Insect diversity patterns along environmental gradients in the temperate forests of Northern China

241 Pages·2014·3.2 MB·English

Insect diversity patterns along environmental gradients in
the temperate forests of Northern China
Yi Zou
Department of Geography
University College London
London
WC1E 6BT
This thesis is submitted for the degree of PhD in Ecology
March 2014
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I, Yi Zou, confirm that the work presented in this thesis is my own. Where
information has been derived from other sources, I confirm that this has been
indicated in the thesis.

Yi Zou

13 March 2014
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Acknowledgements
Foremost, I would like to express my sincerely appreciation to my lovely supervisor,
Dr. Jan Axmacher, who has fully supported me in a number of ways during my study.
Jan’s encouragement and guidance enabled me to develop my research ability and
in-depth understanding of this subject, and allowed me to start my career as a research
scientist. I also would like to thank my second supervisor Dr. Simon Lewis, for his
full support in my upgrade and his insightful comments on this thesis. I also want to
thank my two thesis examiners, Dr. Helene Burningham and Professor Simon Leather,
for their constructive comments.
I am grateful for the support from Professor Weiguo Sang from the Institute of Botany,
Chinese Academy of Sciences, who offered advice and also financial support and a
variety of other resources for this research. I would not have been able to complete
my research without help from him and his group members. In this respect, I
particularly want to thank Dr Fan Bai, Dr Shunzhong Wang, Dr Haifeng Liu and Mr
Wenchao Li for providing data and helping me to complete vegetation surveys during
my research.
I also thank Professor Zhenrong Yu and Dr Yunhui Liu and their group members from
the China Agricultural University. They provided valuable assistance in my insect
specimen preparation. I have spent a very good and productive time while working
with their team members, particularly with Mr Meichun Duan.
Professor Hongbin Liang, Dr Hongliang Shi and Mr Ye Liu from the Institute of
Botany, Chinese Academy of Sciences kindly helped me with the sorting and
identification of ground beetles. Professor Dayong Xue and Dr Hongxiang Han
helped me with the identification of geometrid moths.
Many thanks to Mr Haicheng Zhou from the Changbaishan Nature Reserve
Management Centre for his great support during the field work. My sincerest thanks
also go to the members of the Changbaishan Academy of Sciences and the
Changbaishan Natural Museum for their great assistance. I furthermore sincerely
thank the staff of the Changbaishan Forest Ecosystem Research Station and the
Beijing Forest Ecosystem Research Station for their hospitality. In addition, I greatly
appreciate the help with my fieldwork from Eleanor Warren-Thomas, Chao Long,
Min Liu, Xiasai Zhou, Furen Han, Lijia Dong, Xuenan Yao, Mengjie Yang, Xiaoliang
Zhang and Ya Qin.
My final warmest thanks go to my wife, who has always been my joy and given me
the greatest support. Thank you for spending those difficult times with me! I thank my
new-born beloved daughter bringing greatest joy and a great source of inspiration.
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Abstract

Insects, which represent the most species-rich taxa, are extremely important
ecosystem components. The diversity patterns of insects have, however, been widely
ignored in biodiversity research. In my thesis, I aim to establish a basic understanding
of the diversity patterns of insect assemblages in the temperate forest and forest
plantation ecosystems of Northern China, and to investigate how these patterns
correlate with vegetation and environmental conditions. The study aims to give
further insights into the insect diversity status and measures to conserve or even
enhance their diversity in the large secondary and plantation forests which have been
and are currently established throughout northern China.
The study focuses on two distinct insect taxa: ground beetles (Coleoptera: Carabidae)
and geometrid moths (Lepidoptera: Geometridae). In the main study area located
within the Changbaishan Natural Reserve (CNR) in Jilin Province, 4844 individuals
(47 species) of ground beetle and 9285 individuals (155 species) of geometrid moth
were sampled. In addition, 1488 ground beetles (24 species) and 2047 geometrid
moths (165 species) were sampled in the secondary and plantation forest area at
Dongling Mountain (DLM) in Beijing.
A first important result of this work is that the α-diversity of both ground beetle and
geometrid moth assemblages decreased significantly with increasing elevation at
CNR. My results also show that the relationships between phyto-diversity and the
diversity of insects are weak and furthermore likely to be driven by underlying
environmental factors. The significant changes which have recently occurred in the
plant species composition at CNR chiefly related to changes in the climatic conditions
suggest that insect species are also under high pressure in this area. Finally, this study
suggests that in the temperate religions of Northern China, secondary and plantation
forests can potentially harbour high levels of insect α-diversity compared with mature,
more pristine forests.
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Abbreviations
ACE Abundance-based Coverage Estimator
AIC Akaike information criterion
BF birch forest
BFERS Beijing Forest Ecosystem Research Station
BSP Bashang Plateau
CBD Convention on Biological Diversity
CCDB Canadian Centre for DNA Barcoding
CCFP Conversion of Cropland to Forest Programme
CNESS chord-normalized expected species shared
CNR Changbaishan Natural Reserve
DLM Dongling Mountain
FPC family-based principal component
FTP Fast-Growing and High-Yielding Timber Base Construction Programme in
Key Areas
HD herb density
HH herb Shannon diversity
IB-CAS Institute of Botany, Chinese Academy of Sciences
ICE Incidence-based Coverage Estimator
IDH intermediate disturbance hypothesis
IISE International Institute for Species Exploration
IndVal indicator value
IUCN International Union for Conservation of Nature
IV importance value
IZ-CAS Institute of Zoology, Chinese Academy of Sciences
KPF Korea Pine Forest
LDG latitudinal diversity gradient
MCBF mixed coniferous and broad-leaved forest
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MCF mixed coniferous forest
MDE mid-domain effect
MDS multidimensional scaling
MLR multiple linear regression
MM Michaelis–Menten estimator
NFPP Natural Forest Protection Programme
NMDS non-metric multidimensional scaling
PC Principal Component
PCA Principal Components Analysis
RDA Redundancy analysis
SCP Sand Control Programme
SD shrub density
SH shrub Shannon diversity
SHE species richness, Shannon diversity index and evenness
SKFP Six Key Forestry Programmes
SMCF sub-alpine mixed coniferous forest
SPC species-based principal component
TD tree density
TH tree Shannon diversity
TYSDP Yangtze River Basin’ Shelterbelt Development Programme
WNDP Wildlife Conservation and Nature Reserves Development Programme

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Contents
Chapter 1. Introduction............................................................................................15
1.1. Biodiversity.............................................................................................................15
1.1.1. What is biodiversity?.....................................................................................................15
1.1.2. Biodiversity and human well-being...............................................................................17
1.1.3. Biodiversity and ecosystem functioning.........................................................................19
1.1.4. Global biodiversity and its distribution.........................................................................22
1.1.5. Threats to biodiversity...................................................................................................28
1.1.6. Biodiversity and climate change....................................................................................32
1.2. Insect diversity........................................................................................................34
1.2.1. Importance of insects.....................................................................................................34
1.2.2. Implications of knowledge gaps relating to insect diversity..........................................36
1.3. Insect taxa selected in this study...........................................................................39
1.3.1. Carabidae......................................................................................................................39
1.3.2. Geometridae..................................................................................................................41
1.4. China’s temperate forests and their biodiversity................................................43
1.4.1. China’s temperate forests distribution..........................................................................43
1.4.2. Forest plantations in China...........................................................................................47
1.4.3. Plantation and secondary forest and biodiversity.........................................................49
1.5. The insects research gap in Northern China.......................................................51
1.6. Research aim and objectives.................................................................................52
Chapter 2. Methodology...........................................................................................54
2.1. Study area...............................................................................................................54
2.2. Research design......................................................................................................61
2.3. Sampling plot design..............................................................................................66
2.4. Sampling methods for carabids and geometrids.................................................69
2.4.1. Pitfall trapping..............................................................................................................69
2.4.2. Leaf litter collection......................................................................................................70
2.4.3. Window trapping...........................................................................................................71
2.4.4. Light trapping................................................................................................................73
2.5. Treatment and identification of insect specimens...............................................74
2.6. Vegetation survey...................................................................................................75
2.7. Data analysis...........................................................................................................75
Chapter 3. Altitudinal diversity patterns of ground beetles and geometrid moth in
the forests of Changbai Mountain.............................................................................77
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3.1. Introduction............................................................................................................77
3.2. Method and data analysis......................................................................................79
3.3. Results.....................................................................................................................81
3.3.1. Species compositions.....................................................................................................81
3.3.2. Elevational distribution ranges.....................................................................................84
3.3.3. Abundance and diversity change with increasing elevation..........................................87
3.3.4. Insects diversity in different forest types.......................................................................89
3.3.5. Species turnover............................................................................................................92
3.4. Discussion................................................................................................................94
3.4.1. Overall diversity and species composition of carabids and geometrids........................94
3.4.2. α-diversity and species distribution with increasing elevation......................................96
3.4.3. Species composition and abundance-distribution relationships....................................97
3.4.4. Potential threats to insect diversity at CNR..................................................................98
3.4.5. Synthesis........................................................................................................................99
Chapter 4. Statistical models of the links between insect and plant diversity......100
4.1. Introduction..........................................................................................................100
4.2. Sampling method and data analysis...................................................................102
4.3. Results...................................................................................................................104
4.3.1. Insect-plant relationships............................................................................................106
4.3.2. Insect-plant relationship in the different vegetation types...........................................108
4.4. Discussion...............................................................................................................111
4.4.1. Plant diversity and insect diversity..............................................................................111
4.4.2. Plant diversity and insect abundance..........................................................................112
4.4.3. Plant density and insect diversity................................................................................113
4.4.4. Plant density and insect abundance............................................................................114
4.4.5. Synthesis......................................................................................................................115
Chapter 5. Implications of climate and vegetation change for spatial diversity
patterns of insects......................................................................................................117
5.1. Introduction...........................................................................................................117
5.2. Sampling plots and data analysis.........................................................................119
5.3. Results...................................................................................................................121
5.3.1. Vegetation principal components................................................................................121
5.3.2. Models predicting insect α-diversity............................................................................127
5.3.3. Insect species composition and vegetation..................................................................127
5.3.4. Long term vegetation composition changes................................................................133
5.3.5. Long-term Climate change and insect diversity predictions.......................................134
5.4. Discussion..............................................................................................................138
5.4.1. Vegetation composition and insect diversity...............................................................138
5.4.2. Long-term vegetation change and insect diversity......................................................140
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5.4.3. Synthesis......................................................................................................................141
Chapter 6. Insect diversity patterns in secondary and plantation forests in central
Northern China—a case study from Dongling Mountain, West Beijing...............142
6.1. Introduction..........................................................................................................142
6.2. Sampling plots and data analysis........................................................................144
6.3. Results...................................................................................................................145
6.3.1. Comparison of α-diversity between different forests...................................................147
6.3.2. Indicator species..........................................................................................................149
6.3.3. Insect species turnover rates.......................................................................................150
6.3.4. Species composition of plants......................................................................................153
6.4. Discussion..............................................................................................................153
6.4.1. Overall number of species on Donling Mountain........................................................153
6.4.2. α-diversity and species composition for different forests............................................155
6.4.3. Synthesis......................................................................................................................157
Chapter 7. Comparison of insect diversity patterns in mature, secondary and
plantation forests in temperate North China...........................................................158
7.1. Introduction..........................................................................................................158
7.2. Sampling sites selection and data analysis.........................................................160
7.3. Results...................................................................................................................161
7.3.1. α-diversity....................................................................................................................161
7.3.2. Species composition, turnover and similarities...........................................................166
7.4. Discussion..............................................................................................................171
7.4.1. Plants and carabids.....................................................................................................171
7.4.2. Geometrid moths.........................................................................................................172
7.4.3. Species composition overlap.......................................................................................173
7.4.4. Synthesis......................................................................................................................173
Chapter 8. Discussion and conclusion..................................................................175
8.1. General diversity of ground beetles and geometrid moths in study areas......175
8.2. Main findings and lessons for biodiversity conservation..................................176
8.3. Further work........................................................................................................177
8.4. Conclusion.............................................................................................................179
Bibliography..............................................................................................................181
Appendix 1. Expression of used biodiversity measurements...................................226
Appendix 2. Number of carabid individuals for each species sampled in different
forests in Changbai Mountain..................................................................................229
Appendix 3. Number of geometrid moth individuals for each species sampled in
different forests in Changbai Mountain...................................................................231
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Appendix 4. Number of carabid individuals for each species sampled in different
forests in Dongling Mountain..................................................................................236
Appendix 5. Number of geometrid individuals for each species sampled in different
forests in Dongling Mountain..................................................................................237

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