Insect Diversity Patterns Along Environmental Gradients in the Temperate Forests of Northern China
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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 1 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 2 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. 3 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. 4 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 5 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 6 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 .....................................................................................................