Part I Ecological Settings
1 Introduction ................................................. 3
A. Osawa and O.A. Zyryanova
1.1 Permafrost Forest Biome ................................. 3
1.2 The Environment and Ecology ............................. 5
1.3 Natural Regions of Siberia .............................. 7
1.4 Main Region of Study .................................... 8
1.5 Brief History of Investigation ......................... 11
References .................................................. 13
2 Floristic Diversity and its Geographical Background
in Central Siberia .......................................... 17
O.A. Zyryanova, A.P. Abaimov, H. Daimaru, and Y. Matsuura
2.1 Introduction ........................................... 17
2.2 Regional Landforms Near Tura, Central Siberia .......... 18
2.2.1 Geological Setting .............................. 18
2.2.2 Slope Landforms ................................. 18
2.2.3 Fluvial Landforms ............................... 24
2.3 Soils in Permafrost Region of Siberia .................. 25
2.3.1 Permafrost Distribution in Siberia .............. 25
2.3.2 Unique Soil Characteristics ..................... 26
2.3.3 Revised Knowledge on the Circumpolar Biomes ..... 27
2.4 Geographical Patterns of Floristic Diversity in
Central Siberia ........................................ 27
2.5 Plant Species Diversity of Larch Association ........... 34
2.5.1 Description of Species Diversity ................ 34
2.5.2 Observed Patterns and Interpretations ........... 34
2.6 Conclusions ............................................ 36
References .................................................. 36
3 Geographical Distribution and Genetics of Siberian Larch
Species ..................................................... 41
A.P. Abaimov
3.1 Introduction ........................................... 41
3.2 Systematic Position and Present Status of Siberian
Larch Species .......................................... 42
3.3 Geographical Distribution of Siberian Larch Species .... 45
3.3.1 Larix sibirica Ledeb ............................ 45
3.3.2 Larix gmelinii (Rupr.) Rupur .................... 47
3.3.3 Larix cajanderi Mayr ............................ 48
3.4 Morphological and Ecological Features of Siberian
Larch Species .......................................... 49
3.4.1 Larix sibirica Ledeb ............................ 50
3.4.2 Larix gmelinii (Rupr.) Rupr ..................... 52
3.4.3 Larix cajanderi Mayr ............................ 54
3.5 Conclusions ............................................ 55
References .................................................. 55
4 Wildfire Ecology in Continuous Permafrost Zone .............. 59
M.A. Sofronov and A.V. Volokitina
4.1 Introduction ........................................... 59
4.2 Approaches to Study Wildfire Ecology ................... 60
4.3 Vegetation Fuel ........................................ 63
4.4 Seasonal Conditions of Fuel Moistening, Drying, and
Burning ................................................ 64
4.5 Wildfire Spread over the Territory ..................... 65
4.6 Causes of Wildfire and Areas of Wildfire Occurrence .... 66
4.7 Wildfire Impact on Larch Regeneration .................. 67
4.8 Ecological Effects of Wildfires ........................ 69
4.8.1 Soil Temperature ................................ 72
4.8.2 Summer Soil Thawing Depth ....................... 72
4.8.3 Influence of Fires on Growth of Larch Trees ..... 77
4.9 Conclusions ............................................ 79
References .................................................. 80
5 Recovery of Forest Vegetation After Fire Disturbance ........ 83
O.A. Zyryanova, A.P. Abaimov, T.N. Bugaenko, and
N.N. Bugaenko
5.1 Introduction ........................................... 83
5.2 Approaches to Study Vegetation Recovery ................ 84
5.3 Patterns of Vegetation Development After Fire .......... 86
5.3.1 Sites with Complex Microtopography .............. 86
5.3.2 Sites Without Microtopography ................... 88
5.4 Conclusions ............................................ 93
References .................................................. 94
Part II Ecosystem Dynamics and Function
6 Biomass and Productivity of Siberian Larch Forest
Ecosystems .................................................. 99
T. Kajimoto, A. Osawa, V.A. Usoltsev, and A.P. Abaimov
6.1 Introduction ........................................... 99
6.2 Data Source and Analysis .............................. 100
6.2.1 Study Site ..................................... 100
6.2.2 Estimation of Above- and Below-Ground
Biomass ........................................ 103
6.2.3 Estimation of Aboveground Net Primary
Production ..................................... 105
6.3 Biomass ............................................... 107
6.3.1 Aboveground Biomass ............................ 107
6.3.2 Belowground Biomass ............................ 112
6.4 Net Primary Production ................................ 113
6.4.1 Aboveground Production ......................... 113
6.4.2 Belowground Production ......................... 116
6.5 Carbon Allocation Pattern ............................. 117
6.6 Conclusions ........................................... 119
References ................................................. 120
7 Development of Stand Structure in Larch Forests ............ 123
A. Osawa and T. Kajimoto
7.1 Introduction .......................................... 123
7.2 Approaches to Describe Stand Development in Larch
Forests ............................................... 124
7.2.1 Study Site ..................................... 124
7.2.2 Measurement of Chronosequence Plots ............ 124
7.2.3 Measurement of Additional Stands ............... 126
7.2.4 Yield-Density and Yield Table Data ............. 126
7.2.5 Reconstructing Past Stand Structure ............ 127
7.3 Yield-Density Relationship ............................ 128
7.4 Yield Table Data ...................................... 131
7.5 Chronosequence Data ................................... 132
7.6 Reconstructed Stand Structure in the Past ............. 133
7.6.1 Height Growth .................................. 134
7.6.2 Tree Mortality ................................. 135
7.6.3 Reconstructed Stem Slenderness and Stand
Density ........................................ 136
7.6.4 Reconstructed Stem Size Distribution ........... 139
7.6.5 Reconstructed Total Stem Volume and Stem
Volume Growth .................................. 139
7.6.6 Consideration on Accuracy of the
Reconstruction Method .......................... 143
7.7 Conclusions ........................................... 145
References ................................................. 146
8 Soil Carbon and Nitrogen, and Characteristics of Soil
Active Layer in Siberian Permafrost Region ................. 149
Y. Matsuura and M. Hirobe
8.1 Introduction .......................................... 149
8.2 Approaches to Describe Soil Properties ................ 150
8.2.1 Soil Carbon and Nitrogen Storage, and Carbon
Budget ......................................... 150
8.2.2 Soil Properties Along a Toposequence ........... 152
8.3 Soil Carbon and Nitrogen Storage ...................... 153
8.3.1 Carbon storage in Forest Ecosystems ............ 153
8.3.2 SOC Storage and C/N Ratio in Central and
Northeastern Siberia ........................... 154
8.4 Soil Properties Along a Toposequence in a Larch
Forest ................................................ 156
8.4.1 Thickness of Soil Active Layer and Forest
Floor, and Characteristics of Canopy Cover ..... 156
8.4.2 Soil Chemical Properties ....................... 156
8.4.3 Forest Structure and Soil Nutrient
Properties ..................................... 158
8.5 Conclusions ........................................... 161
References ................................................. 161
9 Soil Respiration in Larch Forests .......................... 165
T. Morishita, O.V. Masyagina, T. Koike, and Y. Matsuura
9.1 Introduction .......................................... 165
9.2 Approaches to Study Soil Respiration .................. 166
9.2.1 Study Site ..................................... 166
9.2.2 Measurement of Soil Respiration ................ 167
9.2.3 CO2 Analysis and Calculation of Soil
Respiration Rate ............................... 168
9.2.4 Climate Condition of the Measurement Period .... 168
9.3 Soil Temperature, Moisture, and Respiration Rate ...... 168
9.4 Relationship Between Soil Respiration and Soil
Temperature and Moisture .............................. 170
9.5 Seasonal Changes of Soil Respiration .................. 171
9.6 Comparison of Soil Respiration in the Growing
Season ................................................ 172
9.7 Dynamics of Other Trace Gases in Larch Forests of
Siberia ............................................... 174
9.7.1 Methane (CH4) .................................. 174
9.7.2 Nitrous Oxide (N20) ............................ 175
9.8 Conclusions ........................................... 175
References ................................................. 179
10 Net Ecosystem Exchange of CO2 in Permafrost Larch
Ecosystems ................................................. 183
Y. Nakai
10.1 Introduction .......................................... 183
10.2 Study Site for Micrometeorological Measurements ....... 184
10.3 Meteorological Condition and Features of the
Measurement Site ...................................... 190
10.4 Intensity and Seasonal Variations in Net Ecosystem
Exchange and Larch Tree Phenology ..................... 192
10.5 Conclusions ........................................... 199
References ................................................. 199
11 Behavior of Dissolved Organic Carbon in Larch Ecosystems ... 205
A.S. Prokushkin, S. Hobara, and S.G. Prokushkin
11.1 Introduction .......................................... 205
11.2 Approaches to Study Dissolved Organic Carbon .......... 206
11.2.1 Site Description ............................... 206
11.2.2 Soil and Water Sampling and Analyses ........... 207
11.3 DOC Content and Release in Soils ...................... 210
11.3.1 Soil Organic Matter Stocks and WEOC
Content in Soils ............................... 210
11.3.2 Soil DOC Concentrations and Fluxes ............. 212
11.4 Export of Terrestrial DOC to Riverine System .......... 218
11.4.1 Seasonal Patterns of Riverine DOC
Concentrations ................................. 218
11.4.2 Implication for Global Change .................. 224
11.5 Conclusions ........................................... 225
References ................................................. 226
12 Soil Nitrogen Dynamics in Larch Ecosystem .................. 229
N. Tokuchi, M. Hirobe, K. Kondo, H. Arai, S. Hobara,
K. Fukushima, and Y. Matsuura
12.1 Introduction .......................................... 229
12.2 Approaches to Examination of Soil Nitrogen
Dynamics and Status ................................... 230
12.2.1 Study Sites .................................... 230
12.2.2 Soil N Mineralization, Leaching, and Status .... 231
12.3 Soil Nitrogen Dynamics ................................ 232
12.3.1 Soil Inorganic N Pool .......................... 232
12.3.2 Soil N Mineralization .......................... 234
12.3.3 Controlling Factors on Soil N Dynamics ......... 235
12.3.4 Inorganic N leaching in Soil ................... 237
12.4 Soil Nitrogen Status in Larch Forest in Central
Siberia ............................................... 237
12.4.1 Available N .................................... 237
12.4.2 The Possibility of N Limitation of Larch
Forest in Central Siberia ...................... 237
12.4.3 N Source of Larch Forest in Central Siberia
Based on Isotopic signature .................... 238
12.5 Conclusions ........................................... 240
References ................................................. 241
13 Hydrological Aspects in a Siberian Larch Forest ............ 245
T. Ohta
13.1 Introduction .......................................... 245
13.2 Approaches to Study Stand-scale Hydrological
Characteristics in a Larch Forest of Northeastern
Siberia ............................................... 246
13.2.1 Study Site for the Stand-scale Investigation ... 246
13.2.2 Measurement of Meteorological and
Environmental Variables ........................ 247
13.2.3 Measurement of Water Vapor and Energy Fluxes ... 247
13.2.4 Evaluation of Hydrological Cycles in the Lena
River Basin .................................... 248
13.3 Seasonal and Interannual Variation of Energy
Partitioning above the Siberian Larch Forest .......... 248
13.4 Water Balance of One-dimensional Scale in the
Siberian Larch Forest ................................. 251
13.4.1 Interannual Variation .......................... 251
13.4.2 Annual Evapotranspiration and Environmental
Variables ...................................... 255
13.4.3 Water and Energy Exchange Differences
between Non-permafrost and Permafrost
Areas of Siberia ............................... 257
13.4.4 Water and Energy Exchange in Different
Environments and Climates ...................... 258
13.5 Evaluation of Hydrological Aspects in Northeastern
Siberia ............................................... 263
13.6 Conclusions ........................................... 265
References ................................................. 266
Part III Tree Physiology and The Environment
14 Photosynthetic Characteristics of Trees and Shrubs
Growing on the North- and South-Facing Slopes in Central
Siberia .................................................... 273
T. Koike, S. Mori, O.A. Zyryanova, T. Kajimoto,
Y. Matsuura, and A.P. Abaimov
14.1 Introduction .......................................... 273
14.2 Study Site and Measurement of Foliar Ecophysiology .... 274
14.3 Environmental Conditions .............................. 275
14.4 Photosynthetic Production and Shoot Morphology ........ 276
14.5 Photosynthesis and Respiration of Trees and Shrubs .... 277
14.5.1 Dominant Tree Species .......................... 277
14.5.2 Nutrient Condition in Needles .................. 280
14.5.3 Shrubs ......................................... 280
14.6 Light-Photosynthetic Curves ........................... 281
14.7 Chlorophyll Content ................................... 282
14.8 Future Vegetation ..................................... 283
14.9 Conclusions ........................................... 284
References ................................................. 285
15 Respiration of Larch trees ................................. 289
S. Mori, S.G. Prokushkin, O.V. Masyagina, T. Ueda,
A. Osawa, and T. Kajimoto
15.1 Introduction .......................................... 289
15.2 Approaches and Measurement System ..................... 290
15.2.1 Study Site ..................................... 290
15.2.2 Setting Whole-Plant Chamber .................... 290
15.2.3 Closed Air-Circulation System .................. 292
15.2.4 C02 Scrubber ................................... 293
15.2.5 Temperature Control ............................ 293
15.2.6 Measurement of Whole-Tree Respiration .......... 293
15.3 System Response and Estimated Tree Respiration ........ 294
15.3.1 Temperature Control of the System .............. 294
15.3.2 Temperature Dependency of Whole-tree
Respiration .................................... 294
15.3.3 Size Dependency of Whole-tree Respiration ...... 296
15.3.4 Estimation of Stand-Level Aboveground
Respiration .................................... 297
15.4 Evaluation of Measurement System ...................... 297
15.5 Aboveground Respiration and Production ................ 298
15.6 Conclusions ........................................... 299
References ................................................. 300
16 Root System Development of Larch Trees Growing
on Siberian Permafrost ..................................... 303
T. Kajimoto
16.1 Introduction .......................................... 303
16.2 Data Source ........................................... 304
16.2.1 Study Site ..................................... 304
16.2.2 Methods of Root System Excavation and
Measurements ................................... 305
16.2.3 Parameters of Above- and Below-Ground
Space Occupation ............................... 306
16.2.4 Growth Pattern Analysis ........................ 308
16.3 Spatial Pattern of Individual Root System ............. 308
16.4 Effects of Microscale Soil Condition on Root
Distribution .......................................... 309
16.4.1 Topography and Soil Temperature ................ 309
16.4.2 Topography and Soil Water ...................... 314
16.5 Temporal Pattern of Root System Development ........... 315
16.5.1 Replacement of Root System ..................... 315
16.5.2 Growth Rate and Pattern of Lateral Root ........ 317
16.6 Below-ground Space Occupation by Root System .......... 321
16.6.1 Relationship Between Root System and Crown ..... 321
16.6.2 Stand-Level Root Network ....................... 323
16.7 Linkage with Postfire Permafrost Soil Environment ..... 323
16.8 Below-ground Competitive Interactions ................. 325
16.9 Conclusions ........................................... 326
References ................................................. 327
17 Seasonal Changes in Stem Radial Growth of Larix gmelinii
in Central Siberia in Relation to its Climatic Responses ... 331
K. Yasue, J. Kujansuu, T. Kajimoto, Y. Nakai, T. Koike,
A.P. Abaimov, and Y. Matsuura
17.1 Introduction .......................................... 331
17.2 Approaches to Study Growth Phenology and Tree-Ring
Responses to Climate .................................. 332
17.2.1 Study Sites .................................... 332
17.2.2 Observations of Snow Melting, Needle
Phenology, and Seasonal Radial Growth .......... 333
17.2.3 Analysis of Climatic Response of Radial
Growth ......................................... 334
17.3 Seasonal Changes in Snow Melting, Needle Phenology,
and Radial Growth ..................................... 337
17.4 Climatic Responses of Radial Growth ................... 338
17.5 Conclusions ........................................... 343
References ................................................. 344
18 Dendrochronology of Larch Trees Growing on Siberian
Permafrost ................................................. 347
E.A. Vaganov and A.V. Kirdyanov
18.1 Introduction .......................................... 347
18.2 Experimental Background ............................... 348
18.3 Relationships of Tree-Ring Parameters Obtained
for Larch Dendrochronological Network ................. 352
18.4 Effects of Climatic Factors on Radial Growth
of Larch Trees ........................................ 354
18.5 Reconstruction of Summer Temperature Based
on Regional Chronologies .............................. 357
18.6 Effect of Ground Fires on Radial Tree Growth .......... 357
18.7 Features of Tree-Ring Growth on Siberian Permafrost ... 359
18.8 Conclusions ........................................... 360
References ................................................. 361
Part IV Ecosystem Comparisons and Responses to Climate Change
19 Characteristics of Larch Forests in Daxingan Mountains,
Northeast China ............................................ 367
F. Shi, K. Sasa, and T. Koike
19.1 Introduction .......................................... 367
19.2 Approaches to Study Biomass, Net Primary Production,
and Regeneration ...................................... 368
19.2.1 Study Sites .................................... 368
19.2.2 Estimation of Biomass and Net Primary
Productivity ................................... 370
19.2.3 Examination of Postfire Forest Dynamics ........ 371
19.3 Biomass, Productivity, and Stand Density .............. 371
19.3.1 Biomass and Aboveground Net Primary
Productivityin Different Climatic Zones ........ 371
19.3.2 Aboveground Biomass and Aboveground
Net Primary Productivity in Different Forest
Types .......................................... 372
19.3.3 Tree Density, Aboveground Biomass, and
Aboveground Productivity in Relation to
Forest Age ..................................... 373
19.3.4 Aboveground Biomass and Aboveground
Net Primary Productivity of Larch Plantation ... 373
19.4 Regeneration of Larch-Dominant Forests after Forest
Fires ................................................. 378
19.5 Synthesis ............................................. 379
19.6 Conclusions ........................................... 381
References ................................................. 381
20 Carbon Dynamics of Larch Plantations in Northeastern
China and Japan ............................................ 385
M. Jomura, W.J. Wang, O.V. Masyagina, S. Homma,
Y. Kanazawa, Y.G. Zu, and T. Koike
20.1 Introduction .......................................... 385
20.2 Site Descriptions ..................................... 386
20.3 Biomass and Net Primary Production .................... 389
20.3.1 Estimation Procedures .......................... 389
20.3.2 Biomass, Allocation, and Net primary
production ..................................... 393
20.4 Photosynthesis and Autotrophic Respiration ............ 397
20.4.1 Data Source .................................... 397
20.4.2 Leaf Photosynthesis ............................ 397
20.4.3 Cone Photosynthesis ............................ 399
20.4.4 Stem Respiration ............................... 401
20.4.5 Soil Respiration ............................... 401
20.5 Soil Respiration and Environment ...................... 402
20.5.1 Enriched CO2 experiment ........................ 402
20.5.2 Effects of CO2 ................................. 402
20.5.3 Effects of Plantation Management ............... 404
20.6 Conclusions ........................................... 408
References ................................................. 408
21 The Role of Ectomycorrhiza in Boreal Forest Ecosystem ...... 413
L. Qu, K. Makoto, D.S. Choi, A.M. Quoreshi, and T. Koike
21.1 Introduction .......................................... 413
21.2 Physiology of Ectomycorrhizal Plants .................. 414
21.3 Ectomycorrhizae in Boreal Forests ..................... 415
21.4 Carbon Flux in Ectomycorrhizal Plants ................. 415
21.5 Ectomycorrhizae in Permafrost Soils, and after
Forest Fires .......................................... 417
21.5.1 Ectomycorrhiza in Permafrost Soils ............. 417
21.5.2 Forest Fires and Ectomycorrhizae ............... 419
21.6 Ectomycorrhizae and Elevated Atmosphere CO2 ........... 420
21.7 Conclusions ........................................... 421
References ................................................. 421
22 From Vegetation Zones to Climatypes: Effects of Climate
Warming on Siberian Ecosystems ............................. 427
N.M. Tchebakova, G.E. Rehfeldt, and E.I. Parfenova
22.1 Introduction .......................................... 427
22.2 Background ............................................ 428
22.2.1 Study Area ..................................... 428
22.2.2 Mapping Current and Future Climates ............ 428
22.2.3 Permafrost ..................................... 429
22.2.4 Vegetation Zones ............................... 429
22.2.5 Major Forest-Forming Tree Species of Siberia ... 431
22.2.6 Distributions of Pinus sylvestris and Larix
Species ........................................ 431
22.2.7 Mapping Climatypes of Pinus sylvestris
and Larix Species .............................. 432
22.3 Effects of Global Warming on Vegetation Shifts ........ 433
22.4 Effects of Global Warming on Species Distributions .... 435
22.5 Effects of Global Warming on Number, Size,
and Distribution of Climatypes ........................ 438
22.5.1 Pinus sylvestris ............................... 438
22.5.2 Larix sibirica ................................. 440
22.5.3 Larix dahurica ................................. 440
22.5.4 Larix sukaczewii ............................... 440
22.6 Synthesis ............................................. 441
22.7 Conclusions ........................................... 444
References ................................................. 444
23 Effects of Elevated CO2 on Ecophysiological Responses
of Larch Species Native to Northeast Eurasia ............... 447
T. Koike, K. Yazaki, N. Eguchi, S. Kitaoka, and R. Funada
23.1 Introduction .......................................... 447
23.2 Growth Characteristics of Larch Species ............... 448
23.3 Photosynthetic Adjustment at Elevated [CO2] ........... 448
23.4 Nitrogen and Water Use Efficiency ..................... 450
23.5 Xylem Formation ....................................... 451
23.6 Rehabilitation with Larch Species ..................... 451
23.7 Conclusions ........................................... 454
References ................................................. 454
Part V Synthesis and Conclusion
24 Characteristics of Permafrost Forests in Siberia
and Potential Responses to Warming Climate ................. 459
A. Osawa, Y. Matsuura, and T. Kajimoto
24.1 Introduction .......................................... 459
24.2 Characteristics of Permafrost Forests in Siberia ...... 460
24.2.1 Forest Fire and Dynamics of Ecosystem
Development .................................... 460
24.2.2 Ecosystem Carbon Budget ........................ 461
24.2.3 Comparison to Permafrost Forests of North
America ........................................ 464
24.3 Potential Responses to Warming Climate ................ 467
24.3.1 Ecosystem Structure ............................ 467
24.3.2 Ecosystem Development .......................... 472
24.3.3 Ecosystem Function ............................. 474
24.4 Conclusions ........................................... 477
References ................................................. 478
Color Plates .................................................. 483
Illustration and Table Credits ................................ 491
Species Index ................................................. 495
Subject Index ................................................. 497
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