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Development and application of subfossil chironomid-based methods for late
Quaternary climate reconstructions in eastern Australia
Jie Christine Chang
BE (Chemical Engineering)
GDip NatResSt
A thesis submitted for the degree of Doctor of Philosophy at
The University of Queensland in 2015
School of Geography, Planning and Environmental Management
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Abstract
The lack of tools to quantify past climate has hindered the development of our knowledge of
climate change in the late Quaternary in eastern Australia. This thesis developed two methods using
subfossil remains of non-biting midge larvae (Chironomid, Diptera: Chironomidae) preserved in
lake sediments to reconstruct past changes in the Australian climate system during and since the
Last Glacial Maximum (LGM). Firstly a transfer function model for reconstructing past summer
temperatures based on the temperature tolerance of modern Australian chironomid taxa was
created. Secondly the stable oxygen and hydrogen isotope composition (δ18
O, δ2H) of the chitinous
head capsules was used to determine temperature changes from chironomids from the same lakes.
Both techniques were tested and the complexities of applying these methods as independent
temperature proxies in Australia were explored.
In order to develop the transfer function, forty-five lakes were examined in eastern Australia for
eighteen physical and chemical parameters. The physical and chemical analysis of the lakes
suggests that sub-humid and semi-arid sites are naturally eutrophic, whereas high altitude and
actively managed sites were mostly mesotrophic. Thirty-four lakes were subsequently used in the
transfer function. The transfer function first axis was Mean February Temperature which explains
9.5% of the variance and the secondary axis is pH which also explains 9.5% of the variance.
Despite these low values the function appears robust. The transfer function had an r2
jack-knifed of 0.69
and a root mean squared error of prediction (RMSEP) of 2.33°C.
The transfer function method was then applied to subfossil chironomids from a subtropical site on
North Stradbroke Island, Australia that spans the LGM and the last deglaciation. This has provided
the first quasi-continuous quantitative temperature reconstruction from mainland terrestrial
Australia covering these critical periods (between c. ~ 23.2 and 15.5 cal ka BP). The results of this
reconstruction show a maximum cooling of c. ~6.5°C had occurred at c. ~18.5 cal ka BP during the
LGM. A rapid warming followed and temperatures reached near Holocene values by c. ~17.3 cal ka
BP. The warming trend started at c. ~18.1 cal ka BP and is consistent with the start of deglaciation
from Antarctic records. The records suggest high latitude influencing of subtropical climate during
the LGM.
Stable isotopes (δ18
O and δ2H) from a single genus of chironomid head capsules (Chironomus spp.)
were evaluated to reconstruct past temperatures also. Results suggest that both δ18
O and δ2H are
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potentially valuable tools for reconstructing temperature in humid and low nutrient regions of
Australia (e.g. Tasmania and the southeastern highlands). The correlation between δ18
O of
chironomid head capsules and temperature is consistent with observations from Europe and
indicates that there is potential for this technique. However, the collection of enough material from
ancient sites is challenging and the use of stable isotopes is recommended for late Holocene and
modern studies, perhaps focussed on nutrient changes rather than temperature.
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Declaration by author
This thesis is composed of my original work, and contains no material previously published or
written by another person except where due reference has been made in the text. I have clearly
stated the contribution by others to jointly-authored works that I have included in my thesis.
I have clearly stated the contribution of others to my thesis as a whole, including statistical
assistance, survey design, data analysis, significant technical procedures, professional editorial
advice, and any other original research work used or reported in my thesis. The content of my thesis
is the result of work I have carried out since the commencement of my research higher degree
candidature and does not include a substantial part of work that has been submitted to qualify for
the award of any other degree or diploma in any university or other tertiary institution. I have
clearly stated which parts of my thesis, if any, have been submitted to qualify for another award.
I acknowledge that an electronic copy of my thesis must be lodged with the University Library and,
subject to the policy and procedures of The University of Queensland, the thesis be made available
for research and study in accordance with the Copyright Act 1968 unless a period of embargo has
been approved by the Dean of the Graduate School.
I acknowledge that copyright of all material contained in my thesis resides with the copyright
holder(s) of that material. Where appropriate I have obtained copyright permission from the
copyright holder to reproduce material in this thesis.
Jie Chang
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Publications during candidature
Peer-reviewed Journal articles Incorporated into the Thesis
Chang, J., Woodward, C., Shulmeister, J. (2014). A snapshot of the limnology of eastern Australian
water-bodies spanning the tropics to Tasmania: The land-use, climate, limnology nexus.
Marine and Freshwater Research: 65, 872-883
Chang, J., Shulmeister, J., Woodward, C. (2015a). A chironomid based transfer function for
reconstructing summer temperatures in south eastern Australia. Paleogeography,
Paleoclimatology, Paleoecology: 423, 109-121
Chang, J., Shulmeister, J. Woodward, C., Steinberger, L., Tibby, J., Barr, C. (2015b). A
chironomid-inferred summer temperature reconstruction from subtropical Australia during
the last glacial maximum (LGM) and the last deglaciation. Quaternary Science Reviews:
122, 282-292
Chang, J., Shulmeister, J., Woodward, C., Michalski, G. (Submitted). Can stable oxygen and
hydrogen isotopes from Australian subfossil chironomid head capsules be used as proxies
for past environmental change? Limnology and Oceanography
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Conference Presentations
Chang, J., Woodward, C., Shulmeister, J., Zawadzki, A., Jacobsen, G. 2012. New data from Little
Llangothlin Lagoon, New England Tablelands, eastern Australia, indicate no significant
post-European erosion. In: 15th Biennial Conference of the Australia and New Zealand
Geomorphology Group (ANZGG), Bundanoon, Australia, 2nd
-7th
December, 2012
Chang, J., Shulmeister, J., Woodward, C. 2013. Developing methods that use Australian
chironomid (non-biting midge) larvae as proxies for past climate change. In: Asia-Oceania
Geoscience Society (AOGS) 10th Annual Meeting, Brisbane, Australia, 24th
-28th
June,
2013
Chang, J., Shulmeister, J., Theiling, B. 2013. Application of stable isotopes from Australian
chironomid (non-biting midge) head capsules as proxies for past climate change. In: 12th
Australasian Environmental Isotope Conference (AEIC), Perth, Australia, 10th
-12th
July,
2013 [awarded Australian Nuclear Science Technology Organization (ANSTO) Student
Travel Grant]
Chang, J., Shulmeister, J., Woodward, C. 2014. Development and application of a chironomid
based transfer function for reconstructing summer temperatures in south eastern Australia.
In: Australasian Quaternary Biennial Conference 2014, Mildura, Australia, 30th
June – 3rd
July, 2014
Chang, J., Shulmeister, J., Woodward, C., Steinberger L., Tibby, J. 2014. Using chironomid-based
transfer function and stable isotopes for reconstructing past climate in south eastern
Australia. In: American Geophysical Union Fall Meeting 2014, San Francisco, USA, 15th
–
19th
Dec, 2014
Chang. J., Shulmeister, J., Woodward, C., Steinberger L., Tibby, J. Barr, C. 2015. (Accepted for
oral presentation) A high resolution chironomid-based summer temperature reconstruction
from North Stradbroke Island, Australia, during the last glacial maximum (LGM). In: XIX
International Quaternary Association (INQUA) Congress 2015, Nagoya, Japan, 28th
July –
1st Aug, 2015 [awarded INQUA Early Career Researchers Travel Grant]
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Chang, J., Shulmeister, J., Woodward, C. Michalski, G. (Accepted for poster presentation). The
potential for applying stable oxygen and hydrogen isotopes in subfossil chironomids from
southeastern Australia as proxy for past changes. In: XIX International Quaternary
Association (INQUA) Congress 2015, Nagoya, Japan, 28th
July – 1st Aug, 2015
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Publications included in this thesis
Publication citation – incorporated as Chapter 3.
Chang, J., Woodward, C., Shulmeister, J. (2014). A snapshot of the limnology of eastern Australian
water-bodies spanning the tropics to Tasmania: The land-use, climate, limnology nexus.
Marine and Freshwater Research: 65 (10), 872-883
Contributor Statement of contribution
Jie Chang (Candidate) Research design (70%)
Field work (70%)
Statistical analyses (85%)
Wrote and edited the paper (70%)
Craig Woodward Research design (15%)
Field work (25%)
Statistical analyses (15%)
Wrote and edited the paper (15%)
James Shulmeister Research design (15%)
Field work (5%)
Wrote and edited the paper (15%)
Publication citation – incorporated as Chapter 4.
Chang, J., Shulmeister, J., Woodward, C. (2015). A chironomid based transfer function for
reconstructing summer temperatures in south eastern Australia. Paleogeography,
Paleoclimatology, Paleoecology: 423, 109-121
Contributor Statement of contribution
Jie Chang (Candidate) Research design (70%)
Field work (60%)
Laboratory work (100%)
Statistical analyses (85%)
Wrote and edited the paper (70%)
James Shulmeister Research design (15%)
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Field work (10%)
Wrote and edited the paper (20%)
Craig Woodward Research design (15%)
Field work (30%)
Statistical analyses (15%)
Wrote and edited the paper (10%)
Publication citation – incorporated as Chapter 5.
Chang, J., Shulmiester, J., Woodward, C. Steinberger, L., Tibby, J., Barr, C. (2015). A chironomid-
inferred summer temperature reconstruction from subtropical Australia during the last glacial
maximum (LGM) and the last deglaciation. Quaternary Science Reivews: 122, 282-292
Contributor Statement of contribution
Author Jie Chang (Candidate) Research design (80%)
Laboratory work (80%)
Statistical analyses (95%)
Wrote and edited the paper (70%)
Author James Shulmeister Research design (10%)
Wrote and edited paper (15%)
Author Craig Woodward Research design (5%)
Statistical analyses (5%)
Wrote and edited paper (5%)
Author Lincoln Steinberger Laboratory work (20%)
Author John Tibby Research design (5%)
Field work (50%)
Wrote and edited paper (5%)
Author Cameron Barr Field work (50%)
Wrote and edited paper (5%)
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Publication citation – incorporated as Chapter 6.
Chang, J., Shulmeister, J., Woodward, C. Michalski, G. (submitted to Limnology and
Oceanography). Can stable oxygen and hydrogen isotopes from Australian subfossil chironomid
head capsules be used as proxies for past environmental change?
Contributor Statement of contribution
Author Jie Chang (Candidate) Designed experiments (70%)
Field work (60%)
Laboratory work and analyses (90%)
Statistical analyses (100%)
Wrote the paper (70%)
Author James Shulmeister Designed experiments (20%)
Field work (10%)
Wrote and edited paper (20%)
Author Craig Woodward Designed experiments (10%)
Field work (30%)
Wrote and edited paper (5%)
Author Greg Michalski Laboratory work and equipment training
(10%)
Wrote and edited paper (5%)
Contributions by others to the thesis
Standard contributions were made by the supervision team of James Shulmeister and Craig
Woodward. This includes shared project design and standard editing and critical revision of work,
which substantially improved the quality and clarity of the research. In addition, John Tibby and
Cameron Barr were responsible for the collection and radiocarbon dating of the sediment core from
Welsby Lagoon presented in Chapter 5 (Chang et al., 2015b). Lincoln Steinberger assisted in
laboratory work for picking chironomids from Welsby Lagoon sediment. Greg Michalski was
responsible for the training of using stable isotope equipment at Purdue Stable Isotope Facility.
Statement of parts of the thesis submitted to qualify for the award of another degree
None
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Acknowledgements
There are so many people to thank throughout the course of my PhD candidature. I would like to
start with thanking my principal advisor Jamie Shulmeister. Jamie has played more than just a
supervisor role in the past 3 and half years to me. He has been also a great mentor and a wonderful
friend to me since I started my postgraduate research project with him in early 2011. He has guided
me all the way through till this point and it has not always been easy. Jamie got me interested in this
field and has successfully converted me from being potentially a chemical engineer to a
palaeoecologist. Today, I still think it was one of the best decisions I made. I deeply appreciate and
respect Jamie for his enthusiasm, approachability and vision/insight of our research field. I
particularly thank Jamie for his approach of training me up to a new level to become an independent
researcher and enthusiastic scientist. I would also like to thank Jamie’s family, wife Val and
daughter Niamh for their warm hospitality on our ‘get together’ BBQs/Mexican nights.
I would like to thank Craig Woodward as my other advisor for his substantial input into the project
and paper ideas, but in particular, his input and training for me in the field and the lab. Together
with Jamie, they are an absolutely great supervision team. I was inexperienced with field work
when I started my PhD project, and when I say ‘inexperienced’, I mean ‘never been to any places
out of the city’ to do any work. Craig helped and taught me the art of field work from scratch, from
planning, equipment organisation and moreover, getting the boat out of a lake to get a nice sediment
core out. I’m thankful for Craig’s patience when it comes to species taxonomy. He trained me from
‘how to use the microscope’ and had to put up with questions such as ‘what is this one again? why
does it have extra teeth?’ throughout the first at least six months of my PhD. I would also like to
thank both Jamie and Craig for their thoughtful and careful feedback on papers, manuscript
revisions, and thesis drafts.
I would like to thank John Tibby, Cameron Barr and Lincoln Steinberger for their input into the
work related to Welsby lagoon. I particularly thank John and Cameron for giving me the
opportunity to work on the site and their constructive feedback on my research related to the site.
I’m grateful for the assistance in picking chironomids from Welsby lagoon provided by Lincoln. It
saved me at least a few weeks of time in the lab. I also thank Greg Michalski from Purdue
University for providing stable isotope equipment training to me during my visit at Purdue. I would
also like to thank him and his family for letting me stay at their place and for their warm hospitality
throughout my visit. I would also like to mention Tanya Katzman from Purdue for her assistance in
the stable isotope laboratory. In particular, I would like to thank Matthew Jones from Nottingham
for having fruitful discussions with me about stable isotopes during his visit here at UQ. I would
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also like to thank Andrew Rees for sharing pictures of Tasmanian chironomids with me and this has
greatly helped me to develop my knowledge on taxonomy.
I would like to thank the Department of Primary Industries, Water and Environment (DPIWE), the
Department of Sustainability and Environment (DSE) and the Department of Environment and
Heritage Protection (DEHP) for the permission of sample collection for the project. I have been
very fortunate to have a few wonderful field assistants. I thank Abdollah (Ben) Jarihani, Lydia
Mackenzie for their outstanding assistance during the field seasons in Tasmania and Victoria
respectively. I would also like to mention Adrian Slee for his assistance in lake coring during the
Tasmanian field season. I have been very lucky to have made some wonderful friends through
working on my project also at GPEM and these include Abdollah (who is also a wonderful office
mate), Lydia, Lincoln, Adrian, Dan (Sabrina), David, Nicola, Sui, Emily, Daniel, Dong, Min, Ming
etc.
I am very grateful to have Tim Cohen and Patrick Moss as my academic referees and I would like
to thank them for providing wonderful recommendation letters for me which helped me get a
Postdoc job lined up. I would like to thank my future advisor Patricio Moreno for providing me an
exceptional opportunity to continue my research work in northwestern Patagonia for the next couple
of years. In addition, I would like to thank Martin Williams and Peter Cranston for their
encouragement and interests in my on-going and future research. I would like to thank all the
professional and technical staff at GPEM. They have been a very supportive team. I especially
thank Michael Tobe, Alan Victor and Clwedd Burns for their assistance in setting up laboratory and
field equipment.
I have been very lucky to receive a number of grants to fund my research and to attend conferences.
The ARC discovery grant has provided adequate funding for me to do field work and attend a
number of conferences, including Australia and New Zealand Geomorphology Group meeting in
Bundanoon and Mount Tamborine, Australasian Quaternary Association (AQUA) meeting in
Mildura, the International Quaternary Association (INQUA) early career researchers (ECR)
meeting in Wollongong. The ARC discovery top-up scholarship and the University of Queensland
Research Scholarship allowed me to fully focus on my research. I also received the Graduate
School International Travel Award which allowed me to visit Purdue to do laboratory work. I also
received the postgraduate travel grant provided through Australia Nuclear Science and Technology
Organisation (ANSTO) to attend the Australasian Environmental Isotope Conference in Perth. I
also received the GPEM school travel grant to attend the American Geophysical Union Meeting at
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San Francisco. More recently, I received the INQUA travel grant to attend the congress in Nagoya,
Japan.
Last, I would like to thank my family. My parents have been very supportive and encouraging
throughout my PhD. I am grateful for both of their emotional and financial backing. This PhD
would not be possible without them. I would like to thank my fiancé David McCann and his family
(Dale, Steve, Mel and Dave). I am very lucky to have David to share this journey with me. I believe
this past 3 and half years will be a very special lifetime memory that we will talk about in the
future. I would like to specially thank David for comforting and putting up with me when I was
stressed and frustrated during my PhD. I particularly thank David for being supportive for me to
pursue my career because he knows what makes me happy.
Finally, I would like to thank the two thesis examiners, Professors Ian Walker and Isabelle
Larocque-Tobler for their useful and constructive suggestions.
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Keywords
Southeast Australia, freshwater lakes, subfossil chrionomids, transfer function, temperature,
palaeoclimate reconstruction, last glacial maximum, last deglaciation, stable isotopes
Australian and New Zealand Standard Research Classifications (ANZSRC)
ANZSRC code: 060206, Paleoecology, 45%
ANZSRC code: 040605, Paleoclimatology, 45%
ANZSRC code: 040203, Isotope Geochemistry, 10%
Fields of Research (FoR) Classification
FoR code: 0406, Physical Geography and Environmental Science, 80%
FoR code: 0402, Geochemistry, 10%
FoR code: 0602, Ecology, 10%
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Contents
1 INTRODUCTION ...................................................................................................................... 1
1.1 BACKGROUND ........................................................................................................................ 1
1.2 RESEARCH OBJECTIVES .......................................................................................................... 5
1.3 OVERVIEW OF RESEARCH METHODOLOGY .............................................................................. 6
1.4 THESIS STRUCTURE AND SYNOPSIS ......................................................................................... 9
2 THE BIOLOGY OF CHIRONOMIDS .................................................................................. 11
2.1 THE PRINCIPLES OF APPLYING THE TRANSFER FUNCTION AND STABLE ISOTOPE METHODS IN
PALAEOCLIMATE RECONSTRUCTIONS .............................................................................................. 14
2.1.1 The transfer function method ........................................................................................... 14
2.1.2 The stable isotope method ............................................................................................... 14
3 A SNAPSHOT OF THE LIMNOLOGY OF EASTERN AUSTRALIAN WATER
BODIES SPANNING THE TROPICS TO TASMANIA: THE LAND-USE, CLIMATE,
LIMNOLOGY NEXUS ................................................................................................................... 16
3.1 SUMMARY ............................................................................................................................ 16
3.2 ABSTRACT ............................................................................................................................ 17
3.3 INTRODUCTION ..................................................................................................................... 18
3.3.1 Previous water-chemistry studies of Australian lakes ..................................................... 18
3.3.2 Artificial water bodies ..................................................................................................... 21
3.4 MATERIALS AND METHODS .................................................................................................. 24
3.4.1 Study sites ........................................................................................................................ 24
3.4.2 Sample collection and laboratory analyses ..................................................................... 27
3.4.3 Spatial data ...................................................................................................................... 27
3.4.4 Statistical analyses ........................................................................................................... 28
3.5 RESULTS ............................................................................................................................... 30
3.5.1 Lake-water chemistry ....................................................................................................... 31
3.5.2 PCA results ...................................................................................................................... 33
3.5.3 RDA results ...................................................................................................................... 38
3.6 DISCUSSION .......................................................................................................................... 41
3.6.1 The effect of including reservoirs and other artificial waterways ................................... 41
3.6.2 Salinity, pH and bedrock effects ...................................................................................... 41
3.6.3 Nitrogen and phosphorus limitation ................................................................................ 42
3.6.4 The interaction of human impact and climatic effects on lake nutrient status ................ 42
3.7 CONCLUSIONS ...................................................................................................................... 44
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4 A CHIRONOMID BASED TRANSFER FUNCTION FOR RECONSTRUCTING
SUMMER TEMPERATURES IN SOUTH EASTERN AUSTRALIA ...................................... 45
4.1 SUMMARY ............................................................................................................................ 45
4.2 ABSTRACT ............................................................................................................................ 46
4.3 INTRODUCTION ..................................................................................................................... 47
4.4 MATERIALS AND METHODS .................................................................................................. 51
4.4.1 Study sites ........................................................................................................................ 51
4.4.2 Chironomid collection and analysis ................................................................................ 51
4.4.3 Lake water chemistry and environmental variables ........................................................ 52
4.4.4 Statistical analyses ........................................................................................................... 54
4.5 RESULTS ............................................................................................................................... 56
4.5.1 Chironomid taxa .............................................................................................................. 56
4.5.2 Test for Tasmanian endemism and difference between natural and artificial lakes ....... 56
4.5.3 Selection of environmental variables and model development ........................................ 61
4.5.4 The transfer function ........................................................................................................ 65
4.6 DISCUSSION .......................................................................................................................... 71
4.6.1 Can we include artificial lakes in temperature training sets? ......................................... 71
4.6.2 Endemism and other considerations ................................................................................ 71
4.6.3 Reconciliation and integration with Tasmanian transfer function .................................. 72
4.6.4 Value of the transfer function .......................................................................................... 73
4.7 CONCLUSIONS ...................................................................................................................... 74
5 A CHIRONOMID-INFERRED SUMMER TEMPERATURE RECONSTRUCTION
FROM SUBTROPICAL AUSTRALIA DURING THE LAST GLACIAL MAXIMUM (LGM)
AND THE LAST DEGLACIATION ............................................................................................. 75
5.1 SUMMARY ............................................................................................................................ 75
5.2 ABSTRACT ............................................................................................................................ 76
5.3 INTRODUCTION ..................................................................................................................... 77
5.4 REGIONAL SETTING .............................................................................................................. 78
5.4.1 Climate ............................................................................................................................. 79
5.4.2 Modern circulation of the Eastern Australian Current (EAC) ........................................ 79
5.4.3 Physical environment of Welsby Lagoon ......................................................................... 80
5.5 METHODOLOGY .................................................................................................................... 81
5.5.1 Chironomids sampling and analysis ................................................................................ 81
5.5.2 Chronology ...................................................................................................................... 82
5.5.3 Statistical methods and reconstruction examination ....................................................... 83
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5.6 RESULTS ............................................................................................................................... 84
5.6.1 Chironomid analysis ........................................................................................................ 84
5.6.2 The LGM (~23.2 cal ka BP – 18.1 cal ka BP) ................................................................. 85
5.6.3 The deglacial period (~18.1 cal ka BP – 13 cal ka BP) .................................................. 86
5.6.4 The Holocene (~13 cal ka BP – present) ......................................................................... 86
5.6.5 Relationships between changes in chironomid assemblages and temperature and other
variables ..................................................................................................................................... 89
5.7 DISCUSSION .......................................................................................................................... 93
5.7.1 Precision and reliability of the reconstruction ................................................................ 93
5.7.2 The deglacial and LGM temperatures of North Stradbroke Island and regional climate
97
5.7.3 Wider paleoclimatic considerations ................................................................................ 99
5.8 CONCLUSION ...................................................................................................................... 100
6 CAN STABLE OXYGEN AND HYDROGEN ISOTOPES FROM AUSTRALIAN
SUBFOSSIL CHIRONOMID HEAD CAPSULES BE USED AS PROXIES FOR PAST
ENVIRONMENTAL CHANGE? ................................................................................................. 101
6.1 SUMMARY .......................................................................................................................... 101
6.2 ABSTRACT .......................................................................................................................... 102
6.3 INTRODUCTION ................................................................................................................... 103
6.4 MATERIALS AND METHODS ................................................................................................ 105
6.4.1 Study sites and sample collection .................................................................................. 105
6.4.2 Climatic and stable isotopes in precipitation data ........................................................ 106
6.4.3 Stable isotope sample preparation ................................................................................ 106
6.4.4 Stable isotope analyses .................................................................................................. 108
6.4.5 Statistical analyses ......................................................................................................... 109
6.5 RESULTS ............................................................................................................................. 110
6.5.1 Stable oxygen isotope (δ18
O) analyses results ............................................................... 111
6.5.2 Stable hydrogen isotope (δ2H) results ........................................................................... 119
6.6 DISCUSSION ........................................................................................................................ 124
6.6.1 Analytical considerations .............................................................................................. 124
6.6.2 Relationship between δ18
O of lake water and regional climate .................................... 125
6.6.3 Lake trophic status related δ18
O fractionation process and the metabolism and
respiration of Chironomus spp. ................................................................................................ 126
6.6.4 Relationship between δ18
O of Chironomus spp. and regional climate .......................... 129
6.6.5 The δ2H of precipitation, lake water, Chironomus spp. and temperature ..................... 129
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6.7 CONCLUSIONS AND FUTURE WORK ..................................................................................... 132
7 CONCLUSIONS AND FUTURE WORK ............................................................................ 134
7.1 SUMMARY OF RESEARCH AND MAJOR FINDINGS ................................................................. 134
7.1.1 Connections of modern limnology to climate and land-use .......................................... 134
7.1.2 The transfer function ...................................................................................................... 135
7.1.3 The application of the transfer function: temperature reconstruction covering the Last
Glacial Maximum and the last deglaciation from subtropical Australia ................................. 137
7.1.4 The potential of applying stable oxygen and hydrogen isotopes from Australian subfossil
chironomid head capsules as independent proxies for past change ......................................... 138
7.2 RESEARCH SIGNIFICANCE ................................................................................................... 139
7.3 CRITICAL REFLECTIONS ...................................................................................................... 139
7.3.1 The debate and controversy about transfer function and its application ...................... 140
7.3.2 The limitation of the stable isotope method ................................................................... 141
7.4 RECOMMENDATIONS AND FUTURE WORK ........................................................................... 142
7.4.1 Extension and integration of the modern calibration dataset in Australia ................... 142
7.4.2 Application of the transfer function to quantify long-term (late-Pleistocene to Holocene)
temperature changes from various sites ................................................................................... 143
7.4.3 Further exploration and application of the chironomid δ18
O and δ2H method ............ 144
7.4.4 Regional patterns and hemispheric to global connections ............................................ 144
REFERENCES ............................................................................................................................... 146
APPENDICES ................................................................................................................................ 168
APPENDIX 1. A SUMMARY OF PREVIOUS MODERN CALIBRATION TRAINING SETS
DEVELOPED AND APPLIED FOR PALEO-RECONSTRUCTIONS USING CHIRONOMIDS
...................................................................................................................................................... 168
APPENDIX 2. CHIRONOMID TAXA ENUMERATED FROM THE MODERN
CALIBRATION DATA SET OF SOUTHEASTERN AUSTRALIA .......................................... 175
APPENDIX 3. CHIRONOMID FOSSIL TAXA ENUMERATED FROM WELSBY LAGOON,
NORTH STRADBROKE ISLAND, AUSTRALIA FROM THE LAST GLACIAL MAXIMUM
TO NEAR PRESENT ................................................................................................................... 195
APPENDIX 4. PUBLISHED JOURNAL ARTICLES DURING CANDIDATURE ................... 203
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List of Figures
Figure 1.1 Overview of the framework and project flow .................................................................... 7
Figure 1.2 Overview of the thesis structure ....................................................................................... 10
Figure 2.1 Key morphological characteristics of chironomid larvae (Brooks et al., 2007, Page 4) .. 11
Figure 2.2 Chironomid Life Cycle (Walker, 1987, Page 852) ........................................................... 12
Figure 2.3 A fully developed chironomid head capsule (Chironomus sp.) from Lake Albina, Mt
Kosciuszko, New South Wales, Australia (photo is taken by J. Chang at the Physical
Geography laboratory, School of Geography, Planning and Environmental Management,
University of Queensland) .......................................................................................................... 12
Figure 2.4 Chitin molecule structure.................................................................................................. 13
Figure 3.1 Map of eastern Australia, with the 45 study sites identified; the numbers correspond to
the lake names and numbers in Table 3.1. .................................................................................. 19
Figure 3.2 Tri-plot of major cations, with lakes labelled by type, as follows: CL, coastal lakes
(lowland lakes that are <100 km away from the sea); M, montane lakes (>1000 m asl); IL,
inland lakes (lowland lakes that are >100 km away from the sea). The tri-plot shows cation
concentration as a percentage. All 45 lakes are distributed closer to the world-average seawater
ratio (WASW) than to the world-average freshwater ratio. On average, lowland coastal lakes
are closer to the ratio than inland lakes or those at high elevation. This indicates that the
primary source of the salts in eastern Australian lakes is seawater. ........................................... 32
Figure 3.3 Biplot of pH against conductivity. This demonstrates that pH co-varies with bedrock
type and salinity. All saline lakes are strongly alkaline, whereas there is more variability in the
pH of freshwater lakes. This means that the freshwater lakes are dominantly controlled by
bedrock types. Lakes above log conductivity of 3 are classified as brackish. ............................ 33
Figure 3.4 Initial PCA plot of a) the 45 water bodies constrained to b) the 19 environmental
variables, with land-use, climate, and lake depth variables entered as passive variables to focus
the analysis on in-lake (limnological variables). The first and second axes of this plot are
predominantly specific conductance and pH gradients; it shows a large number of correlated
water conductivity proxies (ion concentrations). ........................................................................ 35
Figure 3.5 Revised PCA plot of a) the 45 water bodies constrained to b) the 12 environmental
variables (ionic variables removed), with land-use, climate, and lake depth variables entered as
passive variables to focus the analysis on the in-lake (limnological) variables. The first axis is
now a nutrient gradient and the second axis is a pH gradient. ................................................... 36
Figure 3.6 Final principal components analysis (PCA) with all environmental data (excluding ion
concentration data) entered as active variables. (a) The 45 water bodies, along with (b)
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morphology, environment, climate and land-use variation patterns. The primary y-axis is a
trophic status (total nitrogen (TN), total phosphorus (TP), Chlorophyll a), conductivity and
agriculture gradient; the second axis is aligned to temperature. The PCA also indicates that
eutrophic saline lakes tend to be shallow lakes located in arid regions. ..................................... 37
Figure 3.7 Principal components analyses (PCAs) to compare the changes in limnology for natural
lakes and artificial lakes along a gradient that reflects lake depth and human impact. The Axis 1
sample scores from plot (a) shows the amount of agricultural activity in the lake catchments.
The Axis 1 sample scores from plot (b) shows the trophic status of the lakes. The Axis 1
sample scores for both PCAs (a and b) are used to visualise the relationship between land use,
aridity and limnology. The regression for (c) all lakes and (d) artificial lakes only. The trend of
the regression is similar between the two graphs, indicating that artificial lakes have traits
similar to those of natural lakes. The lower slope on the artificial lakes may indicate lower
residence times. Overall, these results demonstrate that lakes in agricultural catchments are
more eutrophic; however, see the main text because this trend is overwritten by climatic effects
and lake morphology. ................................................................................................................. 40
Figure 4.1 (a) Plot of observed species richness vs predicted species richness, (b) rarefaction curves
for individual sites indicating estimated species richness with respect to increasing sub-sample
size. Rarefaction curves begin to flatten once true species richness is achieved. Together, these
plots indicate that sample sizes are adequate for most samples to capture all but the most rare
species. Only one site (CTL) with a low head capsule count may possibly underestimate the
true species richness. Minimum sample size varied from site to site and counts as low as 50
may be sufficient to capture the actual species richness (e.g. LPO). .......................................... 53
Figure 4.2 PCA plots for exploring the difference between (a) mainland and Tasmanian lakes and
(b) natural and artificial lakes. PCA axis 1 and axis 2 explains 16.9% and 10.8% of the variance
in chironomid species data respectively. A t-test was performed on the sample score means for
each (Tables 4.1a and 4.1b). The sample size for the Tasmania and artificial lakes is small
(<10). There are no significant differences apparent between axis 1 scores for both tests. There
are no differences in axis 2 scores either for Tasmania vs mainland lakes, but there is for
natural vs artificial lakes. Warm taxa are over-represented in artificial lakes that are not
reservoirs (Fig. 4.3). .................................................................................................................... 57
Figure 4.3 This shows a PCA of chironomid species used to compare natural and artificial lakes. In
a) PCA axis 2 show separation of warm stenotherms (low PCA axis 2 scores, e.g.
Dicrotendipes, Kiefferulus, Cladopelma, Procladius, Paratanytarsus etc.) from cold
stenotherms (high PCA axis 2 scores, e.g. Parakiefferiella). b) shows surface sediment samples
plotted as circles where the circle size is relative to mean February temperature (MFT). Large
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circles = high MFT, small circles = low MFT. Values for MFT for each site are also provided.
Open circles = natural water bodies, closed circles = artificial water bodies. Note that some
artificial water bodies (not reservoirs) with low MFTs have low PCA axis 2 scores, which
indicates that warm stenotherms are more common in these sites compared to natural sites with
similar MFTs. ............................................................................................................................. 58
Figure 4.4 CCA biplots of (a) sample and (b) species scores constrained to seven environmental
variables that individually explain a significant (p ≤ 0.05) proportion of the chironomid species
data. Sites codes correspond to site names in Tables 3.1 and 3.2 (lake numbers 1-34). Taxon
numbers correspond to taxa in Table 4.5. Sites and taxa in warmer environments tend to plot in
the upper left quadrant. Taxa typical of warmer sites include Harnischia spp.,
Cryptochironomus spp., Polypedilum spp., Coelopynia pruinosa type, Cladopelma spp.,
Paratanytarsus spp., Procladius spp., and Riethia spp., while sites and taxa in colder
environments tend to plot in the lower right hand quadrant. Taxa typical of cold sites include
Paralimnophyes morphotype 3, Parakiefferiella morphotype 1, Orthoclad type 1, Orthoclad
type 4, Pseudosmittia type 2, and Botrycladius. Eutrophic sites and taxa typical of these
environments tend to plot in the lower left hand quadrant, while oligotrophic sites and taxa
typical of these environments tend to plot in the upper right hand quadrant. ............................ 66
Figure 4.5 Taxon response curves for taxa that show a significant response to temperature (p < 0.05)
using a Generalized Linear Model with Poisson distribution (ter Braak and Šmilauer, 2002). (a)
Taxa which are more common at lower temperatures, such as Paralimnophyes morphotype 1
and Parakiefferiella morphotype 2 respond strongly to cooling in temperature (b) Taxa which
are more common at higher temperatures demonstrate weaker but still significant responses.
Examples include Procladius, Polypedilum spp. and Tanytarsus lactescens type. .................... 67
Figure 4.6 Stratigraphy diagram of the 38 non-rare taxa included in the final model, where observed
mean February temperature is on the y-axis and taxon abundance is in percentage. Taxa such as
Cladopelma, Dicrotendipes and Kiefferulus (*) show high abundance in lowland warm lakes
but are also present in highland artificial lakes (Grey bars). ...................................................... 68
Figure 4.7 Performance of the three component PLS model where (a) shows the predicted versus
observed mean February temperature and (b) displays residuals of the predicted versus
observed mean February temperature. Note that the model has a potential to over predict
temperatures from some very shallow high altitude lakes by up to ~6 °C. These lakes have
increased mean water temperature in summer and chironomid assemblages may resemble
lower elevation sites. .................................................................................................................. 73
Figure 5.1 Location of the study site and other sites discussed in this study. (a) A summary of the
surface currents within the Tasman Sea. The denoted by A, B, C, and D shows the direction of
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flow of the relevant part of the EAC along with where the SST records are mentioned. Note
that B is associated with the Tasman Front (Ridgway and Dunn, 2003). The locations of marine
cores PC-27 (Dunbar and Dickens, 2003), GC-12 (Bostock et al., 2006), GC-25 (Troadson and
Davies, 2001), PC-9 (Troadson and Davies, 2001) along the EAC, terrestrial temperature
estimates from Lake Victoria (Miller et al., 1997), the Snowy Mountains (Galloway, 1965), and
Lake Selina (Fletcher and Thomas, 2010) are displayed (b) Location of North Stradbroke
Island (NSI) and Lake Mackenzie, Fraser Island (c) Location on North Stradbroke Island of
Welsby Lagoon and other sites mentioned in the text, including Tortoise Lagoon (Petherick et
al., 2008; Moss et al., 2013), Blue Lake (Barr et al., 2013), Eighteen Mile Swamp (sampling
location) and Native Companion Lagoon (McGowan et al., 2008; Moss et al., 2013). ............. 81
Figure 5.2 Radiocarbon and calibrated ages for Welsby Lagoon with respect to depth. Open shapes
represent calibrated ages calibrated using SHCal13 (Hogg et al., 2013). .................................. 82
Figure 5.3 Chironomid stratigraphy from Welsby Lagoon plotted with respect to the radiocarbon
chronology. Chironomid inferred mean February temperatures (MFTs) are shown in (a) and the
number of head capsules counted for each sample are shown in (b). The dashed line in (a)
indicates the current mean February temperature at Welsby Lagoon and the dashed line in (b)
indicates the minimum number of head capsules for a quantitative reconstruction from a
sample. All fossil taxa present in the Welsby Lagoon record are included in the diagram. Age
in calibrated years is on the y-axis and taxa were grouped according to their optima in the
modern training sets based on Chapter 4. Grey areas represent sediment layers where
chironomids were absent. Note that Dicrotendipes, a benthic taxon that does not have a
significant correlation (p > 0.05) to MFT in the modern calibration data set, is dominant in all
four Holocene samples. Each of the taxon was photographed and included in Appendix 3. ..... 88
Figure 5.4 Reconstruction diagnostics for Welsby Lagoon samples. (a) Reconstructed mean
February temperatures (MFTs) with the RMSEP (2.2 °C) from the modern training set
(Chapter 4) plotted in dashed horizontal lines for each sample. (b) Goodness-of-fit statistics
where the fossil samples are passively fitted to the CCA ordination axis derived from the
modern training set constrained to MFT. The vertical dashed line represents the 95th
percentiles
of modern squared residual lengths beyond which fossil samples are considered to have poor
fits to MFT. (c) Rare taxa plot where closed circles represent fossil samples with well-
represented taxa in the modern training set and open circles represent fossil samples with rare
taxa (Hill's N2 ≤ 5) summing to greater than 10% abundance. (d) Non-analogue plot where
closed circles indicate fossil samples with either close or good modern analogues, whereas
open circles indicate fossil samples with no good modern analogues (at the 5th
percentile cut-
off level). The vertical lines represent the 2nd
, 5th
and 10th
percentiles respectively. ................. 90
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Figure 5.5 The Welsby Lagoon chironomid record was split into three zones using the constrained
incremental sums of squares (CONISS) function in Psimpoll (Bennett, 2002). W-1 represents
the last glacial maximum (LGM), W-2 represents the deglacial and W-3 represents the
Holocene. This zonation is consistent with the observed changes in chironomid assemblages
shown in Fig. 5.3. ....................................................................................................................... 91
Figure 5.6 Trajectory changes of trend through time of fossil samples (black circles) in the Welsby
Lagoon record, passively plotted in a CCA of the training set lakes (open circles, where site 1–
33 are waterbodies from (Chapter 4, Table 4.1) and site 34 is Swallow Lagoon, a waterbody
near Welsby Lagoon with similar physical characteristics) against significant variables (p <
0.05). (a) Samples covering the last glacial maximum (LGM) and deglacial period, where
changes in trend are parallel to the mean February temperature (MFT) gradient, indicating that
MFT is a primary controlling variable and drives the changes of chironomid assemblages in the
samples from c. ∼23.2 to 16.6 cal ka BP. (b) samples covering the period of deglacial and
Holocene from c. ∼16.3 to 0.2 cal ka BP. The trend of changes of chironomid assemblages in
these fossil samples are parallel to the secondary gradient that is represented by nutrients,
conductivity, pH and lake depth, indicating that non-climatic factors are likely dominant in
Holocene changes. ...................................................................................................................... 92
Figure 5.7 Chironomid mean February temperature (MFT) transfer function based reconstruction
from Welsby Lagoon (WEL) (in plot d) compared to (a) the δ18
O and CO2 records from
Antarctica (Stenni et al., 2011; Pedro et al., 2012); (b) insolation curve for seasonality (Berger
and Loutre, 1991); (c) δ18
O measured from G. ruber from marine cores along the East
Australian Current (EAC) and (d) temperature estimated based on branched glycerol dialkyl
glycerol tetraether (GDGT) from Fraser Island (Woltering et al., 2014). The chironomid results
from WEL (d) indicate a maximum cooling relative to the modern MFT along the Southeast
Queensland coast of c. ∼6.5 °C with much lower cooling at other times during the late LGM
and a rapid recovery at about 18.1 cal ka BP. The cooling is significantly less than the 9–12 °C
for the adjacent uplands from periglacial landforms from Galloway (1965), but is in line with
estimates from nearby marine cores shown in (c) and lacustrine record from Lake Mackenzie
(LM), Fraser Island shown in (d) (note the reconstructed temperature from Fraser Island is
lower than Welsby Lagoon because the GDGT is based on mean annual temperatures
calibration whereas chironomid transfer is based on MFT). It is also noteworthy that the
chironomids provide an estimate of summer temperatures whereas the periglacial landforms are
indicators of winter temperature. This suggests a larger seasonal temperature contrast at the
LGM which is consistent with the insolation curves (b). Finally the observed timing and pattern
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is very similar to both the δ18
O curve and CO2 record from Antarctica (a) (Stenni et al., 2011;
Pedro et al., 2012). ...................................................................................................................... 96
Figure 6.1 Sample lakes are distributed across southeast Australia but there is a strong concentration
(ten out of 16 lakes) in western Victoria. This reflects Chironomus spp. abundance in
waterways with the tolerant Chironomus spp. strongly represented in the brackish and saline
lakes of western Victoria. ......................................................................................................... 107
Figure 6.2 Chironomus spp. head capsules from (a) Freshwater Lake, Victoria, 20 ×. (b) Lake
Albina, Mount Kosciuszko, New South Wales, 10 ×. Photos of head capsules were taken by J.
Chang at the Physical Geography Laboratory of School of Geography, Planning and
Environmental Management, University of Queensland .......................................................... 109
Figure 6.3 (a)Plot of δ18
O of precipitation against δ18
O of lake water. The δ18
O of precipitation and
lake water is correlated but significant enrichment of lake water is observed at δ18
O of
precipitation of around -5 ‰ V-SMOW, which the offset is possibly due to high regional
evaporation rate and prolonged residence time or local hydrological conditions (b) the plot of
δ18
O of lake water against Chironomus spp. showed no signifcant correlation (P = 0.13)
between the two data sets, suggesting that not all oxygen stable isotopic composition of
Chironomus spp. head capsules were reflecting changes of their ambient water .................... 110
Figure 6.4 (a) Plot of aridity index (AI) against δ18
O of Chironomus spp.. AI has the best correlation
and is the independent variable that explains the largest percent variance (Table 1) of
Chironomus spp. δ18
O data. At semi-arid and sub-humid areas (e.g. when AI <1), the variation
in δ18
O of Chironomus spp. data is large. (b) Total Nitrogen (TN) against fractionation factor
(α[δ18O(chiro-water)]). TN explains an independent and large percent of variance in the fractionation
factor between Chironomus spp. and host lake water (Table 6.2), suggesting lake
eutrophication is an important consideration for the interpretation of stable isotope data for
Chironomus spp. ....................................................................................................................... 111
Figure 6.5 (a)Plot of δ2H of precipitation against δ
2H of lake water. The δ
2H of precipitation and
lake water is closely correlated, with r2 = 0.92 (b) Plot of δ
2H of lake water aginst δ
2H of
Chironomus spp.. δ2H of lake water and Chironomus spp. showed a close correlation (r
2= 0.69)
suggesting that a large proportion of δ2H stable isotopic composition of Chironomus spp. head
capsules that were directly derived from their ambient water or indirectly from feeding on
planktonic algae, where algae also contain δ2H stable isotopes that are derived from lake water.
.................................................................................................................................................. 112
Figure 6.6 (a) Plot of Mean annual temperature (MAT) against δ2H of Chironomus spp.. MAT had
the best correlation and is the independent variable that explains the largest percent variance
(Table 6.5) of Chironomus spp. δ2H data. The correlation could possibly be complicated by the
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fractionation between δ2H of Chironomus spp. and δ
2H of lake water (α[δ2H(chiro-water)]), which
was influenced by other environmental variables. (b) Plot of ln(COND) against fractionation
factor α[δ2H(chiro-water). Specific conductance (COND) explained an independent and large
percent of variance in the fractionation factor between δ2H of Chironomus spp. and host lake
water (Table 6.6), lake water COND had a close correlation (r2 = 0.695) with the fractionation
factor (α) between δ2H of Chironomus spp. and lake water. .................................................... 116
Figure 6.7 Plot of mean annual temperature (MAT) against δ18
O of precipitation for the 16 study
lakes. There is a positive correlation between MAT and δ18
O of precipitation, but since the
differences within the spatial distribution for both study sites and precipitation sources are
large, the data points are scattered. ........................................................................................... 126
Figure 6.8 Plot of the experimental chironomid δ18
O values against δ18
O of host water in Wang et al
(2009) compared to values derived from this study. Eight of the lakes from this study (CTL,
BL, LA, WP, LEA, LLL, LSP, LTK), fall along the regression line of Wang et al (2009) these
are oligotrophic and mesotrophic lakes from Kosciuszko and Tasmania, and two western
Victorian lakes that are less enriched in δ18
O, i.e. less affected by prolonged residence time.
The other eight lakes (FWL, LEM, LCT, LTP, SWL, LFY, LMB, NGR) produced similar
Chironomus spp. δ18
O values. .................................................................................................. 127
Figure 6.9 (a) Plot of mean annual temperature (MAT) against δ18
O of Chironomus spp. with all
sixteen lakes included (b) Plot of mean annual temperature (MAT) against δ18
O of Chironomus
spp. with only the eight lakes that have the δ18
O enrichment of lake water less than ~8‰
VSMOW relative to precipitation δ18
O are included. There is a close correlation (r2 = 0.78)
between MAT and Chironomus spp. δ18
O for this subset of study lakes. The slope and intercept
values were very similar to those observed by Verbruggen et al. (2011). ................................ 130
Figure 6.10 Plot of ln(COND) against mean annual temperature (MAT). Eight out of ten lakes from
Western Victoria are considered saline. Lake Fyans (LFY) and Nuggetty Gully Reservoir
(NGR) were the only lakes that had a specific conductance value < 500 μs/cm from this region.
Lake order effect and residence time are potentially affecting the observed relationship between
temperature and specific conductance and the overall pattern is presumably due to that regional
evaporation is a function of temperature .................................................................................. 131
Figure 6.11 Eight lakes that had a ln(COND) < ~ 6 were plotted for mean annual temperature
against δ2H of Chironomus spp.. A close correlation was obtained with an r
2 of 0.72. These
include two lakes (Lake Fyans and Nuggetty Gully Reservoir) from western Victoria and six
lakes from non-arid environments. These two western Victorian lakes are freshwater bodies
presumably due to short residence times. ................................................................................. 132
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List of Tables
Table 3.1 Summarised information of the 45 water bodies sampled ................................................ 23
Table 3.2 Selected limnological and land-use/land cover variables for the forty-five water bodies . 29
Table 3.3 List of the 19 variables used, with mean, minimum and maximum values cited and the
appropriate units highlighted ...................................................................................................... 31
Table 3.4 Partial redundancy analysis (RDA) results for trophic status as limnological variable. This
table demonstrates that the effect of climate and water depth is more significant than the
agricultural status of the basin. In short, shallow water bodies in semi-arid settings are more
eutrophic than are other lakes. AI, aridity index; Chl a, Chlorophyll a; TN, total nitrogen; TP,
total phosphorus .......................................................................................................................... 38
Table 4.1 Selected climatic and environmental variables for the thirty-four water bodies sampled
from southeast Australia used for transfer function development .............................................. 50
Table 4.2 Statistical t-Tests for two samples assuming unequal variances performed on (a) natural
against artificial lakes and (b) Tasmania against mainland lakes. PCA scores of axis 1 and axis
2 were used from both datasets from Fig. 4.2a and Fig. 4.2b respectively. ............................... 58
Table 4.3 Wright and Burgin (2007) identified 5 genera, 4 species and 14 morpho-species of
endemic chironomids in Tasmania. These are arranged in alphabetic order, below. We
performed a literature search and show that all the 5 genera and 4 species were previously
recorded and described in mainland Australia, therefore endemism is not supported at these
taxonomic levels. The 14 morpho-species are not described and we are unable to assess the
reliability of the endemism claim for these taxa. (*) Indicates undescribed morpho-species. ... 60
Table 4.4 CCA summary of the seven significant variables including canonical co-efficients and t-
values of the environmental variables with the ordination axes including 33 lakes and 38 non-
rare species .................................................................................................................................. 62
Table 4.5 Partial CCAs of the seven significant (p ≤ 0.05) environmental variables alone and with
the effects of other significant variables partialled out for 33 lakes with 38 non-rare species
included. ...................................................................................................................................... 63
Table 4.6 List of chironomid taxa enumerated in this study along with data on distribution and
environmental significance. ........................................................................................................ 69
Table 4.7 Performance of partial least squares (PLS) model for reconstructing mean February
temperature of southeast Australia using 33 lakes and 38 non-rare chironomid species. The
bold indicates the model of choice. ............................................................................................ 74
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Table 5.1 Radiocarbon Dates for Welsby Lagoon. Data shown includes sample depth, laboratory,
materials used for dating, 14
C age and calibrated age (SHCal13; Hogg et al., 2013) for each of
the samples. (*) indicates samples excluded in the final age depth model. ................................ 83
Table 6.1 Summary of information available for the 16 lakes included in this study. The complete
data-set is available in Chapter 3. ............................................................................................. 113
Table 6.2 Stable isotope analyses (δ18
O and δ2H) results of southeastern Australian modelled
precipitation, lake water, Chironomus spp. and the fractionation factor between chironomus
head capsules and lake water (α). ............................................................................................. 115
Table 6.3 Redundancy Analysis (RDA) of δ18
O of Chironomus spp. head capsules and
environmental variables shows that Aridity Index (AI) has the strongest explanatory power
(53.7%) among the five variables that have a significant correlation (P < 0.05). It is not
independent of MAT but it is impossible that it would be independent as temperature plays a
key role in potential evapotranspiration (PET). AI explains significantly more variance than
MAT and MAT is reduced to very low explanatory power when AI is removed. ................... 117
Table 6.4 Redundancy Analysis (RDA) of the fractionation factor (α) (δ18
O between Chironomus
spp. and lake water) and environmental variables shows that Total Nitrogen (TN) is the most
independent variable that explains the largest variance (64.4%) among all seven variables that
are significantly correlated (P < 0.05) for the stable oxygen isotope (δ18
O) fractionation factor
(α[δ18O(Chironomus spp - water)]) (Table 6.2) between Chironomus spp. and host lake water ............. 118
Table 6.5 Redundancy Analysis (RDA) of δ2H of Chironomus spp. head capsules and
environmental variables shows that for δ2H there are no independent variables. MAT explains
the largest variance (55.2%) in stable hydrogen isotope (δ2H) data of Chironomus spp. but
interacts with aridity index (AI), specific conductance (COND) and total nitrogen (TN). ...... 121
Table 6.6 Redundancy Analysis (RDA) of the fractionation factor (α) (δ2H between Chironomus
spp. and lake water) and environmental variables shows that specific conductance (COND) is
the most independent variable that explains the largest amount of variance (68.4%) among all
seven variables that are significantly correlated (P < 0.05) in the hydrogen isotope (δ2H)
fractionation factor (α) between Chironomus spp. and host lake water ................................... 123
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List of equations
Equation 6.1.
Equation 6.2.
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xxviii | P a g e
List of abbreviations, acronyms, symbols and units
14C Radiocarbon
a.s.l Above sea leve
AAR Amino acid racemisation
ACR Antarctic cold reversal
AI Aridity index
ArcGIS Arc Geographic Information System
AUS-INTIMATE Australasian - INTegration of Ice-core, MArine and Terrestrial rEcords
AveBiasjack Average bias (jack-knifed)
BOM Bureau of Meteorology
BP Before Present (1950)
cal. Calender year
CCA Canonical correspondence analysis
CGIAR-CSI Consortium for Spatial Information
Chl a Chlorophyll a
COND Specific conductance
CONISS Constrained incremental sums of squares
DCA Detrended correspondence analysis
DCCA Detrended canonical correspondence analysis
DEM Digital elevation model
DIN Dissloved inorganic nitrogen
DO Dissolved oxygen
DRP Dissolved reactive Phosphorus
EAC Eastern Australian current
ENSO El Niño Southern Oscillation
GC Gas chromatography
GDGT Glycerol dibiphytanyl glycerol tetraethers
GF/F Glass fibre filters
GLMs Generalized linear models
GNIP Global Network of Isotopes in Precipitation
G-ruber Globigerinoides ruber
HydroSHEDS Hydrological data and maps based on SHuttle Elevation Derivatives at
multiple Scales
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IAEA International Atomic Energy Agency
INQUA INternational QUaternary Association
INTIMATE INTegration of Ice-core, MArine and Terrestrial rEcords
IPO Interdecadal Pacific Oscillation
IRMS Isotope ratio mass spectrometry
ka Thousand years
KOH Potassium hydroxide
LGM Last glacial maximum
LM Lake Mackenzie
MAE Mean annual evaporation
MAP Mean annual precipitation
MAT Mean annual temperature
MaxBiasjack Maximum bias (jack-knifed)
MCR Mutual climatic range
MFT Mean February temperature
NOx Nitrogen oxides
NSW New South Wales
ORP Oxidation reduction potential
P:EP Precipitation : Evapotranspiration
PCA Principal component analysis
pCCA Partial canonical correspondence analysis
PET Potential evapotranspiration
PLS Partial least square
PMIP Paleoclimate Modelling Intercomparison Project
PSI Purdue stable isotope
RDA Redundancy analysis
RMSEP Root Mean Square Error of Prediction
SAL Salinity
SAM Southern Annular Mode
SHAPE Southern Hemisphere Assessment of PaleoEnvironment
SHCal13 Southern Hemisphere Calibration curve for radiocarbon dates updated to
2013
SLAP Standard Light Antarctic Precipitation
SqRL Squared residual length
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SST Sea surface temperature
STAB Subtropical anti-cyclonic belt
TALDICE TALos Dome Ice CorE
TC/EA Temperature Conversion Elemental Analyzer
TDS Total dissloved solids
TN Total nitrogen
TP Total phosphorous
TURB Turbidity
USGS U.S. Geological Survey
VIF Variance inflation factor
VSMOW Standard Mean Ocean water
WAFW World average freshwater ratio
WA-PLS Weighted Average - Partial least square
WASW World average seawater ratio
WEL Welsby Lagoon
WMO World Meteorological Organization
YD Younger Dryas
α
β
λ
°C
Fractionation factor for stable isotopes
Model coefficient in Partial Least Square models
Eigen values in multivariate statistical analyses
Degree Celcius
cm Centimeters
km Kilometers
μg
μg/L
Micrograms
Micrograms per liter
μs/cm Microsiemens per centimeter
m Meters
mg/L
mm
Milligrams per litre
Millimeters
δ2H Stable hydrogen isotope
δ18O Stable oxygen isotope
Page 32
Background
1 | P a g e
1 INTRODUCTION
1.1 Background
Climate change is one of the most serious issues facing humanity (Heltberg et al., 2009).
Continuing debates around this issue stem partly from our limited insight into the differences
between natural and anthropogenically driven climate variability (Rosenzweig et al., 2008). There is
also limited understanding on the possible impacts of future environmental change on ecosystems
(Walther et al., 2002). In addition, climate change can be regional and in order to understand global
climate, knowledge on regional climate change is essential (WMO, 2015). In order to address these
gaps in knowledge, we need to understand past climate variations and their effects on ecosystems
and the environment (Caseldine et al. 2010; Schmidt 2010). Most of the instrumental measurements
of climate are available for no longer than a few hundred years and in many countries, such as
Australia, these records are often only available for a few decades. Because of their limited
duration, these data are not ideal to be used to determine long term climate change or to understand
abrupt climate events that have occurred in the geological past. Instead, an alternative
understanding of the climate system can be achieved from the development and application of
proxies that have the potential to provide long term palaeoclimate records. These climate proxies
are often derived from natural archives (Zachos et al., 2001) and are extremely diverse, including
but not limited to lake sediment, speleothems, tree rings, sand dunes, ice cores, and corals.
Proxy based paleoclimate reconstructions with adequate temporal and spatial resolution will help us
gain a more detailed understanding of climate change and ecosystem responses as a whole (e.g.
Alloway et al., 2007). Proxy based paleoclimate reconstructions are also very useful to validate
results from climate models and to improve the capability of global climate models to better predict
future changes (e.g. Riebeek, 2006). One of the key variables that has been targeted for validating
global climate modelling is temperature (Riebeek, 2006). In this thesis, proxies for inferring past
climate change are developed based on the species distribution and stable isotopic composition of
non-biting midges (Diptera: Chironomidae) from southeastern Australia. The method is applied to
late Quaternary sedimentary records from the subtropics to cool mid-latitudes to generate mean
annual and summer temperature reconstructions.
The INTIMATE project (INTegration of Ice-core, Marine and Terrestrial rEcords) initiated by the
International Union for Quaternary Science (INQUA) in 1995 aims to generate high quality
paleoclimate records from various regions around the globe. It also aims to integrate these regional
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Background
2 | P a g e
records into climate stratigraphies, to gain a better understanding of the changes of the global
climate system through time. A number of synthesis articles of the INTIMATE project have been
published throughout the years of 1998 to 2014 (e.g. Blockley et al., 2012). The project has
significantly advanced the knowledge of paleoclimate and paleoenvironmental changes especially
in the North Atlantic where the project is primarily focussed. Compared to the Northern
Hemisphere, the development of proxies and proxy based paleoclimate records in the Southern
Hemisphere has received less attention and has only started to emerge in the last decade or so. This
was partly associated with the establishment of the Australasian INTIMATE (AUS-INTIMATE)
project in 2003 (Alloway et al., 2007; Turney et al., 2007; Reeves et al., 2013; Petherick et al.,
2013). More recently, the Southern Hemisphere Assessment of PaleoEnvironment (SHAPE) project
has been initiated to further improve the palaeoclimate records from the Southern Hemisphere. This
thesis was initiated as part of a project titled ‘The last glacial maximum (LGM) conundrum and
environmental responses of the Australian continent to altered climate states’ funded through an
Australian Research Council (ARC) Discovery grant. That project was framed in the context of
both AUS-INTIMATE and the SHAPE project. This thesis is intended to contribute to the
following objectives of the SHAPE project:
provide robust interpretations of proxies
generate regional reconstructions of past environmental and climatic change
and if possible;
integrate the results into a hemispheric-wide story for key time slices, highlighting the
changes and testing hypotheses for their causes
Geographically, the distribution of proxy based records is rather sparse and non-uniform in
Australia. However, studies (Reeves et al., 2013; Turney et al., 2007; Williams et al., 2009)
suggested that Australia can provide high quality information on likely patterns and directions of
the future changes in the Southern Hemisphere. Firstly, it covers a large climate range from the cool
temperate zone to the tropics and incorporates broad humid, sub-humid, arid and semi-arid zones.
Secondly, it is situated between two regions that are critical to global climate; the Indonesian
Maritime Continent to the north, and Antarctica and the Southern Ocean to the south. The former is
considered one of the engines of global climate, while the latter plays a major role in inter-
hemispheric climate teleconnections. In addition, the Australian continent contains a diversity of
Quaternary landforms and sediments in a wide range of environments which provides a great
opportunity to create an integrated regional record of climate and environmental change (Turney et
al., 2007). Proxies that can be used to generate long-term records and also to quantify the changes,
from Australia are sparse. Therefore, further investigation of past climate changes in Australia is
Page 34
Background
3 | P a g e
urgently required (Turney et al., 2007) and paleoecological research is one of the key components
(Birks, 2008).
Most of the commonly applied palaeoclimate proxies in Australia such as tree rings and borehole
measurements are unable to provide records that extend as far back as the LGM. The LGM
(traditionally 21-17 ka yr) is one of the key time intervals for climate reconstruction as this interval
represents the most significantly different global climate in the recent geological past.
Understanding long-term climate change that extends back to the LGM will help us separate
anthropogenic impacts on climate from natural internal variability and radiative forcing of climate
change. From these insights we can further investigate the long-term effect of climate change on
ecosystems.
Australian proxy based reconstructions for the time interval from the LGM to the start of the present
interglacial (c. ~11,000 years ago) include temperature reconstructions based on pollen (e.g.
Fletcher and Thomas 2010), periglacial landforms (Galloway 1965), Amino Acid Racemization
(AAR) from Emu eggs (Miller et al., 1997) and more recently branched glycerol dialkyl glycerol
tetraether (GDGT) (Woltering et al., 2014). With the exception of pollen these proxies are relatively
unconventional for the application of temperature inferences and these records have provided
limited or contradicting estimates of temperature change in Australia. These results may reflect
either regional variability, or the unreliability of one or more of the techniques, or both.
In summary, despite efforts through the AUS-INTIMATE project to improve palaeoclimate data
sets, a critical gap remains in techniques that are available to determine past changes in temperature
in mainland Australia. Further developing and refining both conventional and novel proxy based
methods that can reconstruct the LGM temperatures is required. Records derived from applying
these methods will provide valuable information on the connection between Australian terrestrial
temperature changes and adjacent oceans and Polar Regions during the LGM. Such data will
advance our knowledge on hemispheric and global climate teleconnections.
Transfer function technique has been widely applied for quantitative reconstructions of past climate
changes using a number of biological proxies (Birks, 1998). Proxies that have been applied in
Australia include mainly diatoms (Tibby, 2004), pollen (Cook and van der Kaars, 2006; Fletcher
and Thomas, 2010) and chironomids (Rees et al., 2008). Among these, diatoms are primarily used
for salinity and other in-lake variable reconstructions. Pollen and chironomids are essentially
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Background
4 | P a g e
commonly applied climate indicators but with respective to the transfer function technique, more
successful applications are demonstrated for the use of chironomids.
A number of studies show that chironomids can also be used as a proxy for nutrient status
(Woodward and Shulmeister, 2006), pH (Rees and Cwynar, 2010b), hypoliminia conditions
(Broderson et al., 2001) and salinity (Zhang et al., 2007) inferences. However, the most widely and
traditionally applied chironomid based transfer functions are for palaeo-temperature reconstructions
(Walker 1987; Olander et al., 1999; Lotter et al., 1999). A number of long term, high resolution
temperature records have been derived using the transfer function method from high latitudes of the
Northern Hemisphere (e.g. Brooks and Birks, 2000; Larocque-Tobler and Finsinger, 2008;
Larocque-Tobler et al., 2010). In the last decade, chironomid based transfer functions have been
developed and successfully applied from New Zealand (Woodward and Shulmeister, 2006, 2007;
Dieffenbacher-Krall et al., 2007), Tasmania (Rees et al., 2008; Rees and Cwynar, 2010a) and South
America (Massaferro et al., 2014). All of these applications have provided sensible quantitative
temperature reconstructions. This thesis will present the development of the first subfossil
chironomid based transfer function for mainland Australia and trial its application to an LGM
record.
More recently, Wooller et al. (2004, 2008) and Verbruggen et al. (2010, 2011) showed that oxygen
isotopes measurements on chironomid head capsules are a promising proxy that can be used to
independently reconstruct past temperatures. The method of using stable isotope on chironomid
head capsules has not been applied in the Australasian region and has limited investigation (Mayr et
al., 2014) in the Southern Hemisphere. Many aspects associated with the use and interpretations of
this new proxy are insufficiently understood. This thesis will include the first investigation on the
potential of applying stable oxygen and hydrogen isotopes in subfossil chironomids from Australia
as a proxy for past climate and environmental changes.
The paleoecological record generated from this study from the Australian continent will (hopefully)
provide critical quantitative data concerning climate change in the region during the late
Quaternary. The proxy based method developed in this thesis will have wide future applications, for
example, long-term quantitative, high resolution climate reconstructions will be possible from
different climate zones of eastern Australia.
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Research objectives
5 | P a g e
1.2 Research objectives
The general aim of this thesis is to understand the past climate system and changes in eastern
Australia with a focus on the LGM and late glacial by developing and applying proxy (subfossil
chironomid) based methods for past climate reconstructions.
Objective 1: To provide an overview of the physical and chemical properties of water bodies in
eastern Australia, extending from the tropics to Tasmania to provide the background data for the
chironomid transfer function and stable isotope investigations.
To acquire and analyse water chemistry data of samples collected from 45 water bodies
To acquire and analyse remote sensed data of the catchment land-use and climate variables
of the water bodies in ArcGIS
To examine the differences and similarities of the responses to stressors in the catchments
between natural lake systems and reservoirs
To examine the inter-relationship of the catchment land-use, climate and environmental
variables within eastern Australia freshwater systems
To summarize the current status and identify the major environmental controls on the
Australian freshwater bodies
Objective 2: To create a transfer function(s) based on the chironomid assemblages across
southeastern Australia that can be used to reconstruct past climate or other environmental proxies.
To extract chironomids from the water – sediment surface samples
To perform taxonomic analyses on the extracted chironomids
To determine the key environmental and climatic controls on chironomid species
distribution
To develop transfer function(s) that model the relationship between chironomid species
assemblages using the key variable(s) determined
To examine the effects of including artificial waterways for the temperature transfer
function development
To examine the previously claimed endemic taxa effects of including Tasmanian lakes
Objective 3: To apply method(s) for down-core analyses of a selected paleosite(s) with records that
extend back to the last glacial maximum (LGM) for climate reconstruction
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Research objectives
6 | P a g e
To extract chironomid head capsules from the targeted section of a well-dated sediment core
of a selected site
To perform taxonomic analyses on extracted subfossil chironomids throughout the sediment
core
To perform a reconstruction applying the transfer function
To perform the reconstruction diagnostics and interpret the results while consulting other
climatic and environmental reconstructions from adjacent sites
To infer changes in the climate system covering the LGM of eastern Australia based on the
results obtained
Objective 4: To explore the opportunity of using the δ18
O of subfossil chironomid head capsules for
reconstructing past changes.
To obtain sufficient chironomid head capsules (minimum dry weight of 100 µg) from a
selected genus of modern water – sediment lake samples for stable isotope analyses
To analyse the stable oxygen and hydrogen isotopes from chironomid head capsules samples
and from lake water samples
To examine the relationships of the stable isotopes of chironomid head capsules against
stable isotopes of lake water and precipitation, and other significant climatic and
environmental variables
To calculate the fractionation factor of stable isotopes of chironomid head capsules and lake
water
To examine and understand the environmental controls on the chironomid stable isotope
fractionation
To determine if stable isotopic composition of chironomid head capsules has the potential to
be used as a proxy for paleoclimate or other paleoenvironmental reconstructions in Australia
1.3 Overview of research methodology
Australia is generally a dry continent and therefore, there are a limited number of water bodies to
choose from. The aim was to sample as many natural lakes as possible from the southeast
Australian mainland. However, artificial lakes and reservoirs are also included due to the scarcity of
permanent water bodies in some critical climate and elevation spaces. The effect of including
artificial water-bodies in the dataset is investigated in Chapter 3 and 4, respectively. An overview of
the flow and framework of this project and a project timeline is shown in Figure 1.1.
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Research objectives
7 | P a g e
Figure 1.1 Overview of the framework and project flow
Obj 1.
Obj 4.
Obj 2.
Primary
Field Data
Develop
transfer
function(s)
Data screening Statistical
analyses – To understand the
modern limnology, climate
and land-use relationships
Chironomids
extracted and
pre-treated
Chironomid
taxonomy
analyses
Chironomid
stable isotopes
analyses
Field Work 1
(10 Lakes)
Field Work 2a
(10 Lakes)
Field Work 2b
(14 Lakes)
Field Work 3
(Paleo-site)
Collected water
samples to be
analysed for stable
isotopes and
chemistry
Collected
sediment samples
to be processed in
GPEM labs
All other site
related data
collected
Site
Selection
and initial
information
collection
Statistical analyses
for chironomid
stable oxygen
isotopes data
Develop
relationships based
on chironomid
stable isotopic
composition
Down-core application
to selected site for
reconstruction
If
applicable
If
applicable
Obj 3.
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Research objectives
8 | P a g e
Project timeline
Page 40
Thesis structure and synopsis
9 | P a g e
1.4 Thesis structure and synopsis
This thesis uses ‘Thesis by publication’ format. Fig. 1.2 shows the overall thesis structure and
visualizes how each of the three objectives is addressed in relevant publications or submitted
manuscripts.
At the time of thesis submission, Chapter 3 is published in Marine and Freshwater Research
(Chang et al., 2014); Chapter 4 is published in Palaeogeography, Paleoclimatology, Paleoecology
(Chang et al., 2015a); Chapter 5 is published in Quaternary Science Reviews (Chang et al., 2015b)
and Chapter 6 is submitted for publication in Limnology and Oceanography (Chang et al.,
submitted).
As opposed to a traditional thesis, each chapter (paper) contains a relevant literature review,
detailed methodology, results, discussion and a conclusion sections. The final chapter comprises
conclusions from this thesis drawing together the overall outcomes and contributions to the research
context. Recommendations for future work are also discussed in this chapter. In addition,
appendices incorporate supporting materials (Appendix 1) that are not presented in publications,
and photographs of modern southeastern Australian chironomid head capsules (Appendix 2) and
for the fossil taxa from the Welsby Lagoon palaeo-record (Appendix 3). An identification key
based on the photographic materials will be submitted for publication as an electronic resource in
the near future along with work from other colleagues. Published journal articles during the
candidature are incorporated as Appendix 4.
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Thesis structure and synopsis
10 | P a g e
Figure 1.2 Overview of the thesis structure
Chapter 5: A chironomid-inferred
summer temperature
reconstruction from subtropical
Australia during the last glacial
maximum (LGM) and the last
deglaciation
Chapter 7: Conclusions and future work
Chapter 6: Can stable oxygen and
hydrogen isotopes from Australian
subfossil chironomid head capsules be
used as proxies for past
environmental change?
Objective 4 Objective 3
Chapter 1 and 2: Introduction and background
Objective 1
Chapter 3: A snapshot of the
limnology of eastern Australian
water bodies spanning the tropics
to Tasmania: the land-use,
climate, limnology nexus
Chapter 4: A chironomid
based transfer function for
reconstructing summer
temperatures in south
eastern Australia
Objective 2
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The Biology of Chironomid
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2 THE BIOLOGY OF CHIRONOMIDS
Chironomids (Chironomidae, known commonly as non-biting midges) are a family of two-wing
flies (Insecta: Diptera) (Cranston and Martin, 1989). More than 5,000 species have been identified
world-wide, however, it is estimated that up to 15,000 species may exist in total (Cranston 1995).
Chironomid larvae range from 2-30 mm in length and most are pale yellow, green or red in colour.
The larvae are worm-like, having three thoracic segments and a nine-segmented abdomen (Fig. 2.1)
(Brooks et al., 2007). Most chironomid larvae are detritivores, i.e. obtain nutrients from grazing
bacteria and algae (Cranston and Martin, 1989).
Figure 2.1 Key morphological characteristics of chironomid larvae (Brooks et al., 2007, Page 4)
The chironomid eggs are laid in a hygrophilous jelly mass dropped into the water or in a jelly-
string, attached by one end to the substrate. The hatching rate and survivability are temperature
dependant (Cranston and Martin, 1989). Most of the larvae are tube-dwelling, apart from subfamily
Tanypodniiae, which are free-living. The silken tube is constructed from silk-spinning organs
located in the ventro-mentum and protects larvae from predators, and also performs a respiratory
function.
There are four larval stages of chironomids (from 1st instar to 4
th instar) (Fig. 2.2). The development
time is influenced by the combination of temperature and food availability (Johannsson, 1980).
Most of the temperate species are multivoltine, univoltine or bivoltine (voltinism: used in biology to
indicate the number of broods or generations of an organism in a year) (Tokeshi, 1995), for
example, large species living in deep lakes may take more than one year to develop, whereas small
species living in warm, shallow water bodies may complete 3-4 generations in one year (Tokeshi,
1995). In southeast Australia, apart from the high mountain lakes, it is reasonable to assume most
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The Biology of Chironomid
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species may complete 2-3 generations in one year from lowland lakes. The pupal stage usually lasts
3-4 days and the timing of emergence is dependent on water temperature and light intensity
(Kureck, 1979). Adults of most species live for just a few days, at most for few weeks (Oliver,
1971).
Figure 2.2 Chironomid Life Cycle (Walker, 1987, Page 852)
Figure 2.3 A fully developed chironomid head capsule (Chironomus sp.) from Lake Albina, Mt
Kosciuszko, New South Wales, Australia (photo is taken by J. Chang at the Physical Geography
laboratory, School of Geography, Planning and Environmental Management, University of
Queensland)
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The Biology of Chironomid
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Chironomid head capsules (Fig. 2.3) are derived from dead larvae and also largely from head
capsule moults between larval instars. They are usually the most abundant insect remains in lake
sediments, in most freshwater situations, from the tropics to the arctic, as the chironomid larvae
occupy a huge range of aquatic habitats (Cranston and Martin, 1989). The head capsules of
chironomid larvae are made of chitin (Fig. 2.4), usually fully developed, globular, and pigmented
dark brown or black. The subfamily Tanypodinae has a more elongate and un-pigmented head
capsule. The head capsule of all chironomids cannot be retracted into the thorax (Brooks et al.,
2007). Chironomid head capsule samples are usually dominated by third and fourth instar capsules
as the earlier instars (first and second) are less durable.
Figure 2.4 Chitin molecule structure
Chironomid larvae have several qualities that make them useful as a proxy in environmental and
climatic palaeo-studies. They are stenotopic, which means different elements of chironomid
assemblage respond in a distinctive way to particular environmental perturbations (Brooks, 2003).
They are also abundant and ubiquitous, therefore, there is usually a large number of them that can
be found in practically all permanent or semi-permanent freshwater bodies. Chironomids are also
rich in species that make the assemblages sensitive to environmental changes (Brooks, 2003). The
response of chironomids to any change in conditions is local and fast due to their short life cycle.
They are also identifiable as the head capsules are usually well-preserved, although, it is still
challenging to identify most of them beyond genus or species-group level.
More recently, Wooller et al (2004, 2008) and Verbruggen et al. (2011) have demonstrated that the
chitin contained in the head capsules can be analysed for stable isotopes (δ18
O and δ2H). Chitin is a
long-chain polymer of an N-acetylglucosamine, a derivative of glucose (Fig. 2.4), and is the main
component of the exoskeletons of chironomids. This has provided new opportunities for
paleoecologists to investigate the novel methods to be used in paleoclimate and paleoenvironmental
reconstructions.
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The Biology of Chironomid
14 | P a g e
Chironomid taxa have been used as a proxy in paleoclimate and paleoenvironmental studies since
1980s (Boubee, 1983; Hofmann, 1986; Walker, 1987; Walker and Mathewes, 1989). Using
chironomid taxa as quantitative climate indicators were derived from various parts of the world as
early as 1990s (e.g. Walker et al., 1991a, b; Wilson et al., 1993; Levesque et al., 1993; Walker et al.,
1995; Heinrichs et al., 1997; Olander et al 1997; Walker et al., 1997), but all focused on Northern
Hemisphere. The first chironomid based transfer function from the Southern Hemisphere was
developed by Woodward and Shulmeister (2006) from New Zealand. A detailed literature review of
the development and uses of transfer functions is presented in Chapter 4 (Section 4.1). The current
available modern calibration data sets and transfer functions is also summarized in Appendix 1.
2.1 The principles of applying the transfer function and stable isotope methods in
palaeoclimate reconstructions
2.1.1 The transfer function method
As discussed above, chironomid-based transfer function, as a technique to quantify past climate and
environmental variables has been developed and applied as long as thirty to forty years. The basic
and simplified idea of the technique following Sachs et al. (1977) is: Assume that summer lake
temperature (Ts) (which is close to the mean air temperature) will be a function of the chironomid
taxa abundance. If the abundances measured at a particular point were represented by the
probability series (for instance, there are 30 different taxa in the modern training set) p1, p2,
p3…p30 (where Ʃp = 1), here ‘p’ represents the probability of occurrence of a taxon and ‘a’
represents the coefficient factor, then we can express the equation as follows:
Ts = a1p1 + a2p2 + a3p3 + . . . + a30p30 + a31p1p1 + a32 p1p2 + . . . + a60p1p60 + a61p1p2p1 +
a62p1p2p2 ...
If we used all first and second-order terms, we would have to develop 496 parameters in the above
equation. So to simplify, instead of using all of the chironomid taxa abundance data, factor analysis
was used to group taxa which behave in a similar fashion together into a single factor. The
minimum number of factors to represent an acceptable amount of variance will be used. Then the
expression for temperature is simplified and becomes:
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The Biology of Chironomid
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Ts = a1f1 + a2f2 + a3f3 + . . . + a6f6 + a7f1f1 + a8f1f2 + . . . +a27f6f6 + a28. (for instance, in this
example it comes down to 6 factors)
Values for the coefficient factors (a1 to a28) were then found by multiple regression (usually by
applying a Monte Carlo simulation).
After the modern calibration set is established, fossilised chironomids found at different levels of a
dated core can be examined. Different taxa of chironomids from each level should be counted and
then the assemblages converted into factors determined by the modern calibration data set. Then we
apply the factor loading scores to the transfer function and calculate the summer temperatures at
different levels at the site of the dated core.
2.1.2 The stable isotope method
The basic idea for using the δ18
O of subfossil chironomid head capsules to infer mean annual air
temperature (MAAT) changes in the past is that we assume the δ18
O values analysed on chitin from
the chironomid head capsules gathered from the surface sediments have a strong linear relationship
with the MAAT. This premise is based on the observed strong correlation between δ18
O of
chironomids and δ18
O of lake water, δ18
O of lake water and δ18
O of local precipitation, as well as
δ18
O of precipitation and MAAT. A similar principle is applied to the hydrogen isotope (δ2H).
Verbruggen et al’s (2011b) study from northern Europe has shown strong linear relationships
between MAAT and δ18
O of Precipitation (r2=0.94), similar to δ
18O of chironomids and δ
18O of
lake water (r2=0.9), however, the linear correlation between δ
18O of precipitation and δ
18O of lake
water is weaker (r2=0.64). This is likely due to the δ
18O of the lake water reflects both the source of
precipitation and evaporative processes in the lake. In addition, the δ18
O of precipitation can also
reflect changes in source and seasonality among other variations as well as MAAT. In this study,
these correlations will be tested from southeastern Australian lakes and we should expect
complexities among these relationships. Same principle is applied to the hydrogen isotope (δ2H).
For the application of the stable isotope method to a dated core, firstly we convert the measured
δ18
O/ δ2H values from chironomid fossils to temperature (analogous to modern δ
18O/ δ
2H
chironomid inferred temperature values) using the model developed based on δ18
O/ δ2H values
measured on modern chironomids from the lake surface sediment. Secondly, it is assumed that the
MAAT reconstructed using this method represent and quantify the major temperature variations and
fluctuations in the past.
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Land-use, climate and limnology nexus of eastern Australia
16 | P a g e
3 A SNAPSHOT OF THE LIMNOLOGY OF EASTERN AUSTRALIAN
WATER BODIES SPANNING THE TROPICS TO TASMANIA: THE
LAND-USE, CLIMATE, LIMNOLOGY NEXUS
Chapter 3 is published in Marine and Freshwater Research (Chang et al., 2014)
3.1 Summary
The present study is the first comprehensive overview of the physical and chemical properties of
lakes in eastern Australia, extending from the tropics to Tasmania. The results showed that climate
is the dominant control on the lakes. The results suggested that high nutrient concentrations in lakes
in drier areas may be the result of natural processes of concentrating nutrients from high
evaporative flux in eastern Australia, rather than due to human activities.
Highlights:
It provides an insight into how lake water chemistry, climate and human impact factors
interact and correlated with each other from eastern Australian lakes in the modern age. It is
critical to understand these correlations and connections before we try to use any proxies
(not limited to chironomids) from lake sediment to reconstruct past lake conditions,
environment or climate of the region or to use these data to predict any future changes.
This study highlights that lake responses to stresses can vary in predictable ways across the
Australian landscape, and that management and restoration would benefit from considering
factors such as geology, climate, etc..
This study will help to interpret the transfer function results, the transfer function based
paleo-reconstructions and stable isotope results from modern lakes
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Land-use, climate and limnology nexus of eastern Australia
17 | P a g e
3.2 Abstract
The present study investigates 45 natural and artificial water bodies extending across the whole of
eastern Australia from the tropics to Tasmania. A broad variety of physical and chemical, land-use
and climatic parameters were measured. Reservoirs and other artificial water bodies responded to
stressors in their catchments in a similar fashion to natural lakes, but tended to be less nutrient rich,
possibly because of shorter residence times and active management. Salinity and pH were strongly
correlated in the dataset. Bedrock had a strong influence on pH in freshwater lakes, whereas all
highly saline lakes were alkaline, irrespective of bedrock. High concentrations of anions in saline
lakes precluded the existence of acid conditions by binding available hydrogen ions. Almost all
lakes fell on salinity axes that indicated marine origin for their salts. An assessment of the total
nitrogen to total phosphorus molar ratios from the lakes in the present dataset indicated that
productivity in Australian lakes could be limited by both nitrogen and phosphorus. Future research
using macro-nutrient enrichment experiments should be pursued to confirm this preliminary
observation. There was a strong positive correlation between regional aridity and lake
eutrophication. This is typical of semi-arid and seasonally arid environments and reflects the
concentration of nutrients owing to evaporative flux in shallow basins with high residence times.
Key words: eutrophication, evapotranspiration, freshwater, lakes, reservoirs, lithology.
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Land-use, climate and limnology nexus of eastern Australia
18 | P a g e
3.3 Introduction
Apart from a review of Australian lake geomorphology by Timms (1992), which focused primarily
on the formation of lakes, and broad overviews by Brookes and Hamilton (2009) and Bridgman and
Timms (2012) of all Australian lakes, which contained little detail, there is no summary of physical
and chemical data from mainland eastern Australian lakes. The aim of the present paper was to
provide baseline physical and chemical, land-use and climatic data for eastern Australian lakes, and
to provide a preliminary evaluation of the primary controls on lake chemistry in eastern Australia.
The paper emerged from a study designed to determine whether subfossil chironomid head capsules
could be used as a basis for reconstructing temperature and nutrient status in Australian water
bodies. The study questions related to whether and how climate, geology and land use affect lake
water chemistry in eastern Australia. Given that there is little summary work in this area, the present
basic descriptive survey was expected to generate findings that can lead to testable hypotheses
about controls on lake water chemistry.
Globally, freshwater lakes are concentrated at high latitudes. Canada has the largest number of
lakes, with an estimated 2 million water bodies, of which 31 752 are larger than 3 km2 (Renwick
2009). Northern Europe also has abundant lakes. In the Australasian region, New Zealand has 3820
lakes and Tasmania has more than 4000 lakes (Brookes and Hamilton 2009). In comparison, the
Australian mainland has very few natural permanent freshwater bodies (Bridgman and Timms
2012). This is despite the fact that an area of ~1.5 million km2 (roughly six times the size of New
Zealand) is humid (precipitation ≥ evaporation). Therefore, the lack of permanent freshwater lakes
is not simply explained by a lack of precipitation. Geomorphology and, in particular, the limited
extent of past glaciations in Australia appear to be the primary reason for this paucity of lakes
(Bridgman and Timms 2012).
3.3.1 Previous water-chemistry studies of Australian lakes
Previous studies have recognised eight types of lakes in Australia, on the basis of their mode of
formation (Timms 1992). These are glacial lakes, volcanic crater lakes, coastal lagoons, coastal
dune lakes, tectonic lakes, fluvial lakes, solution lakes (sink-hole lakes) and lakes formed by
landslides. We chose not to review tectonic lakes because most of the lakes in this category are not
permanent (e.g. Lake George) and landslide-blocked lakes are rare, and we added an upland-lagoon
category to cover shallow lakes on the tablelands (Fig. 3.1).
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Land-use, climate and limnology nexus of eastern Australia
19 | P a g e
Figure 3.1 Map of eastern Australia, with the 45 study sites identified; the numbers correspond to
the lake names and numbers in Table 3.1.
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Land-use, climate and limnology nexus of eastern Australia
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Glacial lakes in mainland Australia are limited to alpine areas of the Australian Alps (Fig. 3.1).
Williams et al. (1970) demonstrated that the Kosciuszko lakes (Fig. 3.1) were ‘extremely fresh’ and
that salinity, and nitrate and phosphate concentrations were very low. Green (2012) summarised the
seasonal chemical characteristics of five Kosciuszko lakes that are ice-covered in winter. He
suggested that the salts and nutrients of the five Kosciuszko lakes were mainly derived from
bedrock erosion and Aeolian inputs and nutrient deposition from precipitation was low. Most lakes
in the Tasmanian highlands and central plateau (Fig. 3.1) are of glacial origin (Jennings and Banks
1958). The majority of these lakes have ionic composition consistent with aerosol sources deposited
in the lake through both dry and wet deposition and originally relate to a seawater source (Buckney
and Tyler 1973; Vanhoutte et al. 2006). The major water-chemistry gradients are conductivity, pH
and gilvin (i.e. Chloraphyll) (Vanhoutte et al. 2006) and these relate primarily to bedrock geology
and climate.
The second major lake type is the volcanic crater lake. Two regions in Australia have significant
concentrations of crater lakes, namely, the Atherton Tablelands of northern Queensland and the
basalt province of western Victoria (Fig. 3.1). Some isolated crater lakes (e.g. Coalstoun Lakes in
southeastern Queensland) occur outside these main fields. In northern tropical Queensland, the
paleoecology of these lakes is rather better studied than the modern lake chemistry, with extensive
work on human settlement history, palaeoclimate and palaeoenvironmental reconstructions
(e.g.Dimitriadis and Cranston 2001; Turney et al. 2001; Haberle 2005; Kershaw et al. 2007). In
western Victoria and adjacent regions of South Australia, the crater lakes are more saline. These
lakes were heavily investigated and the results of these studies were summarised in De Deckker
(1983) and De Deckker and Williams (1988). These studies showed that sodium is the most
common cation, nitrogen is the limiting plant nutrient and that the lakes are turbid because of high
algal concentrations.
Coastal lagoons are widely distributed around the eastern and southeastern coasts of Australia. The
literature on lake water chemistry of Australian coastal lagoons is scarce, or at least not fully in the
public domain, largely comprising government monitoring data and reports. Scanes et
al. (2007) evaluated 22 coastal New South Wales (NSW) lagoons and concluded that controls on
the lagoons were highly variable and that the lagoons were minimally affected by human activity.
Dune lakes are the most common lake type in eastern Australia. The greatest concentration of dune
lakes occurs on Fraser Island (Fig. 3.1), which has over 100 freshwater bodies and the limnology of
these lakes has been heavily investigated since the 1960s (Bayly 1964; Bayly et al. 1975; Bowling
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Land-use, climate and limnology nexus of eastern Australia
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1988; Hembrow and Taffs, 2012). They occur as either perched or window lakes. Overall, these
lakes are oligotrophic. Dune lakes in Gippsland, western Victoria and northeastern NSW were
investigated by Timms (1973, 1977, 1982).
We present upland ‘lagoons’ of the NSW tablelands (Fig. 3.1) and particularly northern NSW
tablelands as a distinct lake category (e.g. Little Llangothlin Lagoon, Mother of Ducks). There is no
consensus on the origin of these shallow (< 3 m deep) lakes and they may have formed by sapping
or deflation, although some are clearly created by drainage impoundment (Bell et al. 2008). As with
other lakes, sodium is the dominant cation and the ionic concentration relates to the
precipitation : evapotranspiration (P : EP) ratio and the season of precipitation (Briggs 1980).
In lowland Australia, there are numerous lakes that are formed by meander cut-offs (Constantine
and Dunne 2008) and also intermittent lakes that form as part of a chain-of-ponds style drainage.
We do not consider these types of lakes. In limestone areas, e.g. parts of northwestern and southern
Tasmania, sink-hole lakes are occasionally present, such as e.g. Lake Chisholm, Tasmania
(Bowling and Tyler 1988). Lakes formed by landslides, such as e.g. Lake Tali Karng (Timms
1974), are not covered in our dataset.
3.3.2 Artificial water bodies
Because natural water bodies do not occur in all regions, the present study included several
reservoirs and other artificial water bodies. The construction of Australian reservoirs began in the
early 1880s. Reservoirs are concentrated in the eastern and southeastern coastal regions of Australia
where the population density is highest. Hydro-electric schemes are the other major source of
artificial lakes. Tasmania has ~60 major reservoirs across the state for this purpose and other
reservoirs are located in southeastern Australia as part of the Snowy Hydro Scheme. A few lakes
have been created for recreational purposes, notably for fishing, and two of these lakes were
included in the dataset.
Many studies have addressed the water quality issues that are important in reservoirs, including
turbidity problems (Chanson and James 1999), problems with eutrophication and floating
macrophytes (Harris 2001; Smith 2003; Davis and Koop 2006), deep water anoxia (Beutel and
Horne 1999; Townsend 1999) and the presence of toxic cyanobacteria (Baker and Humpage
1994; Jones and Poplawski 1998). Eutrophication from nutrient rich rivers (e.g. Murray River) may
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also be a risk (Baker and Humpage 1994). The biggest difference between reservoirs and natural
lakes is the shorter residence time for water in reservoirs (Søballe and Kimmel 1987).
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Table 3.1 Summarised information of the 45 water bodies sampled
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3.4 Materials and methods
3.4.1 Study sites
The present dataset comprises 45 lakes and artificial water bodies located in the eastern and
southeastern Australia (Table 3.1, Fig. 3.1). The transect covers a distance of 3700 km along the
eastern coast of Australia, from Crater Lakes National Park, Queensland, to Mount Field National
Park, Tasmania (17.24°S to 42.67°S, 140.18°E to 153.26°E; Table 3.1, Fig. 3.1). The climate ranges
from monsoonal tropical in the north, to cool temperate in the south and, hence, there are large
temperature (20.4°C for mean annual temperature) and precipitation (2476 mm for mean annual
rainfall) gradients in the datasets. Elevation of the sites ranges from sea level to 2048 m above mean
sea level (asl; Table 3.1). The dataset comprised 29 natural lakes and 16 artificial water bodies.
Climate
The climate system of eastern Australia is governed by the major global zonal circulation systems,
from the monsoon tropics to the southern hemisphere westerly belt (Sturman and Tapper 2006).
Climate regions can be summarised as follows:
1. The quasi-monsoonal (Gentilli 1971) north. This region, which extends from coastal
northern Queensland to northern NSW, is dominated by southeastern trade winds in the
winter and by the northern Australian monsoon in the summer. The climate in the vicinity of
the coast is significantly modified by persistent moisture-laden onshore winds and
orographic rainfall as these winds rise over the eastern highlands (Fig. 3.1).
2. Coastal southern NSW and Victoria are dominated by the subtropical anti-cyclonic belt
(STAB) in summer and by a westerly flow in the winter. These areas are dry in summer, wet
in winter, but the intermittent influence of easterlies and their interaction with topography
modifies the coastal and near coastal climates.
3. A true Mediterranean climate exists in coastal South Australia and western Victoria.
Rainfall predominantly occurs during winter in this area, under the influence of the
westerlies, and there is sparse rainfall during summer under the STAB; however, this region
is typically unmodified by eastern coast lows or other moisture sources.
4. Tasmania is the only dominantly all year westerly climate, although here again, there is a
weak summer dry period in eastern and northern districts.
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This overall pattern is modified by oscillatory climate phenomena embedded in the circulation
systems. These range from short duration Madden–Julian cycles (~60 days) to long duration cycles
such as the interdecadal Pacific oscillation (IPO). The main oscillatory systems that are effective at
climate scales are El Niño-Southern Oscillation (ENSO) (18 months to 7 years), the Indian Ocean
dipole, which operates on timescales similar to those of ENSO, the IPO (15–30 years and 50–70
years), and the southern annular mode (SAM), which is annual and inter-annual.
Altitude is the other major modifier of climate. The Snowy Mountains region in the southeast
(Fig.3.1) is characterised by cool to cold weather all year around and snow falls in winter. The
record lowest temperature in Australia of –23°C was recorded at Charlotte Pass
(http://www.bom.gov.au/climate/extreme/records.shtml, accessed 18 August 2013). At similar
latitudes at sea level, the climate is warm temperate.
Geology
Australia is an old cratonic plate, with mainly granitoid rocks and sedimentary rocks derived from
them. Locally hot-spot volcanism has generated basalts.
Granite. Granite batholiths are common along the high country in eastern Australia. Granites
dominate the high country from Victoria to northern Queensland. Granite rocks are high in silica
(>70%), and, consequently, lakes on these rocks tend to be acidic.
Basalt. Hotspot volcanism has occurred locally from northern Queensland to western Victoria
over the past 30 million years. The youngest volcanic fields occur in western Victoria and the
Atherton Tableland of northern Queensland and are Holocene in age. Basaltic rocks are mafic and
lakes on basalts tend to have high ionic values, including bicarbonate (HCO3–) and calcium (Ca
2+).
They tend to be alkaline.
Meta-sedimentary and metamorphic rocks. Limnologically, Tasmania is bisected by a line that
runs northwest by southeast, along the 146th
meridian, called Tyler’s Line. Western Tasmania
consists of mainly Precambrian siliceous meta-sedimentary and metamorphic rocks, including
quartzite, schist, phyllite and limestones. Lakes west of Tyler’s Line are typically acidic with pH
ranges from 4.5 to 5, except for lakes on limestone. Central and eastern Tasmania is underlain by
Permian mudstones and Triassic siltstones that were partly metamorphosed by Jurassic doleritic
intrusions. These lakes are less acidic (pH of > 6). Sedimentary rocks occur on the mainland also
(mainly sandstones) and these tend to be neutral to acidic.
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Vegetation
In eastern Australia, there are four broad structural types of vegetation. These are rainforest
(notophyllous vine thicket), wet and dry sclerophyll forest or woodland, shrubland and heathland
and grassland. In the highest country, subalpine herbland occurs. There are many subcategories of
these structural vegetation types.
Rainforest. Rainforest is confined to areas of high annual rainfall close to the eastern coast and
along the dividing range. Rainforests prefer fire protected sites and fertile locations (basalt areas
and river plains). The rainforests are dominated by hardwood taxa with emergent conifers.
Sclerophyll woodland. In areas with poorer soil and relatively lower rainfall, Eucalyptus woodland
becomes dominant. The understorey contains small broadleaved trees, vines, ferns and shrubs. On
the drier eastern side of Tasmania and inland from the main divide on the Mainland, dry sclerophyll
eucalyptus forests and woodland dominate. They contain an understorey of small trees including
Casuarina, and Acacia.
Shrubland and heathland. Shrublands, heaths and sandplain vegetation characterise the coastal
regions, especially on sandy soils. The dominant species are mostly from the families Ericaceae,
Proteaceae and Myrtaceae. Melaleuca spp. are dominant in swampy areas.
Grassland. The lowland country of Victoria (generally below 700 m) with 400–1000 mm annual
rainfall is dominated by grassland ecosystems on relatively nutrient-rich loam, clay-loam and
alluvial soils. Most of the grassland has been removed or altered for agriculture and urban
development. The ground layer is dominated by perennial, tussock-forming grasses, with some
rhizomatous or stoloniferous species. In parts of the high country of Victoria (subalpine areas) and
on other montane areas, tussock grasslands occur. In western Tasmania, communities of the
Cyperaceae, the button grass Gymnoschoenus sphaerocephalus, are extensive as a consequence of
fire regimes (Harris and Kitchener 2005). In alpine areas, tall alpine-herbfield and heathland
communities dominate (Costin et al. 2000).
Forty-five lakes and reservoirs were chosen to span as much as possible the variety of lake types in
eastern Australia (Fig. 3.1, Table 3.1). These lakes span the latitude and elevation of eastern
Australia and gradients of climate, water depth, trophic status and agricultural activity (Table 3.1
and Table 3.2 and 3.3).
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3.4.2 Sample collection and laboratory analyses
Lakes and reservoirs were sampled in eastern Australia during the summers (January to February)
of 2011–2012 and 2012–2013 (Fig. 3.1, Table 3.1, Table 3.2). Water for chemical analyses was
sampled from the lake centre at ~30 cm below the water surface, into four pre-washed and labelled
polyethylene bottles (two 1000 mL, one 250 mL and one 125 mL). Untreated water samples were
collected for the analysis of major ions (Na+, K
+, Ca
2+, Mg
2+, HCO3
–, Cl
-, SO4
2–), total nitrogen and
total phosphorus (TN and TP) analysis. These samples were kept frozen until analysis. A 1000-mL
water sample was filtered for Chlorophyll a (Chl a) on a 4.7-cm-diameter GF/F filter (0.45-µm pore
size). The Chl a filter was wrapped in foil and kept frozen for subsequent analysis. The 125-mL
water sample was filtered using a syringe and a syringe-filter Whatman (0.45-µm pore size, Supor
Membrane) and the filtered water was frozen in the bottle for later analysis for dissolved reactive P
(DRP) and reactive N (NOX). Total dissolve solids (TDS), pH, specific conductance (COND) and
turbidity (TURB) were also obtained from water-chemistry analysis, which was carried out by the
Forensic and Scientific Services in Brisbane, Queensland. In the field, water temperature, oxidation
reduction potential (ORP), pH, dissolved oxygen (DO), COND, TDS, salinity (SAL) and TURB
were recorded using an Aquaread multiparameter meter. The meter also recorded latitude, longitude
and elevation. Lake depths at the sampling points were measured using a Speedtech portable depth
sounder.
3.4.3 Spatial data
WorldClim for lake climatic data: Climate variables were obtained using the combination of the
WorldClim program (available from http://www.worldclim.org/bioclim, accessed 20 August 2013)
and ArcGIS 10.1. For the present study, mean annual temperature (MAT) was selected because it
represents the regional annual average temperature. The aridity index (AI) is defined as AI =
MAP/MAE, where MAP = mean annual precipitation and MAE = mean annual evaporation
(Trabucco and Zomer 2009). The AI values were downloaded from the CGIAR–CSI GeoPortal at
http://www.csi.cgiar.org (accessed 20 August 2013).
Watershed boundaries and catchment land use ArcGIS 10.1 was used to define the lake
catchment area and investigate the catchment land use of the sampled lakes. A digital elevation
model (DEM) was extracted from the US Geological Survey (USGS) HydroSHEDS site (available
at http://hydrosheds.cr.usgs.gov, accessed 22 August 2013), with a resolution of 15 arc-seconds
(equivalent to 304 m). Catchment areas were delineated with the DEM, and percentage land cover
within each catchment was determined using the Australian dynamic land-cover (for every 250 m
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by 250 m area) overlay from the Geoscience Australia website (available at
http://www.ga.gov.au/earth-observation/landcover.html, accessed 22 August 2013). There were
four broad categories of land cover used in the analysis, including bushland, grassland, agriculture
and urban. The bushland category included open and closed forest and woodland; the grassland
category included sparse and scattered forest, shrubland, sedgeland, tussock and alpine grassland;
the agriculture category included all forms of agricultural land use.
The dominant rock type in the lake catchment was determined using an ArcGIS data layer with
1 : 100 000 surface-geology map from Geoscience Australia (2012 edition, available at
http://www.ga.gov.au/meta/ANZCW0703016455.html, accessed 25 August 2013).
3.4.4 Statistical analyses
Prior to ordination, selected climatic and lake water-chemistry data were assessed for normality
using the Shapiro–Wilk normality test (Shapiro and Wilk 1965), and through measurements of
skewness and kurtosis (Zar 1999) in Minitab 16 (Ryan et al. 2004). All variables apart from MAT
and pH were normalised using a log10 transformation.
Both indirect and direct ordinations were used to analyse the data. Principal components analysis
(PCA) was performed using CANOCO version 4.5 (ter Braak and Smilauer 2002), with samples
being constrained by 18 limnological and land-use variables. Geology was excluded, because it was
represented by a nominal scale (categories of rock type). Climate (aridity index), lake-morphology
(water depth) and land-use variables were initially selected as passive variables. Passive variables
do not directly affect the analysis. They are visible in the results so that their pattern can be
observed; however, they are kept passive so that the study can focus the analysis on in-lake
(limnological) variables.
Redundancy analysis (RDA) was used to explore the relationship among lake depth, aridity, land
use and lake trophic status. Partial RDAs were performed in CANOCO v 4.5 (ter Braak and
Smilauer 2002).
To investigate the origins of major cations and anions in the lake water, tri-plots were generated in
Excel following Graham and Midgley (2000). Lake distance from the closest coastal area was
measured using the Google Earth ruler tool.
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Table 3.2 Selected limnological and land-use/land cover variables for the forty-five water bodies
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3.5 Results
The lakes covered a wide range of climatic and physical characteristics. The mean annual
temperature ranged from 3.7°C in Blue Lake, Kosciuszko, to 24.1°C in Horseshoe Lagoon, northern
Queensland (Table 3.3). Lakes were distributed from the semi-arid areas (aridity index <0.5) of
north-western Victoria, to the very humid areas (aridity index = 3.7) of western Tasmania (Table
3.3). Lake depth ranged from 0.2 m (Eagle Tarn, Tasmania) to the 65-m-deep crater lake (Lake
Barrine, Queensland; Table 3.3).
The lakes were also very diverse in terms of limnology. The pH of lake water ranged between 4.7
and 9.2 (Table 3.3). Most of the lakes of western central Tasmania were acidic, whereas those in
western Victoria were strongly alkaline. Montane lakes had low specific conductance (5–6 µS cm–1
for Kosciuszko lakes), whereas some Victorian coastal lakes had very high specific conductance
(>15 000 µS cm–1
; Table 3.3). Algal biomass (Chl-a concentration) ranged from 1 µg L–1
in the
alpine lakes to 90 µg L–1
in the Victorian lakes. TP concentrations were lowest in the alpine lakes
and some Tasmanian lakes (<0.01 mg L–1
) and highest (3.6 mg L–1
) in the Victorian lakes. TN
concentration ranged from 0.11 mg L–1
in the alpine lakes to 36 mg L–1
in the Victorian lakes
(Table 3.3). The trophic state of the lakes ranged from oligotrophic to hypereutrophic (Table 3.1),
following the same trends. Lowland natural lakes of mainland Australia were mostly eutrophic to
hypereutrophic (Table 3.1), which contrasts strongly with the situation in New Zealand and
Tasmania.
Limits for N and P limitation were derived from Guildford and Hecky (2000). TN : TP ratios ranged
from as low as 1, which is indicative of severe N limitation (Coalstoun Lakes), to 139, which is
indicative of severe P limitation (Table 3.1). Most (56%) of the studied lakes were co-limited
(TN : TP between 20 and 50), whereas only four lakes (9% of our dataset) were N-limited
(Coalstoun Lakes and UQ Lakes) (Abell et al. 2010). The remaining 35% (16 lakes) were P-limited.
The land-use patterns in the lake catchment were highly variable. Alpine and highland lakes that are
within national parks or other reserves were, generally, dominated by native woodland and
grasslands. Lakes on private land and around some large reservoirs in northern Queensland were
dominated by agricultural land, with both extensive grazing and irrigated agriculture (cropping and
intensive grazing) occurring (Table 3.3).
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Table 3.3 List of the 19 variables used, with mean, minimum and maximum values cited and the
appropriate units highlighted
3.5.1 Lake-water chemistry
A ternary plot of major cations (Na+ and K
+, Ca
2+ and Mg
2+; Fig. 3.2) showed that the 45 lakes were
distributed close to the world-average seawater ratio (WASW). Unsurprisingly, coastal lakes
(lowland lakes that are <100 km away from the sea) were closer to the ratio than were lakes far
inland or at high elevation. Fig. 3.3 bi-plot demonstrates that pH co-varied with bedrock type and
salinity, but that salinity was a stronger control on alkalinity than was the pH of the bedrock.
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Figure 3.2 Tri-plot of major cations, with lakes labelled by type, as follows: CL, coastal lakes
(lowland lakes that are <100 km away from the sea); M, montane lakes (>1000 m asl); IL, inland
lakes (lowland lakes that are >100 km away from the sea). The tri-plot shows cation concentration
as a percentage. All 45 lakes are distributed closer to the world-average seawater ratio (WASW)
than to the world-average freshwater ratio. On average, lowland coastal lakes are closer to the ratio
than inland lakes or those at high elevation. This indicates that the primary source of the salts in
eastern Australian lakes is seawater.
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Figure 3.3 Biplot of pH against conductivity. This demonstrates that pH co-varies with bedrock
type and salinity. All saline lakes are strongly alkaline, whereas there is more variability in the pH
of freshwater lakes. This means that the freshwater lakes are dominantly controlled by bedrock
types. Lakes above log conductivity of 3 are classified as brackish.
3.5.2 PCA results
An initial PCA plot of the 45 water bodies constrained to the 19 environmental variables, with land-
use, climate and lake-depth variables entered as passive variables, is presented in Fig. 3.4. The first
and second axes of this plot were predominantly specific conductance and pH gradients. Axis 1 and
Axis 2 explained 75.1% and 9.6%, respectively, of the variation in the environmental data. This
result is unusual in both the high explanatory power of the first axis and the low explanatory power
of the second axis and is an artefact of the large number of correlated water conductivity proxies.
So as to address this, we removed all ions from the analysis and then re-ran the PCA. The revised
primary y-axis was predominantly a nutrient gradient and the second axis was a pH gradient. Axes 1
and 2 explained 70.1% and 18.0% of the variance, respectively. This indicated that nutrient status
and pH were the longest gradients in the lake-chemistry variables (see Fig. 3.5). In Fig. 3.6, we
display the results of a final PCA, with all environmental data (excluding ion concentration data)
entered as active variables. The primary y-axis explained 46% of the variance and was a trophic
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34 | P a g e
status (TN, TP, Chl a), conductivity and agriculture gradient. A further 17.2% of the variance was
explained on the second axis, which was primarily a temperature gradient. This PCA plot also
indicated that eutrophic saline lakes tend to be shallow lakes located in arid regions.
To summarise the information from the PCAs, the first axis is always a nutrient and conductivity
related axis showing that shallow lakes, especially those located in agricultural catchments, are
affected by eutrophication. Unsurprisingly, given that our data extended over a latitudinal range of
more than 2000 km and altitudinal range of over 2000 m, temperature was also a significant
gradient. To further examine the implications of these findings, we applied a RDA to investigate the
interactions among water depth, aridity, agriculture and nutrient status.
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-1.0 1.5
-1.0
1.0
CTL
LA
BL
LLL
LCN
ET
WP
LPA
LEA
LE
LS
LD
LBA
JUL
SEL
PL
CHD
CLU
LMO
TL
FWL
NGR
LFYLTK
LPO
GRL
LEM
LRD
LTP
LSP
BR
LCT
FHD
GBL
KIDLAW
RRD
COD
SWL
UQ
LMB
LH
HOL
GHL
WL
Axis 1
Axis
2
-1.0 1.5
-1.0
1.0
Depth MAT
AI
TP
TN
Chla
Na+K+
Ca2+
Mg2+
HCO3-
Cl-SO42-
PH
COND
Bushland
Grasslan
Pasture
Cropping
UrbanAgricult
Figure 3.4 Initial PCA plot of a) the 45 water bodies constrained to b) the 19 environmental variables, with land-use, climate, and lake depth variables
entered as passive variables to focus the analysis on in-lake (limnological variables). The first and second axes of this plot are predominantly specific
conductance and pH gradients; it shows a large number of correlated water conductivity proxies (ion concentrations).
AX
IS 2
AXIS 1
a. b.
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-1.0 1.0
-1.0
1.0
CTL
LA
BLLLL
LCN
ET
WP
LPA
LEA
LE
LS
LD
LBA
JUL
SEL
PL
CHD
CLU
LMO
TL
FWL
NGR
LFY
LTK
LPO
GRL
LEM
LRDLTP
LSPBR
LCT
FHD
GBL
KIDLAW
RRD
COD
SWL
UQ
LMB
LH
HOL
GHL
WL
AXIS 1
AX
IS 2
-1.0 1.0
-1.0
1.0
Depth
MAT
AI
TP
TN
Chla
PH
COND
Bushland
Grasslan
Pasture
Agricult
Figure 3.5 Revised PCA plot of a) the 45 water bodies constrained to b) the 12 environmental variables (ionic variables removed), with land-use,
climate, and lake depth variables entered as passive variables to focus the analysis on the in-lake (limnological) variables. The first axis is now a
nutrient gradient and the second axis is a pH gradient.
AXIS 1
AX
IS 1
a. b.
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-1.0 1.0
-1.0
1.0
Depth
MAT
AI
TP TN Chla
pH
COND
Bushland
Grassland
Urban
Agriculture
AXIS 1
AX
IS 2
b.
-1.0 1.0
-1.0
1.0
CTL
LABL
LLL
LCNET
WP
LPA
LEA
LE
LS
LD
LBA
JUL
SEL
PL
CHD
CLU
LMO
TL
FWL
NGR
LFY
LTK
LPO
GRL
LEM
LRD
LTP
LSP
BR
LCT
FHD
GBL
KIDLAW
RRDCOD
SWLUQ
LMB
LH
HOL
GHL
WL
Figure 3.6 Final principal components analysis (PCA) with all environmental data (excluding ion concentration data) entered as active variables. (a)
The 45 water bodies, along with (b) morphology, environment, climate and land-use variation patterns. The primary y-axis is a trophic status (total
nitrogen (TN), total phosphorus (TP), Chlorophyll a), conductivity and agriculture gradient; the second axis is aligned to temperature. The PCA also
indicates that eutrophic saline lakes tend to be shallow lakes located in arid regions.
AXIS 1
AX
IS 2
a.
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We undertook additional PCA analyses to compare the changes in limnology for artificial lakes and
natural lakes along a gradient that reflects lake depth and human impact (see Fig. 3.7a, b). The Axis
1 sample scores from the first PCA (Fig. 3.7a) indicated the amount of agricultural activity in the
lake catchments. The Axis 1 sample scores for the second PCA (Fig. 3.7b) indicated the trophic
status of the lakes. Plotting the Axis 1 scores for both PCAs as a bi-plot allowed us to visualise the
relationship between stressors (agriculture) and response (trophic status). Figure 3.7c displays the
regression of the PCA axis scores for all lakes, whereas Fig. 3.7d displays the artificial lakes only.
The trend of the regression was similar on both graphs, indicating that artificial lakes were similar
to natural lakes in response to land use in the catchment. The conclusion is perhaps unsurprising;
artificial lakes in agricultural catchments also tend to be more eutrophic.
3.5.3 RDA results
RDA results showed that ~53.2% of the variation in trophic status was explained by the combined
AI, depth and agriculture data (see Table 3.4). Individually, AI and lake depth (depth) each
accounted for 29.8% of the variation in trophic status, whereas agriculture accounted for 22.6% (for
all, P < 0.001). When combined, lake depth and aridity accounted for 50.1% of the total variation.
When the effect of lake depth and aridity was partialled out, agriculture accounted only for 6.3% of
the variation and this was not statistically significant.
Table 3.4 Partial redundancy analysis (RDA) results for trophic status as limnological variable.
This table demonstrates that the effect of climate and water depth is more significant than the
agricultural status of the basin. In short, shallow water bodies in semi-arid settings are more
eutrophic than are other lakes. AI, aridity index; Chl a, Chlorophyll a; TN, total nitrogen; TP, total
phosphorus
Limnological variable Environmental variable Co-variable Explanatory power
(%) P-value
Trophic (TN,TP,Chla) Agriculture None 22.6 0.001
Trophic (TN,TP,Chla) Depth None 29.8 0.001
Trophic (TN,TP,Chla) AI None 29.8 0.001
Trophic (TN,TP,Chla) Depth, AI None 50.1 0.001
Trophic (TN,TP,Chla) Agriculture, depth, AI None 53.2 0.001
Trophic (TN,TP,Chla) Agriculture Depth, AI 6.3 0.082
Trophic (TN,TP,Chla) Depth, AI Agriculture 42.0 0.001
Trophic (TN,TP,Chla) Depth AI, agriculture 26.6 0.001
Trophic (TN,TP,Chla) AI Depth, agriculture 22.0 0.001
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a) b)
AX
IS 2
AXIS 1 AXIS 1
AX
IS 2
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c) d)
Figure 3.7 Principal components analyses (PCAs) to compare the changes in limnology for natural lakes and artificial lakes along a gradient that
reflects lake depth and human impact. The Axis 1 sample scores from plot (a) shows the amount of agricultural activity in the lake catchments. The
Axis 1 sample scores from plot (b) shows the trophic status of the lakes. The Axis 1 sample scores for both PCAs (a and b) are used to visualise the
relationship between land use, aridity and limnology. The regression for (c) all lakes and (d) artificial lakes only. The trend of the regression is similar
between the two graphs, indicating that artificial lakes have traits similar to those of natural lakes. The lower slope on the artificial lakes may indicate
lower residence times. Overall, these results demonstrate that lakes in agricultural catchments are more eutrophic; however, see the main text because
this trend is overwritten by climatic effects and lake morphology.
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3.6 Discussion
There are four major points to highlight from these results. These relate to (1) the contrast between
natural and artificial water bodies, (2) salinity, pH and rock type, (3) N and P limitation and (4) the
interaction among climate, lake morphology and human impacts.
3.6.1 The effect of including reservoirs and other artificial waterways
The trophic status and chemistry of both artificial waterways and natural lakes reflected land cover
and land use in the catchment. However, exploring the data further revealed that the key difference
between the regressions in Fig. 3.7c, d was the range of the data. The artificial lakes did not include
sites that were strongly eutrophic or strongly affected by agriculture. This finding was entirely
predictable. Typically, reservoirs and recreation lakes are managed so as not to be susceptible to
algal blooms and, thus, nutrient fluxes from the catchment are normally designed or managed to be
low.
The other factor that may affect artificial lakes is residence time (Søballe and Kimmel 1987), with
an expectation that the residence time in artificial lakes is lower than in natural lakes. Søballe and
Kimmel (1987) suggested that, for the same nutrient input, reservoirs tend to have lower
productivity because turnover times are faster. Our data showed a weak indication that this might be
the case, because the slope on our regression was lower for the artificial lakes, although the shorter
range of the data limited our ability to make a strong statement. Nevertheless, it is highly likely that
residence times in artificial lakes in Australia are shorter than in natural lakes and we predict that
this effect will also be significant.
3.6.2 Salinity, pH and bedrock effects
The primary source of the salts in eastern Australian lakes is from the ocean and old marine bedrock
and there were higher concentrations in drier regions (Figs 3.2, 3.3). Salt concentration is highest in
lakes in arid zones with high evaporative flux, and in maar craters where there is no outflow and
long residence times.
Lakes on basalt and limestone are on average relatively alkali. However, the alkalinity varies quite
strongly with climate (aridity). The northern Queensland maars are much less alkali than the
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western Victorian ones. This is an effect of the higher precipitation and higher P : EP ratio in
montane near coastal Queensland than in semi-arid western Victoria.
Overall, we conclude that salinity is a stronger control on pH than is bedrock type. This can be
observed in the Fig. 3.3 bi-plot. This figure demonstrates the buffering effect of salinity in that all
highly saline lakes are alkali, whereas there is more variability in the pH of freshwater lakes. These
freshwater lakes are controlled by bedrock type and, in particular, by the abundance of HCO3-. In
more saline lakes, H+ is buffered by Na
+ and other cations.
3.6.3 Nitrogen and phosphorus limitation
In the present paper, we used the molar ratio of TN : TP as a guide to the dominant nutrient limiting
algal productivity in each lake. Assigning each lake to a TN, TP, or co-limited (TN and TP)
category was based on the suggested ranges published in Guildford and Hecky (2000). On the basis
of the criteria of Guildford and Hecky (2000), 56% of the lakes in our dataset are co-limited, 35%
are P-limited, and only 9% are N-limited (Table 3.1). Inferences of nutrient limitation based solely
on TN : TP ratios should be considered a preliminary assessment and should be followed up by
macronutrient enrichment experiments (laboratory bioassays or in situ studies). This is especially
important because the reliability of the TN : TP ratio as a proxy for nutrient limitation has been
challenged (White et al. 1985). Although the TN : TP ratio has commonly been used as an indicator
of nutrient limitation in lakes (e.g. Abell et al. 2010), some studies (such as Morris and Lewis 1988;
Bergström 2010) have suggested that the dissolved inorganic N (DIN) : TP ratio is a better indicator
for nutrient limitation.
3.6.4 The interaction of human impact and climatic effects on lake nutrient status
The nutrient status of most mainland Australian lakes ranges from eutrophic to hypereutrophic, with
only the Kosciuszko lakes and some northern Queensland crater lakes classed as oligotrophic.
McComb and Davis (1993) identified key factors that caused water bodies in southwestern
Australia to become eutrophic, and we argue that these factors also apply widely in eastern
Australia. First, Australian soils frequently have poor nutrient-retention capacities. When fertilisers
are added (containing N and P), they are poorly absorbed by the soils and are washed into the
waterways. Second, the coincidence of the wet season with minimum ground cover as a result of
ploughing and/or harvesting, especially in the Mediterranean climate zones of southern Australia,
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enhances run-off and compounds problems of nutrient flows to the lakes. And finally, many
Australian lakes are small and ephemeral. They tend to have few or no outflows (endorheic basins)
and this results in the concentration of nutrients and ions by evaporative flux. McComb and Davis
(1993) concluded that nutrients are derived mainly from fertiliser applications in the catchments and
so inferred that the trophic status is primarily a human artefact.
In contrast, Beklioglu et al. (2011) investigated the eutrophication of shallow lakes on a global
scale. For lakes in Mediterranean climates, they concluded that nutrient concentrations in these
lakes could rise rapidly during drought periods, as a result of evaporative flux and internal
processes. They also noted that changes in lake levels suppress macrophyte growth and ‘tip’ the
lake toward algal production, thus enhancing eutrophication.
Beklioglu et al. (2011) concluded that these effects could control the concentration of lake nutrients
and productivity, even where human impact was low. We used forward selection to identify the
most important variables in explaining nutrient variability and then undertook partial RDAs on
these. Interestingly, when we partialled out the elements that relate to high nutrient status in lakes
during a series of RDAs (Table 3.4), we observed that whereas water depth and aridity are both
significant as independent variables, agricultural land use is not significant after the effects of water
depth and aridity are partialled out (see Table 3.4). Conversely, lake water depth and aridity (or
combined effect) retained their significance level after agriculture land use was partialled out. What
this suggests is that shallow Australian lakes in semi-arid climate are susceptible to eutrophication
irrespective of the surrounding land use and that agricultural practice simply enhances a process that
would naturally occur anyway. Paleoecological studies to examine pre-agricultural systems would
be an obvious method to test this slightly unexpected observation.
On a world scale, eutrophication is recognised as a major impact on lakes and is most often
attributed to anthropogenic activities (Moss 1998; Smith et al. 1999; Rovira and Pardo 2006).
Clearly, in most settings, agriculture and industrial effluent are the major sources of eutrophication.
However, in much of semi-arid and arid Australia, it appears that agriculture is at best a secondary
effect and that the lakes are likely to be naturally moderately to strongly eutrophic. Beklioglu et al.
(2011) made a point of implicating droughts and variable lake levels in the alteration of lake
behaviour. Australia has the highest inter-annual variability in rainfall of any continent on earth and
if fluctuating water levels and intermittent droughts are indeed the key to natural eutrophication,
then perhaps the provisional results presented here are unsurprising. Clearly, suggesting that
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eutrophication in many Australian systems is ‘natural’ is a significant conclusion and we propose it
as a testable hypothesis rather than a demonstrated occurrence at this point.
3.7 Conclusions
The main conclusions from this study are as follows:
(1) Most mainland Australian lakes are naturally moderately to significantly eutrophic. Oligotrophic
lakes are limited to high alpine and perennially wet settings.
(2) Reservoirs and other artificial lakes behave similarly to natural lakes, except that they tend to be
less eutrophic. This is most likely due to catchment management and a shorter residence time.
(3) The primary source of the salts in east Australian lakes is seawater, either derived directly from
aerosols or indirectly via marine bedrock.
(4) Significant number of lakes in this dataset that are limited by N and P, suggesting that both of
these nutrients should be considered in addressing eutrophication in Australian lakes.
(5) Mainland lakes are susceptible to becoming highly alkaline as a result of long term increases in
salinity that are related to the concentration of cations during drought periods.
(6) We propose as a testable hypothesis that climatically related eutrophication is more significant
than nutrient loading by agriculture in the eastern Australian lakes that we have examined. This
is particularly likely for shallow endorheic basins in semi-arid regions.
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4 A CHIRONOMID BASED TRANSFER FUNCTION FOR
RECONSTRUCTING SUMMER TEMPERATURES IN SOUTH
EASTERN AUSTRALIA
Chapter 4 is published in Palaeogeography, Palaeoclimatology, Palaeoecology (Chang et al.,
2015a)
4.1 Summary
This chapter reports the development of a chironomid based summer temperature transfer function
from a training set spanning a latitudinal gradient from the subtropics to Tasmania in southeastern
Australia.
Highlights:
First subfossil chironomid based transfer function from mainland Australia
Reconstructs mean February temperatures and will give new tool for quantitative
palaeoclimate estimates from Australia
Reservoirs were included in the development of the training set.
Integrating the existing chironomid transfer function from Tasmania with this new model is
a real possibility.
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4.2 Abstract
We present a new chironomid based temperature transfer function which was developed from a
training set of 33 natural and artificial lakes from southeast Australia from subtropical Queensland
to cool temperate Tasmania. Multivariate statistical analyses (CCA, pCCA) were used to study the
distribution of chironomids in relation to the environmental and climatic variables. Seven out of
eighteen available variables were significantly (p < 0.05) related to chironomid species variation
and these were mean February temperature (9.5%), pH (9.5%), specific conductance (8.2%), total
phosphorous (8%), potential evapotranspiration (8%), chlorophyll a (6.9%) and water depth (6.2%).
Further pCCA analyses show that mean February temperature (MFT) is the most robust and
independent variable explaining chironomid species variation. The best MFT transfer function was
a partial least squares (PLS) model with a coefficient of determination (r2
jackknifed ) of 0.69, a root
mean squared error of prediction (RMSEP) of 2.33 °C, and maximum bias of 2.15 °C. Chironomid
assemblages from actively managed reservoirs appear to match assemblages from equivalent
natural lakes in similar climates and therefore can be included in the development of the chironomid
transfer function. Although we cannot completely rule out some degree of endemism in the
Tasmanian chironomid fauna, our analyses show that the degree of endemism is greatly reduced.
Therefore, integrating the existing chironomid transfer function for Tasmania (Rees et al., 2008)
with this new model is a real possibility.
Key words: chironomids, southeast Australia, summer temperature, transfer function,
palaeoclimate, lakes
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4.3 Introduction
In temperate Australia, high-resolution continuous palaeoenvironmental records are scarce due to
the relative paucity of permanent water bodies (see Chapter 3, section 3.1). As a result,
palaeoenvironmental research is focussed on a few regions where these records exist (Petherick et
al., 2011). Continuous records that extend back to the last glaciation maximum (LGM: c. 21,000
years ago) are rare and geographical coverage is poor (Petherick et al., 2013). Pollen is the most
widely used proxy for these palaeoenvironmental reconstructions, and most of the reconstructions
of climate in Australia are qualitative. This has limited the climate inferences that can be made from
these records, which is unfortunate, as there are few estimates of the absolute scale of temperature
change between glacial and interglacial times. Bioclimatic modelling has been used with some
success (D'Costa and Kershaw, 1997) but the technique does not allow biotic effects to be easily
separated from climate change (Jeschke and Strayer, 2008). Statistical approaches using pollen
transfer functions have been developed for mainland Australia (Cook and Van der Kaars, 2006) and
for Tasmania (Fletcher and Thomas, 2010). The former has not been widely applied and the latter is
appropriate only to Tasmania. Transfer functions have also been developed using other organisms,
such as diatoms (Tibby, 2004; Tibby and Haberle, 2007), but diatoms are used primarily for salinity
and other limnological variables rather than temperature estimates. Molluscs (Edney et al., 1990;
D'Costa et al., 1993) and beetles (Porch et al., 2009; Sniderman et al., 2009) have also been applied
but a critical gap remains in the tools that are available to determine past changes in temperature in
Australia.
Chironomids (Diptera: Chironomidae) have been widely used as a proxy in palaeoclimate and
palaeoenvironmental studies (Walker and Paterson, 1985; Hofmann, 1986; Walker, 1987). Since
temperature is a dominant factor in every aspect of the chironomid life cycle (e.g. egg hatching,
larval and pupal development, adult emergence) (Armitage, 1995), many studies have focused on
the influence of temperature on the distribution and abundance of chironomids, in the context of
past climate change (Walker, 2002; Porinchu and MacDonald, 2003; Walker and Cwynar, 2006).
Most chironomid-based reconstructions have been carried out in temperate and sub-polar regions of
the Northern Hemisphere, including areas in central and northern Europe (Olander et al., 1999;
Larocque et al., 2001; Brodersen and Anderson, 2002; Luoto, 2009; Heiri et al., 2011) and northern
North America (Porinchu et al., 2002, 2009; Barley et al., 2006; Brodersen et al., 2008; Medeiros
and Quinlan, 2011). There are only few applications in the Southern Hemisphere and these are from
New Zealand (Boubee, 1983; Woodward and Shulmeister, 2006; Dieffenbacher-Krall et al., 2007),
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Tasmania (Rees et al., 2008), east Africa (Eggermont et al., 2010) and South America (Massaferro
and Larocque-Tobler, 2013). A summary of the training sets on a global scale is presented in
Appendix 1.
Early palaeoecological and palaeoclimate investigations of chironomids from the Southern
Hemisphere were restricted to semi-quantitative interpretations because the numerical techniques
for creating transfer functions were still in an early stage of development. For example, Boubee
(1983) and Schakau (1993) investigated the modern chironomid distribution from New Zealand
lakes using cluster analysis and ordinations. Classification techniques were applied down-core to
interpret the fluctuations in fossil chironomid abundances and species from a 6000 year record from
Lake Grasmere in New Zealand (Schakau, 1991) and a glacial transition to Holocene record from
Blue Lake in alpine Mount Kosciusko (Chapter 3, Fig. 3.1), Australia.
Dimitriadis and Cranston (2001) performed the first quantitative reconstruction based on
chironomids from eastern Australian lakes. Instead of using chironomid head capsules in the
training set, they used the presence and abundance of chironomid exuviae from 68 water bodies in
eastern Australia. Dimitriadis and Cranston (2001) then used the mutual-climate-ranges (MCRs) of
the pupal exuviae in the training set to create a Holocene climate reconstruction from Lake Barrine
(Atherton Tableland, northeast Queensland, Chapter 3, Fig. 3.1) based on the down-core
chironomid head capsule record. They inferred up to 6 °C temperature change in the Holocene.
The first Southern Hemisphere transfer function based on chironomid head capsules was developed
by Woodward and Shulmeister (2006) for New Zealand. Both summer temperature and chlorophyll
a (Chl a) transfer functions were presented. The summer temperature transfer function was
successfully applied to a record spanning the marine oxygen isotope stage 3/2 transition (~26,600–
24,500 cal yr BP) from lake deposits in Lyndon Stream, New Zealand (Woodward and Shulmeister,
2007). A second independent chironomid model for New Zealand was produced by Dieffenbacher-
Krall et al. (2007), which also yields satisfactory reconstructions (Vandergoes et al., 2008).
In Australia, a head capsule based chironomid transfer function was developed by Rees et al. (2008)
for summer temperature from Tasmanian lakes. This transfer function was applied to produce
lateglacial (~16,000 cal yr BP) to late Holocene summer temperature reconstructions from Eagle
Tarn and Platypus Tarn in Mount Field National Park, Tasmania (Chapter 3, Fig. 3.1) (Rees and
Cwynar, 2010). Although the Tasmanian chironomid transfer function appears to be robust,
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biogeographical controls on the Tasmanian chironomid taxa may prevent the application of the
Tasmanian transfer function to mainland sites.
Concerns over the influence of biogeography on Tasmanian and mainland Australian chironomid
taxa stem from a chironomid exuviae survey of eastern Australian lakes by Wright and Burgin
(2007). Wright and Burgin argue for the presence of 23 endemic taxa from Tasmania, including 5
genera, 4 species and 14 undescribed morpho-species. Despite this assertion, Wright and Burgin's
study does not completely rule out the possibility of applying the Tasmanian transfer function to the
mainland or producing a combined mainland and Tasmanian transfer function. Wright and Burgin
(2007) did not include mainland alpine lakes in their study and the degree of Tasmanian endemism
may be over-estimated. The level of taxonomic resolution provided by exuviae is typically higher
than for chironomid head capsules. Even if there are endemic chironomid species in Tasmania, their
presence might not dramatically affect the composition of sub-fossil chironomid head capsule
assemblages.
Here we present a temperature transfer function based on sub-fossil assemblages of Chironomidae
(non-biting midges), from 33 southeast Australian lakes. We included 7 lakes from Tasmania in this
total as an initial test of the feasibility of producing a chironomid training set combining both
mainland and Tasmanian lakes. We assessed Wright and Burgin's argument for Tasmanian
chironomid endemism using our new training set and published information on the distribution of
Wright and Burgin's endemic taxa. A combined training set would be desirable because it is
difficult to find non-impacted, permanent freshwater lakes spread continuously along a long
temperature gradient on mainland Australia. Due to the rarity of freshwater bodies on the mainland,
we also investigated the possibility of including artificial water bodies in the training set.
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Table 4.1 Selected climatic and environmental variables for the thirty-four water bodies sampled from southeast Australia used for transfer function
development
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4.4 Materials and Methods
4.4.1 Study sites
This data set comprises 25 natural lakes and 8 artificial water bodies located in southeast Australia
(Chapter 3, Fig. 3.1, lake numbers 1-34 and Table 4.1). The transect covers a distance of 2500 km
along the east coast of Australia from Kureelpa, Queensland to Mount Field National Park,
Tasmania (25.96°S to 42.67°S, 140.18°E to 153.26°E) (Chapter 3, Fig. 3.1, lake numbers 1-34 and
Table 4.1). The climate ranges from subtropical in the north, to cool temperate in the south, and
hence there are large temperature and precipitation gradients in the data set. Elevation of the sites
ranges from sea level to ~2000 m above mean sea level (a.s.l) (Table 4.1), corresponding to
estimated mean February air temperatures (MFT) of 10.7–24.7 °C (Table 4.1). A detailed
description of the climate, vegetation and geology of the study area is provided in Chapter 3, section
3.
4.4.2 Chironomid collection and analysis
The lakes were sampled during the Southern Hemisphere summer (January or February) of 2012
and 2013 (Chapter 3, Fig. 3.1 and Table 4.1). Lakes were selected along an altitudinal and
latitudinal range to ensure a long temperature gradient. Where possible we sampled shallow to
medium depth (between 1 and 10 m) lakes to ensure a close relationship between bottom water
temperature and air temperature. The sampling of very deep ( >30 m), stratified lakes was avoided
to eliminate the effect of hypolimnetic anoxia on the chironomid species assemblages (Little and
Smol, 2001). A minimum of three sediment cores were collected using a Glew Mini Corer (Glew,
1991) at the deepest point or lake centre where bathymetry was not available. The top 2 cm of each
core were extruded on site and packaged at 0.5 cm intervals in Whirlpak® sample bags. Sediment
samples were refrigerated prior to analysis.
Surface sediment core (top 1-2 cm) samples were processed for chironomid analysis following the
method outlined in Hofmann (1986) with the following modification. Samples were deflocculated in
warm 10% KOH for 20 min and washed on a 90 µm mesh with distilled water. Samples were
transferred to a Bogorov counting tray and examined under a dissection microscope at 50 ×
magnification. Chironomid head capsules were hand-picked using fine forceps onto a glass slide,
until a minimum of 100 head capsules were obtained (when possible). Chironomid head capsules
were mounted on glass slides in a drop of Euparal® and covered with a glass coverslip. Head
capsules were mounted ventral side up to assist identification. Chironomid species were identified
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using a compound light microscope at 400 × magnification, following the published identification
guides by Freeman (1961), Cranston (2000a, 2010), Brooks et al. (2007) and Dieffenbacher-Krall et
al. (2008).
Several studies have examined the minimum number of chironomid head capsules that should be
extracted to provide representative samples. Larocque (2001) found that 50 head capsules is the
minimum required to provide an accurate temperature estimate, but counts of 90 head capsules or
above will give much better representation of taxa in the assemblage. Quinlan and Smol (2001)
concurred with this finding, whereas Heiri and Lotter (2001) argued that the minimum count size is
model and location dependent. We therefore used rarefaction analysis (Birks and Line, 1992) in R
(version 2.11.1, R Development Core Team, 2010) and the Vegan package (version 2.0-10,
Oksanen et al., 2010) to test how representative different sample sizes were in our training set. A
plot of observed species richness vs predicted species richness was derived using the ‘estimateR’
function which uses Chao's method (Chao, 1987) to estimate actual richness. We also created
multiple rarefaction curves for each site based on multiple random subsamples from the full species
pool. This allowed us to visualise the effect of simulated increased sampling intensity on the
observed species richness.
4.4.3 Lake water chemistry and environmental variables
Water samples for chemical analyses were collected from 30 cm below the water surface at the
location where the core samples were taken. Untreated water samples were collected for the
analysis of major ions (Na+, K
+, Ca
2+, Mg
2+, HCO
3-, Cl
-, SO4
2- (Chapter 3, Table 3.2) and total
nitrogen/total phosphorus (TN/TP) analysis. These samples were kept frozen until analysis. A 1000
ml water sample was filtered for Chl a through a 4.7 cm diameter GF/F filter (0.45 µm pore
size).The Chl a filter was wrapped in foil and kept frozen for subsequent analysis. The 125 ml water
sample was filtered using a syringe and a Whatman® syringe filter (Supor® Membrane 0.45 µm
pore size) and the filtered water was frozen in the bottle for later analysis for dissolved reactive
phosphorus (DRP) and reactive nitrogen (NOX ). Total dissolved solids (TDS), pH, specific
conductance (COND) and turbidity (TURB) were also obtained from water chemistry analyses
which were carried out by the Forensic and Scientific Services, in Brisbane, Queensland. In the field,
water temperature, oxidation reduction potential (ORP), pH, dissolved oxygen (DO), specific
conductance (COND), total dissolved solids (TDS), salinity (SAL), and turbidity (TURB) were
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recorded from 30 cm below the water surface using an Aquaread® multi-parameter meter. Lake
depth at the sampling point was measured using a Speedtech® portable depth sounder.
Figure 4.1 (a) Plot of observed species richness vs predicted species richness, (b) rarefaction curves
for individual sites indicating estimated species richness with respect to increasing sub-sample size.
Rarefaction curves begin to flatten once true species richness is achieved. Together, these plots
indicate that sample sizes are adequate for most samples to capture all but the most rare species.
Only one site (CTL) with a low head capsule count may possibly underestimate the true species
richness. Minimum sample size varied from site to site and counts as low as 50 may be sufficient to
capture the actual species richness (e.g. LPO).
Climate variables were obtained using the combination of the WorldClim program (available from
http://www.worldclim.org/ bioclim, accessed 20 August 2013) and ArcGIS 10.1. WorldClim data
for Australia is based on climate surfaces derived from around 600 nation-wide weather stations
that have climate records spanning the years 1950–2000
(http://www.bom.gov.au/climate/data/stations/, accessed 20 January, 2014). For this study, mean
February temperature (MFT), mean annual temperature (MAT) and mean annual precipitation
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(Precip) were considered. The potential evapotranspiration (PET) values were obtained from the
Global Potential Evapo-Transpiration (Global-PET) and Global Aridity Index (Global-Aridity) data
set (CGIAR-CSI, available from http://www.cgiar-csi.org/data/globalaridity-and-pet-database,
accessed 20 August 2013).
4.4.4 Statistical analyses
Test for Tasmanian endemism and difference between natural and artificial lakes
A principal components analysis (PCA) of the chironomid taxa data was run for the 33 lakes. We
tested to determine if the chironomid assemblages were significantly different in mainland lakes
compared to Tasmanian lakes to look for potential biogeographical effects. We also tested for
significant differences between natural and artificial lakes. The distribution of both sets, PCA axis
scores was assessed for normality using the Shapiro–Wilk test (Shapiro and Wilk, 1965)
respectively. The data were normally distributed around the mean so that a Student t-test was
appropriate to test significance of the results. We first performed a t-test (α = 0.05, two sample
assuming unequal variances) on sample axis scores from a principal components analysis (PCA) of
the chironomid taxa based on head capsule assemblages for Tasmania vs. mainland lakes. This is
not an exhaustive test for endemism as it is based on the taxonomic resolution possible with head
capsules. In order to further test for endemism in the Tasmanian chironomid taxa we performed a
literature search for records of Wright and Burgin's (2007) endemic Tasmanian taxa, and searched
the Australian National Insect Collection records (Atlas of Living Australia database:
http://bie.ala.org.au/species/CHIRONOMIDAE, accessed 20 July 2014). The same t-test was
performed on natural vs. artificial lakes.
Selection of environmental variables and model development
Constrained ordinations were performed using CANOCO version 4.5 (ter Braak and Šmilauer,
2002) to determine which variable(s) explained a significant proportion of the variation in the
chironomid species data. Prior to ordination, climate and lake water chemistry data were assessed
for normality in Minitab 16® using the Shapiro–Wilk test (Shapiro and Wilk, 1965), and through
measurements of skewness and kurtosis (Zar, 1999). All variables apart from MAT, MFT and pH
were normalized using a log10 transformation. Chironomid species were used in the form of square
root transformed percentage data.
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A detrended correspondence analysis (DCA) (with rare taxa down-weighted) of the chironomid data
was used to determine whether linear or unimodal methods were appropriate for selecting the best
candidates for model construction. The gradient length for DCA axis 1 was 2.067 standard
deviation units, which means that a unimodal technique (Canonical correspondence analysis, CCA)
is appropriate (Birks, 1998). CCAs were run to determine which environmental variables explained
the highest and most significant amount of the variation in the chironomid species data. The
environmental variable with highest variance inflation factor (VIF) was removed after each CCA
and the CCA was repeated until all VIFs were less than 20 (ter Braak and Šmilauer, 2002). The
ability of the remaining environmental variables to explain a statistically significant amount of the
variation in the chironomid species data was determined using a series of CCAs with manual
forward selection and a Monte Carlo permutation test (999 unrestricted permutations) (ter Braak
and Šmilauer, 2002). Variables that were significant (p ≤ 0.05) were retained for further analyses.
To test the strength of the explanatory power of each of the significant variables for the chironomid
distribution, a series of partial canonical correspondence analyses (pCCAs) were performed with
the remaining significant variables included as co-variables. This step was used to distinguish
between indirect and direct relationships between the environmental variables and the chironomid
species data. Only environmental variables that retained their significance after this step were
considered for transfer function development. CCA bi-plots of sample and species scores were
generated using CanoDraw (ter Braak and Šmilauer, 2002). Chironomid taxon response curves for
significant variables (p ≤ 0.05) were generated in CanoDraw using Generalized Linear Models
(GLMs) with a Poisson distribution.
Transfer functions for the significant environmental variable(s) selected in the CCAs and pCCAs
were developed in the computer program C2 (Juggins, 2005). A detrended canonical
correspondence analysis (DCCA) was used to determine whether chironomids were responding in a
linear or unimodal fashion along the gradient of the environmental variable selected for model
development (Birks, 1998). Leave-one-out, cross-validation was used as this technique is more
robust for data sets with fewer than 80 sites (Kim and Han, 1997).
Transfer functions were selected based on the performance of the jack-knifed coefficient of
determination (r2
jackknifed), average bias of jack-knifed predictions (AveBias jack), maximum bias of
jack-knifed predictions (MaxBias jack), and root mean square error of prediction (RMSEP) (Birks,
1998). Additional components were only included in the model if the addition of an additional
component reduced the RMSEP by at least 5% (Birks, 1998).
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4.5 Results
4.5.1 Chironomid taxa
The average sample size in the training set is 128 head capsules and only two samples (Chaffey
Dam (n = 0) and Lake Cootapatamba (n = 59)) produced fewer than 100 head capsules. Counts of
over 100 are generally regarded as reliable (Heiri and Lotter, 2001). Rarefaction analysis (Fig. 4.1)
indicated that the sample size for most sites was adequate for including all of the common
chironomid species. There is a significant correlation between observed and predicted species
richness (Fig. 4.1a) and only one site (Lake Cootapatamba) produced a low head capsule count (59)
which possibly under-represents the full chironomid species pool. All head capsules from the top 2
cm of the Cootapatamba sediment core were used. The plot of multiple rarefaction curves (Fig.
4.1b) indicates that the minimum number of counted head-capsules that is sufficient to capture the
actual species richness varies from site to site.
One site (Chaffey Dam) had no head capsules and was removed from the dataset for further
analyses. Forty-three (43) chironomid taxa were identified and counted from the training set. Five
rare species were removed from further analyses as they have a maximum abundance of less than
2% and/or occurred in fewer than two lakes (Brooks and Birks, 2001).
4.5.2 Test for Tasmanian endemism and difference between natural and artificial
lakes
Chironomid species assemblages for the seven Tasmanian lakes show no significant difference to
mainland lakes based on t-test results (Fig. 4.2a, Table 4.2a). Fourteen (14) of Wright and Burgin's
23 endemic taxa are undescribed morpho-species (Table 4.3), so it is not possible to assess these
taxa for endemism. The remaining 9 endemic taxa comprise 5 genera and 4 species. Head capsules
from one of Wright and Burgin's (2007) endemic Tasmanian taxa (Thienemanniella sp.) were found
on the Australian mainland in Blue Lake and Lake Albina (Mount Kosciuszko), from our training
set. Head capsules from Wright and Burgin's (2007) other 4 “endemic” Tasmanian genera
(Apsectrotanypus sp., Pentaneurini genus E, Nanocladius sp., Orthocladiinae “MO5” (Now =
Echinocladius sp. Cranston)) were not found in our training set but have been previously recorded
from mainland sites by Marchant et al. (1999), Cranston (2000a), Cranston (2000b), and Krosch
(2011) respectively (Table 4.3). Wright and Burgin's (2007) four endemic Tasmanian species
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(Botryocladius australoalpinus, B. grapeth, Riethia plumosa, Tanytarsus liepae) were not identified
in the training set, but have been previously recorded from the Australian mainland by Cranston and
Edward (1999), Cranston and Edward (1999), Cranston (Australian National Insect Collection,
available from http://bie.ala.org.au/species/CHIRONOMIDAE, accessed 20 July 2014), and
Cranston (2000a) respectively (Table 4.2).
Figure 4.2 PCA plots for exploring the difference between (a) mainland and Tasmanian lakes and
(b) natural and artificial lakes. PCA axis 1 and axis 2 explains 16.9% and 10.8% of the variance in
chironomid species data respectively. A t-test was performed on the sample score means for each
(Tables 4.1a and 4.1b). The sample size for the Tasmania and artificial lakes is small (<10). There
are no significant differences apparent between axis 1 scores for both tests. There are no differences
in axis 2 scores either for Tasmania vs mainland lakes, but there is for natural vs artificial lakes.
Warm taxa are over-represented in artificial lakes that are not reservoirs (Fig. 4.3).
The t-test of PCA axis scores shows that there is no significant difference in the chironomid
assemblages from natural and artificial water bodies on PCA axis 1, but there is on PCA axis 2 (Fig.
4.2b and Table 4.2b). PCA axis 2 mainly separates warm stenotherms (low PCA axis 2 scores, e.g.
Dicrotendipes, Kiefferulus, Cladopelma) from cold stenotherms (high PCA axis 2 scores, e.g.
Parakiefferiella) (Fig. 4.3a). Warm stenotherms are more common in three high altitude, shallow
artificial lakes (not reservoirs) (Highland Waters (LD), Lake Samuel (LS) and Lake Cantani (LCN))
than other high altitude, natural lakes (Fig. 4.3b).
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Figure 4.3 This shows a PCA of chironomid species used to compare natural and artificial lakes. In
a) PCA axis 2 show separation of warm stenotherms (low PCA axis 2 scores, e.g. Dicrotendipes,
Kiefferulus, Cladopelma, Procladius, Paratanytarsus etc.) from cold stenotherms (high PCA axis 2
scores, e.g. Parakiefferiella). b) shows surface sediment samples plotted as circles where the circle
size is relative to mean February temperature (MFT). Large circles = high MFT, small circles = low
MFT. Values for MFT for each site are also provided. Open circles = natural water bodies, closed
circles = artificial water bodies. Note that some artificial water bodies (not reservoirs) with low
MFTs have low PCA axis 2 scores, which indicates that warm stenotherms are more common in
these sites compared to natural sites with similar MFTs.
Table 4.2 Statistical t-Tests for two samples assuming unequal variances performed on (a) natural
against artificial lakes and (b) Tasmania against mainland lakes. PCA scores of axis 1 and axis 2
were used from both datasets from Fig. 4.2a and Fig. 4.2b respectively.
(a)
Tasmania VS Mainland Lakes
AXIS 1
Variable 1 Variable 2
Mean 0.538314286 -0.144930769
Variance 1.105297491 0.951724631
Observations 7 26
Hypothesized Mean Difference 0
df 9
t Stat 1.549215177
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Tasmania VS Mainland Lakes
AXIS 1
Variable 1
Variable 2
P(T<=t) one-tail 0.077869722
t Critical one-tail 1.833112933
P(T<=t) two-tail 0.155739445
t Critical two-tail 2.262157163 No Difference
AXIS 2 Variable 1 Variable 2
Mean 0.6701 -0.180415385
Variance 2.580589327 0.541095722
Observations 7 26
Hypothesized Mean Difference 0
df 7
t Stat 1.362846194
P(T<=t) one-tail 0.107574164
t Critical one-tail 1.894578605
P(T<=t) two-tail 0.215148328
t Critical two-tail 2.364624252 No difference
(b)
Natural VS Artificial Lakes
AXIS 1
Variable 1 Variable 2
Mean -0.05321 0.017028
Variance 0.86037 1.122792
Observations 8 25
Hypothesized Mean Difference 0
df 13
t Stat -0.17989
P(T<=t) one-tail 0.430005
t Critical one-tail 1.770933
P(T<=t) two-tail 0.860011
t Critical two-tail 2.160369 No difference
AXIS 2
Variable 1 Variable 2
Mean -0.6306 0.201788
Variance 0.139766 1.159287
Observations 8 25
Hypothesized Mean Difference 0
df 31
t Stat -3.29437
P(T<=t) one-tail 0.001237
t Critical one-tail 1.695519
P(T<=t) two-tail 0.002473
t Critical two-tail 2.039513 Difference
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Table 4.3 Wright and Burgin (2007) identified 5 genera, 4 species and 14 morpho-species of
endemic chironomids in Tasmania. These are arranged in alphabetic order, below. We performed a
literature search and show that all the 5 genera and 4 species were previously recorded and
described in mainland Australia, therefore endemism is not supported at these taxonomic levels.
The 14 morpho-species are not described and we are unable to assess the reliability of the
endemism claim for these taxa. (*) Indicates undescribed morpho-species.
Name Genus Species Morphotype
species*
Comments
Apsectrotanypus sp. X Paratypes from mainland Australia (originally
Anatopynia maculosa Freeman 1961).
Apsectrotanypus also collected from the
mainland Marchant et al. (1999)
Botryocladius
australoalpinus
X Described only from Pupae (Cranston and
Edward 1999)
Botryocladius
grapeth
X Botryocladius spp. head capsules collected
from Blue Lake. Larvae collected from Blue
Lake (Cranston and Edward 1999)
Chironominae sp. 1 1
Chironominae sp. 2 1
Cricotopus sp. 1 1
Cricotopus sp. 6 1
Nanocladius sp. X Not found in the training set in Tasmanian or
mainland sites, however endemism to
Tasmania can be elimated since this genus has
been found in Western Australia (Cranston
2000a)
Orthocladiinae sp. 1 1
Orthocladiinae sp. 2 1
Orthocladiinae sp. 3 1
Orthocladiinae
"MO5"
(now Echinocladius
sp. Cranston 2000b)
X Echinocladius is distributed along the east
coast of Australia in streams from the tropics
in the north to Tasmania (Krosch 2011)
Parakeifferiella sp. 3 1
Pentaneurini genus
E (Cranston 2000a)
X Actually states in Cranston (2000a) that this
genus has also been found in New South
Wales
Polypediulum sp. 3 1
Polypediulum sp. 8 1
Riethia sp. 1 1
Riethia sp. 2 1
Riethia plumosa X Riethia plumosa is also found on the mainland.
Australian National Insect Collection e.g.
catalogue number 29-002080-767, pupal
exuviae, Mongarlowe River. Records
available from The Atlas of Living Australia
database
Tanytarsus sp. 11 1
Tanytarsus sp. 12 1
Tanytarsus liepae X Collected from Victoria and Southern NSW
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Name Genus Species Morphotype
species*
Comments
Thienemanniella sp. X Head capsules found in Blue Lake and Lake
Albina in the training set.
SUM 5 4 14
4.5.3 Selection of environmental variables and model development
Individual ions (see Chapter 3, Table 3.2), precipitation (Precip), and mean annual temperature
(MAT) were excluded prior to ordination. Cation and anion gradients are correlated in PCA space
and are better represented by specific conductance (see Chapter 3, Fig 3.4). Rainfall is usually a
secondary effect on chironomids species distribution where its effect on chironomids is through
influencing or altering the lake water chemistry by dilution and in-lake macrophyte composition and
structure through changes in water depth. However, potential evapotranspiration (PET) was
included in this dataset since evaporative balance drives salinity and nutrient gradients in sub-humid
and semiarid areas, especially for shallow lakes with endorheic basins (Chapter 3). In Chapter 3, we
concluded that PET is a much stronger climate driver for lake water chemistry and nutrient status
changes of the east coast Australian water-bodies than rainfall alone. The choice of summer
temperatures as a control on chironomids is routine for the alpine lakes and those in more temperate
settings. Only two of our sites (UQ lakes, Lake Poona) are not located in temperate settings as
defined by the Köppen–Geiger climate classification (Kottek et al., 2006), so it is reasonable to
assume that summer is the prime breeding period for chironomids in this study also.
Total nitrogen (TN) produced the highest VIF in a CCA with all selected environmental variables
and chironomid species and it was therefore not considered for further analyses. The remaining
seven variables individually accounted for a significant portion (p ≤ 0.05) of the variance (Fig. 4.4).
In order of explanatory power, these were mean February temperature (9.5%), pH (9.5%), specific
conductance (8.2%), total phosphorus (8%), potential evapotranspiration (8%), Chl a (6.9%) and
water depth (6.2%).
Southeastern Australian lakes in this training set cover large productivity and temperature gradients.
The first four axes of the CCA constrained by the seven significant environmental variables (Fig.
4.4) account for 28% of the variance in the chironomid species data (Table 4.4). Depth, MFT and
pH are significantly correlated to the first axis and Depth, TP, pH and specific conductance are
correlated with the second axis. Mean February temperature (MFT) shows the strongest correlation
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with the first axis (Table 4.4). Total phosphorus (TP) shows the strongest correlation with the
second axis (Table 4.4).
Partial CCAs (pCCAs) were then undertaken to determine the direct and indirect effects of each of
the seven significant variables. The pCCA results show that lake water depth (DEPTH), nutrient
variables (TP, Chl a) and specific conductance (COND) are confounded, while potential
evapotranspiration (PET) is correlated with specific conductance. pH retained 7.7% of variance
explained and remained significant (p ≤ 0.05) after the pCCAs with all other significant variables
partialled out (Table 4.5). Although MFT and PET appear to be confounded, PET is the dependent
variable because evaporation is partially a function of temperature and not vice-versa. In summary,
pH and MFT are the two primary independent parameters that should be considered for model
construction. They both explained the largest amount of variance (both 9.5%) (Table 4.5).
Table 4.4 CCA summary of the seven significant variables including canonical co-efficients and t-
values of the environmental variables with the ordination axes including 33 lakes and 38 non-rare
species
Axis 1 Axis 2 Axis 3 Axis 4
Eigenvalue 0.199 0.109 0.078 0.042
Cum % var. spp. 13.0 20.2 25.3 28.0
Cum % var. spp. - env. relation 38.5 59.7 74.8 83.0
Regression/canonical co-efficient
Variable
Depth 0.384 -0.463 0.704 -0.813
MFT -0.745 0.523 0.869 -0.193
PET 0.262 0.230 -0.645 0.336
TP 0.277 -1.009 0.002 -1.034
Chl a -0.302 -0.402 0.097 0.432
pH -0.650 -0.654 0.506 0.804
COND 0.115 0.912 -0.545 -0.991
t-values for regression co-efficients
FR explained 0.385 0.212 0.151 0.082
Variable
Depth 2.136 -2.432 3.453 -3.316
MFT -2.336 1.547 2.401 -0.442
PET 0.824 0.684 -1.787 0.775
TP 0.835 -2.871 0.005 -2.285
Chl a -1.032 -1.297 0.291 1.083
pH -3.207 -3.045 2.200 2.904
COND 0.443 3.316 -1.850 -2.797
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Although pH is also a good candidate for a transfer function, the focus here is on developing a
palaeoclimate proxy and MFT is a more useful parameter. The taxon response curves and a plot of
chironomid species turnover with respect to temperature (Figs. 4.5 and 4.6) show that the major
taxa (p ≤ 0.05 and N2 ≥ 5) which are dominant at warm sites are Polypedilum spp.,
Parachironomus spp., Coelopynia pruinosa type, Paratanytarsus spp., Tanytarsus lactescens type
and Procladius spp. Taxa that are typical of cool sites include Paralimnophyes morphotype 1and
Botryocladius. However, many taxa may respond to other environmental variables as well (e.g.
Cladopelma, Dicrotendipes and Kiefferulus, see Fig. 4.6 and Table 4.6). A few taxa such as
Chironomus, Procladius, Pentaneurini and undifferentiated Tanytarsini contain many species each
of which is likely to have different environmental responses but cannot be separated from similar
morphotypes.
Table 4.5 Partial CCAs of the seven significant (p ≤ 0.05) environmental variables alone and with
the effects of other significant variables partialled out for 33 lakes with 38 non-rare species
included.
Variable covariable λ1 λ
1/ λ
2 % Variance
explained
P-value
Depth none 0.095 0.302 6.2 0.016
MFT 0.100 0.412 7.2 0.002
PET 0.092 0.347 6.5 0.014
TP 0.062 0.207 4.4 0.163
Chl a 0.064 0.221 4.5 0.133
pH 0.077 0.304 5.6 0.031
COND 0.067 0.240 4.7 0.089
MFT, PET,TP, Chl a, pH, COND 0.071 0.353 6.5 0.034
MFT None 0.145 0.562 9.5 0.001
Depth 0.151 0.621 10.5 0.001
PET 0.062 0.240 4.4 0.150
TP 0.125 0.508 8.9 0.001
Chl a 0.112 0.446 7.9 0.001
pH 0.100 0.450 7.2 0.004
COND 0.095 0.380 6.8 0.007
Depth, TP, Chl a, pH, COND 0.069 0.332 6.2 0.029
Depth, PET,TP, Chl a, pH, COND 0.057 0.284 5.3 0.139
PET None 0.123 0.449 8.0 0.010
MFT 0.040 0.155 2.9 0.468
Depth 0.120 0.453 8.3 0.009
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Variable covariable λ1 λ
1/ λ
2 % Variance
explained
P-value
PET TP 0.100 0.377 7.1 0.012
Chl a 0.082 0.306 5.8 0.035
pH 0.081 0.343 5.8 0.044
COND 0.069 0.261 4.9 0.075
MFT, COND 0.036 0.145 2.7 0.523
Depth, MFT, TP, Chl a, pH,
COND
0.034 0.169 3.2 0.529
TP None 0.122 0.408 8.0 0.003
MFT 0.102 0.439 7.4 0.004
PET 0.099 0.374 7.0 0.009
Depth 0.089 0.297 6.2 0.025
Chl a 0.058 0.201 4.1 0.173
pH 0.066 0.261 4.8 0.083
COND 0.086 0.308 6.1 0.012
Chl a, pH 0.063 0.278 4.8 0.107
Depth, MFT, PET, Chl a, pH,
COND
0.046 0.229 4.4 0.282
Chl a none 0.106 0.367 6.9 0.004
MFT 0.073 0.291 5.3 0.032
PET 0.066 0.246 4.7 0.080
Depth 0.075 0.260 5.2 0.050
TP 0.042 0.145 3.0 0.517
pH 0.065 0.261 4.7 0.068
COND 0.046 0.167 3.2 0.456
Depth, MFT, PET, TP, pH, COND 0.029 0.144 2.8 0.765
pH None 0.146 0.574 9.5 0.001
MFT 0.101 0.462 7.3 0.003
PET 0.105 0.445 7.4 0.004
Depth 0.129 0.514 9.0 0.002
TP 0.091 0.360 6.4 0.005
Chl a 0.106 0.426 7.4 0.002
COND 0.091 0.361 6.4 0.011
Depth, MFT, PET, TP, Chl a,
COND
0.085 0.423 7.7 0.013
COND None 0.125 0.448 8.2 0.002
MFT 0.075 0.300 5.4 0.044
PET 0.071 0.269 5.1 0.055
Depth 0.097 0.348 6.8 0.004
TP 0.089 0.319 6.3 0.015
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Variable
covariable λ
1 λ
1/ λ
2 % Variance
explained
P-value
COND Chl a 0.065 0.236 4.5 0.127
pH 0.069 0.274 5.0 0.053
PET, Chl a, pH 0.059 0.251 4.7 0.114
Depth, MFT, PET, TP, Chl a, pH 0.058 0.289 5.4 0.125
4.5.4 The transfer function
The DCCA results (gradient length = 1.07 standard deviation units) suggest a linear response of
chironomid taxa along the mean February temperature gradient (Birks, 1998), therefore, a partial
least squares (PLS) model was appropriate for the transfer function construction (ter Braak and
Juggins, 1993) for MFT (Table 4.7) in C2 (Juggins, 2005). The third component of the PLS model
(with 3 components, jack-knifing, including 33 lakes and 38 non-rare species, Table 4.7) was
selected based on the criteria of Birks (1998). It produced a coefficient of determination (r2
jackknifed)
of 0.69, RMSEP jack of 2.33 °C, maximum bias jack of 2.15 °C and an AveBias jack of 0.07 °C
(Table 4.7, Fig. 4.7).
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Figure 4.4 CCA biplots of (a) sample and (b) species scores constrained to seven environmental variables that individually explain a significant (p ≤
0.05) proportion of the chironomid species data. Sites codes correspond to site names in Tables 3.1 and 3.2 (lake numbers 1-34). Taxon numbers
correspond to taxa in Table 4.5. Sites and taxa in warmer environments tend to plot in the upper left quadrant. Taxa typical of warmer sites include
Harnischia spp., Cryptochironomus spp., Polypedilum spp., Coelopynia pruinosa type, Cladopelma spp., Paratanytarsus spp., Procladius spp., and
Riethia spp., while sites and taxa in colder environments tend to plot in the lower right hand quadrant. Taxa typical of cold sites include
Paralimnophyes morphotype 3, Parakiefferiella morphotype 1, Orthoclad type 1, Orthoclad type 4, Pseudosmittia type 2, and Botrycladius. Eutrophic
sites and taxa typical of these environments tend to plot in the lower left hand quadrant, while oligotrophic sites and taxa typical of these environments
tend to plot in the upper right hand quadrant.
a. a.
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Figure 4.5 Taxon response curves for taxa that show a significant response to temperature (p <
0.05) using a Generalized Linear Model with Poisson distribution (ter Braak and Šmilauer, 2002).
(a) Taxa which are more common at lower temperatures, such as Paralimnophyes morphotype 1
and Parakiefferiella morphotype 2 respond strongly to cooling in temperature (b) Taxa which are
more common at higher temperatures demonstrate weaker but still significant responses. Examples
include Procladius, Polypedilum spp. and Tanytarsus lactescens type.
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Figure 4.6 Stratigraphy diagram of the 38 non-rare taxa included in the final model, where observed mean February temperature is on the y-axis and
taxon abundance is in percentage. Taxa such as Cladopelma, Dicrotendipes and Kiefferulus (*) show high abundance in lowland warm lakes but are
also present in highland artificial lakes (Grey bars).
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Table 4.6 List of chironomid taxa enumerated in this study along with data on distribution and environmental significance.
No. Taxa name N Hill's N2 Maximum Mean PLS β -
coefficent
(Jack-knifed)
Environmental variables that
have a significant relationship to
each taxon’s response (at p ≤
0.05) based on the GLM results
1 Chironomus Meigen 31 19.8 63.7 19.3 -0.2 -
2 Polypedilum nubifer Skuse 27 19.6 15.8 6.0 0.9 MFT
3 Cladopelma Kieffer 24 12.0 23.2 4.2 0.3 Depth
4 Dicrotendipes Kieffer 24 14.3 25.8 6.6 -0.1 Chla, Depth, TP, COND
5 Polypedilum spp. Kieffer 6 5.1 4.4 0.5 0.9 MFT
6 Cryptochironomus Kieffer 8 5.7 4.7 0.4 0.2 COND
7 Parachironomus Lenz 11 6.9 10.9 1.2 0.4 Chla, MFT, TP
8 Kiefferalus martini Freeman 16 11.7 18.5 3.5 0.3 Chla, Depth
9 Procladius Skuse 33 24.9 32.8 15.3 0.1 -
10 Coelopynia pruinosa Freeman 8 5.9 4.7 0.6 1.2 MFT
11 Pentaneurini undifferentiated 17 10.0 12.3 1.8 -0.7 COND
12 Riethia Kieffer 21 9.8 28.9 4.8 -0.3 COND
13 Tanytarsus lugens type 29 16.8 23.1 6.6 -0.3 -
14 Tanytarsus pallidicornis type 28 17.7 27.0 7.2 -0.3 TP
15 Tanytarsus glabrescens type 18 8.2 20.9 2.6 -0.4 Chla, TP, pH
16 Paratanytarsus Skuse 22 17.2 14.3 4.0 0.6 Chla, MFT, COND
17 Tanytarsus undifferentiated 22 17.3 6.4 1.9 0.1 MFT
18 Tanytarsus lactescens type 10 5.2 27.0 2.4 0.5 Depth, MFT, TP, pH
19 Tanytarsus nr chiyensis 5 3.8 3.6 0.3 -0.3 MFT, pH
20 Stempellina Thienemann & Bause 2 1.8 44.6 2.1 0 Chla, MFT, TP, pH
21 Harnischia Kieffer 4 2.9 3.1 0.2 1.2 MFT
22 Paralimnophytes type 1Unofficial morphotype 13 5.5 15.8 1.3 -0.5 Chla, MFT, COND, pH
23 Paralimnophytes type 2Unofficial morphotype 13 8.6 5.4 1.0 -0.3 Chla
24 Paralimnophytes type 3 Unofficial morphotype 9 6.9 3.6 0.5 -0.3 -
25 Parakiefferiella undifferentiated 7 6.0 4.3 0.6 -0.8 Depth
26 Parakiefferiella type1Unofficial morphotype 3 3.0 1.6 0.1 -0.3 -
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No. Taxa name N Hill's N2 Maximum Mean PLS β -
coefficent
(Jack-knifed)
Environmental variables that
have a significant relationship to
each taxon’s response (at p ≤
0.05) based on the GLM results
27 Parakiefferiella type 2 Unofficial morphotype 4 3.1 13.7 0.9 0 Chla, MFT, TP, COND, pH
28 Parakiefferiella type 3 Unofficial morphotype 4 3.2 2.9 0.2 -1.7 Chla, MFT, COND, pH
29 Botrycladius Cranston & Edward 9 5.4 7.0 0.7 -0.6 MFT
30 Smittia Holmgren 3 1.9 7.8 0.3 0.3 -
31 Gymnometriocnemus type 1 Unofficial morphotype 7 3.2 8.8 0.6 0.1 -
32 Genus Australia 2 2.0 1.2 0.1 -1.1 -
33 Thienemanniella Kieffer 4 2.7 5.8 0.3 -0.3 Chla, Depth, MFT, TP, COND
34 Ortholcad type 1 Unoffical morphotype 3 2.5 1.8 0.1 -0.8 Chla
35 Orthoclad type 4 Unoffical morphotype 8 7.3 3.4 0.5 -0.3 -
36 Pseudosmittia type 2 Unofficial morphotype 2 1.6 2.3 0.1 0.2 MFT, TP, COND
37 Kosciuszko Orthoclad type 1 Unoffical morphotype 3 2.4 2.7 0.1 -0.4 TP
38 Cricotopus ‘Parbicintus’ 2 2.0 1.4 0.1 -0.1 Chla, Depth, MFT, TP, pH
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4.6 Discussion
4.6.1 Can we include artificial lakes in temperature training sets?
Recent study (Chapter 3) showed that reservoirs and other artificial water bodies respond to
stressors in their catchments in a similar fashion to natural lakes. Despite the general preference
for natural lakes for temperature training sets, this observation is not unexpected as human impacts
that could change water quality in reservoir catchments are generally limited. We might expect
chironomid species composition in reservoirs and natural lakes in environmentally equivalent
settings to be similar.
However, we also note that for three high elevation artificial lakes (not reservoirs) (LD, LS and
LCN), chironomid assemblages resemble lowland eutrophic lakes, with high values of
Cladopelma, Dircrotendipes and Kiefferiulus (see Fig. 4.3, Fig. 4.6 and Table 4.6). These three
artificial water bodies comprise two nutrient rich trout stocked lakes in Tasmania and a shallow,
productive recreation lake on Mt Buffalo (Table 4.1, lake number 5, 9, 10). If the two Tasmanian
lakes are excluded, the distinction between warm and cold lakes is markedly diminished (Fig. 4.6).
On Mt Buffalo, the shallowness of Lake Cantani increases its mean temperature in summer,
making it resemble a lower elevation lake due to the increased abundance of Cladopelma.
In summary, chironomid assemblages from true reservoirs appear to match assemblages from
equivalent natural lakes in similar climates. We should be cautious about including other types of
artificial water bodies, especially those where the food web or trophic status might be significantly
altered.
4.6.2 Endemism and other considerations
In our dataset there was no significant difference between the chironomid assemblages from
Tasmania and the Australian mainland (Fig. 4.2a, Table 4.2a) for the taxonomic resolution that is
available for subfossil head capsules. This test can be considered a first order test of the possibility
for combining Tasmanian and mainland training sets. As we will mention below, we did detect a
genus in our dataset (Thienemanniella sp.) that was previously considered endemic to Tasmania
(Wright and Burgin, 2007), but this does not fully rule out some degree of Tasmanian endemism.
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To do this we need to have chironomid distribution records with the same taxonomic resolution
that was provided in Wright and Burgin's (2007) study on Australian chironomid biogeography.
Five genera (including Thienemanniella sp.) and four species that Wright and Burgin (2007)
identified as Tasmanian endemics have actually been reported from the Australian mainland
(Table 4.3). This means that the basis for Tasmanian endemism now relies on the presence of 14
undescribed morpho-species in Wright and Burgin's (2007) dataset. It was not possible for us to
further test for endemism in records of adult and larval chironomid distributions for morpho-
species that are based on exuviae alone. Further collection of exuviae from mainland lakes and
rearing of chironomid larvae to possibly associate morpho-species with other life stages of
described species is required to rule out endemism. In the absence of these data, alternative
methods for testing for endemism would include combining our dataset with Rees et al. (2008)
and splitting the pooled dataset into Tasmanian and Mainland sites. Mainland sites can then be
used to reconstruct temperatures from Tasmanian sites and vice-versa. This technique has been
used to compare trans-Atlantic chironomid datasets (Lotter et al., 1999). At this stage we can only
conclude that claims for endemism are greatly diminished, but we do not expect future efforts to
combine datasets to be thwarted by endemism.
4.6.3 Reconciliation and integration with Tasmanian transfer function
We recognize that the temperature error in this transfer function is relatively high in comparison to
other transfer functions. This reflects the long scalar length of the temperature gradient which
extends from sub-tropical to sub-alpine locations, some 14 °C. The RMSEP is 2.33 °C which
represents 16% of the scalar length. This is comparable with the recently developed western Irish
chironomid-based calibration set (Potito et al., 2014). The Potito et al. (2014) dataset has a
RMSEP of 0.57 °C, but the temperature range this dataset covers is only 3.8 °C; so the error
represents 15% of the scalar length. Furthermore, it has been observed that data sets with large
temperature gradients naturally have larger errors (Walker and Cwynar, 2006) but this in no way
diminishes the value of the reconstruction.
The number of water bodies used in the study is small (33), but this is a function of the rarity of
lakes on continental SE Australia. There are additional natural lakes to sample but they are
exclusively in areas and elevations that we have already sampled. When we established this study
we attempted to focus on natural water bodies. It is clear that in order to extend the training set,
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more reservoirs will need to be included. Even allowing for this, there is a gap between summer
temperatures of c. 13.3–14.8 °C for which there are no ideal candidate lakes or reservoirs. There
are some reservoirs at these temperatures (e.g. Lake Jindabyne) but they are exceptionally large
and deep lakes, and require both alternative sampling strategies and some analyses and
consideration before including in the data set. The other alternative is to integrate this model and
data set with the Tasmania model and data set of Rees et al. (2008). We have deliberately
replicated some of the sites (e.g. Eagle Tarn) from Rees et al. (2008) so that the models can be
compared and harmonised in due course and this is an obvious next step for this research.
Figure 4.7 Performance of the three component PLS model where (a) shows the predicted versus
observed mean February temperature and (b) displays residuals of the predicted versus observed
mean February temperature. Note that the model has a potential to over predict temperatures from
some very shallow high altitude lakes by up to ~6 °C. These lakes have increased mean water
temperature in summer and chironomid assemblages may resemble lower elevation sites.
4.6.4 Value of the transfer function
This transfer function has relatively large errors for detecting change during periods of relative
climate stability (i.e. in the Holocene). However, on longer time scales, the expected change from
Last Glacial Maximum (LGM) to the Holocene in southeastern Australia is between 8 and 10 °C
(Galloway, 1965; Miller et al., 1997). The precision of the current transfer function is ample to
constrain climate change of this magnitude. Transfer functions with relatively large errors may
still be able to provide a reliable indication of variation in temperature through time and this
should be tested using the method developed by Telford and Birks (2011).
a. b.
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Table 4.7 Performance of partial least squares (PLS) model for reconstructing mean February
temperature of southeast Australia using 33 lakes and 38 non-rare chironomid species. The bold
indicates the model of choice.
Method r2
(apparent)
r2
Jack RMSEP
(Jack)
MaxBias
(Jack)
Av
Bias
(Jack)
%
reduced
PLS Component 1 0.67 0.45 3.15 4.39 0.13 -
Component 2 0.87 0.57 2.80 3.28 0.18 10.85
Component 3 0.93 0.69 2.33 2.15 0.07 16.35
Component 4 0.94 0.67 2.45 2.07 -0.01 -4.30
Component 5 0.95 0.66 2.56 1.83 -0.05 -4.56
4.7 Conclusions
We constructed a February mean temperature transfer function based on the modern distribution
of chironomid (Diptera: Chironomidae) species in southeast Australia. The training set comprises
33 natural and artificial lakes in locations that span the subtropics to the alpine zone. The February
mean temperature model is statistically robust with an r2
jackknifed of 0.69, a RMSEP of 2.33 °C and
a maximum bias of 2.15 °C. The transfer function is suitable for the reconstruction of summer
temperatures during the last glacial maximum (LGM) and the late glacial to Holocene transition in
southeastern Australia. In a context where there are few reliable and no continuous estimates of
palaeo-temperature available from mainland Australia, the transfer function represents a
significant advance for palaeoclimatological studies.
We also conclude that chironomid assemblages in actively managed reservoirs (non-impacted)
show no significant difference to natural lakes in the same climate and vegetation zones, and
therefore can be included for transfer function development. Although we cannot completely rule
out some degree of endemism in the Tasmanian chironomid fauna, our analyses show that the
degree of endemism indicated by earlier studies is greatly reduced. This raises the real possibility
of integrating the existing chironomid transfer function for Tasmania with this new model for the
southeast Australian mainland.
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5 A CHIRONOMID-INFERRED SUMMER TEMPERATURE
RECONSTRUCTION FROM SUBTROPICAL AUSTRALIA DURING
THE LAST GLACIAL MAXIMUM (LGM) AND THE LAST
DEGLACIATION
Chapter 5 is published in Quaternary Science Reviews (Chang et al., 2015b)
5.1 Summary
This chapter reports the first application of the newly developed chironomid based transfer function
in Chapter 4, for temperature reconstructions from Welsby Lagoon, North Stradbroke Island,
Australia, detailing the period of the last glacial maximum and the last deglaciation (c. ~ 23.2 and
15.5 ka cal yr BP).
Highlights:
A chironomid based summer temperature reconstruction from Australia.
Reconstruction covers the LGM and part of deglaciation.
Cooling is similar to that derived from nearby marine records.
Deglaciation is synchronous with Antarctica.
Suggests climate link from Australian subtropics to high latitudes.
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5.2 Abstract
A chironomid-based mean February temperature reconstruction from Welsby Lagoon, North
Stradbroke Island, Australia covering the last glacial maximum (LGM) and deglaciation (between c.
~23.2 and 15.5 cal ka BP) is presented. Mean February temperature reconstructions show a
maximum inferred cooling of c. ~6.5 °C at c. ~18.5 cal ka BP followed by rapid warming to near
Holocene values immediately after the LGM. The inferred timing, magnitude and trend of
maximum cooling and warming display strong similarities to marine records from areas affected by
the East Australian current (EAC). The warming trend started at c. ~18.1 cal ka BP and is consistent
with the start of deglaciation from Antarctic records. Near Holocene values are maintained through
the deglaciation to 15.5 cal ka BP. These records suggest that changes in the Australian subtropics
are linked to southern high latitudes.
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5.3 Introduction
Australia is the second most arid continent after Antarctica and, as a consequence, high quality
terrestrial palaeoenvironmental records suitable to investigate late Quaternary climate changes are
rare (Reeves et al., 2013; Petherick et al., 2013). A number of paleoecology records come from the
Atherton Tablelands in North Queensland (Kershaw, 1986; Haberle, 2005; Turney et al., 2006), an
area that is subject to tropical climate systems and is unlikely to provide good palaeoclimate
information for non-tropical southeastern Australia. Similarly, significant research has occurred in
western Victoria (reviewed in Gouramanis et al., 2013) but again this is in a climatically different
environment to subtropical eastern Australia. One of the key time intervals for climate
reconstruction is the last glaciation maximum (LGM; 21–17 ka yr, Barrows et al., 2002; Williams et
al., 2009), as this interval represents the most significantly different global climate in the recent
geological past and has been a focus for palaeoclimatic modelling (e.g. PMIP2: Braconnot et al.,
2007). Australia has traditionally been poorly represented by these models, because there is a dearth
of validation data. This remains true, despite concerted efforts through the Australasian INTIMATE
project (Turney et al., 2007; Reeves et al., 2013) to improve the data sets. Traditional biological
proxies such as pollen are widely applied (PalaeoWorks: Australian National University. Dept. of
Archaeology and Natural History (2000)) but the records, at least away from the Atherton
Tablelands and Victoria (e.g. Caledonia Fen and Tower Hill) (Kershaw et al., 1991, 2007; D'Costa
et al., 1989), are mostly intermittent and have not yielded quantitative palaeoclimatic information
(Kershaw et al., 1994; Kershaw and Bulman, 1996).
Existing quantitative reconstructions of climate at the LGM suggest the climate in southeastern
mainland Australia was significantly cooler, by 8–12 °C (Galloway, 1965 from periglacial
landforms, Miller et al., 1997 from emu shell amino acid racemisation) and with increased aridity
and seasonality (Galloway, 1965) inferred but not quantified. Recently, an LGM mean annual
temperature (MAT) reconstruction from Lake Mackenzie on Fraser Island in Southeast Queensland
(Fig. 5.1; 25°26′51″S, 153°03′12″E, 90 m a.s.l) was developed using branched glycerol dialkyl
glycerol tetraethers (GDGTs), which suggested a cooling of only 4.1 °C at the LGM (c. ∼18.8 ± 0.5
cal ka BP) (Woltering et al., 2014). This lower value is consistent with results from Tasmania (Fig.
5.1) where Fletcher and Thomas (2010) estimated only about 4.2 °C cooling at the LGM from a
pollen transfer function. In summary, there is both a remarkable paucity of LGM
paleothermometers and a clear disagreement between the available reconstructions, which may
reflect either regional variability, or the unreliability of one or more of the techniques, or both.
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Southeast Queensland lies at the northern limit of the Southern Hemisphere winter westerlies (Fig.
5.1) (McGowan et al., 2008). This is the interface between climate driven by the Inter-Tropical
Convergence Zone derived from the Indonesian tropics and the southwest Pacific convergence
zone, and the high latitudes driven by Antarctica and the Southern Ocean. Changes in this region
are particularly valuable as they are likely to give insights into the re-organisation of the major
circulation systems during the LGM.
Subfossil chironomids (Order: Diptera, Family: Chironomidae) have been proved as an applicable
proxy for palaeo-temperature reconstructions since the late 1980s (Hofmann, 1986; Walker, 1987).
Chironomid-based transfer functions have been successfully applied to Southern Hemisphere late
Quaternary palaeoclimatic data (Woodward and Shulmeister, 2007; Rees and Cwynar, 2010;
Massaferro et al., 2014) and have provided quantitative reconstructions. Here, we present the
application of a newly developed subfossil chironomid-based transfer function for the
reconstruction of mean February temperatures (MFTs) (in Chapter 4) to an LGM to Holocene
record from subtropical Australia. We preferred MFT to Mean Summer Temperature because it
provided the best model coefficient (r2) value and conceptually late January and February are the
most stable and warm period in the summer at higher latitudes. The site is Welsby Lagoon in
Southeast Queensland (27°26′12″ S, 153°26′ 56″ E) (Fig. 5.1). Pollen is applied to reconstruct the
moisture balance from the site (Moss et al., 2013), whereas diatoms are absent from the core
sediment of Welsby Lagoon. Therefore, chironomid based reconstructions from the site will provide
valuable complementary information.
5.4 Regional setting
North Stradbroke Island is located on the east coast of Australia at 27.58° S, 153.47° E (Fig. 5.1). It
is 32 km long and has a maximum width of 11 km at its northern end. It is 239 masl (metres above
sea level) at its highest point (Mt Hardgrave). North Stradbroke Island is one of several large sand
islands that form the distal end of the world's largest down drift sand system. This sedimentary
system is fed by rivers on the mid-New South Wales coast, a thousand kilometres to the south, and
the longshore drift transports roughly 500,000 m3 of sand per metre of beach front per year north
(Roy and Thorn, 1981). The sand drift system has been operative for at least the last c. 750,000 ka
yr (Tejan-Kella et al., 1990) and is likely much older. The island is composed of numerous sets of
parabolic dunes that can be traced to beach terraces and are inferred to relate to cycles of sea-level
rise (Ward, 2006). There is a large aquifer underlying the island and dune lakes and wetlands relate
to either breaches of the aquifer (window lakes) or the impoundment of dune hollows by organics
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(perched lakes/wetlands) (Laycock, 1975). Because of the antiquity of the dune fields, many of the
lakes are partly infilled by organics and provide outstanding targets for paleoenvironmental and
paleoclimatic work.
On North Stradbroke Island, research has focussed on aeolian records from Native Companion
Lagoon (McGowan et al., 2008; Petherick et al., 2009) and a broader study of paleoecology from
lagoon and swamp systems across the island including Welsby Lagoon, Tortoise Lagoon (Moss et
al., 2013) and Blue Lake (Barr et al., 2013). The Welsby Lagoon record was selected as a target for
this study as it contains thick gyttja covering the duration of the LGM and the last glacial –
interglacial transition (Moss et al., 2013).
5.4.1 Climate
North Stradbroke Island lies in the east-coast subtropical zone under the Kӧppen – Geiger (Cfa:
humid subtropical climate zone) classification but close to the border with the temperate zone to the
south. The island has warm and wet summers with a mean summer temperature of ∼24 °C and mild
winters with mean temperatures of ∼14 °C (BOM, 2015). The island has a mean annual rainfall of
between 1500 (west coast) and 1700 mm (east coast) and precipitation is summer dominated (BOM,
2015). A number of modern climatic drivers influence precipitation in the region including the El
Niño/Southern Oscillation (ENSO) and the Pacific Decadal Oscillation (PDO). In general, these
oscillatory systems affect the climate by modifying sea surface temperatures (SST). During positive
(negative) phases of ENSO and negative (positive) phases of the PDO, SSTs of the west Pacific are
higher (lower) and there is a positive (negative) rainfall anomaly (BOM, n.d.). The coast is also
more susceptible to cyclones and ex-tropical cyclones during the positive phases of ENSO and
negative phases of the PDO. ENSO and PDO interact and ENSO either amplifies or damps the
background PDO signal (BOM, n.d.).
5.4.2 Modern circulation of the Eastern Australian Current (EAC)
North Stradbroke Island is situated adjacent to the mid-section of the East Australian Current (EAC)
which is formed between 10° and 15°S, and in between the two anti-cyclonic recirculation cells
(Fig. 5.1). The EAC is known to be a highly variable and energetic system with large mesoscale
eddies dominating the flow (Bowen et al., 2005; Mata et al., 2006). The EAC is both the western
boundary of the South Pacific Gyre and the linking element between the Pacific and Indian Ocean
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gyres (Speich et al., 2002). The current is accelerated southward along the coastal boundary, and
then separates into north-eastward (Subtropical Counter Current), eastward (Tasman Front) and a
residual southward (EAC Extension) component at around 31°S (Fig. 5.1, Ridgway and Dunn,
2003). The EAC has an important role in removing heat from the tropics and releasing it to the mid-
latitude atmosphere (Roemmich et al., 2007). It has the strongest southward flow during the austral
summer and the location of separation also migrates along the coast seasonally (Ridgway and
Godfrey, 1997). Observations for the past 60 years show that the EAC has strengthened and
extended further southward (Ridgway and Hill, 2009) in response to the modern warming of
climate.
5.4.3 Physical environment of Welsby Lagoon
Welsby Lagoon is one of several perched lagoons on North Stradbroke Island and is situated near
the northwest coast at 27°26′12″ S, 153°26′ 56″ E and 29 masl (Fig. 5.1). In 2012, it had a
maximum water depth of c. ∼1.3 m and was dominated by sedges and reeds, with large areas of
Melaleuca woodland fringing the lagoon. The surrounding vegetation consists of Eucalyptus forest
and woodlands and Casuarinaceae woodlands, with a predominantly heath understorey (Moss et al.,
2013).
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Figure 5.1 Location of the study site and other sites discussed in this study. (a) A summary of the
surface currents within the Tasman Sea. The denoted by A, B, C, and D shows the direction of flow
of the relevant part of the EAC along with where the SST records are mentioned. Note that B is
associated with the Tasman Front (Ridgway and Dunn, 2003). The locations of marine cores PC-27
(Dunbar and Dickens, 2003), GC-12 (Bostock et al., 2006), GC-25 (Troadson and Davies, 2001),
PC-9 (Troadson and Davies, 2001) along the EAC, terrestrial temperature estimates from Lake
Victoria (Miller et al., 1997), the Snowy Mountains (Galloway, 1965), and Lake Selina (Fletcher
and Thomas, 2010) are displayed (b) Location of North Stradbroke Island (NSI) and Lake
Mackenzie, Fraser Island (c) Location on North Stradbroke Island of Welsby Lagoon and other sites
mentioned in the text, including Tortoise Lagoon (Petherick et al., 2008; Moss et al., 2013), Blue
Lake (Barr et al., 2013), Eighteen Mile Swamp (sampling location) and Native Companion Lagoon
(McGowan et al., 2008; Moss et al., 2013).
5.5 Methodology
5.5.1 Chironomids sampling and analysis
One 450 cm core from Welsby Lagoon was collected by Moss et al. (2013) in October, 2009 using
a Russian corer (Jowsey, 1966). Half of the core was used for pollen, micro-charcoal and
radiocarbon dating analyses (Moss et al., 2013), while the remaining half core was sub-sampled for
this study. Two-centimetre sections of sediment were collected for chironomid analyses with a
sampling resolution of one sample every 30 cm from the top to 340 cm, and every 10–20 cm from
340 cm to the base of the core.
Chironomid analysis followed the standard procedure outlined by Walker and Paterson (1985).
Samples were deflocculated in warm 10% KOH for 20 min and washed on a 90 µm mesh with
distilled water. Samples were transferred to a Bogorov counting tray and examined under a
dissecting microscope at 50× magnification. Chironomid head capsules were hand-picked using
fine forceps and placed ventral-side up on a glass slide, then mounted in a drop of Euparal® with a
cover slip. We classify this as a minimum of 50 head capsules that are fully identified (Heiri and
Lotter, 2001; Quinlan and Smol, 2001) (Fig. 5.3b). Chironomid taxa were identified using a
compound microscope at 400 × magnification following Cranston (2000, 2010) and Brooks et al.
(2007).
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Zonation of the chironomid record was achieved using the constrained incremental sums of squares
(CONISS) function in Psimpoll (Bennett, 2002). This software also allows the determination of the
optimal number of zones using the broken stick model. All chironomid taxa were included in the
analysis and species data was square-root transformed.
5.5.2 Chronology
Chironomid samples for this study were taken from the same core as the pollen record described in
Moss et al. (2013). We therefore utilise the chronology developed by Moss et al. (2013), which is
based on 11 radiocarbon dates on a mix of pollen, charcoal and peat samples. However, we have
updated the model by calibrating the radiocarbon ages against the more recent SHCal13 calibration
curve (Hogg et al., 2013) (Table 5. 1). The resulting age-depth model is presented in Fig. 5.2.
Figure 5.2 Radiocarbon and calibrated ages for Welsby Lagoon with respect to depth. Open shapes
represent calibrated ages calibrated using SHCal13 (Hogg et al., 2013).
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Table 5.1 Radiocarbon Dates for Welsby Lagoon. Data shown includes sample depth, laboratory,
materials used for dating, 14
C age and calibrated age (SHCal13; Hogg et al., 2013) for each of the
samples. (*) indicates samples excluded in the final age depth model.
Depth
(cm)
Lab Code Sample Code Material
Dated
AGE (14
C yr) Calibrated
Age (Cal BP)
80 Wk-29036 Welsby 80cm charcoal 1803 +/- 30 BP 1710
125 Wk-29037 Welsby 125cm charcoal 4070 +/- 30 BP 4494
165 Wk-30668 Welsby 165-166cm charcoal 5556 +/- 26 BP 6311
251 Wk-29038 Welsby 251cm charcoal 8043 +/- 30 BP 8852
287 Wk-30669 Welsby 287cm charcoal 9903 +/- 38 BP 11295
304 Beta - 306711 WL p304 pollen 14250 +/- 50 BP 16970*
342 Wk-29039 Welsby 342cm Peat 18696 +/- 88 22245*
398 Beta - 306712 WL p398 pollen 16350 +/- 70 BP 19446
414 Wk-30670 Welsby 414cm charcoal 29096 +/- 250 33787*
435 Beta - 270553 Welsby-435cm peat 18320 +/- 90 21868
438 Beta - 306713 WL p438 pollen 18980 +/- 80 BP 22644
5.5.3 Statistical methods and reconstruction examination
The southeastern Australian chironomid based transfer function developed from a 33-lake training
set for temperature (mean February temperature, MFT) (Chapter 4, Table 4.1) was applied to the
chironomid data. The transfer function training set did not include any lakes from North Stradbroke
Island so one adjacent site, Swallow Lagoon (c. ∼7 km south of Welsby Lagoon) was added to the
original training set to provide a local analogue site. MFT of each site in the modern calibration data
set was obtained using the combination of the WorldClim program (available from
http://www.worldclim.org/bioclim, accessed 20 August 2013) and ArcGIS 10.1. WorldClim data
for Australia is based on climate surfaces derived from around 600 nation-wide weather stations
that have climate records spanning the years 1950–2000
(http://www.bom.gov.au/climate/data/stations/, accessed 20 January, 2014) (see Chapter 4). The
model was created using partial least squares (ter Braak and Juggins, 1993) in C2, v1.5 (Juggins,
2005) where percent values of fossil taxa were square-root transformed. The addition of Swallow
Lagoon did not alter the parameters of the previously established transfer function. The model has a
jack-knifed coefficient of determination (r2
jack ) of 0.69 and a root mean square error of prediction
(RMSEP) of 2.2 °C.
To evaluate the reliability of the chironomid-inferred MFT reconstruction, we used the
reconstruction diagnostics outlined in Birks (1995). Goodness-of-fit (Birks et al., 1990) was
assessed by passively fitting fossil samples to the canonical correlation analysis (CCA) ordination
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axis derived from the modern training set constrained to MFT in CANOCO version 4.5 (ter Braak
and Šmilauer, 2002). The squared residual length (SqRL) was used to assess the quality of fit by
comparing the distance between modern samples and the constrained axis to the distance between
fossil samples and that same axis. Any fossil sample whose residual distance was equal to or larger
than the residual distance of the extreme 5% of the training set is considered to have a ‘very poor’
fit to temperature, and those with values equal to or larger than the extreme 10% were deemed to
have a ‘poor’ fit (Birks et al., 1990). The chi-square distance (dissimilarity coefficient) to the closest
modern analogue for each fossil sample was calculated in C2 (Juggins, 2005) using the modern
analogue technique (MAT) (Birks et al., 1990). Fossil samples with a chi-square distance to the
closest sample in the modern calibration data set larger than the 5th
percentile of all chi-square
distances in the modern assemblage data were identified as samples with ‘no good analogues’
(Anderson et al., 1989; Larocque-Tobler et al., 2010). The percentage of rare taxa in the fossil
samples was calculated, where a rare taxon has a Hill's N2 ≤ 5 in the modern data set (Hill, 1973) in
C2 (Juggins, 2005). Fossil samples that contain high percentage (>10%) of these rare taxa were
likely to be poorly estimated (Brooks and Birks, 2001). Finally, the fossil assemblages were plotted
passively to a CCA of the training set with all significant variables (p < 0.05) in CANOCO version
4.5 (ter Braak and Šmilauer, 2002) to determine the direction of change through time and the
influence of the environmental variables on the fossil chironomid assemblages.
5.6 Results
5.6.1 Chironomid analysis
A total of 42 sub-samples were taken throughout the 450 cm core where the first sample was
undertaken at the depth of 55 cm. Chironomids were absent from the depth between 277 cm and
340 cm of the core which corresponds to the period between c. ∼10.5 cal ka BP and c. ∼15 cal ka
BP. Discontinuous chironomid counts were obtained between 55 cm–277 cm, which covers roughly
the last ∼10.5 ka calendar years. Altogether, 17 samples yielded enough chironomids for reliable
quantitative reconstructions. These include four from the top 277 cm (0 – c. ∼10.5 cal ka BP) (Fig.
5.3), eight from the depths between 340 cm and 377 cm (c. ∼15.5–17.8 cal ka BP) and five from the
depths between 385 cm and 442 cm (c. ∼18.5–23.2 cal ka BP) (Fig. 5.3).
Goodness-of-fit analyses of the 17 samples from depths of 55 cm to the bottom of the core (442 cm)
show that only one sample (at the depth of 275–277 cm) does not have a good fit to temperature
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(Fig. 5.4a and b). Rare taxon analysis of these samples indicate 4 of the samples between 375 and
442 cm (c. ∼18.1–23.2 cal ka BP) have abundances of taxa that are considered rare (Hill's N2 ≤ 5)
totalling to greater than 10% (Fig. 5.4c). Rare taxa Tanytarsus chinyensis type, Parakiefferella
morphotype 3 and Stempellina (with Hill's N2 = 4.62, 3.74 and 1.95 respectively) which occurred in
these samples are currently restricted to a small number of lakes from Tasmania and one lake in
western Victoria in the modern training set. This is nearly 2000 km south of North Stradbroke
Island. The analysis of dissimilarity between fossil and modern assemblages using the modern
analogue technique (MAT) shows that ten out of the 17 samples have ‘no good’ analogue in the
modern calibration set (Fig. 5.4d). These include three samples from the top 277 cm and seven of
the depths between 340 cm and 442 cm.
A total of 21 taxa were identified from the 17 samples (Fig. 5.3). Each of the chironomid taxa that
have marked temperature preferences in south eastern Australia were highlighted and discussed in
Chapter 4. The Holocene component of the record is intermittent and more samples in the c. ∼15.5–
23.2 cal ka BP range contained sufficient head capsules (>50) for quantitative reconstructions. The
record is split into three zones based on the constrained incremental sums of squares (CONISS)
analysis of the fossil chironomid assemblages (Fig. 5.5) and these zones correspond to the LGM
(from the beginning of this record c. ∼23.2 to ∼18.1 cal ka BP) (Barrows et al., 2002; Clark et al.,
2009), deglaciation (c. ∼18.1 to ∼13 cal ka BP) and the Holocene (c. ∼13 cal ka BP to near
present). The break for the Holocene based on CONISS analysis is ∼13 cal ka BP for this record
which is different from the well recognized 11.5 cal ka BP is likely due to the low sample resolution
in this part of the record. These zones (Fig. 5.5) are consistent with the observed changes in the
chironomid stratigraphy (Fig. 5.3).
5.6.2 The LGM (~23.2 cal ka BP – 18.1 cal ka BP)
The LGM (as defined in Barrows et al., 2002 and Clark et al., 2009) from Welsby Lagoon is
characterised by the presence and increased abundance of cold indicators such as Parakiefferiella
morphotype 3, Paralimnophyes morphotype 1, Tanytarsus glabrescens type and Paralimnophyes
morphotype 3. The cooling also corresponds to an apparent decrease in the abundance of warm
adapted taxa (Chapter 4, Fig. 4.5, Table 4.6) including Cladopelma, Parachironomus,
Paratanytarsus, Tanytarsus lactescens type and Procladius (Fig. 5.3).
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The reconstructed MFT at the start of the record (at c. ∼23.2–21.7 cal ka BP) is between 19.9 and
21.7 °C. This represents a cooling between 2.3 and 4.1 °C relative to the present (Fig. 5.4a).
Decreasing inferred temperature is observed immediately after c. ∼21 cal ka BP. The mean
reconstructed MFT at 20.6 cal ka BP is 18.4 °C which is 5.6 °C cooler than the present (Fig. 5.4a).
Temperature continued to fall and by c ∼18.5 cal ka BP, an MFT minimum of 17.4 °C was obtained
from this site which is a ∼6.5 °C cooling relative to the present day. A rapid warming trend is then
observed, as MFT started rising from c. ∼18.1 cal ka BP and by 17.3 cal ka BP, it rose 3 °C to 20.4
°C (Fig. 5.4a).
5.6.3 The deglacial period (~18.1 cal ka BP – 13 cal ka BP)
During the deglacial, the reconstructed MFTs are warm and highly stable (Fig. 5.4a). This period is
characterised by the high abundance of warm indicating taxa Cladopelma, Procladius,
Paratanytarsus, Tanytarsus lactescens type, Pentaneurini, Tanytarsus lugens type while cold
stenotherms, such as Paralimnophyes morphotype 1 and 3, Parakiefferiella morphotype 3 are
absent during this time period (Fig. 5.3). The average temperature is between 20. 4 and 21.4 °C
from ∼17.3 to ∼15.5 cal ka BP, where fluctuations between samples do not exceed ∼1 °C. This
indicates that local MFT at this time is ∼2.6–3.6 °C cooler than the present.
One sample from the depth of 350–352 cm (c. ∼15.9 cal ka BP) shows a reconstructed MFT as low
as 17 °C at the site (Fig. 5.4a). This sample is characterized by exceedingly high abundances of
Dicrotendipes (31%) and Procladius (25%) (Fig. 5.3). No other warm or cold indicators (apart from
warm stenotherms Pentaneurini and Tanytarsus lugens) are present in this sample (Fig. 5.3). The
composition of this sample is very similar to the Holocene samples that are discussed below.
5.6.4 The Holocene (~13 cal ka BP – present)
The four Holocene samples at the depths of 55–57 cm, 85–87 cm, 257–259 cm and 275–577 cm,
which represent c. ∼0.2, 2, 9.3 and 10.5 cal ka BP respectively, have assemblages dominated by
one taxon – Dicrotendipes (at abundances of 65%, 37%, 62% and 66% in each of the four samples
respectively). Dicrotendipes was widespread in mainland Australian training set and occurred in
lakes at MFTs ranging from 12 °C to 24 °C (Chapter 4, Fig. 4.6). Chironomus sp. is the other
dominant taxon in the Holocene samples and is also present in lakes spanning a long temperature
gradient. Only a small percentage (<5%) of warm stenotherms composed the assemblages of these
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samples. These include Cladopelma, Polypedilum nubifer, Paratanytarsus, Tanytarsus lactescens
type, Pentaneurini, Tanytarsus lugens type (Fig. 5.3).
Quantitative MFT reconstructions of these samples provide generally lower values than expected
(e.g Dimitriadis and Cranston, 2001) from this site for the Holocene (19.2 °C, 18.5 °C, 15.8 °C and
16.3 °C at the age of c. ∼0.2, 2, 9.3 and 10.5 cal ka BP respectively) (Figs. 5.3 and 5.4a). These
show a cooling between 4.8 and 8.2 °C compared to the present in the Holocene at the site.
However, previous studies show that Dicrotendipes at least in the Holarctic occurs in the littoral of
lentic environments (Pinder and Reiss, 1983) where it is often associated with macrophytes
(Brodersen et al., 2001) and mesotrophic to eutrophic waters (Brodin, 1986). The temperature
reconstructions of these samples are highly dependent on the abundance of one taxon,
Dicrotendipes, which is discussed below and this factor may cause inaccurate MFT reconstructions
in these four samples.
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Figure 5.3 Chironomid stratigraphy from Welsby Lagoon plotted with respect to the radiocarbon chronology. Chironomid inferred mean February
temperatures (MFTs) are shown in (a) and the number of head capsules counted for each sample are shown in (b). The dashed line in (a) indicates the
current mean February temperature at Welsby Lagoon and the dashed line in (b) indicates the minimum number of head capsules for a quantitative
reconstruction from a sample. All fossil taxa present in the Welsby Lagoon record are included in the diagram. Age in calibrated years is on the y-axis
and taxa were grouped according to their optima in the modern training sets based on Chapter 4. Grey areas represent sediment layers where
chironomids were absent. Note that Dicrotendipes, a benthic taxon that does not have a significant correlation (p > 0.05) to MFT in the modern
calibration data set, is dominant in all four Holocene samples. Each of the taxon was photographed and included in Appendix 3.
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5.6.5 Relationships between changes in chironomid assemblages and temperature and
other variables
Passive plots of the fossil samples (Fig. 5.6) in a canonical correspondence analysis (CCA) of the
modern chironomid assemblages against the significant (p < 0.05) environmental variables (MFT,
depth, Total Phosphorus (TP), Chlorophyll a (Chl a), Conductivity, pH) indicate the directional
change of trend of fossil assemblages through time. In Fig. 5.6, sample numbers 1–33 represent
modern sites from Chapter 4, Table 4.1 and sample 34 is Swallow Lagoon. Fossil samples are
labelled by their respective ages in Fig. 5.6a and b respectively. Changes in trend of fossil
assemblages through most of the deglacial and LGM samples (Fig. 5.6a and c. ∼23.2–16.6 cal ka
BP) follow and are parallel to the mean February temperature (MFT) gradient, except for two
samples (Fig. 5.6a, at c. ∼17.8 and c. ∼23.2 cal ka BP) that are possibly influenced by the
secondary gradient. The trend of changes of the fossil assemblages in the deglacial through to the
Holocene samples (Fig. 5.6b and c. ∼16.3–0.2 cal ka BP) are parallel to the secondary gradient that
is driven by changes of lake depth, nutrient variables (TP, Chl a), conductivity and pH. It was
impossible to distinguish among these four driving variables, since these parameters are highly
correlated in the south eastern Australian modern lake training set (see Chapter 3 and Chapter 4).
This implies that the MFT reconstructions from the LGM and the deglacial samples are reliable as
the trend of changes in the fossil assemblages is less influenced by a secondary gradient. While only
one sample from c. ∼16.3–0.2 cal ka BP does not have a good fit to temperature, MFT
reconstructions may still be inaccurate through this period. This is due to the fact that the trend of
changes in fossil sample assemblages is overwhelmed by the changes in variables that are driven by
the secondary gradient.
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Figure 5.4 Reconstruction diagnostics for Welsby Lagoon samples. (a) Reconstructed mean
February temperatures (MFTs) with the RMSEP (2.2 °C) from the modern training set (Chapter 4)
plotted in dashed horizontal lines for each sample. (b) Goodness-of-fit statistics where the fossil
samples are passively fitted to the CCA ordination axis derived from the modern training set
constrained to MFT. The vertical dashed line represents the 95th
percentiles of modern squared
residual lengths beyond which fossil samples are considered to have poor fits to MFT. (c) Rare taxa
plot where closed circles represent fossil samples with well-represented taxa in the modern training
set and open circles represent fossil samples with rare taxa (Hill's N2 ≤ 5) summing to greater than
10% abundance. (d) Non-analogue plot where closed circles indicate fossil samples with either
close or good modern analogues, whereas open circles indicate fossil samples with no good modern
analogues (at the 5th
percentile cut-off level). The vertical lines represent the 2nd
, 5th
and 10th
percentiles respectively.
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Figure 5.5 The Welsby Lagoon chironomid record was split into three zones using the constrained incremental sums of squares (CONISS) function in
Psimpoll (Bennett, 2002). W-1 represents the last glacial maximum (LGM), W-2 represents the deglacial and W-3 represents the Holocene. This
zonation is consistent with the observed changes in chironomid assemblages shown in Fig. 5.3.
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Figure 5.6 Trajectory changes of trend through time of fossil samples (black circles) in the Welsby Lagoon record, passively plotted in a CCA of the
training set lakes (open circles, where site 1–33 are waterbodies from (Chapter 4, Table 4.1) and site 34 is Swallow Lagoon, a waterbody near Welsby
Lagoon with similar physical characteristics) against significant variables (p < 0.05). (a) Samples covering the last glacial maximum (LGM) and
deglacial period, where changes in trend are parallel to the mean February temperature (MFT) gradient, indicating that MFT is a primary controlling
variable and drives the changes of chironomid assemblages in the samples from c. ∼23.2 to 16.6 cal ka BP. (b) samples covering the period of
deglacial and Holocene from c. ∼16.3 to 0.2 cal ka BP. The trend of changes of chironomid assemblages in these fossil samples are parallel to the
secondary gradient that is represented by nutrients, conductivity, pH and lake depth, indicating that non-climatic factors are likely dominant in
Holocene changes.
a. b.
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5.7 Discussion
5.7.1 Precision and reliability of the reconstruction
There are three factors that may complicate the temperature reconstruction and may have influenced
the reliability of the record. First, in the Holocene samples and one sample at 15.9 cal ka BP, the
dominance of one taxon, Dicrotendipes, which has a lentic habit and is highly correlated to
macrophyte abundance in the lake (Cranston and Dimitriadis, 2004), led to a lower than expected
reconstruction of temperature for the Holocene (4.8 to −8.2 °C compared to present). Dicrotendipes
is neither a warm nor a cold stenotherm and it does not show a significant correlation (p > 0.05) to
temperature in the modern training set of Australian lakes based on the Generalized Linear Model
(GLM) results (Table 4.6 in Chapter 4). Instead it has a significant correlation to nutrient variables,
conductivity and lake depth. In a study of 21 Danish lakes Brodersen et al. (2001) found that
Dicrotendipes has a high co-occurrence with macrophytes and is found predominantly in shallow
and eutrophic lakes. Luoto (2009) also points out that Dicrotendipes is not distributed along a
temperature gradient in the 77 lakes of a modern training set from Finland. Consequently, the
temperature inferences based on Dicrotendipes as a warm stenotherm in some cases (e.g. Chase et
al., 2008 in cold temperate Canadian lakes) is probably an artefact, although, some studies (e.g.
Dimitriadis and Cranston, 2001; using mutual-climate-ranges technique from eastern Australian
lakes) recognized that Dicrotendipes can be a warm temperature indicator. In this data set, the
presence of Dicrotendipes in eutrophic lakes in southern areas eliminates any association with
warmth.
Secondly, changes in nutrients, conductivity, pH and lake depth may all have had an impact on
chironomid assemblages through time. However, most of the deglacial and LGM samples followed
the MFT gradient closely (Fig. 5.6a), suggesting these samples are not strongly influenced by
variables other than MFT changes. Fig. 5.6b shows the lagoon had experienced large fluctuations in
lake water chemistry from the deglacial throughout the Holocene (c. ∼16.3–0.2 cal ka BP) and
these changes are characterised by significant shifts in pH and the concentration of nutrients, salts
and fluctuations of lake level. Most of these samples are also coincident with the high occurrence of
Dicrotendipes (Fig. 5.3). In contrast, more stable lake conditions were observed during the early
deglaciation and the LGM (c. ∼23.1–16.6 cal ka BP) (Fig. 5.6a). Welsby Lagoon had a relatively
more consistent water level and was not significantly influenced by the secondary gradient during
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this period of time compared to the Holocene. MFT reconstructions from these time intervals are
consequently regarded as reliable.
Finally, the modern calibration training set contains a relatively small number of lakes (33) and this
may be the cause of the ‘no good’ modern analogues situation for a number of samples in the
deglacial and LGM as larger training sets generally allow better estimates of species' optima and
tolerances in relation to important environmental variables (Lotter et al., 1999). A similar situation
was observed in the temperature reconstructions of Eagle Tarn and Platypus Tarn from Tasmania
(Rees and Cwynar, 2010). However, this is less problematic than in some reconstructions (Heiri and
Millet, 2005; Larocque-Tobler et al., 2010), where an abundant taxon (Corynocera ambigua) was
present in a large number of fossil samples and was fully absent from the modern training set. In
many cases, samples with ‘no good’ modern analogues still provide reliable and robust trends in
temperature reconstructions (e.g. Lotter et al., 1999; Rees and Cwynar, 2010) because the (WA)
PLS model performs relatively well in no analogue situations (Birks, 1998; Lotter et al., 1999) and,
consequently, the temperature reconstructions are considered reliable from the deglacial and LGM
samples. Merging a regional training set with Rees et al. (2008) may increase the likelihood of
finding good modern analogues for fossil assemblages from Welsby Lagoon.
Overall, the transfer function model has the tendency to underestimate temperatures at warm sites
and overestimate from colder sites (see Chapter 4), which is not unusual for a model of this type
(Barley et al., 2006). Welsby Lagoon (MFT = ∼24 °C at the present) is located among the warmest
sites of the modern calibration set (Fig. 4.7 in Chapter 4), and consequently the reconstructed mean
February temperatures are likely to be under-predicted.
Finally, one of the more serious limitations of the transfer function is the relatively large RMSEP of
2.2 °C. Nevertheless, the trends between the samples are likely to be very robust, possibly due to
the consistency of local setting (Lotter et al., 1999). In addition, we note that more than two thirds
of the modern calibration sites fall with MFTs between 18 °C and 24 °C and since most
reconstructed temperature estimates fall within this envelope (Chapter 4, Table 4.1), this reinforces
the suitability of the model for application to Welsby Lagoon data.
In summary, the maximum cooling observed in our data set occurred at c. ∼18.5 cal ka BP with a
reconstructed MFT of between 15.2 and 19.6 °C. This gives a range of cooling between 4.4 and 8.8
°C but with cooling considering the errors likely to be close to the midpoint estimate at about 6.5
°C. This cooling times well with Antarctica (Stenni et al., 2011) and is also compatible with the late
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termination of glaciation in Australia (e.g. Barrows et al., 2002; Hughes et al., 2013). It is clear that
most of the LGM and all of the deglaciation period were much warmer than this maximal cooling.
Pollen work from Welsby Lagoon by Moss et al. (2013) shows rainforest taxa (mainly Araucaria
and palms) present through the LGM. A cooling of c. ∼6.5 °C would change the composition of the
local rainforest but would not of itself eliminate rainforest from the site. Consequently, these
findings are compatible with the persistence of rainforest on North Stradbroke noted in Moss et al.
(2013).
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Figure 5.7 Chironomid mean February temperature (MFT) transfer function based reconstruction
from Welsby Lagoon (WEL) (in plot d) compared to (a) the δ18
O and CO2 records from Antarctica
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(Stenni et al., 2011; Pedro et al., 2012); (b) insolation curve for seasonality (Berger and Loutre,
1991); (c) δ18
O measured from G. ruber from marine cores along the East Australian Current
(EAC) and (d) temperature estimated based on branched glycerol dialkyl glycerol tetraether
(GDGT) from Fraser Island (Woltering et al., 2014). The chironomid results from WEL (d) indicate
a maximum cooling relative to the modern MFT along the Southeast Queensland coast of c. ∼6.5
°C with much lower cooling at other times during the late LGM and a rapid recovery at about 18.1
cal ka BP. The cooling is significantly less than the 9–12 °C for the adjacent uplands from
periglacial landforms from Galloway (1965), but is in line with estimates from nearby marine cores
shown in (c) and lacustrine record from Lake Mackenzie (LM), Fraser Island shown in (d) (note the
reconstructed temperature from Fraser Island is lower than Welsby Lagoon because the GDGT is
based on mean annual temperatures calibration whereas chironomid transfer is based on MFT). It is
also noteworthy that the chironomids provide an estimate of summer temperatures whereas the
periglacial landforms are indicators of winter temperature. This suggests a larger seasonal
temperature contrast at the LGM which is consistent with the insolation curves (b). Finally the
observed timing and pattern is very similar to both the δ18
O curve and CO2 record from Antarctica
(a) (Stenni et al., 2011; Pedro et al., 2012).
5.7.2 The deglacial and LGM temperatures of North Stradbroke Island and regional
climate
To further examine this new seasonal temperature reconstruction at the LGM, we compared our
chironomid-based mean February temperature record to Antarctica ice cores (TALDICE: Stenni et
al., 2011 for δ18
O isotope and Byrd: Pedro et al., 2012 for CO2 concentrations) (Fig. 5.7a),
summer/winter insolation curves (Fig. 5.7b , Berger and Loutre, 1991), marine reconstructions from
near the EAC (Troedson and Davies, 2001; Dunbar and Dickens, 2003; Page et al., 2003; Bostock
et al., 2006) (Fig. 5.7c) and a lacustrine temperature record from Lake Mackenzie, Fraser Island
(Woltering et al., 2014) (Fig. 5.7d) respectively.
Previous terrestrial proxies (block stream, AAR and most recently branched glycerol dialkyl
glycerol tetraether (GDGT)) used in mainland Australia to estimate temperatures have not been
widely applied. Pollen based transfer functions for temperature reconstructions in mainland
Australia were tested (Cook and Van der Kaars, 2006) but have not been successfully applied to
reconstruct LGM temperatures, in contrast to the Tasmanian pollen transfer function (Fletcher and
Thomas, 2010). For our study, the recent GDGT record from Lake Mackenzie (LM) (25°26′51″S,
153°03′12″E, 90 m asl), Fraser Island (Woltering et al., 2014) provides the best comparison
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although the record has a hiatus between 18.3 and 14 cal ka BP, which coincides with the time of
best detail in our study. At Lake Mackenzie, the lowest mean annual temperature occurred at 18.8 ±
0.5 cal ka BP, with MAT estimated to be ∼4.1 °C lower than modern day temperature (Fig. 5.7d).
The timing of this maximum cooling is in good agreement with our Welsby Lagoon record (Fig.
5.7d), and is also consistent with the offshore SST estimates based on G. ruber δ18
O values from the
north (GC-12, 23°34.37′S, 153°46.94′E; Bostock et al., 2004, 2006), and south (GC-25. 26°35′S,
153°51′E ; Troedson, 1997; Troedson and Davies, 2001) of Fraser Island, and further south (GC-9,
33°57′S; 151°55′E; Troedson, 1997; Troedson and Davies, 2001) along the East Australian Current
(EAC) (Fig. 5.7c). Our record shows good agreement with the long term temperature trends with
marine records GC-25 and GC-9 (Fig. 5.7c) and is particular similar to the reconstructed SST from
GC-25 (at 26°35′S) (Bostock et al., 2006) (Figs. 5.1 and 5.7c), which suggested a ∼6 °C of mean
annual SST cooling. All these records show a maximum cooling at c. ∼18.5 cal ka BP, followed by
rapid warming to (near) Holocene values by c. ∼17.3 cal ka BP (Fig. 5.7c and d). Lastly, the timing
of this cooling from all these records show a coincidence with the AIM2 event, which marked the
start of deglaciation at 18.2 ± 0.7 cal ka BP (Stenni et al., 2011) represented in the δ18
O TALDICE
ice core record (72°49′S, 159°11′E) (Fig. 5.7a) and the increase of CO2 concentration estimated
from Byrd ice core record (80°S, 119°49′W), Antarctica. The TALDICE and Byrd records also
indicate a rapid warming trend observed immediately after the maximum cooling, similar from
Welsby lagoon and the southern marine sites (GC-25 and GC-9) near the EAC.
The scale of cooling in the Fraser record (4.1 °C, Woltering et al., 2014) is less than we observe at
Welsby Lagoon (6.5 °C) but the results are within the errors of the respective techniques and it is
possible that GDGT data also reflect summer temperatures, assuming that the microbes producing
branched GDGTs are most active during the period of peak photosynthetic activity in summer
(Pearson et al., 2011). However, though Welsby Lagoon is only 2° of latitude south of Fraser
Island, it is possible that the difference reflects changes in the EAC offshore as the degree of
summer cooling from Welsby Lagoon and both of these are similar to a southern site GC-9 (at
33°57′) (Figs. 5.1 and 5.7c). In contrast, the Fraser island record is closer to the reconstruction from
the northern site GC-12 (23°34′S) (Figs. 5.1 and 5.7c). Bostock et al. (2006) interpreted the
difference of SST amplitude (∼3–4 °C temperature change across ∼3° of latitude) of the LGM
between cores GC-12 (23°34′S) and GC-25 (26°35′S) as reflect a northwards shift in the separation
of the Tasman Front (Fig. 5.1) from the EAC during the LGM, resulting in warm waters from the
EAC not reaching as far south as GC-25 and Welsby Lagoon. More marine and terrestrial proxy
data from the LGM are required from subtropical Australia to allow this hypothesis to be further
tested.
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The reconstructed cooling is substantially less than earlier terrestrial reconstructions from
periglacial landforms (8–12 °C) (Galloway, 1965) and amino acid racemization (AAR) values from
Emu egg shells (Miller et al., 1997). This does not necessarily imply a direct disagreement between
the techniques as the chrionomids and the GDGT data may both reflect temperatures from summer
months, whereas the periglacial landforms are interpreted as winter temperature estimates, and the
AAR reconstructions come from semi-desert environment several thousand kilometres west and
south of this region. These differences may be reflecting enhanced seasonality during the LGM, in
which large temperature differences between summers and winters would be expected (Fig. 5.7b).
In addition, it is unsurprising that coastal records resemble the thermal history of adjacent marine
sites.
5.7.3 Wider paleoclimatic considerations
One of the key observations from the Welsby Lagoon chironomid record and other temperature
records from subtropical Australia (including marine records), is that whatever the scale of cooling,
the timing of the deglaciation is synchronous with changes in the Antarctic (Fig. 5.7). This is
noteworthy on two counts. Firstly, Welsby lagoon sits on the transition between the tropics and the
temperate zone. The record clearly responds directly to changes at high latitudes and implies that
high latitude forcing controls climate change at least as far north as 27°S. This is unexpected,
because this is an austral summer month reconstruction and at the present day, while winter climate
is affected by the mid-latitude westerlies, modern summer climate is dominated by tropical air
masses (Ledru and Stevenson, 2012). The possibility that tropical air masses did not penetrate this
subtropical area in the summer during the glacial period needs to be considered. Secondly, there is a
disconnection between the temperature trend we observe and insolation in the Southern
Hemisphere. The last glacial maximum (LGM) coincides with a summer insolation maximum while
the deglaciation corresponds to a period of reduced summer insolation (Fig. 5.7 b–d). This suggests
that regional insolation trends are overwhelmed by hemispheric changes driven from high latitudes.
Complex feedback mechanisms through either greenhouse gases or the thermohaline circulation are
implied.
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5.8 Conclusion
A summer temperature reconstruction based on the recently developed chironomid transfer function
from southeastern Australia (developed in Chapter 4) is presented from Welsby Lagoon of North
Stradbroke Island. This is the first detailed seasonal reconstruction of LGM and the last deglacial
temperatures from terrestrial Australia. Factors that may affect the reliability of the inferred
temperatures include changes in lake chemistry and lake level, which had an impact on chironomid
assemblages, particularly in the Holocene. The relatively small modern calibration training set also
limits the ability to fully understand the environmental controls on some of the fossil samples.
Despite these caveats, the temperature reconstruction from the LGM to the deglacial period of this
site is sensible and a stable lake condition (in relation to lake level and lake chemistry changes) is
observed. The overall reconstruction displays strong similarities to marine records from areas
affected by the East Australian current (EAC), both in terms of timing and magnitudes of cooling
during and immediately after the LGM.
The results show a maximum mean February temperature cooling of c. ∼6.5 °C at the late LGM at
c. ∼18.5 cal ka BP, followed by rapid warming to (near) Holocene values by 17.3 cal ka BP. This
warming is consistent with the AIM2 event, which marked the start of deglaciation at 18.2 ± 0.7 cal
ka BP (Stenni et al., 2011). It is also compatible with the increase of CO2 concentration estimated
from the Byrd ice core record in West Antarctica (Pedro et al., 2012). It suggests that Antarctic
climate influence reached the Australian subtropics during the LGM.
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6 CAN STABLE OXYGEN AND HYDROGEN ISOTOPES FROM
AUSTRALIAN SUBFOSSIL CHIRONOMID HEAD CAPSULES BE USED
AS PROXIES FOR PAST ENVIRONMENTAL CHANGE?
Chapter 6 is submitted to Limnology and Oceanography
6.1 Summary
This chapter reports the investigation of the potential of using stable oxygen (δ18
O) and hydrogen
(δ2H) isotopes of subfossil chironomid head capsules as independent proxies for past climate and
environmental reconstructions from southeastern Australia.
Highlights:
This is the first of such study in the Australasian region and among one of the very few
similar investigations worldwide.
Previous studies have focussed on the relationship of temperature and δ18
O of chironomid
head capsules. The effects of in-lake variables were overlooked.
This work suggests that nutrients and salinity are important to consider for both δ18
O and
δ2H analyses of chironomid head capsules, at least for the genus of Chrionomus, when
applied as proxies for past changes.
This study concludes and re-affirms that both δ18
O and δ2H are potentially valuable tools for
reconstructing temperature in low nutrient and dilute lakes in humid areas.
This work is complementary to the previous work included in Chapter 3 and 4 but also
completely independent.
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6.2 Abstract
Results of stable isotopes δ18
O and δ2H analysed on head capsules of chironomids (Chironomus
spp.) from sixteen southeastern Australian lakes are presented. Lakes that are located in semi-arid
and sub-humid areas have high evaporation, prolonged water residence time and experience
modified local hydrological conditions including increased salinity and nutrient status and
evaporative enrichment of δ18
O values in lake water. These lakes do not appear suitable to be used
for temperature calibration in paleoclimate reconstructions. High nutrient and salt concentrations
create an ecologically challenging aquatic environment for chironomids, however, haemoglobin
helps regulate the oxygen level in Chironomus spp. and allows this taxon to thrive in these
environments. For these lakes, Chironomus spp. was not in equilibrium with δ18
O lake water due to
strong vital effects and we recommend against its use to infer climatic variables in these settings.
For oligotrophic to mesotrophic lakes, the relationship between δ18
O of Chironomus spp. and
temperature appears robust (r2 = 0.78) and very similar to results from European lakes (e.g.
Verbruggen et al., 2011). Similar results were also obtained for δ2H and temperature, with an r
2 of
0.72. The overall findings of this study are that both δ18
O and δ2H are potentially valuable tools for
reconstructing temperature in cooler, low nutrient and low salinity regions of Australia. Chironomus
spp. is an obvious target for stable isotope work because it is both large and ubiquitous but its
ability to adapt to low oxygen conditions makes it a poor climate indicator in non-humid settings.
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6.3 Introduction
Development of proxies that have the potential to provide long-term and reliable palaeoclimate
records is a key to reconstructing and understanding past climate systems. Ideally these proxies
should be widely distributed and common in the environment. Chironomids (Diptera:
Chironomidae) are non-biting midges whose larvae occur in virtually all permanent and semi-
permanent terrestrial water bodies (Cranston, 1995). The fossilized head capsules of chironomid
larvae are preserved in lake sediment and the structure of these chitinous exoskeleton fragments is
usually well preserved (Walker, 1987). The growth of chironomid larvae is strongly controlled by
water temperature (Armitage, 1995). Based on these characteristics, transfer functions based on
chironomid species assemblages have been developed from various parts of the world for
paleoclimate and environment inferences (e.g. Lotter et al., 1999; Larocque et al., 2001; Woodward
and Shulmeister, 2006; Walker and Cywnar, 2006; Rees et al., 2008; Massaferro et al., 2014 and
Chang et al., 2015a – Chapter 4). These have been successfully applied to Late Quaternary
palaeoclimatic data and have provided quality quantitative seasonal temperature reconstructions
(e.g. Woodward and Shulmeister, 2007; Rees and Cwynar, 2010; Samartin et al., 2012; Chang et
al., 2015b – Chapter 5).
In addition to the traditionally applied chironomid-based transfer functions, other studies (e.g.
Wooller et al., 2004, 2008; Gröcke et al., 2006) have shown that the δ18
O value of chironomid and
aquatic beetle chitin is largely a reflection of lake water δ18
O which correlates with the δ18
O of local
precipitation and therefore, has the potential to be used for regional paleoclimate reconstructions.
More recently, δ18
O variations in subfossil chironomid head capsules were applied to quantitatively
characterize the late glacial lake water δ18
O changes and, indirectly, past air temperature changes
near Rotsee, Switzerland (Verbruggen et al., 2010b).
There are two key assumptions that need to be considered before stable isotope measurements from
chironomid head capsules in lake sediment can be used as proxies for climate. Firstly, it is often
assumed that the modern and past isotopic composition of lake water reflects mean annual
precipitation (Rozanski et al. 1993). While this is generally true in humid regions, evaporation
affects the stable isotope composition of water via evaporative enrichment of the heavy isotope
(Leng et al., 2006) and this assumption needs to be tested for sub-humid and semi-arid areas. In
addition, the lake examined needs to be volumetrically large enough but un-stratified for its isotope
composition to reflect the average isotope values in mean annual precipitation (Leng et al., 2006).
But, stratification may protect a large portion of the water from evaporative enrichment
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(Verbruggen et al., 2011). Lake water δ18
O values may approximate those of precipitation in
stratified lakes if there is at least one mixing event per year.
Secondly, it is sometimes assumed that the isotopic composition of the head capsules is in
equilibrium with the lake water and that there are no vital and micro-environmental effects that
might modify head capsule isotope values. Vital effects could be induced by chironomid metabolic
processes and diet. Such effects have been examined in foraminifera (Erez, 1978), ostracods (von
Grafenstein et al. 1999), and cocoliths (Ziveri et al., 2003) but there has been very little work on
chironomid vital effects (Grey et al, 2004a, b; van Hardenbroek et al., 2014).
In an attempt to address these issues in chironomids, Wang et al (2009) quantified how the δ18
O and
δ2H of water and diet influenced the δ
18O and δ
2H of chironomid larvae. Their results revealed that
both water and diet affect the δ18
O and δ2H isotope composition of chironomid larvae, whereby
~70% of the oxygen in the total organic composition is derived from the water of the larval habitat.
In contrast, diet dominates the hydrogen isotope ratios of chironomid larvae (~70%), and only 30%
of hydrogen in the chironomid larvae is derived from the ambient water (Wang et al.
2009). However, this laboratory experiment is based on feeding the chironomid larvae powdered
Spirulina algae as controlled diet. In natural aquatic environments, the total proportion of hydrogen
in the chironomid larvae that are derived from ambient water could be higher if indirect uptake of
δ2H in the natural diet is considered. Their results also suggested that the majority of the hydrogen
in lipids, protein and keratin are derived from diet rather than water, and these indicated vital effects
may have a large influence on the δ2H values and to some degree on the δ
18O values of
chironomids. Differences in metabolic pathways also likely influenced δ18
O and δ2H fractionation
in different taxa.
Here we present the first study of stable isotope (δ18
O and δ2H) analyses on chironomid head
capsules from southeastern Australian lakes. Based on previous research (Wang et al., 2009 ; Grey
et al., 2004a, b; van Hardenbroek et al., 2014), it is likely that different chironomid taxa exhibit
differences in stable isotopic composition (via vital effects) and will thus display different
relationships to climatic or environmental variables. We therefore performed the δ18
O and δ2H
analyses on head capsules from a single genus (Chironomus spp.) because it is the most abundant
genus in southeastern Australian lakes (Chapter 4). We investigate the relationship between the
stable isotope composition (δ18
O, δ2H) of Chironomus spp. head capsules and how this relates to the
composition of lake water and local precipitation. The main goal of this study was to examine the
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feasibility of the application of stable isotopes of Chironomus spp. head capsules as a proxy to
reconstruct past changes in climate in southeastern Australia.
6.4 Materials and Methods
6.4.1 Study sites and sample collection
Fifteen natural lakes and one reservoir located in southeast Australia (Table 6.1, Fig. 6.1) were
included in this dataset. The dataset covers the latitudes between 30˚S and 41.6˚S (Table 6.1, Fig.
6.1) and elevation of the sites ranged from sea level to c. ~2000 m above mean sea level (a.s.l)
(Table 6.1). The sixteen waterbodies included in this dataset had a large spatial distribution across
eastern Australia (Table 6.1). One lake (LLL) was located on top of New England Tablelands (Fig
6.1), which lies in the subtropics but maintains a cool temperate climate due to high elevation
(Table 6.1). Summer is the dominant season for the source of precipitation for this site. Three lakes
were from Mt Kosciusko in the Australian Alps (Fig. 6.1), which is the coolest and highest (Table
6.1) area of Australia and the precipitation (with significant snowfall) is winter and spring
dominant. Ten lakes were from western Victoria, where a true Mediterranean climate exists with
warm dry summers and cool wet winters. The region consists of semi-arid to sub-humid regions and
the lakes are highly susceptible to both natural and agricultural related eutrophication and
salinization (as concluded in Chapter 3). Two lakes were from northwestern Tasmania (Fig. 6.1)
and this is the region that is humid and precipitation is winter and spring dominant (similar to
Australian Alps). Detailed descriptions of the climate, vegetation and geology of the study area
were presented in Chapter 3.
All lakes were sampled during the summer (January and February) of 2012 and 2013 (Table 6.1). A
minimum of three sediment cores were taken using a Glew Mini Corer (Glew 1991) at the
geographical centre of each lake. The top 2 cm of each core were extruded on site and packaged at
0.5 cm intervals into Whirlpaks®. Lake water for stable isotope analyses was sampled from the
location where the core samples were taken, at approximately 30 cm below the water surface. They
were collected into pre-washed and labelled polyethylene bottles. All bottles were sealed to prevent
evaporation. Sediment and water samples were refrigerated until analysis.
Lake water samples were collected at the same time for the analysis of the concentrations of major
ions, total nitrogen/total phosphorus (TN/TP), Chlorophyll a, pH and specific conductance
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(COND). In the field, water temperature, oxidation reduction potential (ORP), dissolved oxygen
(DO), total dissolved solids, salinity and turbidity were also recorded using an Aquaread multi-
parameter meter (AQUAREAD, Kent, UK). Detailed methodology for water chemistry analyses
was outlined in Chapter 3 and analytical results are presented in Table 6.1.
6.4.2 Climatic and stable isotopes in precipitation data
Climate variables were obtained using the combination of the WorldClim program (available from
http://www.worldclim.org/bioclim, accessed 20 January, 2014) and ArcGIS 10.1. WorldClim data
for Australia is based on climate surfaces derived from around 600 nation-wide weather stations
that have climate records spanning the years 1950–2000
(http://www.bom.gov.au/climate/data/stations/, accessed 20 January, 2014). For this study, only
mean annual temperature (MAT) and precipitation (Precip) were considered (Table 6.1), because
the spatial variation in δ18
O in precipitation is usually strongly related to MAT along latitudinal
gradients (Rozanski et al. 1993). The development of chironomid larvae in lowland Australia can
span seasons of spring, summer and autumn, therefore the incorporated chironomid chitin may
reflect annual average isotopic variations. Potential evapotranspiration (PET) and Aridity Index
(AI) values were obtained from the Global Potential Evapo-Transpiration (Global-PET) and Global
Aridity Index (Global-Aridity) dataset (CGIAR-CSI, available from http://www.cgiar-
csi.org/data/globalaridity-and-pet-database, accessed 20 March 2015) (Table 6.1). Stable isotopes in
precipitation data were obtained from the Global Network of Isotopes in Precipitation (GNIP) data
set (IAEA/WMO, 2015) and interpolated using ArcGIS 10.1. for each site (Table 6.2).
6.4.3 Stable isotope sample preparation
Sediment treatment and stable isotope sample preparation were performed in the School of
Geography, Planning and Environmental Management, The University of Queensland laboratories.
Both sample preparation protocols developed and used in Wang et al. (2008) and Verbruggen et al.
(2011) were considered and employed with some modifications. Sediment samples were
deflocculated in cold 10% solution of potassium hydroxide (KOH) for two hours at room
temperature and subsequently sieved with a 180 μm mesh-size sieve (van Hardenbroek et al. 2010).
The top 2 cm was processed for chironomid head capsules from three surface core samples. Sieved
residues were stored in vials with distilled water. Head capsules of the genus of Chironomus
(Chironomus spp.) (Fig. 6.2), were picked from the sieved residue under a dissection microscope at
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50 × magnification using fine forceps and placed into pre-labelled vials. Verbruggen et al (2010a)
suggested that chemical pre-treatment of chironomid head capsules could affect the resulting δ18
O
values. Therefore, a physical treatment was used instead where after a minimum of 150 head
capsules (average count is 185) was picked into the vials, they were placed in an ultrasonic bath for
20-30 seconds to separate contaminants. The head capsules were then examined for contaminants
under the microscope and later transferred to a pre-weighed silver cup of 3.5 × 5 mm (Costech
Analytical Technologies, INC., code: 041066). This was based on a modified two-step transfer
protocol (Wang et al. 2008), which had an advantage of allowing any contaminants to be detached
and manually separated during the process. Silver cups containing the head capsules were allowed
to dry for several days in a covered Petri dish at room temperature. The silver cups were again
weighed and, if a minimum of 100 µg was present, folded and shape-trimmed (Verbruggen et al.
2011). Samples were subsequently measured for stable oxygen (δ18
O) and deuterium (δ2H) isotopes
in Purdue Stable Isotope laboratory (PSI) facility, West Lafayette, USA.
Figure 6.1 Sample lakes are distributed across southeast Australia but there is a strong
concentration (ten out of 16 lakes) in western Victoria. This reflects Chironomus spp. abundance in
waterways with the tolerant Chironomus spp. strongly represented in the brackish and saline lakes
of western Victoria.
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6.4.4 Stable isotope analyses
Stable oxygen isotope analyses of the lake water samples were prepared by vacuum pump filtration
and samples were placed in 10 ml Gas Chromatography (GC) vials. A volume of 0.3 µl aliquots
were auto-injected on the PSI lab high Temperature Conversion Elemental Analyzer (TC/EA,
Thermo Fisher Scientific) by a GC-PAL auto-sampler. The injected water sample was pyrolyzed
under reducing conditions at 1400 °C to produce H2 and CO gases. These were separated
chromatographically in a helium carrier gas stream and introduced sequentially into the ion source
of an isotope ratio mass spectrometer (IRMS, Thermo Fisher Scientific) (Delta V Plus,
TheremoFinnigan) for isotope ratio determination (Gehre et al., 2004). Six sequential injections
were made for each sample and the reported values represent the average of three final injections
(Nielson and Bowen, 2010). All lake water stable isotope data were calibrated using repeated
analysis of three internal standards (PT, UT and PZ) relative to Vienna-Standard Mean Ocean
Water and Standard Light Antarctic Precipitation (VSMOW/SLAP) (Coplen, 1995). All data were
reported as per mil (‰) relative to the V-SMOW standard. Average uncertainties for lake water
were ± 0.44‰ for δ18
O and ± 3.6‰ for δ2H.
The Chironomus spp. head capsule samples were stored in the PSI facility at room temperature for
seven days prior to analysis for the samples to equilibrate with the analytical environment. The
TC/EA coupled to IRMS were used to determine ratios of stable isotopes for Chironomus spp. head
capsules (sample weight > 100 µg). Samples were transferred to a Zero Blank Auto-sampler
(Costech Analytical) interfaced with the TC/EA. They were pyrolyzed at 1400 °C in an oxygen-free
environment to produce H2 and CO gases, which were chromatographically separated and
introduced sequentially to the source of the IRMS (Gehre et al., 2004). Two blanks were measured
at the start of every run. Three commercial keratin laboratory reference materials (powdered KHS,
FH and UH) were used to normalize the results. Oxygen isotope δ18
O data and δ2H values were
calibrated against primary reference materials (Nielson and Bowen, 2010) and all data were
reported as per mil (‰) relative to the VSMOW standard (Coplen, 1995). Average precisions of ±
0.8‰ and ± 4.0‰ for δ18
O and δ2H of Chironomus spp. head capsules were achieved respectively.
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Figure 6.2 Chironomus spp. head capsules from (a) Freshwater Lake, Victoria, 20 ×. (b) Lake
Albina, Mount Kosciuszko, New South Wales, 10 ×. Photos of head capsules were taken by J.
Chang at the Physical Geography Laboratory of School of Geography, Planning and Environmental
Management, University of Queensland
6.4.5 Statistical analyses
Regression analyses and partial redundancy analysis (RDA) in CANOCO version 4.5 (ter Braak and
Šmilauer, 2002) were used to test the correlation significance, regression coefficient and the
independence of the variable correlations between the stable oxygen (δ18
O) and hydrogen (δ2H)
isotope data of Chironomus spp. head capsules against climatic and environmental variables. The
purpose of these analyses was also to determine if fractionation or vital effects were constant or
varied with changes in the environment.
Changes between isotope compositions of two substances are given as fractionation factor (α). The
α for Chironomus spp. and lake water of oxygen (δ18
O) and hydrogen (δ2H) isotopes is defined as:
Eq. 6.1
Eq. 6.2
respectively.
For testing correlations between fractionation factors (α) and environmental and climatic variables,
the natural logarithm of α (lnα) was used by convention (Clark and Fritz, 1997) and was multiplied
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by 103 to adapt to the ‰ notation for δ values. Both regression and RDA (CANOCO version 4.5)
were used to test the correlation significance, regression coefficient and the independence of the
variable correlations between α[δ18O(Chironomus spp. - water)] and α[δ2H(Chironomus spp. - water)] against climatic
and environmental variables, respectively.
6.5 Results
Stable oxygen (δ18
O) and hydrogen (δ2H) isotope analytical results on head capsules of Chironomus
spp. and the respective host lake water were obtained from sixteen southeastern Australian
waterbodies (Table 6.2). Fractionation factors (α) between Chironomus spp. and host lake water
were calculated applying Eq. 1 and Eq. 2 for δ18
O and δ2H, respectively and results are presented in
Table 6.2.
Figure 6.3 (a)Plot of δ18
O of precipitation against δ18
O of lake water. The δ18
O of precipitation and
lake water is correlated but significant enrichment of lake water is observed at δ18
O of precipitation
of around -5 ‰ V-SMOW, which the offset is possibly due to high regional evaporation rate and
prolonged residence time or local hydrological conditions (b) the plot of δ18
O of lake water against
Chironomus spp. showed no signifcant correlation (P = 0.13) between the two data sets, suggesting
that not all oxygen stable isotopic composition of Chironomus spp. head capsules were reflecting
changes of their ambient water
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Figure 6.4 (a) Plot of aridity index (AI) against δ18
O of Chironomus spp.. AI has the best
correlation and is the independent variable that explains the largest percent variance (Table 1) of
Chironomus spp. δ18
O data. At semi-arid and sub-humid areas (e.g. when AI <1), the variation in
δ18
O of Chironomus spp. data is large. (b) Total Nitrogen (TN) against fractionation factor
(α[δ18O(chiro-water)]). TN explains an independent and large percent of variance in the fractionation
factor between Chironomus spp. and host lake water (Table 6.2), suggesting lake eutrophication is
an important consideration for the interpretation of stable isotope data for Chironomus spp.
6.5.1 Stable oxygen isotope (δ18
O) analyses results
The annual averaged of δ18
O of precipitation varied between -4.65 and -7.20‰ and was positively
correlated (r2 = 0.7) with the lake water δ
18O (Fig. 6.3a), which varied between -6.17 and 12.97‰.
Lake water δ18
O in a number of western Victorian lakes was significantly enriched relative to
precipitation (Table 6.2, Fig. 6.3a). The measured δ18
O values from the chitinous head capsules of
Chironomus spp. from the surface sediments of all sixteen southeastern Australian lakes showed no
significant correlation (p > 0.05) with lake water isotopic composition (Fig. 6.3b).
Regression analyses of δ18
O of Chironomus spp. values against climatic and environmental
variables showed that five variables were significantly correlated (p < 0.05). These were mean
annual temperature (MAT), Aridity index (AI), potential evapotranspiration (PET), pH and specific
conductance (COND). Redundancy analyses (RDAs) were then performed using the δ18
O of
Chironomus spp. values against these five variables (Table 6.3). Results showed that AI was the
variable that explained the largest percent (53.7%) of the total variance in the δ18
O Chironomus spp.
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data, although it did not retain its significance when MAT was partialled out (Table 6.3). However,
the RDA on MAT showed that when AI was partialled out, it only explained 0.9% of the total
variance (Table 6.3). AI and MAT are correlated and this is due to the effect of temperature on
evaporation of the lake waters. AI includes a potential evapotranspiration (PET) component which
is dependent on MAT, as PET is primarily controlled by temperature (MAT). However, AI was
selected because it has the most powerful explanatory of variance in the δ18
O of Chironomus spp.
data. The regression plot of δ18
O of Chironomus spp. against AI showed a relatively strong positive
correlation (r2 = 0.54, p < 0.05) (Fig. 6.4a). The variation in the δ
18O of Chironomus spp. values
was large for sites with an AI value of less than one (i.e. semi-arid and sub-humid sites) (Fig. 6.4a).
Figure 6.5 (a)Plot of δ2H of precipitation against δ
2H of lake water. The δ
2H of precipitation and
lake water is closely correlated, with r2 = 0.92 (b) Plot of δ
2H of lake water aginst δ
2H of
Chironomus spp.. δ2H of lake water and Chironomus spp. showed a close correlation (r
2= 0.69)
suggesting that a large proportion of δ2H stable isotopic composition of Chironomus spp. head
capsules that were directly derived from their ambient water or indirectly from feeding on
planktonic algae, where algae also contain δ2H stable isotopes that are derived from lake water.
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Table 6.1 Summary of information available for the 16 lakes included in this study. The complete data-set is available in Chapter 3.
No Lake Name Lake Coordinates ALT Depth PET Precip AI MAT TP TN Chl a pH COND Nutrient
Limitation Trophic status
m m mm/Year mm/Year
˚C mg/L mg/L µg/L - µs/cm
1 Little
Llangothlin
Lagoon LLL S30.09
E151.78 1361 3 1142 944 0.83 11.6 0.16 1.5 31 8.11 212 N,P Eutrophic
2 Blue Lake BL S36.41
E148.32 1901 28 724 1708 2.36 3.9 0.007 0.11 2 6.44 5 N,P Oligotrophic
3 Lake Albina LA S36.42
E148.27 1919 9 728 1693 2.33 4.4 0.007 0.12 1 6.6 6 N,P Oligotrophic
4 Lake
Cootaptamba CTL S36.46
E148.26 2048 3 673 1691 2.51 3.7 0.011 0.15 1 6.47 5 N,P Oligotrophic
5 Nuggety
Gully
Resrvoir NGR S37.00
E143.73 221 1.2 801 537 0.67 14.1 0.041 1.6 11 7.02 400 P Eutrophic
6 Lake Fyans LFY S37.14
E142.62 219 2.7 1367 595 0.44 13.8 0.019 0.64 3 8.79 206 P Eutrophic
7 Freshwater
Lake FWL S37.59
E142.32 227 2.1 802 660 0.82 13.2 0.56 5.1 60 6.74 564 N,P Hypereutrophic
8 Lake
Tooliorook LTK S37.98
E143.28 167 3 1539 591 0.38 13.4 0.12 2.2 1 9.17 2470 P Hypereutrophic
9 Lake
Surprise LSP S38.06
E141.92 107 6 1208 763 0.63 13.3 0.031 0.88 9 8.08 698 P Eutrophic
10 Lake
Mombeong LMB S38.13
E140.18 21 4.8 1096 793 0.72 13.8 0.021 0.79 5 8.03 1140 P Eutrophic
11 Lake
Terangpom LTP S38.14
E143.33 133 0.5 1239 649 0.52 13.5 3.6 36 90 8.68 15700 N,P Hypereutrophic
12 Swan Lake SWL S38.21
E141.31 23 1.1 1058 797 0.75 13.7 0.044 1.3 10 7.61 744 P Eutrophic
13 Lake
Cartcarrong LCT S38.24
E142.45 90 1.1 1241 835 0.67 13.4 1.1 11 24 8.33 4700 N,P Hypereutrophic
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No Lake Name Lake Coordinates ALT Depth PET Precip AI MAT TP TN Chl a pH COND Nutrient
Limitation Trophic status
m m mm/Year mm/Year ˚C mg/L mg/L µg/L - µs/cm
14 Lake
Elingamite LEM S38.35
E143.00 147 1.2 1241 891 0.72 13.3 0.11 4.4 22 7.98 6420 P Eutrophic
15 Lake Lila WP S41.65
E145.96 957 13 720 2207 3.07 6.7 0.019 0.29 1 4.91 23 N,P Mesotrophic
16 Lake Lea
Pond LEA S41.53
E145.91 837 0.75 1484 2095 1.41 7.4 0.028 0.44 5 4.83 33 N,P Mesotrophic
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Table 6.2 Stable isotope analyses (δ18
O and δ2H) results of southeastern Australian modelled precipitation, lake water, Chironomus spp. and the
fractionation factor between chironomus head capsules and lake water (α).
No Lake Name Lake
δ18
O of
modelled
precipitation
δ18
O of
lake
water
δ18
O of
Chironomus
spp.
α
[δ18O(Chironomus
spp - water)]
δD of
modelled
precipitation
δD of
lake
water
δD of
Chironomus
spp.
α
[δ2H(Chironomus
sp - water)]
VSMOW VSMO
W VSMOW VSMOW
VSMO
W VSMOW
1 Little Llangothlin
Lagoon LLL -6.70 -1.25 18.70 19.78 -39.13 -34.04 -91.70 -61.55
2 Blue Lake BL -7.20 -7.76 12.27 19.99 -44.27 -50.69 -84.82 -36.61
3 Lake Albina LA -7.20 -7.02 13.27 20.23 -44.27 -54.33 -101.17 -50.80
4 Lake Cootaptamba CTL -7.20 -6.53 13.10 19.56 -44.27 -44.49 -94.87 -54.17
5 Nuggety Gully Resrvoir NGR -4.80 11.27 15.77 4.44 -26.61 57.81 -63.29 -121.58
6 Lake Fyans LFY -4.91 3.57 15.28 11.61 -27.47 28.15 -48.18 -77.13
7 Freshwater Lake FWL -5.01 5.57 15.94 10.26 -28.30 40.80 -54.70 -96.25
8 Lake Tooliorook LTK -4.91 1.05 16.88 15.69 -27.61 27.87 -73.17 -103.47
9 Lake Surprise LSP -4.91 1.18 16.80 15.47 -27.70 27.48 -75.40 -105.50
10 Lake Mombeong LMB -4.65 3.34 16.28 12.82 -26.71 28.63 -72.82 -103.83
11 Lake Terangpom LTP -4.81 10.54 13.91 3.33 -26.94 56.67 -55.32 -112.04
12 Swan Lake SWL -4.72 4.02 13.63 9.52 -26.46 43.26 -69.18 -114.04
13 Lake Cartcarrong LCT -4.74 12.97 15.09 2.09 -26.52 61.63 -70.00 -132.38
14 Lake Elingamite LEM -4.85 5.52 16.09 10.45 -27.27 51.33 -50.93 102.33
15 Lake Lila WP -6.21 -6.17 12.43 18.54 -38.17 -34.55 -79.53 -47.71
16 Lake Lea Pond LEA -5.73 2.43 15.22 12.67 -34.60 17.26 -75.81 -95.95
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Regression analyses of the δ18
O fractionation factor between the chironomid head capsules and lake
water (α[δ18O(Chironomus spp. - water)] ) with respect to climatic and environmental variables showed that
seven variables: MAT, Total Nitrogen (TN), Total Phosphorus (TP), Chlorophyll a (Chl a), depth,
specific conductance (COND) and AI were significantly correlated (p < 0.05). Redundancy
analyses (RDAs) were performed using the α[δ18O(Chironomus spp. - water)] values against all seven
variables (Table 6.4). TN was the variable that explained the largest percent (64.4%) of the total
variance in the α[δ18O(Chironomus spp - water)] data, although it did not retain its significance level when
COND was partialled out (Table 6.4), the RDA on COND showed that when TN was partialled out,
it only explained 0.1% of the total variance (Table 6.4). Thus, there is a strong interaction between
COND and TN. But clearly TN was the more independent variable. The regression plot of the
values of ɑChironomus spp. – water for δ18
O against TN showed a close negative correlation (with
regression coefficient r2 = 0.64) (Fig. 6.4b).
Figure 6.6 (a) Plot of Mean annual temperature (MAT) against δ2H of Chironomus spp.. MAT had
the best correlation and is the independent variable that explains the largest percent variance (Table
6.5) of Chironomus spp. δ2H data. The correlation could possibly be complicated by the
fractionation between δ2H of Chironomus spp. and δ
2H of lake water (α[δ2H(chiro-water)]), which was
influenced by other environmental variables. (b) Plot of ln(COND) against fractionation factor
α[δ2H(chiro-water). Specific conductance (COND) explained an independent and large percent of
variance in the fractionation factor between δ2H of Chironomus spp. and host lake water (Table
6.6), lake water COND had a close correlation (r2 = 0.695) with the fractionation factor (α) between
δ2H of Chironomus spp. and lake water.
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Table 6.3 Redundancy Analysis (RDA) of δ18
O of Chironomus spp. head capsules and
environmental variables shows that Aridity Index (AI) has the strongest explanatory power (53.7%)
among the five variables that have a significant correlation (P < 0.05). It is not independent of MAT
but it is impossible that it would be independent as temperature plays a key role in potential
evapotranspiration (PET). AI explains significantly more variance than MAT and MAT is reduced
to very low explanatory power when AI is removed.
Variable Co-variable % of Variance
explained
λ1 Correlation p-value
MAT none 42.5 0.425 0.652 0.005
Cond 24.5 0.179 0.495 0.06
pH 23.9 0.179 0.489 0.07
PET 20.5 0.134 0.453 0.091
AI 0.9 0.004 0.093 0.737
ALL 1.4 0.006 0.119 0.707
AI none 53.7 0.537 0.733 0.002
MAT 20.1 0.115 0.448 0.118
Cond 39 0.285 0.625 0.015
pH 38.3 0.287 0.619 0.014
PET 31.4 0.204 0.56 0.029
ALL 9.5 0.044 0.309 0.326
PET none 34.9 0.349 0.591 0.016
AI 3.6 0.016 0.188 0.494
MAT 10 0.057 0.315 0.246
Cond 15.2 0.111 0.389 0.159
pH 20.7 0.155 0.455 0.093
ALL 7.2 0.032 0.268 0.403
pH none 25.1 0.251 0.501 0.055
AI 0.3 0.001 0.056 0.83
MAT 0.9 0.005 0.093 0.743
Cond 5.2 0.038 0.228 0.411
PET 8.8 0.058 0.297 0.28
ALL 0.1 0 0.028 0.93
COND none 26.9 0.269 0.519 0.035
AI 3.8 0.017 0.194 0.497
MAT 4 0.023 0.199 0.468
pH 7.4 0.056 0.272 0.329
PET 4.8 0.031 0.218 0.424
ALL 6.6 0.029 0.257 0.421
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Table 6.4 Redundancy Analysis (RDA) of the fractionation factor (α) (δ18
O between Chironomus
spp. and lake water) and environmental variables shows that Total Nitrogen (TN) is the most
independent variable that explains the largest variance (64.4%) among all seven variables that are
significantly correlated (P < 0.05) for the stable oxygen isotope (δ18
O) fractionation factor
(α[δ18O(Chironomus spp - water)]) (Table 6.2) between Chironomus spp. and host lake water
Variable Co-variable % of
Variance
explained
λ1 Correlation p-value
MAT none 35.7 0.357 0.597 0.006
AI 1.1 0.007 0.107 0.666
TP 10 0.054 0.316 0.244
TN 1.9 0.007 0.136 0.581
Chla 8.8 0.054 0.296 0.29
DEPTH 7.7 0.031 0.278 0.286
COND 0.3 0.002 0.058 0.837
ALL 20.8 0.045 0.456 0.142
TN none 64.4 0.644 0.802 0.001
AI 40.2 0.232 0.634 0.008
MAT 45.7 0.294 0.676 0.006
Depth 29 0.115 0.538 0.03
TP 42.9 0.233 0.655 0.004
Chla 42.4 0.263 0.651 0.004
COND 20.3 0.09 0.45 0.079
ALL 41.9 0.123 0.647 0.036
TP none 45.7 0.457 0.676 0.003
AI 22.9 0.132 0.478 0.061
MAT 23.9 0.154 0.489 0.068
Depth 9.3 0.037 0.304 0.25
TN 12.8 0.046 0.358 0.181
Chla 14.4 0.089 0.379 0.158
COND 6.1 0.027 0.248 0.348
ALL 37.7 0.104 0.614 0.039
Chla none 38.1 0.381 0.617 0.007
AI 11.4 0.066 0.337 0.195
MAT 12.3 0.079 0.35 0.186
Depth 4.8 0.019 0.22 0.433
TP 2.5 0.014 0.159 0.537
TN 0 0 0 1
COND 7.9 0.035 0.281 0.302
ALL 2.3 0.004 0.153 0.681
COND none 55.4 0.554 0.744 0.003
AI 23.9 0.138 0.489 0.06
MAT 30.8 0.198 0.555 0.02
Depth 25 0.099 0.5 0.051
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Variable Co-variable % of
Variance
explained
λ1 Correlation p-value
TP 22.9 0.124 0.479 0.068
TN 0.1 0 0.356 0.916
Chla 33.6 0.208 0.579 0.022
ALL 24.3 0.055 0.493 0.128
DEPTH none 60.4 0.604 0.777 0.001
AI 37.5 0.216 0.612 0.01
MAT 43.1 0.278 0.657 0.005
TP 33.8 0.184 0.582 0.022
TN 21 0.075 0.458 0.084
Chla 39.1 0.242 0.625 0.013
COND 33.4 0.149 0.578 0.024
ALL 15.9 0.032 0.399 0.224
AI none 42.3 0.423 0.65 0.004
MAT 11.3 0.073 0.337 0.208
TP 18.1 0.099 0.426 0.103
TN 3.1 0.011 0.176 0.518
Chla 17.4 0.108 0.417 0.12
COND 1.7 0.008 0.13 0.635
Depth 9 0.036 0.3 0.245
ALL 6.8 0.012 0.26 0.466
6.5.2 Stable hydrogen isotope (δ2H) results
For deuterium (δ2H), the annual average δ
2H of precipitation varied between -44.27 and -26.46 ‰
VSMOW and was closely correlated with the lake water δ2H which varied between -6.17 and
12.97‰ VSMOW (r2 = 0.92) (Fig. 6.5a). The measured δ
2H values from the chitinous head
capsules of Chironomus spp. from the surface sediments of the sixteen southeastern Australian
lakes showed a significant positive correlation (p < 0.05) with host water isotopic composition, with
a regression coefficient (r2) of 0.69 (Fig. 6.5b).
Regression analyses of δ2H of Chironomus spp. values against climatic and environmental variables
showed that seven variables (MAT, TN, TP, Chl a, Depth, COND and AI) were significantly
correlated (p < 0.05). Redundancy analyses (RDAs) were then performed using the δ2H of
Chironomus spp. values against these seven variables (Table 6.5). Results showed that MAT was
the variable that explained the largest percent (55.2%) of the total variance in the δ2H Chironomus
spp. data. The explanatory power of MAT was reduced by partialling out COND, AI and TN (Table
6.5). The reduction in explanatory power was greatest after partialling out COND and AI (Table
6.5). However, MAT appears to be the most independent variable since the explanatory power of AI
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and COND is almost eliminated by partialling out MAT. MAT is clearly more independent than TN
since the explanatory power of TN is much more severely affected by the partialling out of MAT
than MAT by the partialling out of TN (Table 6.5). COND, TN, AI and MAT are correlated and this
is because of the effect of temperature on evaporation and the concentration of ions and nutrients in
the lake waters. The regression plot of δ2H of Chironomus spp. against MAT showed a positive
correlation with a regression coefficient of r2 = 0.55 (Fig. 6.6a).
Regression analyses of the fractionation (ɑChironomus spp. – water) values for δ2H against climatic and
environmental variables showed that TN, TP, Chl a, COND, Depth, MAT and AI were significantly
correlated (p < 0.05). The partial RDAs showed that COND was the sole variable that retained
significance (p < 0.05) when all other variables were partialled out and explained 68.4% of the total
variance in the ɑChironomus spp. – water values for δ2H (Table 6.6). The regression plot of δ
2H
fractionation values against COND has a correlation coefficient of r2 = 0.695 (Fig. 6.6b).
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Table 6.5 Redundancy Analysis (RDA) of δ2H of Chironomus spp. head capsules and
environmental variables shows that for δ2H there are no independent variables. MAT explains the
largest variance (55.2%) in stable hydrogen isotope (δ2H) data of Chironomus spp. but interacts
with aridity index (AI), specific conductance (COND) and total nitrogen (TN).
Variable Co-variable % of
Variance
explained
λ1 Correlation p-value
MAT none 55.2 0.552 0.743 0.001
AI 18 0.098 0.424 0.136
TP 40.2 0.297 0.634 0.016
TN 22.4 0.122 0.473 0.085
Chla 39 0.284 0.624 0.015
DEPTH 35.2 0.209 0.593 0.028
COND 11.6 0.057 0.34 0.24
ALL 5.7 0.014 0.238 0.516
TN none 45.6 0.456 0.675 0.004
AI 13.5 0.073 0.367 0.19
MAT 5.8 0.026 0.242 0.343
Depth 15 0.089 0.387 0.18
TP 55.4 0.41 0.745 0.005
Chla 26 0.189 0.509 0.051
COND 0.7 0.003 0.083 0.777
ALL 38.4 0.14 0.619 0.054
TP none 26.1 0.261 0.511 0.045
AI 4 0.022 0.2 0.5
MAT 1.5 0.007 0.124 0.643
Depth 1.9 0.011 0.138 0.636
TN 39.5 0.215 0.628 0.008
Chla 3.2 0.023 0.178 0.511
COND 1.2 0.006 0.108 0.678
ALL 37.5 0.136 0.613 0.058
Chla none 27.2 0.272 0.521 0.052
AI 1.9 0.011 0.139 0.614
MAT 0.9 0.004 0.092 0.759
Depth 3.3 0.02 0.182 0.526
TP 4.5 0.033 0.212 0.438
TN 0.8 0.004 0.091 0.739
COND 0.4 0.002 0.06 0.817
ALL 9.8 0.024 0.313 0.37
COND none 50.4 0.504 0.71 0.002
AI 14.2 0.077 0.376 0.145
MAT 2.2 0.01 0.147 0.61
Depth 24.8 0.148 0.498 0.06
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Variable Co-variable % of
Variance
explained
λ1 Correlation p-value
TN 9.4 0.051 0.307 0.303
Chla 32.2 0.234 0.567 0.03
ALL 28.2 0.089 0.531 0.112
Depth none 40.5 0.405 0.636 0.007
AI 12.6 0.069 0.355 0.196
MAT 14 0.063 0.374 0.192
TP 21 0.155 0.458 0.074
TN 7 0.038 0.265 0.337
Chla 21.1 0.153 0.459 0.089
COND 9.8 0.049 0.313 0.258
ALL 0 0 0.018 0.959
AI none 45.5 0.455 0.674 0.004
MAT 0.2 0.001 0.045 0.879
TP 29.1 0.215 0.54 0.04
TN 13.2 0.072 0.364 0.194
Chla 26.6 0.194 0.516 0.046
COND 5.6 0.028 0.237 0.368
Depth 19.9 0.118 0.446 0.101
ALL 0.1 0 0.033 0.934
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Table 6.6 Redundancy Analysis (RDA) of the fractionation factor (α) (δ2H between Chironomus
spp. and lake water) and environmental variables shows that specific conductance (COND) is the
most independent variable that explains the largest amount of variance (68.4%) among all seven
variables that are significantly correlated (P < 0.05) in the hydrogen isotope (δ2H) fractionation
factor (α) between Chironomus spp. and host lake water
Variable Co-variable % of
Variance
explained
λ1 Correlation p-value
MAT none 58 0.58 0.762 0.001
AI 6.4 0.023 0.253 0.357
TP 38.2 0.236 0.618 0.01
TN 27.7 0.119 0.526 0.036
Chl a 36.4 0.235 0.603 0.013
DEPTH 36.3 0.168 0.603 0.011
COND 12.3 0.039 0.35 0.181
ALL 3.1 0.006 0.175 0.608
TN none 57.1 0.571 0.755 0.001
AI 18.6 0.067 0.431 0.089
MAT 26 0.109 0.51 0.036
Depth 21.7 0.1 0.466 0.062
TP 41.3 0.256 0.643 0.01
Chla 33.7 0.218 0.581 0.021
COND 0.02 0 0 0.945
ALL 0.3 0.001 0.052 0.888
TP none 38 0.38 0.617 0.007
AI 9.2 0.033 0.304 0.262
MAT 8.7 0.036 0.295 0.299
Depth 5 0.023 0.223 0.422
TN 15.3 0.066 0.391 0.119
Chla 8.2 0.053 0.286 0.28
COND 0.2 0.001 0.043 0.857
ALL 1.7 0.003 0.129 0.698
Chla none 35.3 0.353 0.594 0.01
AI 2.6 0.01 0.163 0.544
MAT 1.9 0.008 0.136 0.633
Depth 4.6 0.021 0.215 0.399
TP 4.1 0.025 0.202 0.467
TN 0.1 0 0.031 0.927
COND 2.5 0.008 0.159 0.579
ALL 0.1 0 0.037 0.93
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Variable Co-variable % of
Variance
explained
λ1 Correlation p-value
COND none 68.4 0.684 0.827 0.001
AI 26.4 0.095 0.514 0.024
MAT 33.9 0.142 0.582 0.015
Depth 46 0.213 0.678 0.003
TP 49.1 0.304 0.7 0.003
TN 26.4 0.113 0.514 0.049
Chla 52.4 0.339 0.724 0.002
ALL 2.4 0.005 0.154 0.638
Depth none 53.8 0.538 0.733 0.002
AI 21.5 0.078 0.464 0.089
MAT 29.8 0.125 0.546 0.028
TP 29.1 0.18 0.54 0.034
TN 15.7 0.068 0.397 0.128
Chla 31.8 0.206 0.564 0.022
COND 21 0.066 0.459 0.071
ALL 17.5 0.044 0.419 0.209
AI none 64 0.64 0.8 0.001
MAT 19.7 0.083 0.444 0.089
TP 47.3 0.293 0.688 0.001
TN 31.7 0.136 0.563 0.029
Chla 45.8 0.297 0.677 0.004
COND 16.2 0.051 0.403 0.142
Depth 38.9 0.18 0.624 0.006
ALL 0.9 0.002 0.094 0.806
6.6 Discussion
6.6.1 Analytical considerations
The application of stable isotope analyses on subfossil chironomids is still at an early stage of
development and a standard chitin sample purification protocol for chironomid head capsules is not
yet established. There have been only two studies that reported the measurement of δ2H on
chironomid head capsules (Deines et al., 2009 and Wang et al., 2009) but studies on keratin material
(e.g. Wassenaar and Hobson, 2003) suggested that the use of hydrogen isotope ratios needs to be
considered carefully because of the rapid, partial exchange of loosely bound hydroxyl and/or amine
hydrogen atoms with the environment. Here, hydrogen isotope ratios were not corrected for the
contribution of exchangeable hydrogen. This was because at the time of analysis, a set of exchange-
calibrated chitin standards was not yet established at PSI (Nielson and Bowen, 2010). The amount
of exchangeable hydrogen is usually expressed as percentage of the total amount of hydrogen and is
15.3 ± 2.9% in chitin (Schimmelmann et al., 1993). Hydrogen atoms in exchangeable hydroxyl and
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amine groups should have the same isotopic composition across all chitin samples analysed at the
same time under the same conditions. This concern was discussed in details in Nielson and Bowen
(2010) for the analysis of shrimp chitin from PSI. As all samples reported in this study were
processed, pre-treated and analysed using internally consistent methods, the offsets should be
constant. Consequently while the absolute δ2H values of Chironomus spp. head capsules obtained
from this study will not be suitable to compare directly with other studies, the trends should still
yield useful information and are worthy of discussion.
6.6.2 Relationship between δ18
O of lake water and regional climate
Due to evaporative enrichment in arid areas and wide range spatial distribution of the sites, there is
a disparity in between the δ18
O of source water (i.e. precipitation) and the δ18
O of lake water.
Therefore, it is not surprising that although the δ18
O of precipitation was correlated with lake water
(Fig. 6.3a, r2 = 0.71) and mean annual temperature (MAT) (Fig. 6.7, r
2 = 0.84), the sites are
unevenly distributed around the respective regression lines.
We observe that δ18
O of lake water is closely correlated with δ18
O of precipitation for lakes in
perenially wet settings. However, there is a wide scatter of the δ18
O of lake water values in lakes
with similar precipitation δ18
O values, specifically when δ18
O precipitation is > 5 ‰ VSMOW (Fig.
6.3a). Sites with precipitation δ18
O values > 5 ‰ are located in sub-humid and arid areas (Fig. 6.3a,
Table 6.1). Evaporative enrichment of δ18
O in lake water is obviously important at these sites and
variations in enrichment probably reflect differences in local evaporation rates and residence time
as the lighter isotope (16
O) is preferentially lost due to evaporation (Leng and Marshall, 2004).
Lakes with enriched δ18
O with respect to source water are closed maar lakes and all are located in
western Victoria. In fact, in two of the western Victorian lakes (LTP and LCT) and one reservoir
(NGR), the δ18
O values exceed 10‰ VSMOW which is considered an excessively large value
(Lamb et al., 1999) and difficult to achieve by natural evaporation processes (Gibson et al., 2002).
In this case, very long residence times are likely the primary cause (Barton et al., 2007; Chivas et
al., 1993). In summary, Australia has a high inter-annual variability in rainfall and in seasonally arid
regions, such as western Victoria, where agricultural activities (e.g. water abstraction) are also
predominant, lakes were especially sensitive to water-balance (Herczeg et al., 1992) and changes in
local hydrology induced isotopic variations. This finding indicated that lakes that are highly
enriched in δ18
O may not be suitable to be included in a calibration set for paleoclimate
reconstructions.
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Figure 6.7 Plot of mean annual temperature (MAT) against δ18
O of precipitation for the 16 study
lakes. There is a positive correlation between MAT and δ18
O of precipitation, but since the
differences within the spatial distribution for both study sites and precipitation sources are large, the
data points are scattered.
6.6.3 Lake trophic status related δ18
O fractionation process and the metabolism and
respiration of Chironomus spp.
A few previous studies (Wang et al., 2009; Verbruggen et al., 2011; Mayr et al., in press) suggested
that oxygen stable isotopic composition of chironomid head capsules can serve as a useful tool for
lake water isotope reconstructions for δ18
O. However, these results were not replicated in this study
(Fig. 6.3b). The implication is that insect metabolism and/or respiration in Chironomus spp. may
play a vital part in the fractionation between δ18
O of lake water and Chironomus spp.. Figure 6.8
shows the experimental chironomid δ18
O values against δ18
O of host water in Wang et al (2009)
compared to this study. Sites where there is a strong relationship between δ18
O of lake water and
δ18
O Chironomus spp. are mostly oligotrophic to mesotrophic lakes in humid areas. In most of the
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eutrophic to hypereutrophic lakes in the semi-arid to sub-humid areas, the relationship does not
apply.
Figure 6.8 Plot of the experimental chironomid δ18
O values against δ18
O of host water in Wang et
al (2009) compared to values derived from this study. Eight of the lakes from this study (CTL, BL,
LA, WP, LEA, LLL, LSP, LTK), fall along the regression line of Wang et al (2009) these are
oligotrophic and mesotrophic lakes from Kosciuszko and Tasmania, and two western Victorian
lakes that are less enriched in δ18
O, i.e. less affected by prolonged residence time. The other eight
lakes (FWL, LEM, LCT, LTP, SWL, LFY, LMB, NGR) produced similar Chironomus spp. δ18
O
values.
There was a reasonably strong, positive (r2 = 0.64) correlation between total nitrogen (TN) and the
fractionation factor of Chironomus spp. and lake water (α[δ18O(Chironomus spp - water)]) for δ18
O (Fig.
6.4b), showing that lake trophic status is an important consideration for the interpretation of δ18
O
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data for Chironomus spp.. Since seven out of sixteen of our lakes are phosphorous (P) limited (nine
are N and P co-limited), TN was always an excess nutrient available in these lakes (Table 6.1). High
levels of nutrients (such as TN) fuel algae blooms, which can initially boost dissolved oxygen (DO)
level. However, more algae means more plant respiration, drawing down DO, and when the algae
die, bacterial decomposition spikes, using up much of the dissolved oxygen available (Granéli and
Solander, 1988). The process of the exchange of nutrients across the sediment-water interface in
shallow water also enhances low oxygen available environments (Boström et al., 1998). Shallow
lakes located in semi-arid regions, such as western Victoria, with high residence times are highly
susceptible to this effect (Chapter 3). These lakes are ecologically challenging but Chironomus spp.
are dominant in most lakes of this type. It reached a maximum abundance of 63.7% (e.g. Lake
Toolirook (LTK), Victoria (Table 6.1)) in lowland eutrophic to hypereutrophic lakes and is present
in 94% of the lakes sampled in Chapter 4 in southeastern Australia. This is similar to New Zealand
(e.g. Woodward and Shulmeister, 2006), where in some warm lowland eutrophic lakes, Chironomus
spp. reached a maximum abundance of 80%.
One of the key features that allows Chironomus spp. to thrive in eutrophic and even anoxic
conditions is that they have the ability to metabolise and respire when oxygen availability is low
(Walker, 1987; Brodersen et al., 2004). This is because Chironomus larvae have haemoglobin
which regulates the respiration and oxygen level in the Chironomus spp. body (Osmulski et al.,
1986) and enables them to survive in low oxygen waters.
In summary, stable oxygen isotope values (δ18
O) of Chironomus spp. from eight out of sixteen sites
within the dataset appear to reflect lake water δ18
O and those are mostly lakes from humid and/or
cooler regions. A strong local evaporation signal and prolonged residence time caused the lake
water δ18
O to be enriched by > ~8 ‰ VSMOW with respect to precipitation δ18
O in eight of the ten
western Victorian sites. High lake nutrient status is associated with the dominance of Chironomus
spp. in the species composition of these lakes. We infer that this is an indication of ecological
adaptation of Chironomus spp. to extreme lake conditions. It is likely that although Chironomus
spp. can survive in these extreme environments due to the oxy-regulatory effect of haemoglobin.
Fractionation effects are altered in these extreme (e.g. oxygen limited) environments. Therefore it is
unsurprising that δ18
O of Chironomus spp. head capsules do not reflect the lake water δ18
O for these
eight sites (due to vital effects) and consequently, they are not directly correlated to climatic factors.
We re-affirm the findings of Mayr et al. (in press) that for semi-arid regions where δ18
O of
precipitation is not a priori correlated with mean annual air temperature, δ18
O values in
chironomids cannot be used for temperature reconstructions.
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6.6.4 Relationship between δ18
O of Chironomus spp. and regional climate
The previous discussion highlighted the large vital effects on Chironomus spp. δ18
O in western
Victorian lakes, or lakes subject to eutrophication and evaporative enrichment. When all lakes were
considered (Fig. 6.9a) there was a reasonable correlation between δ18
O and MAT. However, the
correlation was greatly improved when only lakes that have a δ18
O enrichment less than ~ 8‰
VSMOW compared to precipitation δ18
O were considered, δ18
O of Chironomus spp. and mean
annual temperature (MAT) (r2 = 0.78) (Fig. 6.9b). The slope (0.51) and intercept (10.7 ± 0.8‰)
values of the regression obtained in these eight southeastern Australian lakes were very similar to
Verbruggen et al.’s (2011) European dataset, which had a slope of 0.57 and an intercept value of 12
± 0.32‰ (Fig. 6.9b). This is pleasing, but surprising, as the δ 18
O analyses of chironomid head
capsules from Verbruggen et al. (2011) were based on a mixed taxon assemblage, including
Heterotrissocladius subpilosus-type, Cladotanytarsus, and Cricotopus (Verbruggen et al., 2011). At
first inspection, interspecific vital effects may not be as significant as predicted, but this observation
needs further investigation.
Our results confirm that lakes that are not affected by significant δ18
O enrichment caused by high
local evaporation and residence time can be used for temperature reconstructions. Consequently
temperature reconstructions from Chironomus spp. δ18
O in Australia are applicable only to areas
such as Tasmania and the higher altitude areas of the southeastern mainland.
6.6.5 The δ2H of precipitation, lake water, Chironomus spp. and temperature
Our analyses of lake water δ2H revealed a close correlation with δ
2H of precipitation (Fig. 6.5a) and
the enrichment of δ2H in lake water was not as strongly affected as δ
18O by local evaporation and
residence time (Fig. 6.3a). The δ2H values of Chironomus spp. and lake water showed a moderately
strong and significant (p < 0.05) correlation (r2 = 0.69) (Fig. 6.5b), which indicated a large
proportion of Chironomus spp. δ2H was derived from or exchanged with lake water. At first glance,
this observation does not appear to agree with the cultured experiment by Wang et al (2009), who
suggested that only ~30% of the δ2H of chironomids is derived from ambient water and ~70% is
from diet. However, this is expected because in nature, the genus Chironomus is mainly feeding on
planktonic algae, where algae also contain δ2H stable isotopes that are derived from lake water
(Zhang and Sachs, 2007). In addition, the inter-specifc dietary variability has been reduced in our
study as the chironomids samples were composed of a single genus of Chironomus morpho-species.
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Figure 6.9 (a) Plot of mean annual temperature (MAT) against δ18
O of Chironomus spp. with all
sixteen lakes included (b) Plot of mean annual temperature (MAT) against δ18
O of Chironomus spp.
with only the eight lakes that have the δ18
O enrichment of lake water less than ~8‰ VSMOW
relative to precipitation δ18
O are included. There is a close correlation (r2 = 0.78) between MAT and
Chironomus spp. δ18
O for this subset of study lakes. The slope and intercept values were very
similar to those observed by Verbruggen et al. (2011).
The fractionation (α[δ2H(Chironomus spp - water)]) between δ2H of Chironomus spp. and lake water
however, follows a specific conductance (COND) gradient (Fig. 6.6b) with a r2 = 0.69 indicating
that it is an important factor in the Chironomus spp. δ2H and lake water δ
2H relationship. However,
without further laboratory experiments and more testing, we cannot provide an insight of how
specific conductance affects the fractionation process between δ2H in Chironomus spp. and lake
water. In freshwater lakes (specific conductance < ~500 μs/cm, Behar 1997), that support diverse
aquatic life, mean annual temperature (MAT) and COND co-varied (Fig. 6.10) and there are
potentially three reasons for this observation. First, lake order effect may be driving the salinity
gradient where high altitude (cooler temperature) low order lakes are often dilute (e.g. Mt
Kosciusko lakes, Fig. 6.1) and lowland (warmer temperature) high order lakes have higher salinity.
Secondly, when specific conductance is > 500 μs/cm (e.g. Victorian lakes, Fig. 6.1), the changes in
specific conductance are driven by catchment hydrology, e.g. residence time, which affects
evaporative enrichment of salts. Finally, the overall pattern is perhaps due to regional evaporation
which is a function of temperature.
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Figure 6.10 Plot of ln(COND) against mean annual temperature (MAT). Eight out of ten lakes from
Western Victoria are considered saline. Lake Fyans (LFY) and Nuggetty Gully Reservoir (NGR)
were the only lakes that had a specific conductance value < 500 μs/cm from this region. Lake order
effect and residence time are potentially affecting the observed relationship between temperature
and specific conductance and the overall pattern is presumably due to that regional evaporation is a
function of temperature
In summary, high specific conductance obscures any relationship between δ2H of Chironomus spp.
and MAT but for freshwater lakes there is a strong correlation obtained between δ2H of Chironomus
spp. and MAT (r2 = 0.72 (Fig. 6.11)). Since specific conductance is closely related to lake
eutrophication and salinization (Chapter 3), temperature values can only be derived from
oligotrophic and mesotrophic lakes. These lakes are often found in the cold temperate and high
altitude areas in southeastern Australia and are the same lakes as are suitable for δ18
O temperature
estimates.
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Figure 6.11 Eight lakes that had a ln(COND) < ~ 6 were plotted for mean annual temperature
against δ2H of Chironomus spp.. A close correlation was obtained with an r
2 of 0.72. These include
two lakes (Lake Fyans and Nuggetty Gully Reservoir) from western Victoria and six lakes from
non-arid environments. These two western Victorian lakes are freshwater bodies presumably due to
short residence times.
6.7 Conclusions and future work
Stable isotopes δ18
O and δ2H analysed on head capsules of the chironomid genus Chironomus spp.
from sixteen southeastern Australian lakes suggested that the following aspects need to be
considered when applying them as temperature proxies. For δ18
O, regional aridity and its
consequential issues are critical. For lakes that are located in semi-arid and sub-humid areas,
residence time adds to the evaporative enrichment effect and this results in a high concentration of
nutrients that create low oxygen environments. Haemoglobin helps regulate the oxygen level in
Chironomus and allows this genus to dominate in these low oxygen water bodies but their chitin
δ18
O cannot be used to reconstruct temperatures from these lakes. In contrast, for lakes in humid
settings and where nutrient concentrations are low, the relationship between δ18
O of Chironomus
and MAT appears robust.
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Although, the δ2H values obtained on Chironomus spp. head capsules in this study were not
calibrated for the contribution of exchangeable hydrogen and the absolute values are not reliable,
the trends in the data are robust and yield a similar story to δ18
O. For fresh water lakes, the δ2H
values of Chironomus spp. head capsules showed that it is a promising proxy for MAT
reconstructions. Stable hydrogen isotope (δ2H) values of Chironomus spp. have the apparent
advantage that they are less influenced by insect biological controls (i.e. haemoglobin effects). It
also appears that the δ2H of Chironomus spp. values are less strongly affected by evaporative
enrichment. Since salinity and eutrophication are closely related in these Australian lakes (Chapter
3), the overall findings of this paper are that both δ18
O and δ2H are potentially valuable tools for
reconstructing temperature in low nutrient lakes in humid parts of Australia (e.g. Tasmania and the
southeastern highlands).
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7 CONCLUSIONS AND FUTURE WORK
7.1 Summary of research and major findings
This thesis aimed to develop and apply two methods that use subfossil remains of non-biting midge
larvae of chironomids (Diptera: Chironomidae) preserved in lake sediments to reconstruct past
changes in the Australian climate system during and since the Last Glacial Maximum (LGM).
For the first method, a model (transfer function) for reconstructing past summer temperatures based
on the temperature tolerance of chironomid species living in modern eastern Australian lakes was
created. This involved the physio-chemical investigation of 45 water bodies from tropical
Queensland to Tasmania. The transfer function was developed based on 33 water bodies from
southeastern Australia (the eleven tropical lakes are not included). It was applied to data derived
from chironomid remains extracted from lake sediment deposits from Welsby Lagoon, a subtropical
site in North Stradbroke Island, Australia spanning the last LGM and the last deglaciation. A
quantitative reconstruction was produced and this has provided the first quasi-continuous for the
period between 23.2 – 15.5 cal ka BP, detailed seasonal temperature record from mainland
terrestrial Australia covering the critical periods in the late Quaternary.
After the first method was established and applied, a second method which was based on the stable
oxygen and hydrogen isotope composition (δ18
O, δ2H) of the chitinous heads from chironomids
from the same lakes was first explored for the Australasian region. The potential of applying stable
isotopes δ18
O and δ2H analysed on a single genus of chironomid head capsules (Chironomus spp.)
from southeastern Australian lakes as proxies to reconstruct paleo-temperature was investigated.
The applicability and the complexities of using this method as independent temperature proxies in
Australia were discussed.
7.1.1 Connections of modern limnology to climate and land-use
This thesis has first provided an overview of the current status of 45 water bodies and further
investigated the interactions and connections of limnology, climate and land-use of these natural
and artificial water bodies extending across the eastern coast of Australia from the tropics of
Queensland to Tasmania. This research has been presented in Chapter 3 (Objective 1) in the form a
published paper (Chang et al., 2014). The paper comprises a literature review of the past
limnological studies on Australian lakes. A detailed description of the climate, vegetation and
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geology of the study area has also been provided. Measurements of a broad variety of physio-
chemical, land-use and climatic parameters were obtained using field, laboratory and spatial
techniques. Multivariate statistical analyses, notably Principle Component Analyses (PCAs) and
Redundancy Analyses (RDAs) were applied to explore the relationships and inter-correlations of
the measured variables.
The major findings from this chapter were 1) that most mainland Australian lakes are naturally
moderately to significantly eutrophic. Oligotrophic lakes are limited to high alpine and perennially
wet settings. 2) Reservoirs and other artificial lakes behave similarly to natural lakes, except that
they tend to be less eutrophic. This is most likely due to catchment management and a shorter
residence time. 3) The primary source of the salts in east Australian lakes is seawater, either derived
directly from aerosols or indirectly via marine bedrock. 4) Significant number of lakes in this
dataset are limited by nitrogen (N) and phosphorus (P), suggesting that both of these nutrients
should be considered in addressing eutrophication in Australian lakes. 5) Mainland lakes are
susceptible to becoming highly alkaline as a result of long-term increases in salinity that are related
to the concentration of cations during drought periods. Finally, a testable hypothesis was proposed
stating that climatically related eutrophication is more significant than nutrient loading by
agriculture in the eastern Australian lakes that were examined in this study. This is particularly
likely for shallow endorheic basins in semi-arid regions.
The results and findings obtained from this chapter were critical for the development of a
chironomid (Diptera: Chironomidae) based transfer function (Chapter 4, Objective 2) and the
interpretation of a paleo-record (Chapter 5, Objective 3). This is because understanding the
characteristics and the nexus of the geology, climate, water chemistry and the major environmental
controls on modern lakes is essential to identify the primary driving variables of the changes in
chironomid species assemblages and the species distribution in a region for both temporal and
spatial applications. This chapter has also provided valuable information to assist the understanding
of the stable isotope analyses results obtained on chironomid head capsules (Chapter 6, Objective
4).
7.1.2 The transfer function
A chironomid based temperature transfer function developed from a training set of 33 natural and
artificial lakes from southeast Australia spanning the subtropics to the alpine zone has been
presented in Chapter 4 (Objective 2 – Chang et al., 2015a). Eleven lakes from the tropics were not
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included because this thesis is focussed on climate reconstructions from southeastern Australia.
Previously published transfer functions were reviewed from other parts of the globe and a summary
table has been provided in Appendix 1. Chironomid taxonomic analyses were performed for each of
the sampled lakes and a list of taxa identified with corresponding pictures was discussed and shown
in Chapter 4 and Appendix 2 respectively. An ecological note is accompanied with the pictures.
Multivariate statistical analyses, canonical correspondence analysis (CCA) and partial CCA were
used to study the distribution of chironomids in relation to the environmental and climatic variables.
The results showed seven out of eighteen available variables were significant (p < 0.05) with
respect to chironomid species variation. These were mean February temperature (9.5%), pH (9.5%),
specific conductance (8.2%), total phosphorus (8%), potential evapotranspiration (8%), chlorophyll
a (6.9%) and water depth (6.2%). Though the total explained variance was low, Mean February
temperature (MFT) was found to be the most robust and independent variable explaining
chironomid species variation after further pCCA analyses.
A MFT transfer function was then constructed based on the modern distribution of chironomids
species in southeast Australia. The best MFT transfer function was a partial least squares (PLS)
model with a coefficient of determination (r2
jackknifed) of 0.69, a root mean squared error of
prediction (RMSEP) of 2.33 °C, and maximum bias of 2.15 °C. Compared with transfer function
models developed from other parts of the world, this transfer function has comparable r2
(Jackknifed)
but a relatively higher RMSEP. This reflects the long scalar length of the temperature gradient
which extends from sub-tropical to sub-alpine locations, some 14 °C. The RMSEP represents 16%
of the scalar length and this is comparable with most other transfer functions (e.g. Potito et al.,
2014). It has been observed that data sets with large temperature gradients naturally have larger
errors (Walker and Cwynar, 2006) but this in no way diminishes the value of the reconstruction.
The main findings from this chapter were that 1) chironomids assemblages in actively managed
reservoirs (non-impacted) show no significant difference to natural lakes in the same climate and
vegetation zones, and therefore can be included for transfer function development. 2) Although we
cannot completely rule out some degree of endemism in the Tasmanian chironomid fauna, our
analyses show that the degree of endemism indicated by earlier studies is greatly reduced. This
raises the real possibility of integrating the existing chironomid transfer function for Tasmania
(Rees et al., 2008) with this new model for the southeastern Australian mainland. 3) The transfer
function is appropriate (e.g. the RMSEP is relatively too to be used for the late Holocene period) for
the reconstruction of summer temperatures during the last glacial maximum (LGM) and the late
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glacial to Holocene transition in eastern Australia. The transfer model was then applied to an LGM
and deglacial record.
7.1.3 The application of the transfer function: temperature reconstruction covering
the Last Glacial Maximum and the last deglaciation from subtropical Australia
A summer temperature reconstruction based on the newly developed chironomid transfer function
from southeastern Australia in Chapter 4 (Objective 2) has been presented from Welsby Lagoon of
North Stradbroke Island in Chapter 5 (Objective 3 – Chang et al., 2015b). This is the first
quantitative and detailed seasonal reconstruction of the LGM and the last deglacial temperatures
from terrestrial Australia. A review of the past inferences of the LGM temperatures in southeastern
Australia was provided in the chapter. Fossil chironomid taxonomic analyses from the LGM to the
present were performed throughout the 450 cm sediment core, with 13 samples concentrated
between the periods of 23.2 – 15.5 cal ka BP. Each of the identified fossil taxa was listed and
photographed and the pictures have been provided in Chapter 4 and Appendix 3 respectively.
Sediment samples that generated sufficient chironomid head capsules for quantitative temperature
reconstructions detailed the period between 23.2 and 15.5 cal ka BP. Several reconstruction
diagnostics, including goodness-of-fit, modern analogue technique (MAT) and rare taxa analysis
were applied to evaluate the reliability of the reconstruction. Trajectory diagrams were also
constructed to show if the changes of chironomid assemblages in the fossil samples were driven by
variables other than temperature through time.
The major findings from this chapter were 1) the temperature reconstruction from the LGM to the
deglacial period of this site is sensible. 2) Factors that may affect the reliability of the inferred
temperatures include changes in lake chemistry and lake level. These had an impact on chironomid
assemblages, particularly in the Holocene. These Holocene samples are dominated by a temperature
insensitive taxon, Dicrotendipes. 3) The relatively small modern calibration training set also limits
the ability to fully understand the environmental controls on some of the fossil samples. 4) The
overall reconstruction displays similarities to marine records from areas affected by the East
Australian current (EAC), both in terms of timing and magnitudes of cooling during and
immediately after the LGM. 5) A maximum summer temperature cooling of c. ~6.5°C at the late
LGM at c. ~18.5 cal ka BP is shown, followed by rapid warming to (near) Holocene values by 17.3
cal ka BP. 6) the warming is consistent with the start of deglaciation at 18.2 ± 0.7 cal ka BP derived
from ice core records in Antarctica. 7) Results suggest that this coastal record resembles the thermal
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history of adjacent marine sites and the Antarctic climate influence reached the Australian
subtropics during the LGM and the last deglaciation.
This long term temperature record has filled a critical gap in our current knowledge of temperature
changes during the glacial times in the subtropics and the connections to changes of the ocean
surface current and the higher latitudes. The transfer function method developed in Chapter 4
(Objective 2) has been confirmed as a valuable tool to quantify terrestrial temperatures from eastern
Australia during glacial times. However, since there have been very limited independent proxies
that can be used for quantitative temperature reconstructions extending back to the glacial times in
Australia, a second method based on the stable isotope measurements of chironomid head capsules
was explored in Chapter 6 (Objective 4).
7.1.4 The potential of applying stable oxygen and hydrogen isotopes from Australian
subfossil chironomid head capsules as independent proxies for past change
Stable isotopes (δ18
O and δ2H) analysed from head capsules of the chironomid genus Chironomus
spp. from sixteen southeastern Australian lakes were measured and this work has been included in
Chapter 6 (Objective 4 – manuscript submitted). This is the first use worldwide of both δ18
O and
δ2H instantaneously from the same samples using a single genus of chironomid head capsules. It is
also the first exploration of the potential of their application to reconstruct paleo-temperature in the
Australasian region. The relationship between the stable isotope composition (δ18
O, δ2H) of
Chironomus spp. head capsules and how this relates to the composition of lake water and local
precipitation was investigated. The study also inspected whether fractionation or vital effects were
constant or varied with changes in the environment. Regression and redundancy analyses were
applied to examine the relationships of the measured variables.
Results suggested that the following aspects need to be considered when applying them as
temperature proxies. 1) For δ18
O, regional aridity and its consequential issues are critical. For lakes
that are located in semi-arid and sub-humid areas, residence time adds to the evaporative
enrichment effect and this results in a high concentration of nutrients that create low oxygen
environments. 2) Haemoglobin helps regulate the oxygen level in Chironomus spp. and allows this
taxon to dominate in low oxygen water bodies but their chitin δ18
O cannot be used to reconstruct
temperatures from these lakes. 3) For lakes in humid settings and where nutrient concentrations are
low, the relationship between δ18
O of Chironomus spp. and mean annual temperature (MAT)
appears robust. 4) The δ2H values obtained on Chironomus spp. head capsules in this study were
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not calibrated for the contribution of exchangeable hydrogen and the absolute values are not
reliable, but the trends in the data are robust and yield a similar story to δ18
O. 5) For fresh water
lakes (specific conductance measurements of < 500 μs/cm), the δ2H values of Chironomus spp.
head capsules showed that it is a promising proxy for MAT reconstructions. 6) Stable hydrogen
isotope (δ2H) values of Chironomus spp. have the apparent advantage that they are less influenced
by insect biological controls (i.e. haemoglobin effects) and it also appears that the δ2H of
Chironomus spp. values are less strongly affected by evaporative enrichment. 7) The overall
findings of this study are that both δ18
O and δ2H are potentially valuable tools for reconstructing
temperature in low nutrient lakes in humid parts of Australia (e.g. Tasmania and the southeastern
highlands) and this is because, as it has been concluded in Chapter 3 of the thesis, salinity and
eutrophication are closely related in the Australian lakes dataset.
7.2 Research significance
In this thesis, valuable tools to quantify past climate in Australia were explored and established. It
demonstrated that the transfer function can be applied to the late Quaternary data to generate
quantitative summer temperature reconstructions in eastern Australia. The chironomid-based
temperature record from Welsby Lagoon showed cooling and warming trends similar to those
derived from nearby marine records and that deglaciation is synchronous with the records from
Antarctica. This suggests a climate link from terrestrial Australian subtropics to the nearby ocean
and hence to the high latitudes. It has demonstrated that critical gaps in our knowledge on climate
change during the glacial period from Australia can be addressed by applying the chironomid-based
transfer function developed in this thesis. Stable oxygen and hydrogen isotopes were measured on
chironomid head capsules and showed the potential of using chironomid δ18
O and δ2H as proxies
for temperature reconstructions. The application will be likely limited to non-arid and non-impacted
areas of Australia.
The Welsby Lagoon record has also shown that chironomid-based summer temperature
reconstructions from a coastal site can contribute to establishing links among oceanic, terrestrial,
atmospheric, polar and insolation driven changes in temperature. This has further advanced our
knowledge on the hemispheric and global climate teleconnections.
7.3 Critical reflections
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7.3.1 The debate and controversy about transfer function and its application
Recently, the debate about whether chironomid-based transfer functions are reliable tools to
quantify past temperatures has been re-opened. One of the key arguments is that the transfer
function method simplifies the complexity of the real world by maximizing single ecological
gradients, by not taking into account co-varying variables (Velle et al., 2010). This is typical of
empirical or mathematical models ranging from the field of engineering, through ecology to the
social sciences. The basic principle in developing a quantatitive method to model real world
complexity is the need to minimize the degrees of freedom and this is the key issue regarding the
appropriate use of the technique. Velle et al. (2010) raises the problem related to confounding factor
(such as temperature may interact with trophic status, water depth etc.). In reply, to Velle et al.
(2010), Brooks et al. (2012) argue in many cases, temperature is an independent variable. In
Chapter 3 of this thesis, a partial Canonical Correspondence Analysis (pCCA) was applied to the
modern chironomid calibration data and results show that mean Febuary temperature (MFT) is not
confounded by other variables.
Velle et al. (2010) also argues that one of the main reasons for the strong performance of the
chironomid-based transfer function models is because the temperature gradient is maximised in the
calibration data set and the gradients of other environmental variables are minimised. This is true in
many studies (e.g. Diffenbacher-Kraal et al., 2007). This is not true for the data set (Chapter 3)
developed in this thesis. Due to the limited number of lakes in Australia, manipulation of a
temperature gradient is unachievable. Temperature, trophic status, salinity and pH all have a large
range of values. This calibration data set suggests, in line with evidence as argued in Brooks et al.
(2012), that temperature is a particularly significant variable in driving the composition of
chironomid assemblages.
In addition to Velle et al. (2010; 2012), many other studies and reviews (e.g. Hann et al., 1992;
Walker and Matthewes, 1989; Warwick, 1989; Brooks and Birks, 2001) in the literature critically
analysed the application of the chironomid-based transfer function method to the reconstruction of
past temperatures. There are several limitations: first, the environmental preferences of one or more
taxa might change on geological timescales. Second, different taxa might bloom during different
times of the year (although the prime season for chironomids blooming is summer in temperate and
polar climate zones), and this may also change due to evolutionary pressures. Third, taxa
assemblages could be subjected to some confounding effects such as that water depth, temperature
and oxygen concentration can be correlated and the structure of the correlation changes on
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geological timescales. These can result in reconstructions from adjacent sites showing inconsistent
patterns regarding inferred temperature changes, such as the Holocene records from Scandinavia
discussed in Velle et al. (2010).
As recognized in both Velle et al. (2010; 2012) and Brooks et al. (2012), the possible solutions are
as follows: sites for downcore records should be carefully selected, chironomid-based temperature
reconstruction should be undertaken within a multiproxy framework, and fossil assemblages should
be carefully evaluated for modern analogues and tested against random data (Telford and Birks,
2011). Reconstruction diagnostics were adequately applied for the application to Welsby Lagoon
data in Chapter 5. Multiproxy approaches are preferable but not easily available. Diatoms are absent
from my sites so I compared and cross-validated our reconstruction with the pollen record from the
same site and temperature records based on other independent proxies from the region. This is less
desirable as pollen is not strongly correlated with temperature in subtropical Australia (Moss et al.,
2013) and with the exception of aquatics represents regional terrestrial variation rather than changes
within the lake basin.
Despite these caveats, the transfer function technique has been a cornerstone of biologically based
paleoclimatological reconstruction for about thirty to forty years. Palaeoecologists and
paleolimnologists should not dismiss this useful method, but rather should treat the reconstructions
with due caution and continue to refine the understanding of the proxies (such as chironomids) (e.g.
Brooks et al., 2012; Velle et al., 2012).
7.3.2 The limitation of the stable isotope method
A number of limitations of the stable isotope method were discussed in Chapter 6. These include
first, the method is only useful for reconstructing temperatures in low nutrient lakes in humid parts
of Australia (e.g. Tasmania and the southeastern highlands). Second, the analyses of stable isotopes
currently requires a minimum dry weight of 100 μg (equivalent to 150 head capsules), and this has
limited the application to some parts of the record from palaeo-sites as chironomids are simply not
common enough to collect this mass of a single taxon. Third, similar to the application of the
transfer function method, it is challenging to recognize and validate what parts of a chironomid-
based stable isotope record are likely to be reliable.
Stable isotope reconstruction should be undertaken within a multiproxy framework. Chironomid
taxa should be identified from the same part of the record and the transfer function method can be
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applied as a cross-validation for the stable isotope record. Alternatively, other independent proxies
such as pollen which could provide information about positive/negative moisture balance, diatoms
which could provide information about salinity changes and other groups such as zooplankton
which could provide information about trophic status will help to validate the temperature
reconstructions based on chironomid derived stable isotope records.
7.4 Recommendations and future work
7.4.1 Extension and integration of the modern calibration dataset in Australia
In Chapter 4 of this thesis, previously claimed endemism for Tasmanian chironomid taxa was
assessed and results showed that the degree of endemism indicated by earlier studies is much less
than expected, although some degree of endemism in the Tasmanian chironomid fauna cannot be
completely ruled out. Very recent studies using molecular markers (Cranston and Krosch 2015;
Krosch et al., 2015) have demonstrated that the previously suspected endemic Tasmanian
chironomid fauna are not endemic. On the mainland, the previously assumed endemic Tasmanian
taxa are present at several high elevation sites including lakes on Mount Kosciuszko. Therefore, we
can be more confident about integrating the existing chironomid transfer function for Tasmania
with this new model for the southeastern Australian mainland.
In the future, the integration of the lakes incorporated in this thesis with the Rees et al. (2008)
Tasmania dataset is planned. This will increase the modern chironomid calibration set from eastern
Australia to ~100 lakes. The major advantages of including a large number of modern calibration
sites are that it can be expected that most chironomid taxa encountered in the late Pleistocene and
Holocene lake sediment records from both lowland and mountainous sites are well-represented.
Consequently, it will be more widely applicable, both temporally and geographically. This also has
the potential to help with improving the statistical performance of the robustness (model
coefficient) and accuracy (root mean squared error in prediction, RMSEP) of the transfer function
model, which will then further constrain the absolute scale of past temperature variations. Well-
constrained quantitative temperature values reconstructed from the glacial times will be highly
valuable as validation data for palaeoclimate modelling.
To encourage the wider application of the chironomid-based techniques in Australia, a photographic
subfossil chironomid identification guide will be developed for Australia. Pictures of the identified
subfossil chironomid taxa used in this thesis have been provided in Appendix 2 and Appendix 3.
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These materials will be merged with the chironomid taxa identified and photographed from
Tasmanian lakes in Dr Andrew Rees’s PhD thesis (Rees 2013). The eleven tropical lakes that were
sampled and included in Chapter 3 will also be encompassed in the future subfossil chironomid
work.
7.4.2 Application of the transfer function to quantify long-term (late-Pleistocene to
Holocene) temperature changes from various sites
The 450 cm sediment core taken from Welsby Lagoon analysed in this thesis dated back to c. ~ 24
cal ka BP. During recent field work, much longer sediment cores have been recovered from this site
and the record of this site has the potential of extending through the last glacial cycle. If there are
sufficient chironomids throughout the older sections of the record, the current transfer function will
be applied to reconstruct temperature changes of the earlier glacial times.
Early palaeoecological and palaeoclimate investigations of chironomids from the Southern
Hemisphere were restricted to qualitative and semi-quantitative interpretations because the
numerical techniques for creating transfer functions were still in an early stage of development. In
Australia, two previous chironomid species records from Lake Barrine, Atherton Tableland and
Blue Lake, Mount Kosciuszko (Dimitriadis and Cranston, 2001; Schakau 1993 respectively) have
demonstrated the applicability of subfossil chironomids as a proxy for unravelling the regional
paleoclimate history from tropical to alpine climate respectively. With the transfer function
established in this thesis, both of these sites can be revisited and the transfer function can be applied
to the chironomid data provided in the published work. Quantitative reconstructions can be
developed and this will provide valuable information to understand climate changes and to identify
abrupt climate events (if there are any) in the late glacial to the Holocene from these two very
different climate zones of Australia. A two-metre sediment core was recovered from Lake
Cootaptamba, Mount Kosciuszko in December 2014, and radiocarbon dates will be obtained from
this core. Chironomid analysis will be performed under a recently funded project (ANLGRA15008,
in collaboration with CI: Shulmeister, J), to reconstruct the climate history from the site. Other
potential sites that the transfer function could be applied to reconstruct the climate history extend
back to the last glacial maximum (LGM) are Mountain Lagoon, Blue Mountains (New South
Wales), Lake Frome, South Australia (in collaboration with Cohen, T) and Hazards Lagoon, eastern
Tasmania (in collaboration with Mackenzie, L and Moss, P). Sediment cores of the above sites were
obtained from various working groups and have established radiocarbon age chronology.
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7.4.3 Further exploration and application of the chironomid δ18
O and δ2H method
In Chapter 6 (Objective 4) of this thesis, the overall conclusion made was that both δ18
O and δ2H
are potentially valuable tools for reconstructing temperature in low nutrient lakes in humid parts of
Australia (e.g. Tasmania and the southeastern highlands). To further develop and test this method,
future studies should be conducted for lakes located in regions that have similar environmental and
climatic conditions to southeastern Australia, such as some areas in South America, as potential
evapo-transpiration (PET) plays a key role in the application of the stable isotope methods. The
method will be better understood when it is tested in both arid (Precipitation < PET) and non-arid
(Precipitation > PET) conditions. In this thesis it was concluded that δ18
O of Chironomus spp. and
mean annual temperature (MAT) are strongly correlated (r2 = 0.78) and the relationship obtained in
the nutrient low southeastern lakes were very similar to Verbruggen et al.’s (2011) European
dataset, so it is reasonable to apply the method to Mount Kosciuszko or Tasmanian sites. Specific
potential sites for isotope work include Lake Cootaptamba and Eagle Tarn (Rees and Cwynar,
2010), as the transfer function application has been (or will be) established from these lakes. In the
future, the stable isotopes method can be validated for temperature reconstructions, in adjunct with
applying the transfer function method. On the other hand, the collection of enough material from
paleo-sites is challenging and the use of stable isotopes is recommended for late Holocene and
modern studies, perhaps focussed on nutrient changes rather than for paleo-temperature
reconstructions.
7.4.4 Regional patterns and hemispheric to global connections
Many literature reviews have addressed the fact that there are inadequate quantitative and multi-
proxy based paleo-temperature data in the Southern Hemisphere, for us to gain a systematic
understanding of the past climate change over the last 20,000 years. This has hindered us from
advancing our knowledge of hemispheric and global climate teleconnections. Within the
interpretation of the limited paleoecological records, there are divergent views concerning climate
dynamics during key periods of large-scale climate fluctuations, such as the Younger Dryas (YD)
and Antarctic Cold Reversal (ACR). The disagreement stems from the disparities among the
terrestrial records based on pollen, beetles, chironomids and glaciers from the Southern
Hemisphere. This discrepancy may be due to different proxy responses to temperature changes,
chronological issues or sampling resolution. However, these inconsistencies could also reflect the
actual regional climate variability. For instance, there could be differential influences of the
Southern Hemisphere Westerly Winds on the east or west side of the Andes, New Zealand and
Tasmania, at similar latitudes.
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The future constructed chironomid-based high resolution, detailed quantitative temperature records
of the same region and regions from other parts of Southern Hemisphere can be compared and
synthesized to reveal if there are regional and hemispheric coherent patterns. An inter-proxies
approach, such as using chironomid data in conjunction with the pollen and glacial records, could
then be used to elucidate the regional and hemispheric paleoclimate conditions (e.g. seasonality and
rainfall). This has been carried out in New Zealand for the INTIMATE project (e.g. Alloway et al.,
2007) but is not yet possible at a high level of resolution for Australia (e.g. Reeves et al., 2013) and
is definitely worth working towards.
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Appendices
168 | P a g e
Appendices
APPENDIX 1. A SUMMARY OF PREVIOUS MODERN
CALIBRATION TRAINING SETS DEVELOPED AND
APPLIED FOR PALEO-RECONSTRUCTIONS USING
CHIRONOMIDS
Page 200
Appendix 1. Review of past training set
169 | P a g e
Region No. of Lakes Application Other Correlations Number of
Taxa
Model
coefficient
( r2)
Authors and
Year of
publication
Europe
Across north-
western Finland
30 lakes used
in lake surface
water
temperature
transfer
function
Expanded to
53 lakes for
July air
temperature
model
Lake water temperature
transfer function
developed based on
chironomid assemblage
(Olander et al., 1997)
Expanded calibration
model for
inferring lake water and
July air temperatures
from fossil chironomid
assemblages is developed
(Olander et al., 1999)
Maximum lake depth is
strongly correlated (only
limited environmental
variables were considered
in Olander et al., 1997).
Sediment organic content,
maximum lake depth, and
lake water temperature are
the most important
explanatory variables
(Olander et al., 1999)
32 identified and
20 taxa used in
the lake water
temperature
transfer function
Extended to 38
taxa included for
July air
temperature
model
r2= 0.5
(Jackknifing)
RMSEP = 1.5–
1.6°C
(Olander et al.,
1997, Olander
et al., 1999)
Subarctic region
of northern
Sweden
100 Mean July air temperature
transfer function was
developed based on
chironomid assemblage
LOI, maximum lake depth
and mean January air
temperature are also
strongly correlated
85
42 taxa used in
the model
r2= 0.65
(Jackknifing)
RMSEP =
1.13°C
(Larocque et
al., 2001)
Greenland 43 Statistical analysis were
used to relate chironomid
taxa to the environmental
variables
The trophic variables
(total nitrogen and total
phosphorus (TN, TP))and
Temperature
24 - (Brodersen and
Anderson,
2002)
Switzerland
(Swiss Alps)
81 A July air temperature
inference model (transfer
function) was developed
based on chironomid
assemblage
July air temperature is the
only constraining variable
76
50 taxa included
in the model
r2= 0.81
(Bootstrapping)
RMSEP =
1.51°C
(Heiri et al.,
2003)
Iceland 50 Mean July air temperature
transfer function
developed by chironomid
assemblage
The chironomid
assemblages is influenced
by Lake substrate (%TC)
54
53 taxa used in
the model
r2=0.66
(Jackknifing)
RMSEP =
1.095°C
(Langdon et al.,
2008)
(Holmes et al.,
2009).
Page 201
Appendix 1. Review of past training set
170 | P a g e
Region No. of Lakes Application Other Correlations Number of
Taxa
Model
coefficient
( r2)
Authors and
Year of
publication
Finland 77 Mean July air temperature
inference model was
developed based on
chironomid assemblage
Sampling depth, dissolved
oxygen and conductivity
are relatively significant
110
72 taxa used in
the model
r2=0.78
(Jackknifing)
RMSEP =
0.721°C
(Luoto, 2009)
Sweden,
Denmark,
Germany, the
Netherlands,
Austria,
Switzerland,
France and Italy
31 Relationships of
temperatures, trophic
state, and oxygen
between chironomid
assemblages.
Stable oxygen isotopes in
chironomid head capsules
is measured in all sites
and a relationship was
developed between
temperature and stable
oxygen isotopic
composition of
chironomid head
capsules.
Summer surface water
and July air temperature,
as well as total
phosphorus (TP)
concentrations,
hypolimnetic oxygen
availability and
conductivity were found
statistically significant
106 - (Verbruggen et
al., 2011a)
Europe
(Norway and
Switzerland)
274
(Norway:153,
Switzerland:
102)
Mean July air temperature
models were developed
for both Norwegian and
Swiss calibration data sets
respectively, as well as a
combined model of these
two data sets.
Larger range of both lake
types and pH are included
131 taxa found
in the Norwegian
lakes, 120 taxa
found in Swiss
Alps
154 taxa
included
r2= 0.87
(Bootstrapping)
RMSEP = 1.4°C
(Heiri et al.,
2011)
(Largest
calibration set
so far)
North western Asia
Tibetan Plateau
(North-western
China)
42 A chironomid-based
salinity inference model
was built.
Salinity is the most
significant
30
24 taxa included
r2 = 0.80
(Jackknifing)
RMSEP = 0.29
log10 TDS
(Zhang et al.,
2007)
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Appendix 1. Review of past training set
171 | P a g e
Region No. of Lakes Application Other Correlations Number of
Taxa
Model
coefficient
( r2)
Authors and
Year of
publication
North America
South eastern
Ontario
50 Transfer function for the
hypolimnetic oxygen
conditions of lakes, which
measured as average end-
of-summer hypolimnetic
dissolved oxygen
(AvgDO(Summ) ) was
constructed based on
chironomid assemblages.
Dissolved inorganic
carbon, the Anoxic
Factor, maximum depth
and total phosphorus
concentrations were also
correlated with
assemblage composition
67
45 taxa included
in the transfer
function
r2 = 0.58
(Jackknifing)
RMSEP = 0.032
log(x + 19.88)
AvgDO(summ)
(Little and
Smol, 2001)
Central, eastern
Sierra Nevada of
California
57 A taxon response model
was used to assess the
statistical relationship of
each taxon to surface water
temperature. Quantitative
transfer functions were
developed to estimate
surface water temperature
from the chironomid
assemblages.
Surface water
temperature, elevation,
depth, strontium,
particulate organic carbon
is statistically significant
variables.
68
44 taxa included
in the transfer
function
- (Porinchu et al.,
2002)
Canadian Arctic
Islands
50 Chironomid assemblage
was developed in these
lakes and statistical models
were used to relate taxa to
the environment.
lake morphometry, pH,
nutrients (TP) and
temperature are identified
as statistical important
factors
32 - (Gajewski et
al., 2005)
Quebec, Canada 52 Mean August air
temperature transfer
function was developed
based on chironomid
assemblage
Lake depth, dissolved
organic carbon (DOC),
are significant also
variables.
97
64 taxa included
in the model
development
r2=0.67
(Jackknifing)
RMSEP =
1.17°C
Larocque, 2006
Page 203
Appendix 1. Review of past training set
172 | P a g e
Region No. of Lakes Application Other Correlations Number of
Taxa
Model
coefficient
( r2)
Authors and
Year of
publication
Eastern North
America
(Yukon/Northwest
Territories, British
Columbia and the
Canadian
Arctic Islands)
145 Chironomid taxa
assemblage was developed
to the training set. Midge
inference models for mean
July air temperature and
transformed depth were
developed.
Lake depth, arctic tundra
vegetation, alpine tundra
vegetation, pH, dissolved
organic carbon, lichen
woodland vegetation and
surface area are all
significant to midge
distribution
80
62 taxa included
in the model
development
r2=0.82
(Bootstrapping)
RMSEP =
1.46°C
(Barley et al.,
2006)
*Expanded
results of
Gajewski et al.,
(2005)
Central Canadian
Arctic
88 Quantitative midge based
inference model was
developed for average July
air temperature and
summer surface water
temperature.
Maximum depth, pH, total
nitrogen-unfiltered (TN-
UF), Cl and Al are
statistically significant
50 included in
the models
r2 = 0.77
(Jackknifing)
RMSEP =
1.03°C
(Porinchu et al.,
2009)
Eastern Canadian
Arctic
51 (Walker et
al., 1997)
63 (Brodersn
et al., 2008)
Statistical analysis were
used to relate chironomid
taxa assemblages to the
environmental variables
Temperature and trophic
status were found to
strongly influence the
chironomid assemblages
86
32
r2 = 0.88
(Jacknifing)
RMSEP =
2.26°C
(Walker et al.,
1997)
(Walker et al.,
1997;
Brodersen et
al., 2008,
Medeiros and
Quinlan, 2011)
Southern Hemisphere
New Zealand
(North and South
Island)
46 The first chironomid based
transfer function build in
the southern hemisphere. A
temperature inference
model was built with the
most productive lakes
removed and a Chlorophyll
a. transfer function based
on chironomid
assemblages was
Chlorophyll a is also
significantly correlated.
Conductivity, Total
Nitrogen (TN) and pH are
also statistically
significant.
50
Retention of all
species
r2 = 0.77
(Jackknifing)
RMSEP =
1.31°C
For temperature
model
r2 = 0.49
(Jackknifing)
RMSEP = 0.46
Log 10μg l-1
(Woodward
and
Shulmeister,
2006; 2007)
Page 204
Appendix 1. Review of past training set
173 | P a g e
presented. For Chlorophyll
a model
Region No. of Lakes Application Other Correlations Number of
Taxa
Model
coefficient
( r2)
Authors and
Year of
publication
New Zealand
(South Island)
60 Taxon response models
were constructed and a
mean summer air
temperature transfer
function was developed
Chlorophyll a,
conductivity, organic
content of sediment and
mean annual precipitation
are also significant
variables
49 r2 = 0.8
(Jackknifing)
RMSEP =
1.29°C
(Dieffenbacher-
Krall et al.,
2007;
Vandergoes et
al., 2008)
Tasmania
(Australia)
54 Taxon response models
were constructed and a
midge transfer function
was developed for
temperature of the warmest
quarter (TWARM) of
Tasmania.
pH, annual radiation,
magnesium, annual
precipitation, SiO2, and
depth are also significant
factors.
49 non-rare taxa
were used in
model
development
r2 = 0.72
(Jackknifing)
RMSEP = 0.94
(Rees et al.,
2008; Rees and
Cwynar, 2010)
Uganda and
Kenya (Africa)
65 Taxon response models
and transfer functions of
Mean Air Temperature
(MAT) and Surface Water
Temperature (SWT) were
developed.
Water depth, conductivity
(salinity) and pH are
important for EL-EM
calibration data set.
DOC, LOI of the surface
sediments and nutrients
(defined as TN, TP) are
important for the
Rwenzori-only Data set
(29 lakes).
81
68 taxa used in
the models
development
r2 = 0.81–0.97
(Jackknifing) for
Mean Air
Temperature
RMSEP = 0.61-
1.50°C
r2 = 0.77–0.95
(Jackknifing) for
Surface Water
Temperature
RMSEP = 1.39-
1.98°C
(Eggermont et
al., 2010)
Page 205
Appendix 1. Review of past training set
174 | P a g e
South America
(Argentina and
Chile)
46 Transfer function for
warmest months mean
temperature was developed
and applied
Secchi depth and
conductivity are also
significantly correlated
49 r2 = 0.56
(Jackknifing)
RMSEP =
1.69°C
(Massaferro
and Larocque-
Tobler, 2012;
Massaferro et
al., 2014)
Page 206
Appendix 2. Photographs of chironomid taxa identified from modern eastern Australian waterways
175 | P a g e
APPENDIX 2. CHIRONOMID TAXA ENUMERATED FROM
THE MODERN CALIBRATION DATA SET OF
SOUTHEASTERN AUSTRALIA
• Chironomid taxa listed from 1 to 38 correspond to taxa specified in Table 4.5,
Chapter 4. These are the taxa used in the South-eastern Australia transfer
function development.
• The ‘red box’ shows the key part of each taxa to be used for identification
Page 207
Appendix 2. Photographs of chironomid taxa identified from modern eastern Australian waterways
176 | P a g e
a. Nuggetty Gully Reservoir, Victoria, 20 X b. Lake Poona, QLD, 100 X
2. Polypedilum nubifer Skuse
b. Freshwater Lake, Victoria a. Lake Cootapatamba, Kosciuszko, NSW
1. Chironomus Meigen
Ecology: occurs in 94% of the lakes in the training set. It is abundant in both warm eutrophic
and cool oligotrophic lakes; mostly confined to the profundal but may also be present in the
littoral; tolerant of low oxygen concentrations and even anoxia; tolerant of low pH and high
salinity; often an early coloniser after significant environmental change where it may occur
in suboptimal conditions; detritivores and filter-feeders; associated with soft sediments
(Brooks et al., 2007). These descriptions match with the obervations from this training set.
Ecology: occurs in 82% of the lakes in the training set. It is often an indicator of temperate
climatic conditions. It occurs in the litteral of eutrophic lakes and some species occur
amongst vegetation (Brooks et al., 2007). It shows a strong correlation with temperature in
the southeastern Australian training set. However, some morphotypes are also found in
subtropical lakes.
Page 208
Appendix 2. Photographs of chironomid taxa identified from modern eastern Australian waterways
177 | P a g e
3. Cladopelma Kieffer
a. Lake Poona, QLD, 40 X b. Lake Cantani, Mt Buffalo, Victoria, 40 X
4. Dicrotendipes Kieffer
a. Nuggetty Gully Reservoir, Victoria, 20 X b. Lake Terangpom, Victoria, 40 X
Ecology: occurs in 70% of the lakes in the training set. It is a littoral taxon occurring in
muddy and sand/gravel substrates. It is often a warm stenotherm but not particularly tolerant
of high nutrient conditions, typical of mesotrophic lakes (Brooks et al., 2007). These
descriptions match the observations from the southeast Australian training set but it shows a
particularly strong correlation with water depth.
Ecology: occurs in 70% of the lakes in the training set. It occurs in the littoral of lentic
environments where it is often associated with macrophytes and mesotrophic to eutrophic
waters. Most of the training sets indicate that it is a warm indicator (Brooks et al., 2007).
However in this training set, it is more clearly as a eutrophic and shallow water indicator.
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5. Polypedilum spp. Kieffer
a. Thirlmere Lakes, NSW, 40 X
b. Lake Lila, Tasmania, 40 X
6. Cryptochironomus Kieffer
a. Lake Elingamite, Victoria, 20 X b. Lake Tooliorook, Victoria, 40 x
Ecology: it is a morphotype of Polypedilum, occurs in 18% of the lakes in the training set.
The ecology is not clearly known, it seems occur in a few reservoirs and a few natural lakes
with in and outflows in the current training set. It is possibly related to flowing water or
stream species.
Ecology: occurs in 24% of the lakes in the training set. It often occurs in the profundal, in
nutrient rich waters and apparently can be acidophilic. Larvae occur in a variety of substrates
in the littoral and sublittoral (Brooks et al., 2007). It shows a close relationship with
conductivity in the southeastern Australian training set.
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7. Parachironomus Lenz
a. Lake Fyans, Victoria, 40 X b. Lake Tooliorook, Victoria, 40 X
a. Freshwater Lake, Victoria, 20 X b. Lake Samuel, Tasmania, 20 X
8. Kiefferulus martini Freeman
Ecology: occurs in 33% of the lakes in the training set. It usually occurs in the littoral of
standing waters and also running water, some morphotypes are asscociated with the presence
of macrophytes. The genus is descrbed as acidophobic (Buskens 1987; Brooks et al., 2007).
It shows correlation with temperature, macrophytes, lake productivity in the southeastern
Australian training set. It shows as a warm temperature indicator.
Ecology: occurs in 50% of the lakes in the training set. Larvae of Kiefferulus inhabit
sediments of small water bodies. Three species of this genus is known from Australia
(Cranston, 2010). It shows correlation with macrophytes and water depth in the southeastern
Australian training set.
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9. Procladius Skuse
a. Lake Albina, Kosciusko, NSW, 40 X b. Freshwater Lake, Victoria, 40 X
10. Coelopynia pruinosa Freeman
a. Green Lake, Victoria, 10 X b. Grahamstown Lake, NSW, 40 X
Ecology: occurs in 97% of the lakes in the training set. There are many species and
morphotypes within this genus. The genus is abundant in mesotrophic and eutrophic lakes
but may be absent from very cold lakes. It does not tolerant anoxia conditions. It tolerates
acidic conditions (Brooks et al., 2007). These descriptions match the observations from the
southeastern Australian training set in general. It shows as a warm temperature indicator in
the training set.
Ecology: occurs in 25% of the lakes in the training set. Larvae are usually found in deep
lakes (e.g. in Tasmania), billabongs in Australia, and in both standing and slow flowing
waters (Northern Territory and Western Australia) (Cranston, 2010). It shows correlation
with temperature in the southeastern Australian training set and appears as a warm
stenotherm.
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a. Eagle Tarn, Tasmania, 20 X
11. Pentaneurini undifferentiated
b. Lake Hiawatha, NSW, 20 X
Ecology: ocurrs in 50% of the lakes in the training set. Cranston (2010) divided the genus to
A-E. It has a wide range of distribution in Australia, from monsoonal tropics to Tasmania
and west to the east. The ecology of its distribution is not well identified. In the southeastern
Australian training set, it shows a correlation with conductivity.
a. sp. 1 - Grahamstown Lake, NSW, 40 X b. sp. 2 – Grahamstown Lake, NSW, 40 X
12. Riethia Kieffer
Ecology: occurs in 64% of the lakes in the training set. In Australasia Riethia larvae are
found in depositional areas in low order streams, often naturally shaded by riparian
vegetation. Records range from New Zealand’s temperate North Island, southern New
Caledonia, Tasmania and along the eastern margin of Australia to monsoonal tropical
Northern Territory, and in temperate southern Western Australia. Some species occur in
habitats characterised by high amounts of fine particle organic matter in both lotic and
lenthic systems (Cranston, 2010). It occurs in most of the reservoirs in the training set. It
shows a correlation with conductivity in southeastern Australian lakes.
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14. Tanytarsus Pallidicornis type
a. Lake Poona, QLD, 40 X b. Nuggetty Gully Reservoir, Victoria, 40 X
13. Tanytarsus lugens type
a. Lake Cantani, Mt Buffalo, Victoria, 20 X b. Green Lake, Victoria, 40 X
Ecology: it occurs in 85% of the lakes. The lugens type is a cold stenotherm and usually
occurs in the profundal of oligotrophic lakes or in the littoral of cold subalpine and subarctic
lakes (Brooks et al., 2007). The morphotype appears in the southeastern Australian lakes
shows close features to the lugens type, but it does not show clear correlations with any
particular environmental variables.
Ecology: occurs in 80% of the lakes in the training set. Most of the Tanytarsus morphotypes
usually occur in the littoral of relatively warm, productive lakes and will tolerate acidic
condtions (Brooks et al., 2007). The morphotype appears in the southeastern Australian lakes
shows close correlation with lake productivity.
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16. Paratanytarsus Skuse
a. Lake Terangpom, Victoria, 40 X b. Grahamstown Lake, NSW, 20 X
a. Grahamstown lake, NSW, 40 X
15. Tanytarsus glabrescens type
b. Eagle Tarn, Tasmania, 40 X
Ecology: occurs in 50% of the lakes in the training set. As described above, the morphotype
usually occurs in the littoral of relatively warm, productive lakes and will tolerate acidic
condtions (Brooks et al., 2007). The morphotype appears in the southeastern Australian lakes
shows close correlation with nutrients loading, lake productivity and pH.
Ecology: occurs in 67% of the lakes in the training set. It may be abundant in warm or cold
lakes. Some morphotypes occur in cold oligotrophic lakes at high latitude or altitude
whereas, others are typical of warmer conditions. It is often assicated with macrophytes
(Brooks et al., 2007). In the the southeastern Australian lakes, it shows close correlation with
macrophytes, conductivity and appears as a warm indicator.
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17. Tanytarsus undifferentiated
a. Lake Selina, Tasmania, 40 X b. Lake Lila, Tasmania, 40 X
It is required a more detailed identification guide to further refine the ecology of the genus of
Tanytarsus in southeastern Australian lakes.
a. Lake Poona, QLD, 40 X
18. Tanytarsus lactescens type
b. Lake Hiawatha, NSW, 40 X
Ecology: occurs in 27% of the lakes in the training set. It is usually more abundant in
temperate, carbonate lakes (Heiri, 2001). In the the southeastern Australian lakes, the related
morphotype shows close correlation with water depth, lake productivity and pH. It also
shows as a warm temperature indicator.
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a. Lake Selina, Tasmania, 20 X b. Lake Lea Pond, Tasmania, 20 X
20. Stempellina Thienemann & Bause
a. Lake Lila, Tasmania, 40 X b. Lake Plimsoll Pond, Tasmainia, 40 X
19. Tanytarsus nr chinyensis
Ecology: occurs in 15% of the lakes in the training set. It usually appears as a cold
stenotherm from oligotrophic lakes but in Scandinavia it occurs at the warm end of the
thermal gradient (Brooks et al., 2007). In the the southeastern Australian lakes, the related
morphotype shows close correlation with temperature and pH. It appears as a cold
temperature indicator in southeastern Australia.
Ecology: only occurs in two Tasmanian lakes in the training set. It is a warm stenotherm
most abundant in oligotrophic lakes. Since it only appears in two lakes, it is difficult to
summarize the ecology. However, it seems to be abundant in the Tasmanian training set
(Rees et al., 2008).
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21. Harnischia Kieffer
a. Reedy Lake, Victoria, 40 X b. Bamberang Reservoir, NSW, 40 X
Ecology: occurs in 12% of the lakes in the training set. It is usually associated with warm
eutrophic lakes and the genus is described as acidophobic (Brooks et al., 2007). In the the
southeastern Australian lakes, it shows a correlation with temperature and also as a warm
stenotherm.
a. Lake Mombeong, Victoria, 100 X
22. Paralimnophytes type 1 unofficial morphotype
b. Lake Selina, Tasmania 40 X
Ecology: occurs in 40% of the lakes in the training set.
For the genus: It is usually associated with very shallow water in the littoral of lakes and with
streams. The genus can sometimes be a useful indicator of lake level fluctuations. Some
species are associated with aquatic macrophytes, others are terrestrial or semiterrestrial. The
genus usually occurs in temperate waters. Larvae of Paralimnophyes can be found in
eutrophic lowland pools (Brooks et al., 2007).
This morphotype (type 1) described in the southeastern Australian training set is correlated
with macrophytes, conductivity, pH and shows as a cold stenotherm.
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24. Paralimnophytes type 3 unofficial morphotype
b. Green Lake, Victoria, 40 X a. Thirlmere Lakes, NSW, 40 X
a. Lake Plimsoll Pond, Tasmania, 40 X
23. Paralimnophytes type 2 unofficial morphotype
b. Lake Wellums, NSW, 40 X
Ecology: occurs in 40% of the lakes in the training set.
The description of the genus is shown in 22 above. In the the southeastern Australian lakes,
this morphotype (type 2) shows as a close correlation with macrophytes.
Ecology: occurs in 27% of the lakes in the training set.
The description of the genus is shown in 22 above. In the the southeastern Australian lakes,
this morphotype (type 3) does not show any clear correlation with the tested environmental
variables.
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b. Lake Terangpom, Victoria, 40 X
25. Parakiefferiella undifferentiated
a. Green Lake, Victoria, 40 X
Ecology: occurs in 20% of the lakes in the training set.
For the genus: species of Parakiefferiella are often abundant in the littoral of temperate lakes.
Some morphotypes are associated with warmer and more eutrophic lakes than others. The
distribution ranges from profundal to littoral or sublittoral of shallow lakes. Some
morphotypes are cold stenotherms occurring in oligotrophic lakes (Brooks et al., 2007).
The undifferentiated species of the genus Parakiefferiella in the southeastern Australian
lakes shows correlation with water depth.
26. Parakiefferiella type 1 unofficial type
a. Green Lake, Victoria, 40 X b. Lake Mombeong, Victoria, 40 X
Ecology: only occurs in 3 lakes in the training set.
The description of the genus is shown in 25 above. In the the southeastern Australian lakes,
this morphotype (type 1) does not show any clear correlation with the tested environmental
variables.
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b. Green Lake, Victoria, 40 X a. Lake Selina, Tasmania, 40 X
27. Parakiefferiella type 2 unofficial type
Ecology: occurs in 4 lakes in the training set.
The description of the genus is shown in 25 above. In the the southeastern Australian lakes,
this morphotype (type 2) shows correlation with temperature, macrophytes, productivity, pH
and conductitivy. It is identified as a cold stenotherm.
28. Parakiefferiella type 3 unofficial type
a. Lake Lea Pond, Tasmania, 20 X b. Lake Selina, Tasmania, 40 X
Ecology: occurs in 4 lakes in the training set.
The description of the genus is shown in 25 above. In the the southeastern Australian lakes,
this morphotype (type 3) shows correlation with temperature, macrophytes, pH and
conductivity. It is also identified as a cold stenotherm.
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29. Botryocladius Cranston & Edward
a. Green lake, Victoria, 40 X
b. Little Llangothlin Lagoon, NSW, 40 X
Ecology: occurs in 27% of the lakes in the training set. Larval Botryocladius occur in a range
of mostly temperate habitats, ranging from glacial-fed lakes (Patagonia), to moderate
elevation lakes (Patagonia, Tasmania, s.e. Australia), to rivers and creeks (Patagonia and s.e.
Australia) to ephemeral streams (southern and western Australia) (Cranston and Edward,
1999). It shows correlation with temperature in the southeastern Australian training set and
appears as a cold stenotherm.
30. Smittia Homgren
a. Lake Terangpom, Victoria, 40 X b. Green lake, Victoria, 40
Ecology: only occurs in 3 lakes in the training set.
Most species of Smittia are terrestrial although the genus may be aquatic in the littoral and
splash zone of lakes or its presence may be indicative of erosion events (Brooks et al., 2007).
This matches with the observation from the southeastern Australian lakes.
This genus does not show any clear correlation with the tested environmental variables.
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31. Gymnometriocnemus type 1 unofficial morphotype
a. Little Llangothlin Lagoon, NSW, 40 X b. Blue Lake, Kosciuszko, NSW, 40 X
Ecology: occurs in 7 lakes in the training set.
The larvae of the genus is said to be terrestrial, living in woodland soil (Cranston et al.,
1983), so their presence in lake sediment may be indicative of erosion events.
This matches with the observation from the southeastern Australian lakes, such as in Little
Llangothlin Lagoon.
This genus does not show any clear correlation with the tested environmental variables.
32. Genus Australia
a. Blue Lake, Kosciuszko, NSW, 40 X b. Lake Fortuna, Tasmania, taken by Andrew Rees (shared on October, 2013)
Ecology: only occurs in 2 lakes in the training set.
This genus occurs sporadically in running waters, riffles and waterfalls of eastern Australia,
from Far North Queensland south to Tasmania. Pupae have been found in New Zealand, in
several moderate to large rivers in both the South Island and far northern North Island
(Cranston, 2010).
This genus does not show any clear correlation with the tested environmental variables.
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33. Thienemanniella Kieffer
a. Spp 1. Little Llangothlin Lagoon, NSW, 40 X b. Spp 2. Lake Cartcarrong, Victoria, 40 X
Ecology: occurs in 4 lakes in the training set.
The genus usually occurs in streams and rivers but is also found in the sediment of temperate
lakes, often at cool temperature sites and can be relatively common in some Icelandic lakes.
In the southeastern Australian training set, this genus shows correlation with temperature,
macrophytes, water depth, lake productivity and conductivity. It is identified as a cold
stenotherm.
34. Orthoclad type 1 unofficial morphotype
a. Little Llangothlin Lagoon, NSW, 40 X b. Lake Reedy, Victoria, 40 X
Ecology: occurs in 3 lakes in the training set. The three lakes are lowland sites in souteastern
Australia. The species morphotype is not previously well described. The ecology is not well
understood. In the southeastern Australian training set, it shows correlation with
macrophytes.
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36. Pseudosmittia type 2 unofficial morphotype
b. Lake Cantani, Mt Buffalo, NSW, 40 X a. Lake Lea Pond, Tasmania, 40 X
Ecology: only occurs in 2 lakes in the training set.
Many species in the genus are terrestrial or semi-terrestrial but some occur in the littoral and
splash zone of lakes and are associated with aquatic macrophytes.
In the southeastern Australian training set, this genus shows correlation with temperature,
lake productivity and conductivity. It is identified as a cold stenotherm.
35. Orthoclad type 4 unofficial morphotype
a. Lake Cootaptamba, Kosciuszko, NSW, 40 X b. Wombat Pool, Tasmania, taken by Andrew Rees (shared on October, 2013)
Ecology: occurs in 8 lakes in the training set. The species morphotype was not previously
well recognized. It appears in glacial lakes of Tasmania and Kosciuszko, southeastern
Australia. It does not show any clear correlation with the tested environmental variables in
the southeastern Australian training set.
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38. Cricotopus ‘parbicintus’
b. Lake Esperance, Tasmania, taken by Andrew Rees (shared on October, 2013)
a. Blue Lake, Kosciuszko, NSW, 40 X
Ecology: only occurs in 2 lakes in the training set.
At genus level, it is eurytopic and can be found in flowing and standing waters. The genus is
ubiquitous in lake sediment samples. Most lentic taxa are associated with temperate,
relatively eutrophic conditions. Many taxa are littoral and are associated with vegetation,
some being leaf and stem miners. A number of species are cold stenotherms and occur in
oligotrophic cool waters (Brooks et al., 2007).
In the southeastern Australian training set, this genus shows correlation with temperature,
macrophytes, water depth, lake productivity and pH. It is also identified as a cold stenotherm.
37. Kosciuszko Orthoclad type 1
a. Lake Cootaptamba, Kosciuszko, NSW, 40 X b. Lake Cygnus, Tasmania, taken by Andrew Rees (shared on October, 2013)
Ecology: occurs in 3 lakes in the training set. It is recognized and named ‘SO2’ by Cranston
(2010). It was found in eastern Australia and Tasmania (Rees et al., 2008). It appearance is
restricted to montane tarns. The ecology is not described previously. It shows correlation
with lake productivity in the southeastern Australian training set.
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APPENDIX 3. CHIRONOMID FOSSIL TAXA ENUMERATED
FROM WELSBY LAGOON, NORTH STRADBROKE ISLAND,
AUSTRALIA FROM THE LAST GLACIAL MAXIMUM TO
NEAR PRESENT
• Chironomid taxa listed from 1 to 21 correspond to taxa specified in the
chironomid stratigraphy diagram displayed in FIGURE 5.3, Chapter 5.
• The ‘red box’ shows the key part of each taxon to be used for identification.
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3. Cladopelma Kieffer
b. 40 X (Glacial) a. 40 X (Holocene)
1. Procladius
a. 20 X (Holocene) b. 20 X (Glacial)
2. Parachironomus
a. Sp 1. 40 X (Holocene) b. Sp 2. 20 X (Holocene)
It is identified as a warm stenotherm in the modern training set and shows increase in
abundance from the LGM to deglaciation in the Welsby Lagoon record.
It is identified as a warm stenotherm in the modern training set and it is absent in the
samples that were inferred as cooler periods.
It is identified as a warm and shallow water indicator in the modern training set and shows
increase in abundance from the LGM to deglaciation in the Welsby Lagoon record.
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4. Paratanytarsus
b. 40 X (Glacial) a. 40 X (Glacial)
5. Tanytarsus lactescens type
b. 40 X (Glacial) a. 40 X (Glacial)
6. Cryptochironomus Kieffer
b. 40 X (Glacial) a. 20 X (Glacial)
It is identified as a warm stenotherm in the modern training set and shows increase in
abundance from the LGM to deglaciation in the Welsby Lagoon record.
It is identified as a warm stenotherm in the modern training set and shows increase in
abundance from the LGM to deglaciation in the Welsby Lagoon record.
It is identified as correlated with conductivity in the modern training set. It is absent before
18 cal ka BP and appears from the deglaciation in the Welsby Lagoon record.
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7. Tanytarsus glabrescens type
a. 40 X (Holocene) b. 40 X (Holocene)
8. Pentaneurini
20 X (Holocene) b. 40 X (Glacial)
9. Tanytarsus lugens type
a. 40 X (Glacial) b. 40 X (Holocene)
It is identified as correlated with macrophytes, productivity and pH in the modern training
set. It shows apparent increase in abundance from the LGM to deglaciation in the Welsby
Lagoon record.
It is identified as correlated with conductivity in the modern training set and shows
increase in abundance from the LGM to deglaciation in the Welsby Lagoon record.
It does not show clear correlation with the tested variables in the modern training set. It
only appears after 19 cal ka BP in the Welsby Lagoon record.
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12. Dicrotendipes
a. Spp 1. 40 X (Holocene) b. Spp 2. 40 X (Glacial)
10. Tanytarsus pallidicornis type
b. 100 X (Glacial) a. 40 X (Holocene)
11. Chironomus
b. Spp 2. 40 X (Holocene) a. Spp 1. 40 X (Holocene)
It is identified as correlated with lake productivity in the modern training set. It shows low
abundance at 18.5and 22 cal ka BP and high abundance in the rest of samples of the LGM
and deglaciation in the Welsby Lagoon record.
It does not show clear correlation with the tested variables in the modern training set. No
apparent trend is observed from the LGM to the deglaciation in the Welsby Lagoon record,
but the abundance is apparently high in the two Holocene samples.
It is identified as correlated with lake productivity, macrophytes, depth and conductivity in
the modern training set. It shows high abundance and is dominant the assemblages in the
four Holocene samples but much lower abundance in the LGM and deglaciation samples.
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14. Tanytarsus nr Chinyensis
a. 100 X (Glacial)
13. Polypedilum
a. 40 X (Holocene) b. 40 X (Glacial)
a. 100 X (Glacial)
15. Stempellina
b. 100X (Glacial)
It is often identified as a temperate climate indicator in modern training sets although some
morphotype species appear in subtropical lakes in Australia. It shows decrease in
abundance from the LGM to deglaciation in the Welsby Lagoon record.
It is identified as a cold stenotherm in the modern training set. It does not show apparent
trend from the LGM to the deglaciation in the Welsby Lagoon record.
It is identified as correlated with macrophytes, productivity and pH in the modern training
set and also a cold stenotherm. It shows decrease in abundance from the LGM to
deglaciation and is absent from the Holocene in the Welsby Lagoon record. It was not
found in modern subtropical lakes.
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16. Paralimnophyes unofficial morphotype 1
a. 40 X (Glacial) b. 40 X (Glacial)
17. Paralimnophyes unofficial morphotype 2
a. 100 X (Glacial) b. 40 X (Glacial)
18. Paralimnophyes unofficial morphotype 3
a. 40 X (Glacial) b. 100 X (Glacial)
It is identified as correlated with macrophytes, conductivity and pH in the modern training
set and also a cold stenotherm. It does not show apparent changes in abundance from the
LGM to deglaciation in the Welsby Lagoon record, but it is absent throughout the
Holocene and it was not found in modern subtropical lakes.
It is identified as correlated with macrophytes in the modern training set. It shows low
abundance and only in 3 samples from the deglaciation in the Welsby Lagoon record. It is
absent throughout the Holcene and it was not found in modern subtropical lakes.
It does not show clear correlation with the tested variables in the modern training set. It
shows low abundance in 4 samples from the LGM and deglaciation in the Welsby Lagoon
record. It is absent throughout the Holcene and it was not found in modern subtropical lakes.
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20. Pseudosmittia type 2 unofficial morphotype
a. Spp .1. 40 X (Glacial) b. Spp. 2. 40 X (Glacial)
19. Parakiefferiella nr. unofficial morphotype 3
b. Spp .2. 100 X (Glacial)
a. Spp. 1. 40 X (Glacial)
21. Genus Australia
a. 40 X (Glacial) b. 40 X (Glacial)
It is identified as correlated with macrophytes, conductivity and pH in the modern training
set and also a cold stenotherm. It does not show apparent changes in abundance from the
LGM to deglaciation in the Welsby Lagoon record, but it is absent throughout the
Holocene and it was not found in modern subtropical lakes.
It is identified as correlated with productivity and conductivity in the modern training set
and also a cold stenotherm. It only appears from the start of deglaciation in the Welsby
Lagoon record, but it is absent throughout the Holocene and it was not found in modern
subtropical lakes.
It does not show clear correlation with the tested variables in the modern training set. It has
low abundance in 4 of the LGM and deglaciation samples. It is absent throughout the
Holocene.
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Appendix 4. Published peer-reviewed journal articles during candidature
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APPENDIX 4. PUBLISHED JOURNAL ARTICLES DURING
CANDIDATURE
Paper 1, included as Chapter 3:
A snapshot of the limnology of eastern Australian water bodies spanning
the tropics to Tasmania: the land-use, climate, limnology nexus
Jie Christine Chang A B , Craig Woodward
A and James Shulmeister
A
A School of Geography, Planning and Environmental Management, The
University of Queensland, St Lucia, Brisbane, Qld 4072, Australia. B Corresponding author. Email: [email protected]
Marine and Freshwater Research 65(10) 872-883 http://dx.doi.org/10.1071/MF13265
Submitted: 10 October 2013 Accepted: 20 December 2013 Published: 7 July 2014
Hyperlink to full-text:
http://www.publish.csiro.au/view/journals/dsp_journal_fulltext.cfm?nid=126&f=MF13265
Paper 2, included as Chapter 4:
Hyperlink to full-text:
http://www.sciencedirect.com/science/article/pii/S0031018215000486
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Paper 3, included as Chapter 5:
Hyperlink to full-text:
http://www.sciencedirect.com/science/article/pii/S0277379115300147