466.8

15-May-75 783.8 306.0 430.3

15-Jun-75 746.8 349.0 403.8

15-Jul-75 761.5 345.0 412.3

15-Aug-75 816.5 443.0 426.8

15-Sep-75 869.0 240.0 488.1

15-Oct-75 903.4 614.0 442.5

15-Nov-75 903.4 233.0 508.6

15-Dec-75 903.4 322.0 493.2

15-Jan-76 912.0 344.0 494.0

15-Feb-76 920.7 231.0 518.7

15-Mar-76 903.4 277.0 501.0

15-Apr-76 851.0 548.0 427.6

15-May-76 783.8 329.0 426.8

15-Jun-76 746.8 126.0 435.8

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Monthly

Insolation

(cal/cm2/day)

Monthly

Rainfall

(mm)

Total Solar

Irradiance

(cal/cm2/day)

Month QA p Q

15-Jul-76 761.5 184.0 435.9

15-Aug-76 816.5 159.0 471.3

15-Sep-76 869.0 22.0 524.5

15-Oct-76 903.4 120.0 528.2

15-Nov-76 903.4 233.0 508.6

15-Dec-76 903.4 322.0 493.2

15-Jan-77 912.0 244.0 511.6

15-Feb-77 920.7 285.0 509.1

15-Mar-77 903.4 356.0 487.3

15-Apr-77 851.0 211.0 482.7

15-May-77 783.8 480.0 404.1

15-Jun-77 746.8 216.0 422.9

15-Jul-77 761.5 58.0 454.3

15-Aug-77 816.5 64.0 486.2

15-Sep-77 869.0 79.0 515.0

15-Oct-77 903.4 215.0 511.8

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APPENDIX VIII: PRECIPITATION AND GROWTH

Sample Date

Growth Thickness (per

29 day interval; µm)

Precipitation

(mm)

K-133 15-Apr-74 1095.8 434.0

K-133 15-May-74 1109.5 305.0

K-133 15-Jun-74 755.1 338.0

K-133 15-Jul-74 816.1 84.0

K-133 15-Aug-74 1151.2 133.0

K-133 15-Sep-74 1559.3 96.0

K-133 15-Oct-74 961.4 265.0

K-133 15-Nov-74 875.8 279.0

K-133 15-Dec-74 665.6 488.0

K-133 15-Jan-75 571.6 383.0

K-133 15-Feb-75 681.1 449.0

K-133 15-Mar-75 1015.4 352.0

K-133 15-Apr-75 820.7 308.0

K-133 15-May-75 801.4 306.0

K-133 15-Jun-75 1029.5 349.0

K-133 15-Jul-75 797.9 345.0

K-133 15-Aug-75 820.7 443.0

K-133 15-Sep-75 835.4 240.0

K-133 15-Oct-75 635.4 614.0

K-133 15-Nov-75 966.7 233.0

K-133 15-Dec-75 862.8 322.0

K-133 15-Jan-76 646.3 344.0

K-133 15-Feb-76 906.3 231.0

K-133 15-Mar-76 691.9 277.0

K-133 15-Apr-76 637.5 548.0

K-133 15-May-76 826.3 329.0

K-133 15-Jun-76 914.4 126.0

K-133 15-Jul-76 1067.0 184.0

K-133 15-Aug-76 917.9 159.0

K-133 15-Sep-76 1037.5 22.0

K-133 15-Oct-76 713.3 120.0

K-133 15-Nov-76 658.2 233.0

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Sample Date

Growth Thickness (per

29 day interval; µm)

Precipitation

(mm)

K-133 15-Dec-76 693.0 322.0

K-133 15-Jan-77 633.7 244.0

K-133 15-Feb-77 850.9 285.0

K-133 15-Mar-77 655.4 356.0

K-133 15-Apr-77 691.2 211.0

K-133 15-May-77 612.3 480.0

K-133 15-Jun-77 496.5 216.0

K-133 15-Jul-77 575.8 58.0

K-133 15-Aug-77 582.8 64.0

K-133 15-Sep-77 656.8 79.0

K-133 15-Oct-77 520.4 215.0

PT-1 30-Aug-76 1348.8 0.9

PT-1 30-Sep-76 1045.7 3.2

PT-1 30-Oct-76 819.7 22.9

PT-1 30-Nov-76 974.5 70.6

PT-1 30-Dec-76 1338.1 317.8

PT-1 30-Jan-77 929.6 97.6

PT-1 28-Feb-77 925.8 478.4

PT-1 30-Mar-77 1484.5 271.8

PT-1 30-Apr-77 1603.0 84.6

PT-1 30-May-77 1355.3 180.8

PT-1 30-Jun-77 1621.6 0.2

PT-1 30-Jul-77 1361.7 0.4

PT-1 30-Aug-77 1205.8 22.2

PT-1 30-Sep-77 994.1 20.4

PT-1 30-Oct-77 833.1 2.4

PT-1 30-Nov-77 502.1 33.4

PT-1 30-Dec-77 968.6 127.6

PT-1 30-Jan-78 1158.1 436.6

PT-1 28-Feb-78 1353.5 210.2

PT-1 30-Mar-78 1051.4 76.4

PT-1 30-Apr-78 1381.6 115.2

PT-1 30-May-78 827.7 37

PT-1 30-Jun-78 867.4 2.6

PT-1 30-Jul-78 1532.9 21.8

PT-1 30-Aug-78 950.3 29

PT-1 30-Sep-78 1245.0 18.4

PT-1 30-Oct-78 1511.6 13.4

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Sample Date

Growth Thickness (per

29 day interval; µm)

Precipitation

(mm)

PT-1 30-Nov-78 1203.4 49

PT-1 30-Dec-78 977.8 60.8

PT-1 30-Jan-79 744.4 183.6

PT-1 28-Feb-79 1228.2 217.4

PT-1 30-Mar-79 1727.7 309.6

PT-1 30-Apr-79 1329.3 53.2

PT-1 30-May-79 1431.8 5

PT-1 30-Jun-79 930.3 34.4

PT-1 30-Jul-79 913.6 9.4

PT-1 30-Aug-79 651.7 0.4

PT-1 30-Sep-79 863.1 10.8

PT-1 30-Oct-79 621.6 15.8

PT-1 30-Nov-79 752.5 0.4

PT-1 30-Dec-79 770.2 185

PT-1 30-Jan-80 767.7 257.4

PT-1 29-Feb-80 660.9 94.8

PT-1 30-Mar-80 662.5 153.6

PT-1 30-Apr-80 764.3 21.4

PT-1 15-May-80 354.5 41.1

Rainfall data from Madang, Papua-New Guinea (IAEA/WMO (2014)

and Orpheus Island, Great Barrier Reef, Australia (Bureau of

Meteorology, Government of Australia (2014).

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APPENDIX VIX: RAMAN SPECTROSCOPY

Raman Scattering (arbitrary units)

Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

76.62 562.00 330.25 611.25 542.25 1408.50 969.00

77.91 673.00 362.50 634.00 618.00 1632.75 1050.00

79.20 739.00 397.50 721.00 660.25 1841.25 1163.50

80.47 797.00 415.00 735.00 715.00 1980.00 1251.25

81.76 825.33 411.75 765.25 767.25 2164.25 1265.50

83.05 843.67 419.50 770.00 778.25 2250.25 1361.75

84.35 886.00 435.25 798.00 811.25 2328.50 1367.75

85.64 866.00 439.75 807.50 793.75 2423.50 1410.00

86.93 880.33 438.25 846.75 852.25 2481.25 1469.25

88.20 891.33 447.00 853.75 836.00 2563.75 1459.75

89.49 895.00 450.50 852.75 873.25 2586.50 1509.50

90.78 920.33 438.50 868.50 922.50 2719.50 1607.00

92.07 945.00 466.25 919.50 937.00 2788.25 1630.50

93.36 1000.33 464.50 934.25 1017.00 2893.75 1679.50

94.63 1053.00 466.00 983.00 1060.25 2996.25 1764.25

95.92 1089.00 497.25 1031.50 1075.50 3085.00 1846.00

97.22 1158.67 527.50 1080.75 1155.25 3234.25 1952.75

98.48 1203.00 539.00 1138.75 1174.00 3360.00 2055.00

99.78 1237.33 567.25 1176.00 1251.75 3525.50 2116.75

101.07 1308.00 601.50 1249.25 1289.25 3692.00 2244.25

102.36 1347.00 586.50 1281.50 1339.75 3851.00 2278.25

103.63 1331.33 622.25 1296.50 1402.25 3967.75 2370.75

104.92 1382.67 636.75 1353.00 1397.75 4015.75 2467.00

106.21 1419.67 648.25 1416.00 1378.50 4070.00 2483.25

107.48 1413.67 611.00 1378.75 1448.50 4247.75 2485.50

108.77 1421.67 615.25 1408.25 1442.75 4242.00 2538.00

110.06 1399.33 615.25 1394.00 1409.00 4242.50 2534.00

111.33 1390.00 610.00 1398.00 1415.50 4237.25 2507.75

112.62 1408.33 637.50 1400.00 1445.75 4273.50 2510.00

113.91 1413.33 621.00 1394.00 1421.25 4271.50 2510.25

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

115.18 1382.67 642.50 1395.25 1403.00 4232.25 2544.50

116.47 1406.00 624.75 1385.75 1434.00 4265.00 2465.00

117.74 1393.67 613.00 1384.50 1371.50 4188.50 2510.50

119.03 1395.33 601.00 1370.00 1412.25 4207.00 2462.25

120.32 1374.00 603.00 1337.50 1409.00 4195.75 2443.00

121.59 1428.33 615.50 1382.00 1405.25 4254.25 2422.25

122.88 1399.00 609.25 1360.75 1429.75 4206.75 2464.00

124.15 1408.67 596.00 1354.75 1403.50 4274.75 2414.00

125.44 1361.67 611.00 1354.75 1393.50 4222.50 2463.25

126.71 1398.67 589.25 1315.75 1413.00 4248.75 2488.00

128.00 1404.33 611.50 1346.75 1429.50 4211.00 2459.00

129.27 1389.33 620.00 1375.00 1412.75 4239.50 2504.75

130.56 1420.33 614.25 1354.75 1444.25 4257.25 2484.50

131.83 1421.67 623.00 1378.50 1488.00 4347.50 2520.50

133.12 1448.33 622.50 1378.00 1478.75 4316.00 2522.00

134.39 1441.33 625.50 1380.25 1524.75 4396.75 2567.25

135.68 1427.00 639.75 1401.25 1585.75 4390.50 2706.75

136.95 1425.00 700.25 1439.75 1617.00 4525.50 2708.75

138.24 1450.67 701.50 1474.50 1680.50 4550.25 2879.00

139.51 1532.33 756.75 1524.75 1767.50 4635.25 2938.50

140.80 1514.67 770.50 1570.75 1836.75 4819.00 3083.25

142.07 1573.33 801.25 1669.25 1931.25 5061.75 3232.75

143.33 1607.00 854.25 1765.50 2056.25 5179.75 3400.25

144.63 1636.33 918.75 1824.50 2208.25 5403.75 3564.50

145.89 1696.33 983.75 1987.75 2372.00 5771.50 3792.00

147.18 1733.00 1061.25 2117.00 2523.00 6009.75 4120.00

148.45 1738.67 1149.25 2232.75 2727.00 6323.00 4429.00

149.72 1844.00 1256.00 2495.00 2966.75 6685.50 4766.00

151.01 1901.33 1355.50 2708.75 3099.25 6961.25 5018.50

152.28 1929.67 1417.50 2833.25 3003.50 7138.00 5176.25

153.55 1897.00 1347.00 2872.25 2910.00 7129.00 5189.00

154.84 1832.67 1211.50 2798.00 2642.75 7050.50 4918.50

156.10 1718.33 1064.75 2663.50 2418.25 6790.25 4473.25

157.37 1632.00 970.25 2387.75 2147.25 6486.50 4185.00

158.66 1589.33 921.50 2187.75 2015.00 6033.25 3862.25

159.93 1554.33 851.25 1997.75 1869.25 5638.00 3584.50

161.20 1520.00 810.75 1798.00 1776.75 5336.25 3369.00

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

162.49 1518.00 740.50 1664.75 1701.00 5081.00 3108.75

163.76 1484.00 697.75 1555.25 1601.50 4856.50 2935.50

165.02 1422.00 674.25 1471.00 1593.75 4729.25 2767.00

166.29 1433.33 595.75 1417.25 1497.00 4590.50 2599.25

167.58 1441.67 619.75 1355.00 1492.25 4544.50 2550.75

168.85 1472.33 582.50 1313.50 1488.25 4453.00 2471.50

170.12 1431.00 582.75 1297.50 1496.50 4385.75 2470.25

171.39 1428.33 565.25 1288.25 1487.75 4394.00 2469.00

172.68 1420.33 574.00 1277.75 1500.50 4410.75 2472.00

173.94 1446.33 582.50 1303.50 1509.75 4390.25 2438.50

175.21 1449.67 580.00 1278.75 1584.00 4431.25 2555.75

176.48 1507.67 603.25 1297.00 1631.50 4522.75 2566.50

177.75 1478.00 648.75 1374.25 1596.25 4616.25 2642.25

179.01 1493.67 623.50 1352.00 1627.00 4630.25 2622.75

180.31 1516.33 614.00 1389.50 1658.75 4596.75 2674.25

181.57 1478.00 608.00 1385.50 1604.50 4586.00 2644.25

182.84 1472.33 596.25 1361.50 1569.00 4589.75 2572.50

184.11 1484.00 566.50 1336.00 1571.75 4630.50 2601.75

185.37 1450.33 592.75 1343.00 1591.75 4515.00 2567.75

186.64 1474.00 580.50 1330.50 1573.75 4556.75 2495.25

187.91 1502.33 572.00 1312.00 1537.00 4497.50 2528.25

189.20 1505.67 606.50 1289.50 1556.00 4513.25 2505.00

190.47 1426.33 588.75 1284.00 1575.50 4586.00 2541.75

191.74 1507.00 574.00 1302.00 1585.75 4506.00 2517.50

193.00 1481.67 574.25 1305.25 1597.00 4505.50 2546.75

194.27 1444.00 572.00 1277.00 1579.00 4519.00 2520.50

195.54 1443.00 575.75 1316.75 1592.50 4511.25 2542.00

196.80 1474.67 572.00 1324.00 1637.25 4605.25 2597.50

198.07 1490.33 584.25 1360.50 1700.25 4700.50 2643.25

199.34 1470.00 638.75 1396.25 1791.50 4782.50 2782.75

200.61 1524.00 681.00 1481.75 1892.50 4896.00 2947.75

201.87 1559.67 723.00 1568.75 2017.00 5114.25 3087.50

203.14 1623.00 821.00 1678.50 2163.00 5380.00 3316.00

204.41 1686.67 909.50 1842.50 2269.75 5517.75 3577.25

205.67 1733.67 934.25 2007.75 2298.25 5726.00 3750.25

206.94 1724.00 934.75 2069.50 2162.75 5711.50 3745.50

208.21 1662.00 805.50 2080.00 2102.75 5742.75 3579.25

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

209.48 1561.33 734.00 1963.75 1986.25 5603.25 3422.50

210.74 1539.67 683.50 1753.25 1862.25 5412.00 3159.25

212.01 1521.00 659.25 1660.25 1750.75 5213.50 2993.50

213.28 1563.00 660.00 1522.00 1717.50 4992.25 2794.75

214.54 1541.33 641.25 1456.00 1674.50 4861.50 2712.25

215.81 1501.00 602.75 1390.75 1585.25 4761.50 2650.75

217.08 1482.00 608.75 1337.25 1587.50 4624.25 2546.50

218.32 1546.33 599.00 1313.50 1520.50 4550.25 2487.00

219.59 1522.67 585.25 1273.75 1474.00 4511.00 2458.00

220.86 1502.00 558.00 1252.00 1472.75 4436.75 2401.75

222.12 1451.00 556.25 1222.75 1480.25 4305.25 2357.25

223.39 1489.67 554.50 1213.25 1404.50 4256.25 2348.75

224.66 1456.33 547.50 1172.75 1441.00 4325.75 2282.75

225.92 1444.67 536.00 1157.25 1407.25 4262.50 2265.75

227.19 1461.00 544.25 1148.75 1413.00 4280.75 2249.50

228.43 1433.33 521.25 1084.75 1420.75 4171.75 2280.75

229.70 1463.33 551.00 1117.50 1402.00 4128.50 2236.25

230.97 1472.33 517.25 1094.25 1364.50 4171.00 2216.50

232.24 1459.67 524.50 1129.00 1419.00 4094.25 2214.00

233.50 1466.00 505.25 1143.50 1394.00 4148.00 2209.00

234.75 1419.67 506.00 1079.75 1430.25 4180.25 2236.50

236.01 1458.67 509.25 1089.00 1389.50 4150.25 2234.25

237.28 1468.67 499.75 1101.00 1426.00 4127.75 2209.50

238.55 1444.67 518.75 1095.50 1410.75 4099.00 2236.50

239.79 1451.33 509.25 1081.00 1386.25 4178.25 2263.25

241.06 1439.67 506.75 1092.75 1429.00 4146.00 2196.00

242.32 1487.00 505.75 1097.25 1402.00 4208.75 2294.50

243.59 1443.33 509.50 1114.75 1434.00 4247.75 2259.25

244.83 1447.33 524.75 1121.75 1460.50 4213.25 2279.00

246.10 1452.33 511.00 1145.75 1449.50 4223.50 2317.50

247.37 1458.00 511.50 1138.50 1456.25 4218.75 2313.25

248.64 1416.67 489.00 1129.00 1462.50 4247.75 2356.50

249.88 1423.67 519.25 1129.50 1460.25 4250.75 2256.25

251.14 1417.67 499.50 1150.75 1415.50 4273.50 2315.50

252.39 1462.33 484.50 1153.50 1435.25 4199.00 2276.00

253.65 1454.67 501.50 1138.50 1438.75 4276.00 2282.75

254.92 1444.33 492.50 1128.00 1457.75 4178.25 2283.00

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

256.16 1425.00 487.25 1157.00 1463.50 4191.25 2262.75

257.43 1449.00 499.50 1133.75 1483.00 4224.00 2304.75

258.70 1446.67 513.50 1137.25 1457.50 4249.75 2314.00

259.94 1446.33 503.25 1145.00 1453.25 4248.00 2311.25

261.21 1438.67 495.00 1153.25 1460.00 4286.00 2328.00

262.48 1447.00 489.00 1135.00 1459.50 4288.50 2290.50

263.72 1430.33 480.00 1157.00 1480.25 4307.25 2274.50

264.98 1443.00 503.00 1113.00 1427.50 4252.00 2289.75

266.23 1433.33 495.75 1096.50 1473.25 4248.50 2283.50

267.49 1412.00 489.50 1136.25 1431.75 4170.25 2236.00

268.74 1486.67 479.25 1111.75 1455.00 4242.75 2296.50

270.00 1461.33 503.25 1117.00 1483.25 4222.25 2257.75

271.25 1497.33 502.25 1120.75 1473.50 4218.25 2274.25

272.51 1460.33 485.00 1156.75 1439.50 4229.00 2258.75

273.78 1425.67 495.50 1134.75 1448.00 4234.50 2284.75

275.02 1471.00 495.00 1114.25 1471.00 4254.25 2261.25

276.29 1460.33 469.00 1098.00 1437.75 4257.00 2241.75

277.53 1441.00 488.00 1107.75 1458.00 4249.25 2253.25

278.77 1488.33 488.00 1118.75 1438.25 4254.50 2295.75

280.04 1455.33 505.25 1105.00 1424.75 4253.75 2279.00

281.28 1491.67 496.25 1108.50 1441.00 4228.00 2293.25

282.55 1468.00 493.50 1099.50 1448.75 4240.50 2263.75

283.79 1469.33 503.50 1104.25 1406.75 4203.50 2237.75

285.06 1428.33 482.25 1078.25 1435.50 4186.25 2237.00

286.30 1454.33 474.25 1109.00 1425.00 4233.25 2249.25

287.57 1434.67 471.00 1114.50 1409.00 4182.75 2241.00

288.81 1424.33 459.25 1098.75 1399.75 4177.25 2220.00

290.06 1439.33 449.00 1064.50 1423.25 4192.25 2177.75

291.32 1465.00 452.25 1064.75 1425.50 4146.50 2214.00

292.57 1403.67 455.50 1055.00 1394.50 4093.25 2136.75

293.83 1447.33 425.25 1049.50 1381.25 4085.00 2209.25

295.07 1455.67 429.25 1032.00 1431.00 4077.75 2159.25

296.32 1438.00 433.50 1067.25 1396.75 4096.00 2128.25

297.58 1435.67 443.00 1023.75 1367.50 4102.50 2131.00

298.83 1393.00 455.25 1021.25 1373.25 4069.00 2174.25

300.07 1447.33 427.50 1064.00 1377.50 4079.75 2159.50

301.34 1405.00 446.50 1063.00 1400.25 4125.00 2157.25

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

302.58 1406.33 410.75 1018.00 1400.50 4083.50 2110.75

303.82 1397.67 431.75 1002.75 1379.50 4054.75 2135.00

305.09 1411.33 425.00 1034.00 1353.50 4067.25 2160.25

306.33 1407.00 420.00 1029.75 1379.50 4065.50 2149.25

307.57 1404.33 427.75 1026.50 1397.50 4111.50 2118.50

308.84 1410.33 417.00 1045.00 1407.50 4119.50 2147.00

310.08 1392.00 424.50 1028.50 1400.00 4044.75 2152.75

311.32 1432.33 431.75 1014.00 1400.75 4086.25 2134.75

312.57 1386.67 416.25 1037.50 1379.50 4108.00 2165.75

313.83 1399.67 427.25 1000.50 1371.50 4115.75 2127.00

315.08 1392.33 419.00 1036.50 1403.00 4096.00 2102.50

316.32 1410.33 414.25 1013.75 1378.25 4045.25 2114.50

317.56 1399.00 432.50 999.50 1399.75 4053.25 2111.25

318.80 1391.67 398.75 1006.50 1369.50 4029.25 2153.25

320.07 1421.67 419.50 1030.75 1374.50 4029.25 2120.00

321.31 1447.33 410.25 1060.75 1386.00 4039.25 2138.50

322.56 1419.67 416.00 1035.75 1409.25 4008.25 2147.00

323.80 1453.00 410.25 1016.25 1379.00 4078.75 2152.50

325.04 1366.67 411.00 1006.75 1357.00 4113.25 2124.75

326.31 1414.67 418.50 1012.50 1411.50 4083.25 2163.25

327.55 1411.33 397.50 1019.75 1398.50 4088.25 2118.25

328.79 1373.67 413.50 1037.50 1362.25 4072.25 2110.25

330.03 1407.67 414.75 1042.75 1386.00 4042.25 2113.00

331.28 1395.00 410.75 997.25 1405.25 4073.75 2113.00

332.52 1416.33 418.25 1014.75 1412.25 4089.50 2102.50

333.76 1400.33 429.00 1022.50 1357.25 4046.25 2159.00

335.00 1434.33 405.75 1015.50 1401.50 4056.25 2155.00

336.27 1385.00 408.00 1006.00 1368.25 4087.25 2143.00

337.51 1435.67 406.00 1016.75 1375.75 4077.75 2129.00

338.76 1429.67 408.75 1024.75 1396.75 4050.25 2126.25

340.00 1402.33 406.25 1001.50 1363.25 4052.50 2121.75

341.24 1408.00 417.25 1010.00 1377.50 4064.25 2153.50

342.48 1381.00 418.50 1004.25 1388.00 4084.50 2124.25

343.73 1433.00 405.25 1010.00 1397.50 4028.75 2112.25

344.97 1427.00 409.25 998.00 1372.75 4029.25 2091.00

346.21 1402.67 434.50 1027.25 1385.25 4048.50 2104.00

347.45 1377.67 398.00 998.00 1388.25 4146.00 2119.25

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

348.69 1363.00 409.50 1012.50 1385.00 3978.50 2133.25

349.94 1367.67 423.50 1019.50 1389.75 4010.25 2142.00

351.18 1419.67 427.75 985.00 1382.25 4025.75 2133.00

352.42 1414.33 404.75 1027.00 1386.00 4096.00 2127.50

353.66 1375.00 422.50 1004.50 1365.00 4064.50 2106.00

354.91 1437.67 408.00 989.25 1386.25 4062.00 2140.00

356.15 1387.33 395.00 1002.50 1408.25 4070.25 2121.50

357.39 1416.67 412.50 993.75 1374.75 4105.00 2120.50

358.63 1395.33 404.25 994.50 1373.00 4047.25 2083.00

359.88 1418.33 415.75 1005.50 1381.00 3980.75 2134.75

361.12 1452.33 420.50 1000.25 1385.75 4032.75 2113.75

362.34 1408.67 428.00 1014.25 1412.75 4090.00 2121.75

363.58 1401.00 408.75 1030.00 1414.25 4040.50 2058.00

364.82 1370.33 402.00 1013.50 1397.50 4010.00 2140.25

366.06 1391.33 409.25 1033.75 1408.75 4062.75 2129.75

367.31 1447.00 400.50 1001.00 1393.00 4055.00 2128.25

368.55 1387.67 413.00 992.50 1406.00 4062.50 2125.00

369.79 1411.33 419.50 1021.50 1412.25 4103.00 2123.75

371.03 1463.00 428.50 979.75 1397.50 4056.25 2109.25

372.25 1418.33 418.50 1009.00 1364.00 4050.75 2134.75

373.49 1414.33 416.50 1028.50 1383.50 4086.75 2098.00

374.73 1399.00 412.75 1004.25 1429.25 4133.25 2113.75

375.98 1418.67 396.50 1002.75 1430.50 4081.00 2127.75

377.22 1423.00 423.50 1030.50 1392.75 4056.00 2128.25

378.46 1411.00 399.75 1019.75 1376.00 4092.50 2101.00

379.68 1453.33 426.75 1036.50 1383.25 4047.75 2114.75

380.92 1412.00 431.00 1014.75 1394.00 4059.00 2136.75

382.16 1411.67 416.00 1024.00 1421.25 4108.00 2158.25

383.41 1402.00 406.75 1006.75 1409.50 4049.25 2145.00

384.62 1394.67 402.50 1009.25 1366.50 4065.25 2125.25

385.87 1454.33 434.25 1020.75 1344.25 3998.75 2107.25

387.11 1445.67 413.25 1021.75 1399.75 4122.50 2172.25

388.35 1398.33 406.75 1029.00 1387.75 4124.50 2162.50

389.57 1455.67 419.25 1000.00 1389.25 4072.00 2121.50

390.81 1417.33 409.25 1045.00 1420.75 4077.00 2146.75

392.05 1430.67 409.50 1018.50 1384.00 4048.50 2127.50

393.30 1413.33 398.25 998.75 1410.50 4089.75 2144.25

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

394.51 1414.67 415.00 1035.00 1401.50 4081.50 2108.75

395.76 1439.67 411.50 1020.00 1422.25 4119.50 2100.75

397.00 1421.67 414.50 1005.25 1401.00 4082.50 2138.75

398.22 1395.33 406.50 1012.00 1388.25 4075.50 2120.00

399.46 1404.67 406.50 999.25 1410.00 4044.50 2144.25

400.70 1395.33 410.00 988.00 1419.00 4038.00 2130.25

401.92 1419.00 411.75 996.50 1418.00 4016.25 2110.25

403.16 1402.33 410.25 1010.00 1390.75 4119.75 2132.75

404.40 1394.67 419.50 1021.75 1411.75 4048.50 2131.00

405.62 1399.00 410.50 1018.75 1382.00 4058.50 2131.75

406.86 1392.00 416.50 1006.00 1419.50 4074.25 2143.25

408.08 1433.33 415.75 991.00 1417.00 4058.25 2117.50

409.32 1412.00 402.25 1035.25 1432.75 4052.50 2089.75

410.54 1395.00 417.25 1020.75 1386.75 4052.50 2133.00

411.78 1445.33 417.75 1009.25 1409.75 4047.75 2118.75

413.02 1418.67 411.75 996.75 1373.00 4077.25 2133.00

414.24 1395.67 409.50 1010.00 1380.50 4085.25 2082.25

415.48 1429.67 410.75 1014.75 1410.00 4077.25 2093.75

416.70 1442.33 403.00 1012.25 1413.00 4028.50 2149.25

417.94 1410.33 415.75 997.75 1382.00 4091.00 2103.25

419.16 1417.67 415.50 1002.00 1392.50 4121.75 2144.00

420.40 1438.00 411.25 1022.75 1389.00 4056.75 2107.25

421.62 1381.33 393.25 1024.00 1408.00 4087.75 2097.50

422.86 1428.33 401.00 1003.00 1399.50 4091.00 2114.75

424.08 1373.67 414.00 1005.25 1411.50 4107.50 2110.75

425.32 1399.00 409.75 1015.50 1389.00 4061.75 2146.75

426.54 1410.33 404.00 1027.00 1382.00 4043.00 2134.75

427.78 1405.00 428.75 997.50 1406.50 3986.75 2163.00

429.00 1393.00 411.00 1008.75 1395.75 4053.50 2118.75

430.24 1395.00 399.25 988.00 1423.50 4037.00 2153.75

431.46 1414.33 418.00 1047.75 1425.50 3973.75 2122.25

432.70 1388.67 409.50 999.75 1385.75 3998.00 2132.75

433.92 1393.33 416.25 1013.25 1385.75 4014.75 2144.00

435.14 1399.00 406.25 1001.25 1419.50 4059.25 2116.75

436.38 1432.00 412.50 1022.00 1393.25 4037.75 2092.75

437.60 1404.00 418.50 964.50 1407.00 4029.50 2113.00

438.84 1406.00 404.00 1029.75 1375.00 4081.00 2105.75

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

440.06 1400.00 402.75 1013.50 1370.75 4044.00 2123.25

441.28 1408.33 425.00 1024.50 1388.75 4053.50 2099.75

442.52 1411.67 400.50 984.25 1367.50 4042.25 2103.25

443.74 1403.67 387.25 991.50 1392.25 4051.25 2131.00

444.95 1378.00 413.00 1008.00 1425.25 4034.50 2091.75

446.20 1434.00 401.25 989.25 1399.00 4078.75 2096.50

447.41 1412.33 405.00 982.25 1370.50 4035.50 2148.25

448.63 1408.00 398.50 1005.75 1403.75 4049.00 2073.75

449.87 1411.67 403.50 1010.50 1401.00 4024.00 2098.00

451.09 1415.67 417.00 990.75 1389.00 4060.00 2123.75

452.31 1379.67 407.50 1012.00 1412.75 4068.25 2124.25

453.55 1424.67 415.00 1011.25 1386.75 3994.00 2055.00

454.77 1392.00 423.50 1002.75 1396.50 4075.25 2112.25

455.98 1400.00 400.75 1018.25 1403.25 4024.75 2085.75

457.23 1350.00 418.75 990.50 1423.50 4026.50 2124.25

458.44 1421.33 403.25 1009.50 1415.25 4028.00 2087.75

459.66 1430.33 403.75 1018.00 1402.50 4048.25 2117.25

460.88 1415.33 410.75 996.75 1416.50 4050.75 2144.50

462.10 1411.33 418.75 994.50 1417.25 4046.25 2090.25

463.34 1419.67 393.00 1003.25 1421.25 4006.00 2099.00

464.56 1399.00 418.00 1009.75 1408.50 4029.50 2093.00

465.77 1401.00 408.50 996.50 1428.50 3989.25 2112.25

466.99 1389.33 418.25 1000.00 1402.50 4023.00 2069.25

468.23 1417.00 418.00 1006.75 1434.75 4040.50 2125.00

469.45 1401.00 397.75 1011.25 1384.25 4085.00 2125.75

470.67 1405.67 407.00 1018.50 1406.75 3998.00 2137.00

471.89 1420.00 407.25 994.25 1391.25 4074.75 2106.50

473.10 1393.00 410.25 994.25 1391.75 4027.00 2099.00

474.32 1428.00 420.75 1000.25 1412.25 4050.75 2117.75

475.56 1396.00 408.75 1025.50 1408.75 4076.50 2092.50

476.78 1387.67 417.00 982.00 1425.75 4077.25 2134.00

478.00 1396.00 406.75 992.00 1396.00 4039.00 2117.25

479.22 1401.00 419.50 1005.75 1373.00 3999.00 2136.25

480.43 1421.00 407.75 1029.25 1404.00 4004.75 2129.75

481.65 1419.67 394.25 1010.50 1435.00 4044.00 2125.75

482.87 1421.33 398.25 1022.75 1404.00 4057.25 2120.50

484.09 1424.67 433.75 1002.50 1399.75 4012.25 2128.25

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

485.30 1405.67 418.75 1014.75 1427.25 3996.75 2090.75

486.52 1424.67 407.25 1005.50 1391.75 4062.50 2085.50

487.74 1396.00 400.25 1009.50 1433.75 3972.00 2093.25

488.98 1416.33 406.50 1000.25 1373.00 4038.75 2121.50

490.20 1420.33 412.00 1004.75 1404.25 4017.50 2098.50

491.42 1425.00 396.25 1013.00 1437.00 4069.50 2064.00

492.63 1389.67 406.25 991.75 1410.75 4016.25 2116.25

493.85 1400.00 389.25 995.75 1427.00 4046.75 2120.50

495.07 1425.67 408.00 977.50 1397.75 4062.00 2128.50

496.29 1396.33 396.75 991.50 1397.75 3995.50 2127.00

497.50 1380.33 407.75 1008.50 1385.50 4013.25 2103.75

498.72 1417.00 391.75 1003.75 1390.50 3985.75 2110.75

499.94 1421.33 403.50 998.75 1409.25 4071.25 2091.50

501.16 1378.33 391.50 976.50 1398.75 4013.75 2095.00

502.37 1424.67 407.50 1000.00 1408.50 4037.75 2075.75

503.57 1392.00 383.25 971.00 1430.75 4064.25 2110.25

504.78 1407.00 390.75 978.50 1379.50 3929.75 2076.50

506.00 1415.00 407.75 966.50 1375.50 4039.50 2105.25

507.22 1398.33 406.00 983.75 1404.25 3995.25 2086.25

508.44 1379.67 382.00 959.50 1380.25 4010.00 2072.25

509.65 1416.67 388.75 980.75 1384.75 3964.75 2106.00

510.87 1389.33 393.00 945.25 1397.50 4001.50 2044.75

512.09 1388.67 387.50 982.00 1406.75 4012.50 2060.25

513.31 1371.67 393.25 971.25 1364.75 3935.50 2061.50

514.52 1436.67 394.00 993.75 1401.50 3975.50 2037.50

515.74 1389.00 392.00 983.50 1367.75 3936.50 2067.75

516.94 1351.00 409.00 967.25 1402.75 4000.25 2123.75

518.15 1390.33 401.25 960.50 1402.00 4003.25 2084.75

519.37 1370.67 404.00 978.75 1389.50 3979.75 2064.25

520.59 1386.33 406.75 974.00 1393.75 3961.25 2080.50

521.81 1441.33 389.25 946.50 1364.50 3895.25 2060.75

523.02 1398.67 394.25 973.50 1371.50 3967.75 2070.25

524.22 1412.67 386.25 963.25 1353.75 4013.75 2013.75

525.43 1393.33 391.75 963.00 1359.50 3968.00 2041.75

526.65 1383.67 389.50 974.25 1390.25 3964.25 2035.25

527.87 1407.00 398.25 956.25 1367.75 3941.00 2068.75

529.09 1346.33 402.25 957.50 1392.50 3997.00 2065.25

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

530.28 1378.67 391.75 969.25 1360.00 3999.75 2058.25

531.50 1345.00 370.75 962.50 1394.50 3931.00 2051.00

532.71 1385.00 374.75 949.00 1356.75 3964.50 2062.75

533.93 1387.33 391.25 948.50 1322.25 3883.50 2058.25

535.12 1406.67 388.25 966.25 1370.75 3914.25 2051.25

536.34 1402.00 403.25 961.75 1344.75 3932.50 2019.25

537.56 1384.33 390.25 929.50 1363.25 3905.00 2032.75

538.75 1360.67 402.00 962.50 1376.25 3873.50 2037.25

539.97 1393.33 394.50 970.50 1378.50 3891.50 2020.75

541.19 1383.33 379.25 977.25 1356.25 3947.75 2029.50

542.40 1361.67 398.00 966.75 1341.75 3949.25 2076.50

543.60 1358.33 399.25 952.75 1373.50 3906.50 2013.50

544.81 1373.00 376.75 944.25 1389.50 3897.50 2037.50

546.03 1357.67 400.25 945.00 1364.75 3872.00 2047.50

547.22 1394.00 374.25 948.75 1387.75 3955.50 2016.25

548.44 1369.00 387.25 946.50 1383.50 3817.00 2028.50

549.66 1381.33 391.00 962.25 1388.00 3956.00 2072.75

550.85 1385.67 404.25 934.00 1378.75 3942.00 2066.50

552.07 1390.00 397.25 957.25 1391.00 3893.50 2055.50

553.26 1367.67 377.25 920.00 1355.00 3885.75 2069.50

554.48 1400.33 394.50 950.00 1410.25 3923.00 2027.00

555.70 1354.00 377.75 945.50 1359.00 3890.75 2024.00

556.89 1363.33 377.00 963.50 1379.00 3894.00 2041.50

558.11 1367.33 396.50 959.00 1361.00 3871.25 1993.00

559.30 1371.67 376.50 941.00 1404.50 3880.00 2039.00

560.52 1378.67 398.75 937.00 1372.25 3934.75 2020.00

561.71 1362.33 386.00 928.00 1355.50 3901.25 2017.50

562.93 1398.67 384.25 931.50 1384.75 3875.50 2009.75

564.14 1374.67 367.00 962.50 1373.25 3896.25 2043.50

565.34 1357.67 387.00 964.00 1373.00 3874.00 2062.25

566.55 1377.67 402.75 936.75 1383.25 3867.00 2013.75

567.75 1381.33 398.00 965.75 1376.00 3970.25 2032.00

568.96 1377.33 390.25 946.00 1378.50 3890.75 2023.00

570.16 1392.00 383.50 937.25 1372.50 3896.50 2016.75

571.38 1388.00 377.25 961.25 1357.25 3866.50 2021.75

572.57 1357.00 392.00 949.25 1393.25 3911.00 2015.00

573.79 1405.00 401.25 939.50 1367.00 3937.00 2069.50

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

574.98 1390.00 387.25 959.50 1408.75 3935.25 2003.25

576.20 1360.67 390.50 964.75 1389.25 3915.25 2016.25

577.39 1403.33 389.00 950.50 1388.75 3882.75 2031.50

578.58 1399.67 387.50 942.25 1391.00 3948.50 2005.00

579.80 1365.00 379.50 923.50 1361.50 3913.25 2034.00

580.99 1381.33 383.75 973.75 1373.75 3915.25 2023.50

582.21 1420.33 398.50 940.00 1363.00 3925.75 2018.75

583.40 1381.00 401.75 955.75 1410.75 3930.50 2039.25

584.62 1378.33 388.25 928.50 1405.50 3896.25 1991.50

585.81 1351.33 405.75 931.75 1398.00 3859.50 2012.50

587.00 1361.33 407.75 930.75 1367.25 3915.75 1994.25

588.22 1389.67 396.25 939.25 1396.25 3935.25 2047.00

589.41 1416.67 395.00 955.75 1372.50 3918.50 2033.50

590.61 1418.00 391.50 969.50 1401.50 3949.25 2005.50

591.82 1413.00 389.50 959.50 1390.50 3938.00 2043.50

593.02 1408.00 385.25 918.00 1365.00 3915.75 2038.00

594.21 1354.33 392.00 920.75 1390.75 3923.00 2067.50

595.43 1427.00 406.50 958.50 1381.25 3951.50 2013.75

596.62 1423.33 377.50 943.25 1391.00 4008.00 2078.00

597.81 1398.67 379.25 938.50 1400.75 3963.75 2038.75

599.03 1393.33 393.50 953.75 1411.75 3935.00 2028.75

600.22 1370.67 385.00 946.25 1378.00 3927.00 2073.00

601.42 1388.33 386.00 947.00 1379.25 3947.00 1992.25

602.63 1381.67 399.00 952.25 1377.75 3932.50 1997.25

603.83 1413.33 388.50 937.25 1371.50 4004.25 2038.00

605.02 1414.33 387.25 936.75 1393.50 3886.00 2053.25

606.21 1421.00 369.75 952.75 1381.25 3941.75 2055.25

607.43 1385.00 395.75 957.75 1400.25 3892.75 2082.00

608.62 1392.33 381.00 965.25 1395.75 3929.50 2030.50

609.81 1393.33 375.75 934.00 1433.75 3889.50 2064.00

611.01 1417.33 383.00 956.25 1399.25 3879.25 2038.00

612.22 1413.33 375.50 956.25 1396.50 3884.50 2015.25

613.42 1402.00 400.00 937.25 1351.25 3920.75 2011.50

614.61 1433.33 383.25 927.50 1383.50 3880.50 2029.50

615.80 1404.00 372.75 932.75 1408.00 3933.00 2029.75

616.99 1382.33 401.25 949.75 1396.25 3902.50 1992.50

618.21 1395.33 375.25 937.50 1393.25 3921.00 2049.75

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

619.40 1452.67 381.50 926.75 1396.75 3912.75 2027.25

620.60 1394.67 398.25 937.75 1386.50 3851.00 2044.00

621.79 1354.67 387.25 940.00 1375.50 3879.00 2047.00

622.98 1383.67 386.25 939.75 1365.75 3911.00 2064.25

624.18 1404.33 399.75 939.00 1381.50 3939.25 2033.25

625.37 1430.33 391.00 934.00 1377.00 3906.75 2029.00

626.59 1388.00 388.75 957.75 1353.00 3883.50 2038.25

627.78 1425.33 393.00 962.00 1410.50 3929.00 2073.00

628.97 1413.67 372.50 914.00 1395.75 3933.75 2015.75

630.16 1368.00 385.75 949.25 1393.00 3894.00 2057.00

631.36 1422.00 368.00 943.50 1369.75 3886.50 1995.00

632.55 1374.00 378.00 937.75 1361.75 3903.25 2017.25

633.74 1369.00 384.00 939.75 1404.50 3856.25 1998.75

634.93 1457.33 379.00 950.75 1385.25 3902.00 1954.25

636.13 1389.67 393.00 920.75 1408.50 3854.00 2004.50

637.32 1383.67 374.25 926.50 1393.25 3897.00 2023.00

638.51 1410.00 380.50 931.25 1378.00 3886.25 2052.00

639.70 1390.33 388.75 925.25 1391.00 3820.00 1994.75

640.90 1408.33 395.00 916.50 1399.75 3865.50 2009.75

642.09 1413.67 393.00 927.75 1355.00 3911.75 2054.00

643.28 1417.00 380.50 925.00 1373.00 3915.00 2009.75

644.48 1370.33 384.50 927.00 1356.50 3866.75 2013.00

645.67 1406.33 381.75 950.50 1359.25 3906.75 2062.50

646.86 1400.67 368.50 939.25 1381.75 3871.00 2023.00

648.05 1372.67 386.75 948.25 1390.00 3915.00 2039.25

649.25 1379.67 386.75 928.00 1387.50 3911.00 2029.75

650.44 1370.67 380.25 959.25 1388.75 3870.25 1994.75

651.63 1431.33 388.50 951.75 1365.00 3945.00 2045.25

652.82 1418.00 377.00 929.75 1399.25 3836.50 2003.50

654.02 1400.00 386.25 932.50 1354.25 3838.00 2053.25

655.21 1406.67 384.25 950.75 1342.75 3843.25 2039.25

656.40 1428.67 376.50 928.00 1394.00 3843.50 1988.00

657.59 1397.67 378.75 922.50 1373.50 3878.50 1959.50

658.79 1396.00 373.75 940.50 1363.00 3797.75 2022.75

659.98 1401.67 365.00 928.75 1352.50 3803.50 2027.00

661.17 1411.33 387.00 940.00 1370.50 3850.25 2010.75

662.34 1400.67 370.50 941.75 1351.00 3863.50 1999.25

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

663.53 1382.67 374.75 929.25 1364.00 3898.75 1993.25

664.73 1378.00 370.50 952.00 1390.50 3864.50 1995.25

665.92 1382.67 358.00 936.25 1363.00 3815.50 2006.25

667.11 1400.33 392.00 910.50 1374.50 3849.00 2022.00

668.30 1341.67 381.50 928.25 1358.50 3786.75 2002.50

669.50 1397.33 389.25 923.50 1366.50 3820.75 1953.25

670.66 1402.00 381.50 938.75 1360.00 3815.50 2007.25

671.86 1357.00 381.50 932.50 1355.75 3865.25 1997.00

673.05 1406.00 387.75 911.75 1374.00 3847.75 1996.00

674.24 1424.33 379.50 918.25 1371.50 3871.50 1985.25

675.44 1399.67 378.00 934.75 1362.50 3792.25 2031.75

676.60 1386.67 362.75 922.50 1374.75 3840.25 1990.00

677.80 1382.67 369.50 926.75 1389.50 3805.75 2010.50

678.99 1393.33 384.75 940.25 1370.75 3871.00 2012.75

680.18 1371.00 381.25 923.25 1356.25 3849.00 2019.50

681.35 1398.00 381.75 909.00 1357.75 3903.00 2015.50

682.54 1400.33 362.50 936.00 1378.50 3827.75 1981.75

683.73 1384.67 383.75 909.50 1378.00 3825.25 1981.25

684.93 1347.67 386.50 938.25 1386.50 3850.75 2005.25

686.09 1346.00 380.00 913.25 1365.25 3812.50 2002.00

687.29 1438.00 379.50 914.25 1386.50 3849.75 1985.75

688.48 1381.33 371.25 916.25 1408.00 3857.50 2022.75

689.65 1342.33 390.25 932.25 1386.25 3853.75 1981.25

690.84 1400.00 375.25 922.00 1387.75 3850.50 2021.50

692.03 1374.00 388.00 934.75 1395.75 3829.00 2007.50

693.20 1354.00 370.75 920.25 1425.25 3871.25 2019.00

694.39 1406.00 404.00 911.50 1441.25 3899.25 2074.25

695.59 1366.33 411.00 967.75 1462.50 4003.25 2084.75

696.75 1427.67 400.75 937.25 1493.50 4051.00 2168.25

697.95 1463.00 420.25 987.25 1549.75 4167.75 2212.25

699.14 1480.67 452.75 979.75 1641.50 4241.50 2307.25

700.31 1557.67 539.25 1081.50 1729.00 4338.00 2436.25

701.50 1541.33 569.75 1181.00 1751.00 4511.50 2520.25

702.69 1521.33 551.25 1266.75 1766.75 4526.75 2572.50

703.86 1526.67 536.25 1259.50 1755.25 4589.75 2601.00

705.05 1598.67 585.00 1256.50 1715.25 4581.00 2540.00

706.22 1538.00 552.25 1283.00 1671.75 4620.25 2578.00

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

707.41 1465.67 470.00 1263.00 1570.50 4486.25 2483.50

708.58 1415.33 437.00 1193.25 1459.25 4345.50 2303.00

709.77 1464.67 418.25 1085.75 1426.75 4244.00 2219.00

710.97 1379.67 399.00 1024.50 1397.50 4044.75 2158.25

712.13 1366.33 383.50 986.50 1393.50 4070.25 2056.50

713.33 1403.00 389.25 979.00 1399.00 3986.25 2078.00

714.49 1363.00 388.50 942.00 1408.00 3920.00 2024.00

715.69 1413.33 387.25 981.75 1389.50 3929.50 2068.75

716.86 1387.33 388.50 964.50 1394.00 3900.75 2037.50

718.05 1389.00 371.00 937.00 1360.00 3882.25 2067.50

719.22 1377.67 379.25 938.00 1335.25 3853.25 2033.00

720.41 1414.67 387.00 912.50 1385.50 3849.25 1972.00

721.58 1373.33 389.25 922.50 1354.75 3837.75 2016.00

722.77 1393.33 374.75 918.00 1346.25 3812.50 2000.50

723.94 1374.00 383.75 911.50 1364.00 3856.50 1994.25

725.13 1405.00 370.25 908.00 1367.25 3862.00 1969.75

726.30 1383.67 365.00 908.75 1362.25 3785.50 1971.25

727.47 1373.33 375.75 921.25 1363.50 3838.00 1997.00

728.66 1409.00 367.25 907.50 1332.50 3799.00 1984.00

729.83 1368.67 366.75 916.75 1354.50 3750.50 1983.50

731.02 1366.00 356.50 895.50 1338.50 3827.50 1967.75

732.19 1396.67 378.75 907.50 1337.50 3778.25 1924.75

733.38 1375.67 365.50 907.75 1346.75 3816.75 1989.25

734.55 1378.33 363.75 913.75 1340.00 3779.25 1951.25

735.71 1394.00 370.50 915.00 1377.75 3810.25 1954.50

736.91 1345.33 384.25 899.50 1377.50 3849.50 1961.75

738.07 1383.33 378.50 936.00 1343.00 3758.00 1933.50

739.24 1403.33 382.25 898.25 1333.25 3747.50 1995.25

740.44 1358.33 377.25 898.00 1342.00 3740.50 1955.75

741.60 1413.67 359.00 888.00 1337.25 3779.75 1984.00

742.80 1393.00 354.75 894.50 1334.50 3837.25 1984.00

743.96 1370.33 354.00 896.25 1352.00 3752.25 1988.00

745.13 1381.00 369.75 924.50 1344.00 3712.75 1894.75

746.32 1360.67 366.75 894.75 1349.50 3741.75 1955.50

747.49 1384.33 361.00 864.75 1373.25 3776.50 1944.50

748.66 1403.67 376.25 891.75 1326.50 3763.50 1931.25

749.83 1378.00 375.25 894.75 1347.75 3806.25 1932.25

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

751.02 1383.33 366.75 864.50 1342.00 3783.75 1910.25

752.19 1394.00 379.25 887.00 1343.00 3742.50 1957.50

753.36 1404.67 382.00 905.75 1331.75 3812.25 1969.00

754.52 1401.67 369.25 882.25 1318.25 3755.75 1967.25

755.72 1373.67 370.75 904.25 1311.50 3723.25 1939.00

756.88 1406.00 356.00 891.75 1340.75 3739.75 1957.00

758.05 1391.33 368.00 896.00 1329.00 3732.00 1922.00

759.22 1368.00 367.50 863.50 1354.50 3688.25 1913.75

760.41 1390.00 366.25 928.00 1337.00 3799.75 1952.75

761.58 1426.33 371.00 882.50 1328.00 3733.75 1959.75

762.75 1426.67 369.75 887.00 1348.75 3753.25 1933.00

763.92 1376.33 364.75 913.75 1350.00 3752.25 1929.50

765.08 1384.00 375.75 897.25 1328.75 3778.50 1908.75

766.28 1391.33 369.75 907.50 1337.50 3719.75 1936.00

767.44 1375.67 372.25 899.75 1338.00 3738.75 1950.75

768.61 1386.67 352.50 883.00 1368.50 3694.50 1978.75

769.78 1387.00 362.50 908.75 1320.50 3720.75 1933.25

770.95 1381.33 370.75 892.25 1321.25 3728.75 1926.75

772.12 1360.33 358.50 893.00 1315.25 3759.75 1970.50

773.31 1368.33 371.75 858.75 1340.75 3730.75 1961.50

774.48 1395.67 347.75 884.75 1358.25 3752.25 1953.25

775.64 1394.33 373.25 897.25 1310.50 3718.75 1950.00

776.81 1349.00 364.00 870.50 1328.25 3770.25 1970.50

777.98 1368.33 376.25 879.25 1314.75 3773.25 1901.25

779.15 1393.00 367.50 912.50 1355.00 3790.25 1949.00

780.31 1415.00 372.75 876.00 1296.50 3773.75 1953.25

781.48 1368.33 361.50 898.50 1362.00 3735.75 1951.50

782.65 1389.67 352.25 871.50 1303.75 3769.75 1953.25

783.82 1390.67 363.25 891.25 1330.50 3768.00 1951.75

785.01 1385.67 366.25 884.25 1353.75 3722.75 1916.50

786.18 1350.33 360.75 887.00 1340.25 3719.75 1968.50

787.35 1375.33 359.75 891.00 1315.75 3703.25 1934.25

788.51 1389.00 366.50 869.50 1334.50 3742.50 1940.75

789.68 1361.67 366.50 866.50 1300.50 3742.25 1958.25

790.85 1386.00 370.75 877.00 1330.00 3750.50 1890.00

792.02 1383.33 379.75 891.75 1333.50 3743.50 1932.50

793.19 1380.33 361.25 899.50 1318.25 3682.50 1955.75

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

794.35 1382.00 348.25 869.00 1330.50 3683.75 1932.75

795.52 1399.33 367.75 898.25 1326.50 3732.75 1910.00

796.69 1374.00 356.50 872.75 1340.75 3725.25 1878.75

797.86 1395.00 381.25 897.75 1338.50 3657.25 1914.50

799.02 1356.00 377.25 889.25 1336.75 3713.75 1916.75

800.19 1430.33 346.75 892.50 1324.75 3695.25 1904.00

801.36 1395.67 368.25 875.50 1314.25 3728.00 1934.00

802.50 1369.00 368.50 885.50 1321.00 3788.50 1918.25

803.67 1372.33 351.25 880.50 1348.75 3685.25 1878.75

804.84 1364.33 380.00 909.00 1326.25 3707.50 1962.75

806.01 1390.33 379.25 881.75 1318.75 3731.00 1905.25

807.17 1392.00 359.75 899.25 1330.50 3714.50 1908.25

808.34 1390.00 342.25 891.75 1291.75 3707.75 1919.25

809.51 1369.67 356.75 868.25 1307.75 3735.75 1933.50

810.68 1397.33 361.50 875.50 1326.25 3705.75 1890.75

811.85 1368.00 372.50 888.75 1330.75 3772.50 1898.75

813.01 1396.67 347.75 885.50 1319.75 3707.75 1938.50

814.16 1414.00 363.50 868.25 1306.25 3672.75 1906.50

815.32 1383.67 378.75 875.75 1334.00 3698.50 1915.50

816.49 1381.33 358.75 875.75 1323.00 3713.75 1900.00

817.66 1428.67 367.75 869.25 1301.75 3665.25 1928.50

818.83 1379.33 362.75 915.75 1307.50 3686.25 1943.25

820.00 1378.00 349.75 890.50 1307.25 3741.75 1922.00

821.14 1383.00 364.00 895.00 1273.75 3702.00 1946.00

822.31 1381.33 356.50 889.00 1284.75 3705.75 1943.50

823.47 1375.33 358.75 899.00 1315.25 3715.00 1930.75

824.64 1385.00 351.75 892.75 1351.50 3657.00 1884.50

825.81 1417.33 351.00 878.25 1330.75 3685.00 1963.25

826.95 1392.67 364.00 846.50 1315.75 3659.75 1929.00

828.12 1371.00 362.00 870.25 1287.00 3659.00 1934.50

829.29 1373.00 352.75 858.50 1323.00 3653.75 1927.75

830.46 1400.00 373.75 859.25 1314.00 3794.00 1882.25

831.60 1411.67 351.50 891.75 1288.50 3704.50 1965.50

832.77 1402.33 368.25 881.75 1282.50 3725.25 1918.50

833.94 1348.00 342.75 862.50 1320.25 3688.50 1925.00

835.10 1409.33 353.50 876.75 1315.50 3746.75 1938.00

836.25 1382.00 357.25 878.75 1324.00 3711.25 1918.25

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

837.41 1385.67 361.75 861.25 1282.50 3737.25 1895.75

838.58 1347.67 374.50 847.75 1310.75 3736.50 1882.25

839.72 1430.67 358.50 890.25 1296.00 3687.00 1955.50

840.89 1407.00 374.50 874.25 1285.00 3696.75 1903.50

842.06 1385.33 345.25 858.25 1295.00 3666.00 1914.50

843.20 1411.00 359.75 899.50 1304.00 3668.75 1933.00

844.37 1379.67 361.75 866.50 1356.75 3703.25 1952.50

845.54 1392.67 354.25 866.25 1300.00 3631.25 1914.25

846.68 1396.67 345.50 876.50 1309.50 3732.50 1931.25

847.85 1390.67 369.25 862.75 1279.25 3678.50 1905.50

849.02 1363.33 367.00 851.00 1317.25 3667.00 1918.25

850.16 1376.33 349.00 873.75 1275.00 3722.00 1923.50

851.33 1440.00 361.00 862.25 1332.75 3723.00 1934.25

852.50 1397.33 369.50 873.50 1313.25 3691.75 1879.75

853.64 1372.00 369.00 897.50 1316.50 3678.50 1910.25

854.81 1352.33 355.25 892.25 1337.25 3710.25 1949.50

855.95 1376.00 353.00 891.00 1338.50 3724.75 1940.50

857.12 1384.67 353.75 862.75 1343.25 3712.50 1909.50

858.26 1381.00 340.75 883.00 1303.75 3622.75 1946.75

859.43 1425.67 365.75 868.75 1300.25 3635.50 1926.25

860.60 1362.67 356.25 875.75 1302.50 3701.25 1877.50

861.74 1380.00 351.25 872.75 1311.75 3678.25 1936.75

862.91 1376.67 369.75 873.25 1297.00 3650.75 1865.25

864.05 1357.67 362.50 855.00 1308.00 3656.00 1906.50

865.22 1408.00 360.00 843.25 1307.25 3643.25 1884.50

866.36 1388.33 366.00 890.50 1301.50 3701.00 1904.50

867.53 1385.00 355.50 865.75 1308.00 3675.00 1926.50

868.67 1360.67 369.25 867.25 1306.25 3623.75 1913.00

869.84 1361.00 363.00 861.00 1292.00 3631.50 1885.25

870.98 1355.00 370.25 877.25 1295.25 3731.00 1929.25

872.15 1398.00 366.25 842.00 1304.00 3701.00 1863.75

873.29 1379.67 363.00 855.00 1326.50 3654.75 1870.25

874.46 1404.67 356.75 852.50 1301.00 3721.75 1888.25

875.60 1384.33 347.50 853.25 1312.50 3622.25 1880.75

876.77 1374.67 363.00 879.75 1299.75 3649.50 1899.00

877.91 1374.00 358.25 834.75 1272.25 3668.50 1880.75

879.06 1399.00 359.50 859.75 1310.25 3628.75 1882.50

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

880.23 1406.33 371.25 853.00 1298.75 3675.00 1902.50

881.37 1368.67 348.25 868.00 1274.50 3601.75 1898.00

882.54 1365.33 349.00 850.75 1298.75 3645.75 1889.00

883.68 1358.67 357.75 866.75 1311.50 3667.00 1882.75

884.82 1376.67 355.00 861.00 1278.00 3685.50 1847.00

885.99 1405.67 362.00 836.00 1291.50 3627.25 1905.25

887.13 1381.67 356.75 850.50 1310.00 3624.25 1868.75

888.30 1361.33 375.00 875.00 1303.50 3594.25 1899.25

889.44 1394.33 361.00 852.75 1289.00 3602.75 1934.25

890.59 1390.00 342.25 878.25 1306.25 3660.75 1918.75

891.75 1376.33 351.50 871.75 1275.75 3637.00 1876.00

892.90 1393.00 353.00 872.75 1297.00 3659.50 1882.50

894.04 1391.00 361.50 864.00 1278.00 3625.75 1846.25

895.21 1406.33 365.00 867.50 1280.25 3608.50 1911.00

896.35 1381.00 338.50 855.00 1275.00 3611.75 1856.25

897.49 1407.67 342.50 847.50 1311.25 3596.75 1914.00

898.66 1389.00 354.75 861.50 1312.00 3617.50 1866.00

899.81 1376.67 348.25 862.00 1272.50 3605.25 1909.00

900.95 1423.33 351.75 868.00 1273.75 3646.50 1901.50

902.09 1392.00 349.75 844.25 1293.75 3620.25 1884.00

903.26 1375.67 355.00 853.00 1290.50 3643.00 1889.25

904.40 1415.33 358.25 861.25 1312.50 3662.00 1880.25

905.54 1393.67 360.50 841.25 1259.25 3652.75 1888.00

906.69 1409.67 362.00 868.25 1304.50 3618.75 1908.25

907.86 1389.33 369.00 835.25 1280.00 3653.25 1883.75

909.00 1401.67 354.25 863.50 1322.75 3652.50 1880.75

910.14 1415.00 347.75 868.50 1318.75 3662.75 1857.25

911.28 1388.67 359.25 839.75 1312.75 3625.75 1885.00

912.45 1353.33 353.75 881.50 1321.50 3623.50 1896.50

913.60 1371.33 343.50 850.25 1316.50 3585.25 1898.25

914.74 1432.67 348.00 861.50 1310.75 3660.75 1863.50

915.88 1402.00 351.25 864.50 1275.75 3647.00 1875.75

917.02 1348.00 359.25 863.00 1323.25 3626.75 1865.75

918.19 1401.33 353.75 844.25 1258.50 3617.50 1897.75

919.33 1387.00 363.75 863.00 1297.00 3565.75 1872.50

920.48 1380.67 356.75 860.00 1289.50 3681.25 1905.25

921.62 1424.00 355.50 845.25 1306.75 3696.00 1851.25

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

922.76 1393.33 353.75 864.25 1287.50 3668.75 1870.25

923.91 1390.33 341.50 854.25 1301.50 3632.50 1902.25

925.05 1411.67 352.75 859.00 1270.75 3651.75 1872.50

926.19 1392.33 368.00 866.50 1287.75 3637.00 1886.75

927.36 1381.00 348.25 855.50 1296.00 3666.00 1864.00

928.50 1399.00 349.25 863.50 1292.50 3618.00 1879.75

929.65 1414.33 350.00 864.00 1279.50 3660.25 1856.00

930.79 1387.33 342.25 864.50 1273.00 3633.00 1874.50

931.93 1378.00 341.75 888.00 1305.00 3670.50 1914.00

933.08 1410.67 352.25 851.25 1293.50 3619.00 1894.25

934.22 1397.00 354.00 841.00 1299.50 3638.25 1897.50

935.36 1416.67 369.50 855.75 1288.50 3647.00 1837.50

936.50 1405.00 355.50 860.50 1299.00 3627.00 1893.00

937.65 1392.33 356.25 846.25 1261.50 3612.25 1882.50

938.79 1419.33 341.75 849.25 1293.25 3673.75 1856.25

939.93 1403.67 347.00 853.25 1258.25 3646.00 1892.75

941.08 1419.00 355.75 871.00 1290.00 3621.50 1890.00

942.22 1382.33 342.75 870.50 1265.75 3625.75 1887.00

943.36 1379.67 364.25 859.25 1262.50 3652.75 1873.50

944.51 1375.00 348.75 861.25 1305.00 3647.50 1901.25

945.65 1398.00 340.50 845.25 1271.75 3628.75 1878.25

946.79 1401.67 353.00 846.75 1291.75 3618.25 1871.75

947.93 1395.67 346.75 853.00 1270.25 3602.75 1908.75

949.08 1391.33 366.75 845.00 1309.75 3632.50 1914.75

950.22 1411.00 345.25 840.75 1265.25 3625.25 1873.75

951.36 1401.33 355.50 849.75 1271.00 3604.25 1866.25

952.51 1447.00 361.75 837.50 1310.00 3628.25 1863.25

953.65 1404.67 358.50 830.00 1268.50 3615.50 1892.75

954.79 1396.67 355.00 865.25 1305.25 3663.00 1890.75

955.93 1381.33 361.50 832.75 1283.50 3613.25 1880.75

957.08 1370.33 360.50 852.00 1317.25 3579.75 1888.25

958.22 1387.67 355.75 856.50 1282.00 3629.00 1855.50

959.36 1422.00 350.25 877.00 1293.75 3582.25 1859.25

960.48 1404.33 366.25 842.75 1306.00 3617.50 1869.50

961.62 1369.67 350.25 871.00 1241.75 3632.00 1833.00

962.77 1430.00 350.75 849.50 1273.50 3614.50 1891.50

963.91 1407.00 348.25 856.25 1277.00 3643.00 1847.00

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Wavelength External FG LG External FG LG

965.05 1413.33 357.50 843.75 1283.50 3648.00 1879.00

966.20 1421.33 345.75 848.75 1308.25 3630.00 1890.75

967.34 1403.33 357.50 809.25 1287.25 3562.50 1828.50

968.46 1382.67 339.75 838.75 1296.00 3626.50 1860.25

969.60 1401.00 352.50 853.50 1281.75 3607.50 1881.25

970.74 1391.00 335.75 846.00 1274.00 3639.25 1894.75

971.89 1396.00 353.50 829.00 1272.50 3637.25 1883.50

973.03 1378.33 362.75 857.75 1276.25 3635.00 1869.75

974.17 1376.00 347.25 856.00 1304.00 3619.25 1858.50

975.29 1431.33 339.25 824.00 1275.50 3653.00 1856.00

976.43 1393.33 352.25 824.00 1277.50 3662.50 1852.00

977.58 1423.00 352.00 837.50 1252.75 3585.25 1834.25

978.72 1367.33 362.75 821.75 1259.75 3623.75 1868.50

979.84 1382.00 367.50 817.50 1256.00 3622.50 1893.25

980.98 1347.33 346.00 843.25 1255.50 3549.25 1881.00

982.12 1422.67 340.00 832.25 1274.25 3611.75 1873.50

983.27 1411.67 360.50 828.50 1289.75 3559.25 1870.75

984.38 1391.67 363.00 830.50 1255.25 3595.00 1871.25

985.53 1396.67 353.00 837.25 1248.25 3608.00 1870.00

986.67 1386.33 351.00 862.50 1282.25 3588.00 1890.25

987.81 1387.67 355.50 831.00 1246.75 3568.50 1861.75

988.93 1393.33 339.75 859.50 1296.50 3621.00 1847.25

990.07 1407.67 342.25 843.00 1263.00 3589.50 1891.25

991.22 1389.33 335.00 844.50 1266.00 3598.50 1864.25

992.34 1402.00 348.50 820.25 1267.00 3551.25 1842.00

993.48 1386.33 347.25 838.50 1226.50 3508.50 1849.50

994.62 1367.67 336.25 841.25 1256.25 3602.75 1823.50

995.74 1382.00 355.50 800.25 1285.50 3508.75 1868.50

996.88 1405.33 348.75 836.75 1269.25 3547.00 1850.50

998.03 1386.00 336.25 845.00 1264.25 3572.75 1859.00

999.14 1412.00 348.75 828.75 1279.25 3556.50 1839.25

1000.29 1382.00 364.25 841.50 1255.50 3571.25 1853.50

1001.40 1393.67 344.25 838.00 1244.50 3586.25 1853.25

1002.55 1391.67 355.50 843.75 1275.00 3566.50 1845.75

1003.69 1398.00 342.00 859.25 1278.50 3580.00 1851.75

1004.81 1381.00 354.25 851.00 1282.00 3584.25 1853.50

1005.95 1373.33 350.25 840.25 1262.25 3553.75 1860.25

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

1007.09 1366.67 352.50 847.75 1274.50 3497.75 1866.50

1008.21 1399.67 332.75 837.75 1271.25 3603.25 1882.25

1009.36 1455.33 360.75 825.25 1240.50 3645.00 1825.75

1010.47 1394.67 352.00 834.50 1254.50 3517.25 1844.00

1011.62 1375.00 354.25 837.75 1236.75 3557.00 1824.50

1012.73 1409.67 331.75 816.00 1241.75 3490.00 1845.25

1013.88 1393.33 330.00 817.00 1261.50 3575.25 1841.75

1015.00 1352.67 356.75 826.00 1223.75 3636.00 1830.75

1016.14 1355.67 354.25 835.00 1277.50 3550.00 1803.25

1017.26 1413.33 364.75 814.75 1253.25 3589.25 1836.75

1018.40 1380.67 369.25 864.75 1262.75 3534.50 1817.50

1019.52 1397.00 349.75 842.00 1260.25 3575.00 1863.00

1020.66 1426.67 335.75 834.75 1241.75 3530.50 1848.75

1021.78 1401.67 355.25 842.25 1253.75 3590.00 1847.75

1022.92 1404.67 327.75 836.00 1223.00 3562.75 1820.50

1024.04 1397.67 343.75 833.75 1239.75 3483.00 1869.50

1025.18 1387.00 353.50 839.75 1260.00 3530.25 1831.50

1026.30 1425.33 336.75 825.00 1257.25 3593.50 1824.00

1027.44 1395.33 340.50 859.25 1264.00 3627.00 1875.75

1028.56 1396.67 350.75 827.50 1278.25 3539.50 1870.50

1029.71 1391.33 340.00 824.75 1252.25 3509.75 1855.50

1030.82 1434.67 353.50 820.75 1222.00 3521.25 1845.25

1031.94 1387.00 354.75 849.75 1275.50 3589.75 1834.25

1033.08 1386.67 347.50 845.25 1249.25 3610.50 1827.50

1034.20 1387.33 355.25 834.25 1231.50 3568.50 1828.50

1035.35 1384.33 361.00 828.00 1256.75 3521.25 1844.75

1036.46 1369.67 349.50 853.25 1246.50 3499.00 1816.50

1037.58 1394.67 354.00 810.00 1261.75 3582.50 1826.00

1038.72 1404.00 346.00 821.25 1233.25 3483.50 1829.00

1039.84 1405.67 366.50 814.25 1258.00 3538.00 1801.25

1040.96 1394.00 362.00 830.25 1239.50 3544.75 1877.75

1042.10 1423.33 347.25 841.50 1261.50 3597.50 1858.50

1043.22 1402.00 354.50 841.00 1250.75 3592.00 1842.50

1044.37 1428.67 371.50 854.25 1267.75 3588.00 1882.25

1045.48 1407.00 349.75 823.25 1236.25 3553.25 1858.75

1046.60 1484.00 338.00 841.00 1284.75 3607.75 1842.00

1047.74 1400.00 356.75 820.75 1293.25 3547.50 1867.25

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

1048.86 1422.33 341.25 822.50 1270.50 3583.25 1873.25

1049.98 1442.00 355.00 829.50 1244.50 3541.50 1829.50

1051.10 1413.33 337.25 832.25 1264.50 3580.50 1839.25

1052.24 1379.33 348.00 823.75 1245.75 3560.00 1822.25

1053.36 1376.33 360.00 854.50 1254.00 3552.25 1836.00

1054.48 1416.33 322.00 812.25 1269.50 3610.50 1875.00

1055.62 1425.00 363.00 838.75 1279.50 3542.50 1854.00

1056.74 1441.00 355.50 817.75 1251.25 3562.50 1838.25

1057.86 1419.00 367.25 840.25 1239.00 3553.75 1863.50

1058.98 1414.33 363.75 845.50 1302.50 3571.50 1880.00

1060.09 1424.67 367.75 862.50 1258.25 3577.75 1863.50

1061.24 1434.00 365.25 855.50 1261.00 3632.50 1848.00

1062.35 1456.00 366.50 853.00 1290.25 3601.50 1904.50

1063.47 1432.00 364.25 846.75 1253.75 3621.00 1840.00

1064.59 1390.00 375.50 837.50 1288.75 3588.75 1886.25

1065.73 1417.00 368.50 834.50 1263.00 3604.25 1889.50

1066.85 1402.33 363.25 890.00 1276.75 3614.00 1884.00

1067.97 1435.33 361.25 888.50 1289.50 3602.25 1897.00

1069.09 1418.67 377.00 860.75 1311.00 3626.25 1887.25

1070.21 1412.67 381.25 872.00 1287.25 3650.75 1917.25

1071.32 1409.00 395.50 890.25 1296.75 3680.50 1931.50

1072.47 1435.00 379.25 899.50 1317.25 3689.00 1965.50

1073.59 1405.33 392.75 881.00 1324.75 3706.00 1983.50

1074.70 1485.00 394.75 920.50 1392.00 3722.25 2011.75

1075.82 1470.67 424.75 969.50 1432.75 3783.75 2139.25

1076.94 1461.67 425.75 1001.50 1595.25 3887.50 2199.75

1078.06 1481.67 470.75 1066.75 1799.50 4047.75 2455.50

1079.18 1542.67 509.00 1195.25 2140.00 4482.25 2768.50

1080.29 1590.33 618.50 1389.00 2451.75 5129.75 3363.50

1081.44 1767.67 814.50 1704.25 2974.25 5944.50 4042.25

1082.56 2113.33 1219.75 2164.75 3650.25 6794.50 4921.75

1083.67 2840.67 2112.00 2901.25 4539.25 7756.00 6110.75

1084.79 3769.67 3140.75 4050.25 4962.25 8660.75 7468.75

1085.91 3234.00 2739.75 5081.50 4057.00 9321.75 8096.25

1087.03 2084.00 1523.25 5211.00 2931.25 9204.50 6640.75

1088.15 1678.00 851.00 3729.75 2145.00 8058.00 4843.25

1089.26 1548.33 554.75 2456.75 1670.75 6848.25 3609.00

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

1090.38 1452.67 462.50 1764.50 1427.00 5720.75 2812.00

1091.50 1464.00 398.75 1311.25 1355.25 4712.75 2318.75

1092.62 1460.67 385.00 1061.75 1289.25 4143.25 2096.25

1093.74 1415.00 376.00 966.75 1247.00 3817.50 1966.00

1094.85 1417.00 381.75 902.00 1286.25 3693.25 1936.75

1095.97 1432.00 367.25 873.50 1277.75 3657.25 1915.75

1097.09 1461.67 346.50 867.25 1257.25 3596.75 1889.25

1098.21 1440.67 363.75 847.25 1252.25 3593.75 1882.50

1099.33 1450.33 356.00 838.25 1258.75 3538.50 1831.25

1100.45 1437.67 358.50 838.75 1231.25 3581.00 1850.00

1101.56 1471.67 345.75 829.50 1216.50 3601.75 1825.50

1102.68 1384.67 339.00 821.75 1228.50 3566.50 1828.75

1103.80 1374.33 347.50 814.75 1239.50 3550.00 1849.00

1104.92 1418.33 348.00 819.75 1243.25 3590.75 1827.00

1106.04 1412.00 348.75 792.50 1267.00 3525.50 1802.25

1107.15 1451.67 347.50 780.25 1238.25 3537.25 1800.25

1108.27 1437.67 343.25 804.75 1217.50 3513.00 1816.75

1109.37 1451.67 340.50 805.75 1209.75 3536.25 1809.25

1110.48 1416.00 343.00 803.75 1239.75 3487.00 1793.75

1111.60 1411.00 339.25 802.75 1236.50 3572.00 1847.00

1112.72 1434.00 329.50 812.75 1215.00 3511.50 1796.75

1113.84 1431.00 337.00 817.00 1222.25 3440.75 1851.00

1114.96 1432.06 340.08 806.59 1235.20 3506.24 1808.31

1116.07 1379.53 335.03 801.87 1212.74 3495.75 1809.59

1117.19 1391.29 353.98 793.22 1221.93 3478.59 1763.85

1118.29 1392.89 345.06 814.06 1226.44 3495.15 1755.63

1119.40 1433.24 342.48 808.69 1223.01 3530.06 1831.88

1120.52 1413.14 359.98 786.75 1224.64 3470.30 1795.13

1121.64 1442.73 344.02 797.84 1207.83 3536.55 1776.75

1122.76 1387.34 337.08 778.41 1219.20 3475.77 1790.94

1123.88 1418.64 328.42 812.60 1226.01 3453.91 1784.25

1124.97 1411.68 338.14 790.16 1229.47 3496.10 1788.69

1126.09 1402.04 338.70 788.19 1202.39 3473.79 1763.50

1127.21 1434.97 339.80 796.72 1220.56 3476.95 1793.09

1128.32 1412.62 358.61 797.63 1212.32 3482.33 1799.28

1129.44 1408.78 345.61 791.62 1215.18 3439.19 1747.44

1130.54 1414.02 348.28 785.13 1193.42 3460.51 1788.03

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

1131.65 1376.25 345.70 794.00 1217.24 3461.37 1799.03

1132.77 1392.14 340.06 811.91 1191.36 3464.05 1761.97

1133.89 1417.15 339.41 795.03 1203.09 3465.78 1785.97

1134.98 1371.55 337.91 786.47 1193.66 3428.05 1773.69

1136.10 1409.04 339.86 793.16 1212.45 3467.29 1777.22

1137.22 1377.67 334.08 789.19 1184.25 3423.35 1797.22

1138.34 1400.27 333.72 783.06 1215.95 3453.06 1748.38

1139.43 1385.85 339.09 779.59 1198.47 3442.55 1730.28

1140.55 1411.61 327.02 778.19 1196.33 3379.24 1759.28

1141.67 1375.46 331.98 797.56 1194.03 3438.44 1749.25

1142.76 1403.30 327.39 776.31 1195.29 3457.33 1752.41

1143.88 1396.59 342.91 768.81 1221.11 3384.91 1758.53

1145.00 1387.07 340.75 785.47 1203.67 3386.94 1741.75

1146.09 1413.70 334.14 780.50 1220.44 3400.82 1736.97

1147.21 1376.11 336.08 764.28 1191.97 3355.54 1748.88

1148.33 1382.68 334.06 786.44 1199.08 3395.49 1719.07

1149.42 1366.92 338.34 755.09 1197.04 3327.79 1745.81

1150.54 1390.99 340.16 789.00 1208.31 3402.07 1721.31

1151.66 1413.11 331.95 778.56 1181.12 3357.70 1743.97

1152.75 1449.42 334.06 765.31 1200.27 3383.01 1751.91

1153.87 1429.60 319.73 781.60 1201.14 3388.63 1707.85

1154.96 1387.64 329.66 771.25 1203.99 3396.49 1745.78

1156.08 1378.45 332.53 759.72 1182.12 3361.22 1709.94

1157.20 1377.39 336.64 781.00 1178.56 3370.88 1732.16

1158.29 1394.19 330.84 783.16 1157.86 3374.59 1768.50

1159.41 1397.91 329.89 757.44 1201.21 3346.36 1725.53

1160.50 1376.67 314.13 778.88 1188.83 3383.19 1725.09

1161.62 1386.08 328.53 788.81 1169.57 3389.64 1743.22

1162.74 1407.61 326.86 783.25 1210.43 3305.41 1742.44

1163.83 1395.11 324.03 764.25 1185.15 3400.71 1738.97

1164.95 1347.32 320.20 750.09 1181.20 3337.25 1738.13

1166.04 1358.94 339.47 769.00 1188.33 3321.56 1750.53

1167.16 1404.23 316.56 762.94 1185.95 3388.71 1748.19

1168.25 1399.56 336.48 747.75 1167.59 3343.36 1726.63

1169.37 1404.67 322.17 776.56 1186.52 3346.60 1731.10

1170.46 1385.95 334.84 769.16 1175.14 3300.08 1732.50

1171.58 1382.42 330.39 752.03 1192.19 3353.94 1722.47

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Huon Peninsula (K-133) Palm Island (PT-1)

Wavelength External FG LG External FG LG

1172.68 1385.42 325.09 766.41 1197.28 3368.74 1740.72

1173.79 1354.33 327.64 745.72 1186.60 3345.37 1726.06

1174.89 1404.48 325.73 771.03 1161.46 3372.66 1729.34

1176.01 1402.88 324.45 755.56 1187.93 3329.64 1730.85

1177.10 1401.24 320.52 752.28 1190.61 3332.61 1743.81

1178.22 1391.39 333.78 769.00 1172.45 3347.67 1730.47

1179.31 1403.97 330.16 770.88 1177.12 3316.34 1733.28

1180.43 1359.40 330.36 771.53 1184.94 3322.10 1702.16

1181.52 1389.07 325.97 765.44 1208.59 3319.03 1724.28

1182.61 1377.80 324.42 749.28 1169.19 3269.04 1719.91

1183.73 1394.03 315.75 754.03 1180.55 3291.07 1694.63

1184.83 1375.52 323.45 750.09 1169.72 3311.06 1716.88

1185.94 1383.80 322.53 774.09 1153.08 3302.27 1720.25

1187.04 1382.73 329.36 753.34 1192.14 3297.40 1711.00

1188.13 1371.86 325.42 750.47 1167.31 3278.84 1695.06

1189.25 1386.73 329.03 749.50 1175.54 3300.06 1714.60

1190.34 1376.76 322.92 760.19 1180.99 3270.56 1693.10

1191.46 1368.32 325.97 755.10 1183.73 3291.48 1705.60

1192.55 1365.26 318.59 767.56 1181.07 3285.96 1713.47

1193.65 1354.36 317.33 757.94 1168.52 3295.40 1707.94

1194.77 1358.73 323.20 741.19 1162.97 3272.48 1685.44

1195.86 1374.27 314.81 752.09 1173.10 3279.53 1730.91

1196.95 1365.45 326.13 759.38 1171.46 3289.11 1695.72

1198.07 1368.52 332.67 732.31 1170.71 3291.79 1704.94

1199.16 1377.64 324.36 761.84 1159.96 3260.84 1721.84

1200.26 1354.47 319.08 768.00 1173.52 3258.31 1702.78

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119

APPENDIX X: STABLE ISOTOPIC ANALYSIS

Measurements: External: Distance from initial milled trench; Internal: Distance from pallial line

Sample Layer (Fig. 1.1) Distance (µm) δ13

C (‰VPBD) δ18

O (‰VPBD)

K-133 EL -800 2.56 -1.59

K-133 EL -750 2.24 -1.68

K-133 EL -700 2.57 -1.45

K-133 EL -650 2.28 -1.56

K-133 EL -600 2.52 -1.47

K-133 EL -550 2.77 -1.40

K-133 EL -500 2.63 -1.43

K-133 EL -450 2.55 -1.48

K-133 EL -400 2.42 -1.47

K-133 EL -350 2.89 -1.36

K-133 EL -300 3.24 -1.70

K-133 EL -250 3.20 -1.80

K-133 EL -200 3.23 -1.61

K-133 EL -150 2.23 -1.67

K-133 EL -100 2.29 -1.68

K-133 EL -50 2.33 -1.54

Figure A.3, Powder for stable carbon and oxygen isotope analysis was milled from the external

(EL) and internal (IL) layers of modern T. gigas shells: K-133 (PNG; left) and PT-1 (GBR;

right). The pallial line (dashed black line) separates the shell layers.

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120

Sample Layer (Fig. 1.1) Distance (µm) δ13

C (‰VPBD) δ18

O (‰VPBD)

K-133 IL 0 2.23 -1.51

K-133 IL 50 2.19 -1.62

K-133 IL 100 2.19 -1.54

K-133 IL 150 2.11 -1.47

K-133 IL 200 2.14 -1.44

K-133 IL 250 2.20 -1.18

K-133 IL 300 2.20 -1.56

K-133 IL 350 2.16 -1.44

K-133 IL 400 2.25 -1.21

K-133 IL 450 2.15 -1.41

K-133 IL 500 2.27 -1.22

K-133 IL 550 2.20 -1.33

K-133 IL 600 2.19 -1.41

K-133 IL 650 2.18 -1.37

K-133 IL 700 2.17 -1.43

K-133 IL 750 2.18 -1.34

K-133 IL 800 2.17 -1.80

K-133 IL 850 2.22 -1.19

K-133 IL 900 2.12 -1.24

K-133 IL 950 2.19 -1.27

K-133 IL 1000 2.16 -1.15

K-133 IL 1050 2.30 -1.29

K-133 IL 1100 2.13 -1.21

K-133 IL 1150 2.16 -1.12

K-133 IL 1200 2.11 -1.24

K-133 IL 1250 2.22 -1.04

K-133 IL 1300 2.10 -1.22

K-133 IL 1350 2.13 -1.20

K-133 IL 1400 2.10 -1.31

K-133 IL 1450 2.06 -1.27

K-133 IL 1500 2.06 -1.27

K-133 IL 1550 2.09 -1.08

K-133 IL 1600 2.02 -1.23

K-133 IL 1650 2.04 -1.20

K-133 IL 1700 2.12 -1.14

K-133 IL 1750 2.16 -1.15

K-133 IL 1800 2.01 -1.15

K-133 IL 1850 2.04 -1.47

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121

Sample Layer (Fig. 1.1) Distance (µm) δ13

C (‰VPBD) δ18

O (‰VPBD)

K-133 IL 1900 2.17 -1.41

K-133 IL 1950 2.10 -1.12

K-133 IL 2000 2.21 -1.07

K-133 IL 2050 2.27 -1.26

K-133 IL 2100 2.09 -1.36

K-133 IL 2150 2.16 -1.27

K-133 IL 2200 2.04 -1.29

K-133 IL 2250 1.98 -1.20

K-133 IL 2300 2.13 -1.52

K-133 IL 2350 2.21 -1.18

K-133 IL 2400 2.09 -1.19

K-133 IL 2450 2.03 -1.42

K-133 IL 2500 2.17 -1.09

K-133 IL 2550 2.15 -1.48

K-133 IL 2600 2.02 -1.43

K-133 IL 2650 2.05 -1.64

K-133 IL 2700 2.14 -1.41

K-133 IL 2750 2.17 -1.60

K-133 IL 2800 2.12 -1.50

PT-1 EL -950 2.95 -0.35

PT-1 EL -900 2.60 -0.43

PT-1 EL -850 3.06 -0.58

PT-1 EL -800 3.05 -0.33

PT-1 EL -750 3.25 -0.59

PT-1 EL -700 2.17 -0.24

PT-1 EL -650 3.59 -0.59

PT-1 EL -600 3.40 -0.58

PT-1 EL -550 3.22 -0.48

PT-1 EL -500 3.67 -0.61

PT-1 EL -450 3.43 -0.68

PT-1 EL -400 3.57 -0.74

PT-1 EL -350 3.52 -0.55

PT-1 EL -300 3.62 -0.76

PT-1 EL -250 3.42 -0.67

PT-1 EL -200 2.98 -0.55

PT-1 EL -150 3.09 -0.71

PT-1 EL -100 3.04 -0.59

PT-1 EL -50 3.43 -0.67

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122

Sample Layer (Fig. 1.1) Distance (µm) δ13

C (‰VPBD) δ18

O (‰VPBD)

PT-1 IL 0 1.63 -1.14

PT-1 IL 50 1.42 -1.75

PT-1 IL 100 1.78 -1.14

PT-1 IL 150 1.71 -1.04

PT-1 IL 200 1.73 -0.93

PT-1 IL 250 1.70 -1.12

PT-1 IL 300 1.75 -1.02

PT-1 IL 350 1.69 -1.01

PT-1 IL 400 1.71 -0.99

PT-1 IL 450 1.76 -0.86

PT-1 IL 500 1.67 -1.01

PT-1 IL 550 1.60 -0.80

PT-1 IL 600 1.61 -0.88

PT-1 IL 650 1.66 -0.68

PT-1 IL 700 1.55 -0.75

PT-1 IL 750 1.62 -0.71

PT-1 IL 800 1.63 -0.92

PT-1 IL 850 1.50 -0.39

PT-1 IL 900 1.70 -0.30

PT-1 IL 950 1.62 -0.42

PT-1 IL 1000 1.66 -0.30

PT-1 IL 1050 1.64 -0.21

PT-1 IL 1100 1.61 0.06

PT-1 IL 1150 1.67 -0.04

PT-1 IL 1200 1.48 -0.30

PT-1 IL 1250 1.56 -0.04

PT-1 IL 1300 1.60 -0.13

PT-1 IL 1350 1.58 -0.06

PT-1 IL 1400 1.61 0.03

PT-1 IL 1450 1.61 -0.01

PT-1 IL 1500 1.48 -0.05

PT-1 IL 1550 1.60 0.05

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123

APPENDIX XI: TERRACE CORRELATIONS

The Huon Peninsula is not the only location where raised coral reef terraces have been

investigated for paleoclimate studies. Chrono-correlated terraces show the same glacio-eustatic

features through morphological and chemical analyses (Aharon, 1983; Dodge et al., 1983;

Radtke and Schellmann, 2005). Differences arise in the terrace building mechanisms with only

minor discrepancies in geologic age. Along the coastal shores of Kenya in the Kilifi District,

terraces have been associated with times of interglacial, high sea level for the past 25ka. During

these times, significant coral reefs built up causing isostatic loading. When sea level dropped, the

land rebounded due to transgression (Åse, 1981).

Terraces in Haiti, Indonesia, and Barbados are preserved due to tectonic uplift, similar to the

Huon Peninsula. The Northwest Peninsula of Haiti contains reef crests composed of nearly

exclusively Acropora palmate corals, which typically thrive in less than 5m of water. Several

reef crests were dated using U/Th and produced ages of 130ka, 108ka, and 81ka (Dodge et al.,

1983). Barbados terraces are also primarily composed of A. palmate, indicating there were low

tides during the time they built up (Radtke and Schellmann, 2005). These have been

radiometrically dated using electron spin resonance (ESR), concluding the ages for one (N1) of

the many studied sites to be: 106 ± 8ka, 85 ± 5ka, and 76 ± 4ka (Radke and Schellmann, 2006).

Some of these ages are within error of dates from other terraces in Haiti and Papua New Guinea

(Radtke and Schellmann, 2006). The calculated sea level elevations in Haiti correlate to those

seen in Barbados and Huon Peninsula for similar time period (Dodge et al., 1983). Uplifted

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124

Indonesian terraces on Sumba Island record the past 1-million years of sea level. Terraces in this

location were dated using ESR in order to obtain ages for early Pleistocene terraces where

Uranium-Thorium might not have been reliable. This allowed for sea level reconstruction

through MIS 25 (Pirazzoli et al., 1991). While T. gigas do not inhabit the Atlantic regions, their

use in the Huon Peninsula could help further global climate research due to the associations

between preserved terraces in these and other locations.

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125

APPENDIX X: TERRACE AGES

This table represents all of the dates that have been calculated from the major projects assessing

geologic age of the raised coral reef terraces of the Huon Peninsula, Papua-New Guinea. Bolded

entries are thought to be valid and have been included in the age assignment for this study.

Ter

race

(Fig

. 2.1

)

Met

hod

olo

gy

Sou

rce

Date

(k

a)

Err

or

Sam

ple

Nam

e

(Typ

e)

Sam

ple

Typ

e

Note

s

I 14

C

Aharon,

personal

communication

5.35 20 K-134 Tridacna

gigas

I 14

C

Aharon,

personal

communication

5.52 20 K-135 Tridacna

gigas

I U/Th Bloom et al.,

1974 5.5 0.4 28-1351I

Hydnophora

microconos

It is not

possible to

determine

age from

these

samples

though it is

constrained

to between

5-9 ka.

I U/Th Veeh and

Chappell, 1970 6 1 ANU 165 coral

I 14C Veeh and

Chappell, 1970 6.7

0.0

6 ANU 165 coral

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126

Ter

race

(Fig

. 2.1

)

Met

hod

olo

gy

Sou

rce

Date

(k

a)

Err

or

Sam

ple

Nam

e

(Typ

e)

Sam

ple

Typ

e

Note

s

I U/Th Veeh and

Chappell, 1970 6.8 0.1 ANU 153

Tridacna

gigas

Not

reliable

because

uses

Tridacna

I 14

C Aharon,

Chappell 1986 7.2 0.5 Reef I crest

3 Tridacna

gigas; 5

coral

I U/Th Aharon,

Chappell 1986 8.2 1.9 Reef I crest 5 coral

I U/Th Bloom et al.,

1974 9.2 0.6 33b-1351D

Favia

stelligera

It is not

possible to

determine

age from

these

samples

though it is

constrained

to between

5-9 ka.

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127

Ter

race

(F

ig.

2.1

)

Met

ho

do

logy

So

urc

e

Da

te (

ka)

Err

or

Sa

mp

le N

am

e

(Ty

pe)

Sa

mp

le T

yp

e

No

tes

I U/Th Bloom et al.,

1974 9.4 0.6 2-1351H

Goniastrea

retiformis

It is not

possible to

determine

age from

these

samples

though it is

constrained

to between

5-9 ka.

I U/Th Bloom et al.,

1974 9.4 0.6 29-1347H

Leptoria

phygia

It is not

possible to

determine

age from

these

samples

though it is

constrained

to between

5-9 ka.

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128

Ter

race

(Fig

. 2.1

)

Met

hod

olo

gy

Sou

rce

Date

(k

a)

Err

or

Sam

ple

Nam

e

(Typ

e)

Sam

ple

Typ

e

Note

s

I U/Th Bloom et al.,

1974 9.7 0.6 1-1351F

Favia

stelligera

It is not

possible to

determine

age from

these

samples

though it is

constrained

to between

5-9 ka.

I U/Th Bloom et al.,

1974 20 2 1-1351F

Favia

stelligera

It is not

possible to

determine

age from

these

samples

though it is

constrained

to between

5-9 ka.

II U/Th Bloom et al.,

1974 3.3 0.2 21-1353B Favia sp.

Cannot be

confirmed

but 29 ka is

used for

tectonic

purposes

II U/Th Bloom et al.,

1974 5.8 0.4 30-1355A

Leptoria

phygia

Cannot be

confirmed

but 29 ka is

used for

tectonic

purposes

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129

Ter

race

(Fig

. 2.1

)

Met

hod

olo

gy

Sou

rce

Date

(k

a)

Err

or

Sam

ple

Nam

e

(Typ

e)

Sam

ple

Typ

e

Note

s

II 14

C Aharon, Chappell

1986 28.9 0.6 Reef II crest 2 Tridacna

Not using 14

C

II 14C Veeh and

Chappell, 1970 29.3 0.9 ANU 156

Tridacna

gigas

Considered

more

reliable

than Bloom

II U/Th Aharon,

Chappell 1986 31 2.5 Reef II crest coral

II Pa/U Cutler et al.,

2003 35.2 0.9 KNM-T-2 (b) Porites sp.

No

diagenesis

II Pa/U Cutler et al.,

2003 36.3 1.6 KNM-T-2 (a) Porites sp.

No

diagenesis

II U/Th Cutler et al.,

2003 36.76

0.4

2 KNM-T-2 (b) Porites sp.

No

diagenesis

II U/Th Cutler et al.,

2003 36.8 0.2 KNM-T-2 (a) Porites sp.

No

diagenesis

II Pa/U Cutler et al., 2003 44.7 0.6 KWA-I-1 (B) Porites sp. Some

diagenesis

II Pa/U Cutler et al., 2003 45.1 0.9 KWA-I-1 (A) Porites sp. Some

diagenesis

II U/Th Cutler et al., 2003 46.41 0.2 KWA-I-1 (B) Porites sp. Some

diagenesis

II U/Th Cutler et al., 2003 46.64 0.4

5 KWA-I-1 (A) Porites sp.

Some

diagenesis

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130

Ter

race

(Fig

. 2.1

)

Met

hod

olo

gy

Sou

rce

Date

(k

a)

Err

or

Sam

ple

Nam

e

(Typ

e)

Sam

ple

Typ

e

Note

s

II Pa/U Cutler et al., 2003 49 1 KWA-N-1 (A) Porites sp.

No

diagenesis;

Ages are

too old,

maybe

from III?

II U/Th Cutler et al., 2003 50.23 0.4 KWA-N-1 (A) Porites sp.

No

diagenesis;

Ages are

too old,

maybe

from III?

II U/Th Cutler et al., 2003 50.8 0.2

6 KWA-N-1 (B) Porites sp.

No

diagenesis;

Ages are

too old,

maybe

from III?

II Pa/U Cutler et al., 2003 50.8 1.2 KWA-N-1 (B) Porites sp.

No

diagenesis;

Ages are

too old,

maybe

from III?

III U/Th Bloom et al.,

1974 4.9 1.4 39b-1351J

Lobophyllia

corynbosa

Disregarde

d by

author.

III U/Th Bloom et al.,

1974 7.5 5 39b-1351J

Lobophyllia

corynbosa

Disregarde

d by

author.

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131

Ter

race

(Fig

. 2.1

)

Met

hod

olo

gy

Sou

rce

Date

(k

a)

Err

or

Sam

ple

Nam

e

(Typ

e)

Sam

ple

Typ

e

Note

s

III U/Th Veeh and

Chappell, 1970 23 2 AUN 117

Tridacna

gigas

Not

reliable

because

uses

Tridacna.

III 14C Veeh and

Chappell, 1970 30.9 0.9 ANU 150

Tridacna

gigas

Ages too

young,

maybe

from II?

IIIb Pa/U Cutler et al., 2003 31.3 0.9 KAN-C-2 Favia laxa Some

diagenesis

IIIb 14

C Aharon, Chappell

1986 33.8 2.3 Reef IIIb crest

4 Tridacna

gigas

Ages too

young,

maybe

from II?

III U/Th Veeh and

Chappell, 1970 34 4 ANU 116

Tridacna

gigas

Not

reliable

because

uses

Tridacna.

III U/Th Bloom et al.,

1974 35 3 24-1353C

Favia

speciosa

IIIb is

accepted s

41 ka,

maybe

there are

from II

III 14C Veeh and

Chappell, 1970 35.4 1.3 AUN 117

Tridacna

gigas

Ages too

young,

maybe

from II?

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132

Ter

race

(Fig

. 2.1

)

Met

hod

olo

gy

Sou

rce

Date

(k

a)

Err

or

Sam

ple

Nam

e

(Typ

e)

Sam

ple

Typ

e

Note

s

III 14C Veeh and

Chappell, 1970 35.8 1.5 ANU 116

Tridacna

gigas

Ages too

young,

maybe

from II?

IIIb Pa/U Cutler et al., 2003 37.9 0.9 KWA-K-1 (B) Porites sp. Some

diagenesis

IIIb U/Th Aharon,

Chappell 1986 40.3 3.5 Reef IIIb crest 4 coral

III U/Th Bloom et al.,

1974 42 3 25-1353D

Hydnophor

a exesa

IIIb is

accepted

as 41 ka

by author

III U/Th Bloom et al.,

1974 42 3 26-1347D

Goniastrea

parvistella

IIIb is

accepted

as 41 ka

by author

III U/Th Bloom et al.,

1974 42 3 42-1351E

Symphyllia

mobilis

IIIb is

accepted

as 41 ka

by author

IIIb Pa/U Cutler et al., 2003 43.9 1.5 KWA-K-1 (A) Porites sp. Some

diagenesis

III U/Th Veeh and

Chappell, 1970 46 3 ANU 150

Tridacna

gigas

Not

reliable

because

uses

Tridacna

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133

Ter

race

(Fig

. 2.1

)

Met

hod

olo

gy

Sou

rce

Date

(k

a)

Err

or

Sam

ple

Nam

e

(Typ

e)

Sam

ple

Typ

e

Note

s

IIIa Pa/U Cutler et al., 2003 46.3 1.3 KAN-D-4 Porites sp. Some

diagenesis

IIIb U/Th Cutler et al., 2003 47.18 0.2 KWA-K-1 (B) Porites sp. Some

diagenesis

IIIb U/Th Cutler et al., 2003 48.56 0.3

2 KAN-C-2 Favia laxa

Some

diagenesis

IIIa U/Th Cutler et al., 2003 48.76 0.3

6 KAN-D-4 Porites sp.

Some

diagenesis

III U/Th Veeh and

Chappell, 1970 49 3 NG 601 (C) coral

IIIb U/Th Cutler et al., 2003 49.81 0.2 KWA-K-1 (A) Porites sp. Some

diagenesis

IIIa U/Th Aharon,

Chappell 1986 51 2.8

Reef IIIa

transgression 2 coral

III U/Th Veeh and

Chappell, 1970 53 3 NG 600 coral

IIIa U/Th Cutler et al., 2003 60.57 0.2

6 KWA-Q-1

Gardinerose

ris planulata

No

diagenesis;

Ages are

too old,

maybe

from IV?

IIIa Pa/U Cutler et al., 2003 60.8 0.8 KWA-Q-1 Gardinerose

ris planulata

No

diagenesis;

Ages are

too old,

maybe

from IV?

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134

Ter

race

(Fig

. 2.1

)

Met

hod

olo

gy

Sou

rce

Date

(k

a)

Err

or

Sam

ple

Nam

e

(Typ

e)

Sam

ple

Typ

e

Note

s

IV U/Th Bloom et al.,

1974 48 3 6-1351A

Acropora

sp.

Species

thought to

not be

reliable by

author.

IV U/Th Bloom et al.,

1974 57 4 7-1351G

Favia

pallida

IV U/Th Bloom et al.,

1974 58 4 3-1347A

Favia

stelligera

IV U/Th Veeh and

Chappell, 1970 60 6 NG 623

Tridacna

gigas

Judged

reliable.

IV U/Th Aharon,

Chappell 1986 60.4 3.5 Reef IV crest

1 Tridacna

gigas; 4

coral

IV U/Th Bloom et al.,

1974 61 4 4-1351C

Favia

pallida

IV U/Th Bloom et al.,

1974 66 4 45-1347E

Hydnophor

a micronos

IV U/Th Cutler et al., 2003 68.07 0.3

1 SIAL-E-1

Montipora

sp.

Some

diagenesis

IV U/Th Veeh and

Chappell, 1970 74 4 NG 625

Tridacna

gigas

Judged

reliable.

V U/Th Bloom et al.,

1974 61 4 8-1347F

Platygyra

lamellina

Disregarde

d by

author,

probably

from

terrace IV.

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135

Ter

race

(Fig

. 2.1

)

Met

hod

olo

gy

Sou

rce

Date

(k

a)

Err

or

Sam

ple

Nam

e

(Typ

e)

Sam

ple

Typ

e

Note

s

V U/Th Bloom et al.,

1974 84 4 12-1347B

Porites

lutea

Author

accepts

age of 85

ka.

V Pa/U Cutler et al., 2003 84.6 4.9 KWA-U-1 (A) Porites sp. Some

diagenesis

V U/Th Aharon,

Chappell 1986 85 1.4 Reef V crest 2 coral

V U/Th Bloom et al.,

1974 86 4 38-1353E

Goniastrea

pectinata

Author

accepts

age of 85

ka.

V U/Th Cutler et al., 2003 91.61 0.5

2 KWU-U-1 (B) Porites sp.

Some

diagenesis

V U/Th Cutler et al.,

2003 92.57

0.4

5 KWA-S-1 (A) Porites sp.

No

diagenesis

V U/Th Cutler et al.,

2003 92.6

0.5

1 KWA-S-1 (B) Porites sp.

No

diagenesis

V Pa/U Cutler et al.,

2003 92.7 5.2 KWA-S-1 (A) Porites sp.

No

diagenesis

V U/Th Cutler et al., 2003 93.06 1.8

9 KWA-U-1 (A) Porites sp.

Some

diagenesis

V Pa/U Cutler et al.,

2003 94.4 2.3 KWA-S-1 (B) Porites sp.

No

diagenesis

VI U/Th Aharon,

Chappell 1986 107 7.5 Reef VI crest 2 coral

Page 150: TRIDACNA GIGAS - Acumenacumen.lib.ua.edu/content/u0015/0000001/0002339/u0015...The giant clam, Tridacna gigas, is the largest known bivalve and one of the largest members of the Phylum

136

Ter

race

(Fig

. 2.1

)

Met

hod

olo

gy

Sou

rce

Date

(k

a)

Err

or

Sam

ple

Nam

e

(Typ

e)

Sam

ple

Typ

e

Note

s

VI U/Th Bloom et al.,

1974 107 9 14b-1353A

Favia

speciosa

Author

accepts

age of 107

ka.

VI U/Th Bloom et al.,

1974 107 6 20-1347C

Hydnophor

a micronos

Author

accepts

age of 107

ka.

VI Pa/U Cutler et al., 2003 117.6 6.7 SIAL-Q-1 Porites sp. Some

diagenesis

VI U/Th Cutler et al., 2003 119.34 0.7

6 SIAL-Q-1 Porites sp.

Some

diagenesis

VI U/Th Cutler et al., 2003 121.87 0.7

8 SIAL-Q-2 Porites sp.

Some

diagenesis

VI Pa/U Cutler et al., 2003 126.7 8.1 SIAL-Q-2 Porites sp. Some

diagenesis

VIIb Pa/U Cutler et al., 2003 98.7 5.5 KIL-5 Porites sp. Some

diagenesis

VIIb Pa/U Cutler et al., 2003 109.8 2.8 KIL-3 (c) Gardinerose

ris planulata

Some

diagenesis

VIIb U/Th Cutler et al., 2003 113.9 0.6

5 KIL-3 (b)

Gardinerose

ris planulata

Some

diagenesis

VIIb U/Th Cutler et al., 2003 115.36 0.6

6 KIL-3 (D)

Gardinerose

ris planulata

Some

diagenesis

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137

Ter

race

(Fig

. 2.1

)

Met

hod

olo

gy

Sou

rce

Date

(k

a)

Err

or

Sam

ple

Nam

e (T

yp

e)

Sam

ple

Typ

e

Note

s

V* U/Th Veeh and

Chappell, 1970 116 7 NG 618 coral

*Bloom et

al. (1974)

calls this

sample

terrace

VIIb

VIIb U/Th Cutler et al., 2003 116.16 1.8 KIL-3 (c) Gardinerose

ris planulata

Some

diagenesis

VIIb U/Th Stein et al., 1992 116.4 1.8 KIL-5b Porites

lutea

VIIb U/Th Cutler et al., 2003 116.8 1.1

5 KIL-5 Porites sp.

Some

diagenesis

VIIb U/Th Stein et al., 1992 117.6 1.2 KIL-5 (a-2) Porites

lutea

VIIb U/Th Cutler et al., 2003 117.77 0.6

9 KIL-3 (a)

Gardinerose

ris planulata

Some

diagenesis

VIIb U/Th Aharon,

Chappell 1986 118 2

Reef VIIb

crest coral

VIIb U/Th Stein et al., 1992 118.1 1 HP-47

Gardinerose

ris

planulata

VIIa U/Th Stein et al., 1992 118.7 1.4 KAM-A-1 Platygyra

lamellina

KAM-A

are

different

but valid

Page 152: TRIDACNA GIGAS - Acumenacumen.lib.ua.edu/content/u0015/0000001/0002339/u0015...The giant clam, Tridacna gigas, is the largest known bivalve and one of the largest members of the Phylum

138

Ter

race

(Fig

. 2.1

)

Met

hod

olo

gy

Sou

rce

Date

(k

a)

Err

or

Sam

ple

Nam

e (T

yp

e)

Sam

ple

Typ

e

Note

s

V* U/Th Veeh and

Chappell, 1970 119 7 NG 618 coral

*Bloom et

al. (1974)

calls this

sample

terrace

VIIb

VIIb U/Th Stein et al., 1992 119.5 1.2 KIL-5 (a-1) Porites

lutea

VIIc U/Th Stein et al., 1992 123.8 1.3 SIAL-M-3 Cyphastrea

serailia

VIIb U/Th Stein et al., 1992 131.9 1.2 HP-23b Gardinerose

ris

V* U/Th Veeh and

Chappell, 1970 133 10 NG 616 coral

*Bloom et

al. (1974)

calls this

sample

terrace

VIIa

VIIc U/Th Stein et al., 1992 134 1.9 PI-155 Platygyra

n.i.

Substantial

diagenesis

VIIb U/Th Stein et al., 1992 134.7 1.3 HP-23a Gardinerose

ris

VIIb U/Th Stein et al., 1992 135.8 1.9 HP-22

Gardinerose

ris

planulata

VIIa U/Th Stein et al., 1992 136.2 2.5 HP-16c Favia

pallida

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139

Ter

race

(Fig

. 2.1

)

Met

hod

olo

gy

Sou

rce

Date

(k

a)

Err

or

Sam

ple

Nam

e

(Typ

e)

Sam

ple

Typ

e

Note

s

VIIb U/Th Stein et al., 1992 136.5 2.3 KIL-4 Porites

lutea

VIIb U/Th Aharon,

Chappell 1986 138 5

Reef VIIa

crest 2 coral

V* U/Th Veeh and

Chappell, 1970 140 10 NG 616 coral

*Bloom et

al. (1974)

calls this

sample

terrace

VIIa

VIIa U/Th Stein et al., 1992 140.8 1.5 HP-16b Favia

pallida

VII U/Th Bloom et al.,

1974 142 8 15-1347G

Porites

lutea

Author

suggests

VIIa

underlies

VIIb and

grades up

into VIIb.

VIIa U/Th Stein et al., 1992 146.4 2.6 HP-17 Favia

pallida

VIIa U/Th Stein et al., 1992 151.7 2.4 HP-16a Favia

pallida

All HP are

different,

although

from same

specimen.

VIIb U/Th Stein et al., 1992 166.9 3.5 SIAL-M-1 Platygyra

sinesis

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140

Ter

race

(Fig

. 2.1

)

Met

hod

olo

gy

Sou

rce

Date

(k

a)

Err

or

Sam

ple

Nam

e (T

yp

e)

Sam

ple

Typ

e

Note

s

VIIa U/Th Stein et al., 1992 196.1 3.4 KAM-A-2 Hydnophora

microconos

VIII U/Th Stein et al., 1992 198.7 4.6 SIAL-B-1 Plesiastea

curta

VIII U/Th Stein et al., 1992 225.9 3.1 SIAL-D-1 Favia

pallida

Likely not

reliable.