Principles of Terrestrial Ecosystem Ecology F Stuart Chapin III Pamela A Matson Harold A Mooney Springer PTEPR 10/9/2002 7:23 PM Page i Principles of Terrestrial Ecosystem Ecology PTEPR 10/9/2002 7:23 PM Springer New York Berlin Heidelberg Barcelona Hong Kong London Milan Paris Singapore Tokyo Page ii PTEPR 10/9/2002 7:23 PM Page iii F Stuart Chapin III Pamela A Matson Harold A Mooney Principles of Terrestrial Ecosystem Ecology Illustrated by Melissa C Chapin With 199 Illustrations 13 PTEPR 10/9/2002 7:23 PM Page iv Pamela A Matson Department of Geological and Environmental Science School of Earth Sciences Green 355 Stanford University Stanford, CA 94305-2115 USA matson@pangea.stanford.edu F Stuart Chapin III Institute of Arctic Biology University of Alaska Fairbanks, AK 99775 USA terry.chapin@uaf.edu Harold A Mooney Department of Biological Sciences Herrin Hall, MC 5020 Stanford University Stanford, CA 94305-5020 USA hmooney@jasper.stanford.edu Cover illustration: Waterfall and forests on Valean Poas in Costa Rica Photograph by Peter Vitousek Library of Congress Cataloging-in-Publication Data Chapin, F Stuart (Francis Stuart), III Principles of terrestrial ecosystem ecology / F Stuart Chapin III, Pamela A Matson, Harold A Mooney p cm Includes bibliographical references (p ) ISBN 0-387-95439-2 (hc :alk paper)—ISBN 0-387-95443-0 (sc :alk paper) Ecology Biogeochemical cycles Biological systems I Matson, P A (Pamela A.) II Mooney, Harold A III Title QH541 C3595 2002 577¢.14—dc21 2002017654 ISBN 0-387-95439-2 (hardcover) ISBN 0-387-95443-0 (softcover) Printed on acid-free paper © 2002 Springer-Verlag New York, Inc All rights reserved This work may not be translated or copied in whole or in part without the written permission of the publisher (Springer-Verlag New York, Inc., 175 Fifth Avenue, New York, NY 10010, USA), except for brief excerpts in connection with reviews or scholarly analysis Use in connection with any form of information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed is forbidden The use in this publication of trade names, trademarks, service marks, and similar terms, even if they are not identified as such, is not to be taken as an expression of opinion as to whether or not they are subject to proprietary rights Printed in the United States of America SPIN 10866301 (hardcover) SPIN 10866759 www.springer-ny.com Springer-Verlag New York Berlin Heidelberg A member of BertelsmannSpringer Science+Business Media GmbH (softcover) PTEPR 10/9/2002 7:23 PM Page v Preface Human activities are affecting the global environment in myriad ways, with numerous direct and indirect effects on ecosystems The climate and atmospheric composition of Earth are changing rapidly Humans have directly modified half of the ice-free terrestrial surface and use 40% of terrestrial production Our actions are causing the sixth major extinction event in the history of life on Earth and are radically modifying the interactions among forests, fields, streams, and oceans This book was written to provide a conceptual basis for understanding terrestrial ecosystem processes and their sensitivity to environmental and biotic changes We believe that an understanding of how ecosystems operate and change must underlie our analysis of both the consequences and the mitigation of human-caused changes This book is intended to introduce the science of ecosystem ecology to advanced undergraduate students, beginning graduate students, and practicing scientists from a wide array of disciplines We also provide access to some of the rapidly expanding literature in the many disciplines that contribute to ecosystem understanding The first part of the book provides the context for understanding ecosystem ecology We introduce the science of ecosystem ecology and place it in the context of other components of the Earth System—the atmosphere, ocean, climate and geological systems We show how these components affect ecosystem processes and contribute to the global variation in terrestrial ecosystem structure and processes In the second part of the book, we consider the mechanisms by which terrestrial ecosystems function and focus on the flow of water and energy and the cycling of carbon and nutrients We then compare and contrast these cycles between terrestrial and aquatic ecosystems We also consider the important role that organisms have on ecosystem processes through trophic interactions (feeding relationships), environmental effects, and disturbance The third part of the book addresses temporal and spatial patterns in ecosystem processes We finish by considering the integrated effects of these processes at the global scale and their consequences for sustainable use by human soci- v PTEPR 10/9/2002 vi 7:23 PM Page vi Preface eties Powerpoint lecture notes developed by one of the authors are available online (www.faculty.uaf.edu/fffsc/) as supplementary material Many people have contributed to the development of this book We particularly thank our families, whose patience has made the book possible, and our students from whom we have learned many of the important ideas that are presented In addition, we thank the following individuals for their constructively critical review of chapters in this book: Kevin Arrigo, Teri Balser, Perry Barboza, Jason Beringer, Kim Bonine, Rich Boone, Syndonia Bret-Harte, John Bryant, Inde Burke, Zoe Cardon, Oliver Chadwick, Scott Chambers, Melissa Chapin, Kathy Cottingham, Joe Craine, Wolfgang Cramer, Steve Davis, Sandra Diaz, Bill Dietrich, Rob Dunbar, Jim Ehleringer, Howie Epstein, Werner Eugster, Valerie Eviner, Scott Fendorf, Jon Foley, David Foster, Tom Gower, Peter Groffman, Paul Grogan, Diego Gurvich, Bill Heal, Sarah Hobbie, Dave Hooper, Shuijin Hu, Pilar Huante, Bruce Hungate, Jill Johnstone, Jay Jones, Jürg Luterbacher, Frank Kelliher, Jennifer King, Dave Kline, Christian Körner, Hans Lambers, Amanda Lynch, Michelle Mack, Steve MacLean, Joe McFadden, Dave McGuire, Sam McNaughton, Knute Nadelhoffer, Jason Neff, Mark Oswood, Bob Paine, Bill Parton, Natalia Perez, Steward Pickett, Stephen Parder, Mary Power, Jim Randerson, Bill Reeburgh, Peter Reich, Jim Reynolds, Roger Ruess, Steve Running, Scott Rupp, Dave Schimel, Josh Schimel, Bill Schlesinger, Guthrie Schrengohst, Ted Schuur, Stephen Parder Mark Serreze, Gus Shaver, Nigel Tapper, Monica Turner, Dave Valentine, Peter Vitousek, Lars Walker, and Katey Walter We particularly thank Phil Camil, Valerie Eviner, Jon Foley, and Paul Grogan for comments on the entire book; Mark Chapin, Patrick Endres, and Rose Meier for comments on illustrations; Phil Camil for comments on educational approaches; and Jon Foley and Nick Olejniczak for providing global maps F Stuart Chapin III Pamela A Matson Harold A Mooney PTEPR 10/9/2002 7:23 PM Page vii Contents Preface v Part I Context Chapter The Ecosystem Concept Introduction Overview of Ecosystem Ecology History of Ecosystem Ecology Ecosystem Structure Controls over Ecosystem Processes Human-Caused Changes in Earth’s Ecosystems Summary Review Questions Additional Reading 3 10 11 13 16 17 17 Chapter Earth’s Climate System Introduction Earth’s Energy Budget The Atmospheric System Atmospheric Composition and Chemistry Atmospheric Structure Atmospheric Circulation The Oceans Ocean Structure Ocean Circulation Landform Effects on Climate Vegetation Influences on Climate Temporal Variability in Climate Long-Term Changes Interannual Climate Variability Seasonal and Daily Variations 18 18 21 21 22 24 28 28 29 31 32 34 34 38 40 vii PTEPR 10/9/2002 viii 7:23 PM Page viii Contents Relationship of Climate to Ecosystem Distribution and Structure Summary Review Questions Additional Reading 41 44 44 45 Chapter Geology and Soils Introduction Controls over Soil Formation Parent Material Climate Topography Time Potential Biota Human Activities Controls over Soil Loss Development of Soil Profiles Additions to Soils Soil Transformations Soil Transfers Losses from Soils Soil Horizons and Soil Classification Soil Properties and Ecosystem Functioning Summary Review Questions Additional Reading 46 46 47 48 48 49 50 50 50 53 54 54 56 57 58 61 66 67 67 Part II Mechanisms Chapter Terrestrial Water and Energy Balance Introduction Surface Energy Balance Solar Radiation Budget Ecosystem Radiation Budget Energy Partitioning Seasonal Energy Exchange Water Inputs to Ecosystems Water Movements Within Ecosystems Basic Principles of Water Movement Water Movement from the Canopy to the Soil Water Movement Within the Soil Water Movement from Soil to Roots Water Movement Through Plants Water Losses from Ecosystems Evaporation from Wet Canopies Evapotranspiration from Dry Canopies 71 73 73 74 75 77 77 78 78 79 80 81 83 89 89 90 PTEPR 10/9/2002 7:23 PM Page ix Contents ix Changes in Storage Runoff Summary Review Questions Additional Reading 92 93 95 96 96 Chapter Carbon Input to Terrestrial Ecosystems Introduction Overview Photosynthetic Pathways C3 Photosynthesis C4 Photosynthesis Crassulacean Acid Metabolism Photosynthesis Net Photosynthesis by Individual Leaves Basic Principle of Environmental Control Light Limitation CO2 Limitation Nitrogen Limitation and Photosynthetic Capacity Water Limitation Temperature Effects Pollutants Gross Primary Production Canopy Processes Satellite Estimates of GPP Controls over GPP Summary Review Questions Additional Reading 97 97 98 98 102 103 105 105 105 109 110 113 114 115 115 115 117 119 121 121 122 Chapter Terrestrial Production Processes Introduction Overview Plant Respiration Physiological Basis of Respiration Net Primary Production What Is NPP? Physiological Controls over NPP Environmental Controls over NPP Allocation Allocation of NPP Allocation Response to Multiple Resources Diurnal and Seasonal Cycles of Allocation Tissue Turnover Global Distribution of Biomass and NPP Biome Differences in Biomass Biome Differences in NPP Net Ecosystem Production 123 123 125 125 127 127 128 129 132 132 133 134 136 137 137 138 140 PTEIndex 10/9/2002 426 7:40 PM Page 426 Index Dewfall, 76 Diffuse radiation, 73 Diffusion, nutrient movement to roots, 178 Diffusion shell, roots, 178 Dinitrogen, atmospheric, 22 Direct radiation, 73 Disasters See Disturbances to ecosystem Dissolved inorganic carbon (DIC), 335–336 Dissolved organic carbon (DOC), 335 leaching of, 141, 145 Dissolved organic nitrogen (DON), production and cycling of, 202–204 Disturbance regime, 12 species effect on, 272 Disturbances to ecosystem, 285–288 anthropogenic disturbances, 297–298 and biomass loss, 137 and carbon loss, 145–146 and change in ecosystem See Succession disturbance regime, species effects, 272 frequency of, 286–287 intensity of, 286 and landscape heterogeneity, 309–310 and net primary production (NPP), 140 and nutrient loss, 194 resilience of ecosystem to, 282–283, 287 severity of, 286 size of, 287, 310 spread of disturbance, influencing factors, 311–314 timing of, 288 types of events, 285 Diversity of species and climate, 42–43 effects on ecosystem, 274– 277 Doldrums, 27 Down regulation, 109 Downwelling, oceans, 29 Dry adiabatic lapse rate, 24 Dung beetles, and decomposition, 156 E Earth, orbit of, 34–35 Earthquakes, plate collisions, Earth System in Anthropocene epoch, 339 biotic homogenization of, 14 carbon balance, 123–125 carbon and photosynthesis, 97 current extinction event, 265–266 and ecosystem changes, 10 and ecosystem ecology concepts, 6–7 glacial events by geologic time periods, 34 global warming, 36–37 and hydrologic cycle, 71, 73 and nutrient cycling, 176, 197 Earthworms, and decomposition, 156, 272 Eccentricity, 25, 34 Ecological modeling, spatial scaling, 325–330 Ecosystem, first use of, Ecosystem ecology comparative studies of ecosystems, and global information, 6–7 historical view, 7–10 and related disciplines, scope of study, 3–7 state factor approach, 11 systems approach, 8–9, 10 Ecosystem management adaptive management, 364–365 basic concepts in, 357–359 definitions of, 363 ecosystem approach, 362–364 ecosystem goods and services valuation, 366–368 ecosystem restoration, 360–361 of endangered species, 360, 362 of fisheries, 359–360 of forests, 359 integrated conservation and development projects (ICDPs), 365–366 sustainable management, 358, 360, 362 Ecosystem respiration, 143 Ecosystems abiotic components of, 10 biological components of, 10 biotic community, 12–13 carbon budget, components of, 142–144 control variables, 11–13 ecosystem model, 10–11 energy budget, 74–75 energy storage, 75–77 in equilibrium with environment, feedbacks, positive and negative, 13, 14 human-caused changes, 13–16, 19 interactive controls of, 12 interannual variations in, 281, 283 and nitrification, 209 and nitrogen fixation, 200–201 processes, resilience of system, 282–283 size of, 4–5 steady state, time and changes to See Seasons; Temporal dynamics; Time Ectomycorrhizae, 183 E horizon, 58 Emissivity, 74 Endangered species, management of, 360, 362 Endocellulases, 168 Endodermis, 184 Energy budget and atmospheric composition, 21–22 and climate, 18–20 and clouds, 22 ecosystem radiation budget, 74–77 energy partitioning, 75–77 relationship to water properties, 72–73 seasonal energy exchange, 77 solar radiation effects, 20 storage of energy in ecosystem, 75–77 surface radiation budget, 73–74 Energy pyramid, 251, 252 Energy transfers and ecosystem processes, plants and photosynthesis, 8, 10, 98 and succession, 300–301 and trophic structure of ecosystem, 244–246 PTEIndex 10/9/2002 7:40 PM Page 427 Index Entisols, 59 Enzymes in soil, and decomposition, 152, 153, 154, 168–169, 204 Epiphytes, 43 Equator rising air of, 24–25 solar radiation at, 24 Equilibrium of ecosystems, Erosion See Soil loss Estuaries, productivity of, 316–317 Euphotic zone, 225, 229, 231, 234, 237 Eutrophication, lakes, 237–238, 257 Evaporation, water loss process, 89–90 Evapotranspiration, 71 from canopies, 90–92 control factors, 88 Everglades, restoration study, 361 Exocellulases, 168 Exoenzymes, 152, 154, 204 Extensification, 321–323 Extinction coefficient, 108 Extinction event, current in Earth system, 265–266 F Facilitation, 291–292 Feedbacks, and ecosystem dynamics, 13, 14 Ferrell cell, 25 Fertilizers See Agriculture Field capacity, 63, 80–81 Filter feeders, 226–227 Fine particulate organic matter (FPOM), 240 Fires flammability of plants, 272 frequency of, 287 and nitrogen loss, 213 patch dynamics, 310 See also Disturbances to ecosystem Fisheries, management of, 359–360 Fishing, overfishing, 14, 254, 273, 323 Floods See Disturbances to ecosystem Fog, rain forests, 77–78 Food chain defined, 245 length of, 257–258 links in, nutrient transfers, 259–261 organisms in, 244–245 plant-based/detritus-based integration, 261–263 trophic transfers, 244–245 Food webs complexity related to, 263 defined, 246 interconnections in, 245 plant-based/detritus-based, integration of, 261–262, 263 Forests biomass distribution of, 138 canopy systems and gross primary production (GPP), 115–117 and climate, 33 climate of, 41–42 and energy budget, 74–75 evaporation from canopies, 89–90 evapotranspiration from canopies, 90–92 humus of, 170 light capture and leaves, 107–109, 116–119 management of, 359, 363 net primary production (NPP), 130 types of, 41–42 water movement, from canopy to soil, 79–80 See also Boreal forest (taiga); Temperate forests; Tropical forests Fossil fuels, carbon dioxide increase, 337, 339 Fragmentation and decomposition, 151–153 process of, 153 Frankia, 198, 199 Fresh-water producers, 228 Fulvic acids, 170 Functional types, 265 functions of, 12–13 Fungi decomposition by, 137, 153–154, 162, 168 growth of, 153 427 and mycorrhizae, 153–154 in trophic system, 245 Fungivores, 262 G Gap-phase succession, 287, 309 Gas emissions, soil loss from, 58 Gases, atmospheric, 21–22 Gelisols, 60, 61 Generalist herbivores, 255 Geochemical tracers, input identification, 221 Geotropic, root length, 182 Glacial events by geologic time periods, 34 glacial advances See Disturbances to ecosystem and orbit of Earth, 35 and soil loss, 53 Gley soils, 57 Globalization, and biotic homogenization, 14 Global warming climate model indications, 36–37 greenhouse effect, 19 human-based causes, 15–16, 39, 341–342 last millennium, 37 ozone hole, 15–16 sources of evidence for, 38 terrestrial effects, 37 Goods and services of ecosystem, valuation of, 366–367 Grasslands biomass distribution of, 138 climate of, 42 humus of, 170 net primary production (NPP) of, 130, 131 sour versus sweet veldt, 248 Gravity and mass wasting, 52 and water movement, 79, 80 Grazers, 239–240 Grazing lawns, 253 Great Barrier Reef Marine Park Authority (GBRMPA), 365 Greenhouse effect, 19 Greenland, oceans, 29 Gross primary production (GPP), 97–98, 115–121 and canopy systems, 115–117 PTEIndex 10/9/2002 428 7:40 PM Page 428 Index Gross primary production (GPP) (cont.) carbon budget of ecosystem, components of, 142–144 control factors, 98 defined, 115 growing-season, environmental controls, 121 and leaf area variations, 119–120 and plant respiration, 127 post-production carbon cycling, 123–124 relationship to net primary production in forests, 132 satellite estimates of, 117–119 and seasonal variations, 120–121 Ground heat flux, 75 Groundwater carbon dioxide saturation, 141, 145 modification of, 93 size of pool, 93 Guano, 320 Gulf Stream, 31 Gyres, 29 H Haber process, 344 Hadley cell, atmospheric circulation, 25, 26, 31 Halocline, 29 Hardpans, 80 Hartig net, 183 Heat flux, 19 types of, 75–76 Heat of fusion, 72 Heat transfer, and oceans, 29–31 Heat of vaporization, 72 Herbivores assimilation efficiency, 256, 261 food supply depletion, 283, 284 and nutrient loss, 193–194 in trophic system, 245–250 types of, 255 Heterocysts, 199 Heterotrophic nitrifiers, 207 Heterotrophs aerobic heterotrophic respiration, 170–173 anaerobic heterotrophic respiration, 173–174 and carbon balance, 124–125 classification, basis of, 170 in trophic system, 245 Heterotropic respiration, 143 High-nitrogen, low-chlorophyll (HNLC) syndrome, 231 Himalayan Mountains, formation of, 48 Histosols, 59, 61 Homeotherms assimilation efficiency, 256 production efficiency, 256 Homeothermy, 255 Horse latitudes, 27 Hubbard Brook Forest, 214 Human activities in arid regions, 93 and carbon production, 337, 339 direct and indirect ecosystem effects, 15 and ecosystem changes, 13–16, 19 and landscape heterogeneity, 312–313, 321–324 land use modification, 321–324 and leaf area, 120 and nitrogen deposition, 201 and nitrogen production, 344–347 and phosphorus production, 347–348 and soil formation, 50 and sulfur production, 349 Humans, consumption and trophic levels, 246 Humic acids, 170 Humidification, and humus formation, 169 Humin, 170 Humus, 152, 169–170 chemical structure of, 152, 170 ecosystem variations, 170 formation of, 169–170 Hurricanes See Disturbances to ecosystem Hydraulic conductivity, 80, 86 Hydraulic lift, 84 Hydrothermal vents, 230 Hyphae, 153 Hyporrheic zone, 180, 242 I Ice, sea ice, 29–31 Ice cores, climate record, 35–36 Iceland, low-pressure zones, 27 Igneous rocks, 47, 55 Immobilization, nitrogen, 202 Inceptisols, 59 Infiltration, 80 Integrated conservation and development projects (ICDPs), 365–366 Intensification, 323–324 Interactive controls, types of, 12 Interannual climate variability, 37–40 Interception, 79 Intertropical convergence zone (ITCZ), 26–27, 42 Inverse mode, 329 Iron, and nitrogen fixation, 200 Irradiance, 108 Isotopes of carbon and photosynthesis, 104 and soil carbon turnover, 171 Isotopic signatures, of water sources, 84–85 Isotopic tracers, 301 J Jet streams, 26 K Kaolinite, 54 Katabatic winds, 32 Keystone species, 267 L Labile compounds, 152 Lakes, 236–238 carbon cycling, 238 eutrophication, 237–238, 257 nutrient cycling, 238 origin of, 236 photosynthesis, 236–238 seasonal turnover, 237 types of, 236 Land and sea breezes, 31 Landforms, and climate, 31–32 Landscape dynamics, 13 Landscape ecology, study of, 305 Landscape heterogeneity atmospheric transfers, 317–320 basic concepts, 305–307 and community dynamics, 307, 309 disturbance spread, 320–321 PTEIndex 10/9/2002 7:40 PM Page 429 Index and disturbances to ecosystem, 309–310 and human activities, 312–313, 321–324 interaction among causes of, 311–314 and legacies, 307, 309 and patch characteristics, 305–307 plant and animal movement, 320 spatial scaling, 325–330 and state factors, 307 transfer of materials, 314– 317 Landslides, soil loss, 51–52 Land use modification extensification, 321–323 and hydrologic cycle, 352 intensification, 323–324 Latent heat flux, 19, 75–76 Latent heat of vaporization, 75 Lateral transfers, of carbon, 140, 145 Laterite, 57 Latitude, and net ecosystem exchange (NEE), 147–148 Law of minimum, 133 Leaching carbon loss, 141, 145 and cations release and cycling, 219–220 and decomposition, 151, 152 defined, 152 nitrogen loss from, 214–215 and nutrient loss, 193 soil transfers by, 56–57 Lead, cycling of, 220 Leaf area index (LAI) determinants of, 128–129, 139–140 and net primary production (NPP), 138–140 and photosynthesis, 108, 119 Leaves death of See Decomposition decomposition rate, 164 life-span of, 110–113, 135, 136–137 nutrient concentration, 189 photosynthesis process, 105–115, 119–120 water movement through, 88–89 Legacies and landscape heterogeneity, 307, 309 succession, 283–285 Legumes, nitrogen-fixing, 199–200 Lichens, nitrogen-fixing, 199 Life history traits, 291 Light capture, leaves in forests, 107–109 Light compensation point, 108 Light-harvesting reactions, photosynthesis, 99–101 Light saturation, 107 Light use efficiency (LUE), 106, 113, 117–118 Lignin, and decomposition, 165–166, 168–169 LINKAGES model, 327 Lithosphere, 48 Litter, 58, 132, 152 quality and quantity, 163–166 See also Decompositon Litter bags, 157–158 Litterfall, 123 Littoral zone, 227–228, 236 Loam, 61, 62, 80 Loess soils, 62 Logs, decomposition of, 162 Longwave radiation, 19, 22, 74 M Macrofauna and decomposition, 156 types of, 155 Macronutrients, 179 Macropores, 80, 154 Magma, 47 Magnesium, cycling of, 219 Manganese, cycling of, 219 Marine ecosystem, alteration by human activities, 14 Mass flow, nutrient movement to roots, 180 Mass wasting, soil loss, 51–52 Matric potential, 79 Matrix, 306 Mean residence times (MRT), atmospheric gases, 21 Mechanical turbulence, 74 Mediterranean shrublands, 138 climate of, 42 net primary production (NPP) of, 130 429 Mercury, cycling of, 220 Mesofauna and decomposition, 156 types of, 155 Mesophyll cells, 99, 102 Mesosphere, 23 Metamorphic rocks, 47 Metapopulations, 306 Methane global sources of, 343 methane budget, 342–343 production of, 173–174, 272, 306, 324, 343 Methanogens, 173–174 Methanotrophs, 174 Microbivores, in trophic system, 245 Microclimate, 32 Microfauna, types of, 155 Microflora, types of, 155 Micronutrients, 179 Mid-latitude deserts, climate of, 42 Milankovitch cycles, 35 Mineralization of organic matter, 152 See also Nitrogen mineralization Minerals and soils loss from soil, 57–58 primary minerals, 54 secondary, 54, 55–56 and soil formation, 49–50 soil transfer, 57 stability and weathering, 55 Mites, and decomposition, 156 Models, ecological, 325–330 Modulators, 12 Moist adiabatic lapse rate, 24 Moisture and decomposition, 161–162, 170 and nitrogen deposition, 201 Mollisols, 60, 61 Molybdenum, and nitrogen fixation, 200 Monsoon, landform/air flow conditions, 31 Montreal Protocol, 15–16 Mosses, and soil insulation, 271 Mountains and climate, 31–32 formation of, 47–48 and planetary waves, 27 PTEIndex 10/9/2002 430 7:40 PM Page 430 Index Mutualists effects on ecosystem processes, 273 in trophic system, 263 Mycorrhizae, 127, 134 and decomposition, 153–154, 167–168 dissolved organic nitrogen absorption, 204 as mutualists, 263 and nutrient uptake, 182–184 types of, 183 Mycorrhizosphere, 167–168 N NADP, and photosynthesis, 100, 105 Natural disturbances See Disturbances to ecosystem Negative feedbacks, effects on ecosystem, 13, 14 Nematodes, 253, 262 and decomposition, 155–156 Net biome production (NBP) and disturbance of biome, 146 estimation of, 144–145 Net ecosystem exchange (NEE), 146–147 ecosystem respiration, 146–147 estimation of, 144 latitudinal variations, 147–148 seasonal variations, 147–148 Net ecosystem production (NEP), 140–148 and carbon gain and loss imbalance, 145–146 carbon storage, 140–141 components of, 127–128 defined, 140 and disturbance of ecosystem, 145 ecological importance of, 140–141 lateral transfers, 140, 145 leaching, 141, 145 net ecosystem exchange (NEE), 146–147 and succession, 293–295 Net primary production (NPP), 123–124, 127–140 allocation of, 132–135 biomass differences in, 137–140 climate controls, 129–132, 134 defined, 123, 127 of major biomes, 130 and nutrient use, 189 and photosynthesis, 128–129 physiological controls, 128–129 and plant respiration, 126 relationship to gross primary production (GPP) in forests, 132 response to multiple resources, 133–134 and seasonal variation, 134–135, 138 tissue turnover, 136–137 Net radiation, 73, 75 New York City, water purification, 368 Niche, meaning of, La Niña, production of, 38–39, 40 El Niño/southern oscillation (ENSO), production of, 37–39, 40, 281 Nitrates and dentrification, 212 release and cycling of, 219 See also Nitrification Nitric oxide, atmospheric, 22 Nitrification, 202, 203, 207–210 assimilatory nitrate reduction, 209–210 control factors, 209 ecosystem variations, 209 effects on ecosystem, 207 nitrifying bacteria, 207 and nitrogen loss, 211 and oxygen, 208–209 pathways of, 208 rate, determinants of, 208 Nitrobacter, 207 Nitrogen, 198–215 adjustment in photosynthesis, 110–113, 133 atmospheric, 21, 198, 213 C : N ratio, 165, 206 in dead organic matter, 204 human activities, effects on, 201 and humus, 169 and lakes, 237 as limiting factor, 221 nitrogen cycling, 202–211 nitrogen deposition, 201–202 nitrogen fixation, 198–201 nitrogen loss, 211–215 N : P ratio, 187 and ocean nutrients, 231 and plant respiration, 126 Nitrogen-based defenses, plants, 249 Nitrogen cycle, 343–347 anthropogenic changes in, 344–347 sources and sinks of nitrogen, 345 Nitrogen cycling, 202–211 ammonia production, 204– 207 dissolved organic nitrogen (DON), 202–204 mineralization, 202–207 nitrification, 207–210 spatial factors, 210 terrestrial view, 203 time factors, 210–211 Nitrogen fixation, 198–201 in aquatic systems, 199, 200 ecosystem variations, 200–201 limiting elements, 200 Nitrogen-fixing crops, 345 Nitrogen-fixing microbes, 127, 134, 198–200 grazing on, effects of, 200–201 nitrogen-fixing invader, 268–269 types of, 198–200 Nitrogen loss, 211–215 ecological controls, 211–213 and erosion, 215 gaseous loss, 211 solution losses, 214–215 Nitrogen mineralization, 202– 207 and ammonia production, 204–207 C : N ratio for, 206–207 factors affecting process, 206–207 gross and net mineralization, 204–205 process of, 202–203, 205 Nitrogen saturation, 214, 346 Nitrosolobus, 207 Non-steady state mosaics, 309–310 Nonessential elements, cycling of, 220 Normalized difference vegetation index (NDVI), 118–119 PTEIndex 10/9/2002 7:40 PM Page 431 Index Northern Hemisphere atmospheric circulation, 26–27 seasons, 35, 41 N : P ratio, 187 Nuclear testing, negative effects of, 16 Nutrient cycling cations, cycling of, 219–220 element cycle interactions, 220–221 nitrogen inputs to ecosystem, 198–215 nutrient loss from plants, 191–194 nutrient movement to roots, 177–180 nutrient use by plants, 189–191 phosphorus inputs to ecosystem, 215–219 and succession, 296–298 sulfur inputs to ecosystem, 219 uptake mechanisms, 180–189 Nutrient loss, 191–194 and disturbances to ecosystem, 194 and herbivores, 193–194 and leaching, 193 senescence, 192–193 Nutrient productivity, 190 Nutrient uptake, 180–189 controlling factors, 180 and mycorrhizae, 182–184 and nutrient supply, 180–181 and root length, 181–182 and root longevity, 191 root uptake properties, 184–189 species variations, 185–188 Nutrient use, 189–191 and net primary production (NPP), 189 nutrient use efficiency (NUE), 190–191 supply versus concentration, 189 Nutrients sources of plant nutrients, 177 types of, 179 O Occluded phosphorus, 218 Oceans, 28–31, 228–236 and atmospheric circulation, 27–28 bloom, 233 carbon availability, 228–229 carbon cycling, 234–236 carbon dioxide in, 228–229, 335–337 circulation of, 29–31 compared to continents, 226 decomposition, 234–235 euphotic zone, 225, 229, 231, 234 hydrothermal vents, 230 light availability, 229–230 major currents, 30 nitrogen leaching into, 215 nutrient availability, 231–233 nutrient cycling, 233–234 overfishing, 14, 254, 273, 323 photosynthesis, 220 structure of, 28–29 temperature variations, 29 upwelling, nutrient-rich, 233 See also Aquatic organisms; Aquatic systems Ogallala aquifer, depletion of, 93 O horizon, 58 Oligotrophic, 247 Oligotrophic bogs, 48 Olivine, 55 Orographic effects, 31 Osmotic potential, 79 Overland flow, 80 Oxidation, 64 Oxidation-reduction, and soils, 64 Oxisols, 60, 61 Oxygen in aquatic systems, 226–227 atmospheric, 21 and nitrification, 208–209 Ozone atmospheric, 21, 22–23 production of, 346 Ozone hole, causes of, 15–16, 23 P Pacific North America (PNA) pattern, production of, 39–40 Pangaea, 48 Parent material, 11 soil formation, 47–48 Particulate organic carbon (POC), 335 Pastures, and climate, 33 Patches of landscape See Landscape heterogeneity 431 Pelagic ecosystems, 46, 224 See also Aquatic systems Percolation, 80 Periphyton, 238 Permafrost, 42, 61, 80 Permanent wilting point, 63–64, 81 Perturbation, 282 Phagocytosis, 155 Phenological specialization, 270–271 Phenology, 135 Phosphoenolpyruvate (PEP) carboxylase, 102, 104 Phosphorus, 215–219 availability in soils, 217–218 cycling of, 217 as limiting factor, 221 N : P ratio, 187 occluded phosphorus, 218 and ocean nutrients, 231 weathering and inputs, 215, 216, 217 Phosphorus cycle, 347–348 anthropogenic changes in, 347–348 Photo-oxidation, 107 Photoperiod, 135 Photoprotection, 100–101 Photosynthesis biochemical pathways, 98– 103 carbon dioxide fixation and water exchange, 102–103 carbon-fixation reactions, 99–102, 114–115 crassulacean acid metabolism (CAM) photosynthesis, 103–104 in dry environments, 102–103, 113–114, 115 and individual leaves, 101, 105–115 lakes, 236–238 light-harvesting reactions, 99–101 and net primary production (NPP), 128–129 oceans, 220 plant structures, 99, 100, 101, 102, 103 time course, 106 See also Gross primary production (GPP) PTEIndex 10/9/2002 432 7:40 PM Page 432 Index Photosynthesis adjustment mechanisms carbon dioxide gains, 133 carbon dioxide limitations, 105, 109–110, 133 light limitation, 105–109 nitrogen gains, 133 nitrogen limitation, 110–113, 133 to pollutants, 115 to temperature extremes, 114–115 to water limitation, 113–114 Photosynthetically active radiation (PAR), 106–107 Phototrophs, 199 Phreatophytes, 77 Phyllosphere decomposition, 157 Phyllosphere fungi, 137 Physical weathering, 54 Phytoplankton, 224–226 Picoplankton, 225 Planetary boundary layer (PBL), 23 height increase, 23 Planetary waves, 27 Plant respiration, 125–127 functions of, 125 and gross primary production (GPP), 127 maintenance respiration, 126–127 and net primary production (NPP), 126 sugar to biomass conversion, 125–126 Plants carbon inputs See Photosynthesis energy transfer, 8, 10, 13 hydrostatic pressures in, 79, 80 and linking of ecosystems, 320 nutrient use See Nutrient cycling scent of, 124 tissue turnover, 136–137 in trophic system See Trophic systems, plant-based water movement to roots, 81–82 water movements through, 83–89 See also Vegetation Plate tectonics, and rock formation, 48 Poikilothermic animals, 254 production efficiency, 256–257 Polar cell, 25 Pollen, as climate record, 36 Pollutants fertilizers and algae overgrowth, 14 and ozone hole, 15–16, 23 photosynthesis adjustment to, 115 Polyphenol formation, in humus, 169 Positive feedbacks, 13, 14 Potassium, release and cycling of, 219 Potential biota, meaning of, 11 Precession, 34 and seasons, 35 Precipitation process of, 32–33 regional changes in, 352 and runoff, 94 as water input, 77 Predation and ecosystem alteration, 273–274 lakes, 238 negative feedback of, 13 oceans, 233–234 scavengers, 263 selectivity in, 255 Pressure potential, 79 Prevailing winds, 26 Primary detritivores, in trophic system, 245 Primary minerals, 54 Primary producers, 244–245 Primary succession, 286, 288–291, 292–297 defined, 286 See also Succession Production efficiency, 256–257 Proteases, 204 Proteoid roots, 186 Protozoans, and decomposition, 155–156 Q Quantum yield of photosynthesis, 106 Quinone formation, in humus, 169 R Radiation See Solar radiation Radiatively active gases, 19 Rain shadow, 31 Redfield ratio, 187, 231 Redox potential, and soils, 64, 65 Redwood trees, 287, 293 longevity of, 283–284 Reflected radiation, 73 Reforestation, 322 Regolith, 59 Relative humidity, meaning of, 72 Residence time, 157, 190 Resilience of ecosystem, 282–283, 287 and management of ecosystem, 357–358 Resistance of system, 282 Resorption, nutrients, 152, 192 Resources, interactive processes related to, 12 Response of system, 282 Restoration of ecosystem, 360–361 Reynolds numbers, 225, 226 Rhizobium, 198, 199 Rhizosphere, 54 decomposition in, 154, 156, 159, 167 and nutrient uptake, 188–189 Rivers and streams, 238–242 carbon cycling, 240 horizontal flow, 242 nutrient cycling, 240–241 organisms of, 238 river continuum concept, 239–240 seasons and runoff, 94–95 spiraling, 241 terrestrial interface, 240 Rock cycle, events of, 47–48 Rocks as carbon source, 337 as sulfur source, 349 types of, 47 Rock weathering nitrogen inputs, 202 phosphorus inputs, 215, 216, 217 rock cycle, 47–48 soil profiles, 54–55 as sulfur source, 219 types of, 54 PTEIndex 10/9/2002 7:40 PM Page 433 Index Rocky Mountains, rain shadow, 31 Root caps, 189 Root cortex, 183 Root exudation, 123, 127, 128 Root hairs, 182 Roots cross-section of, 184 length, development of, 181–182 nutrient movement to, 177–180 nutrient uptake mechanisms, 182–189, 191 rooting depth, ecosystem effects, 82, 93, 269 water movement through, 83–86 water movement to, 81–82 water roots, depth of, 158 Rotation of Earth, and winds, 26–27 Rubisco, in photosynthesis, 99–100, 102, 104 Ruminants, assimilation efficiency, 256 Runoff meaning of, 93–94 process and effects of, 89, 93–95, 299 and succession, 299–300 S Salinization process of, 57 salt flats, 56 Salt flats, 56 Salt licks, 259 Salt marsh, 64 net primary production (NPP) of, 132 Sampling effects, 275 Sand, 61, 62, 80 Sapwood, 87 Savannas biomass distribution of, 138 climate of, 42 grazing lawns, 253 net primary production (NPP), 130 nitrogen fixation, 200 Scavengers, and food web, 263 Seasons and carbon gain, 109 and climate variability, 40–41 and decomposition, 158 and Earth’s precession, 35 and energy exchange, 77 and gross primary production (GPP), 120–121 lake turnover, 237 and net ecosystem exchange (NEE), 147–148 and net primary production (NPP), 134–135, 138 and nitrogen cycling, 210–211 and plant growth, 135 and resource capture, 270–271 and runoff, 94–95 trophic level interactions, 258–259 Secondary minerals, in soils, 54, 55–56 Secondary succession, 286, 291–292, 295, 297–298 defined, 286 See also Succession Sedimentary rocks, 47, 55, 202 Seed bank, 291 Seedling bank, 291 Senescence defined, 136 leaf life-span, 110–113, 135, 136–137 nutrient loss, 192–193 tissue turnover, 136–137 See also Decomposition Sensible heat flux, 19 Seston, 227 Shade leaves, 107 Shifting steady state mosaics, 309 Shortwave radiation, 18–19 Shredders, 239 Shrublands See Mediterranean shrublands Siderophores, 186 Silt, 61, 62 Sink strength, 128 Slash-and-burn agriculture, 311 Slopes and soil deposition, 49 and soil loss, 52 Slow variables, 302 Snow melt and runoff, 94 precipitation as, 90 Snow (ice) albedo feedback, 74 Soil acidified, 162 433 surface and decomposition, 158–159 water movements within, 80–81 water storage in, 78 Soil animals, decomposition by, 155–157 Soil disturbance, and decomposition, 163 Soil formation, 46–50 and animals, 50 and climate, 48 and human activities, 50 parent material, 47–48 time factors, 49–50 and topography, 48–49 and vegetation, 50 Soil horizons formation of, 56 types of, 58–59, 60 Soil loss, 50–53 and climate, 51 and glaciers, 53 instability of soil mass, 52 mass wasting, 51–52 and nitrogen loss, 215 and water flow, 52–53 and wind, 53 Soil minerals, 10 Soil orders, names/characteristics of, 59–61 Soil organic matter (SOM) carbon concentration, 98 carbon production, 124–125, 142–143 and decomposition, 166– 168 elements of, 64–65 humus, 169–170 litter, 58 soil transfer, 57 soil transformation, 54 and succession, 295 Soil profiles, 53–58 additions to soil, 54 mineral losses, 57–58 process of, 53 secondary minerals, 55–56 soil transfers, 56–57 weathering, 54–55 Soil properties anion absorption, 66 buffering capacity, 66 bulk density, 63 PTEIndex 10/9/2002 434 7:40 PM Page 434 Index Soil properties (cont.) cation exchange capacity (CEC), 65–66 and decomposition, 162–163 organic matter, 64–65 oxidation-reduction, 64 redox potential, 64, 65 structure, 62–63 texture, 61–62 water contents, 63–64 Soils functions in Earth system, 46 parent material, 11 Solar radiation absorption, and stratospheric ozone, 22–23 and atmospheric circulation, 24 emission of, 18–20 and energy balance, 20 energy budget, 73–74 at equator, 24 and evolution of climate, 34–35 light in ocean environment, 229–230 solar irradiance and seasons, 41 wavelength, 18–19 Solstices, time periods of, 35, 41 Solubility pump, 337 South America, El Niño, 38–39 Southern Hemisphere atmospheric circulation, 26–27 seasons, 35, 41 Spatial scaling, 325–330 Specialist herbivores, 255 Species composition, 266 Species effects on ecosystem on climate, 271–272 on disturbance regime, 272 diversity effects, 274–277 on nutrient turnover, 270 and phenological specialization, 270–271 on resources, 268–269 and species interactions, 268, 273–274 Species evenness, 266 Species richness, 266 Specific heat, 72 Specific leaf area (SLA), 111–113, 112 Specific root length (SRL), 182 Spodosols, 60, 61 Stand-replacing disturbances, 287 State factors and ecosystem ecology, 11 and ecosystem management, 358–359 and landscape heterogeneity, 307 types of, 11 Steady state ecosystems, Stem flow, 79, 193 Stems, water movement through, 86–88 Stomata, water loss from, 88–89, 92 Stomatal conductance and photosynthesis, 101–102, 105, 109, 110, 114 Stratosphere, 22–23 Streams See Rivers and streams Strontium, 16 Subduction, plates of Earth, 48 Suberin-coated cells, 184 Sublimation, and snow, 90 Subsidies, 247 Subtropical deserts climate of, 42 diversity in, 43 Succession, 8–9, 288–301 and carbon balance, 292–296 climax, 285 defined, 285 and disturbances to ecosystem, 285 ecosystem dynamics, 8–9 and ecosystem structure/composition, 288–292 energy exchange, 300–301 gap-phase succession, 287 life history patterns, 290 and nutrient cycling, 296–298 primary succession, 286, 288–291, 292–297 secondary succession, 286, 291–292, 295, 297– 298 species changes, 291–292 temperature of site, 300–301 trophic dynamics, 298–299 water exchange, 299–300 Succulents, water storage, 87–88 Sugar conversion, in plant respiration, 125–126 Sulfur atmospheric, 22 cycling of, 219 oceans, 230 weathering as source of, 219 Sulfur cycle, 348–350 and human activities, 349 Sun, radiation emission, 18–20 Sunflects, 106 Sun leaves, 107 Surface conductance, 91 Surface water, oceans, 28–29 Swidden agriculture, 311 Systems approach, ecosystem ecology, 8, 10 T Tamarix, 93 Teleconnections, 39 Temperate forests biomass distribution of, 138 climate of, 42 net primary production (NPP), 130 wet forests, 42 Temperature and decomposition, 159–161 extremes and photosynthesis, 114–115 and soil respiration, 160 of successional site, 300–301 Temperature inversions, 32 Temporal dynamics disturbance to ecosystem, 285–288 interannual variability, 281, 283 legacies, 283–285 long-term change, 283–285 succession, 288–301 temporal scaling, 301–302 Temporal scaling, approaches to, 301–303 Termites, and decomposition, 156, 272 Terrestrial Ecosystem Model (TEM), 326 Thermocline, 29 Thermohaline circulation, 29–30 Thermosphere, 23 Throughfall, 79, 193 Thylakoid membranes, and photosynthesis, 100 Tilt, 34, 35 Time and decomposition, 157–158, 164 PTEIndex 10/9/2002 7:40 PM Page 435 Index and ecosystem, 11 global warming, 36–37, 38 and nitrogen cycling, 210–211 soil formation, 49–50 See also Seasons; Temporal dynamics Time steps, 325 Tissue turnover, plants, 136–137 Top-down controls, 246, 273– 274 Topography and soil formation, 48–49 and soil loss, 51 Toposequence, meaning of, 11 Trade winds, 26, 31 Transpiration, 71 and photosynthesis, 101 Tree rings, as climate record, 36 Trophic cascades, 257–258 Trophic efficiency, 251 Trophic interactions, early research, 7–8 Trophic level example of, 245 production controls, 246 Trophic systems, detritus-based systems, 245, 257, 261 Trophic systems, plant-based, 246–261 assimilation efficiency, 256 biomass distribution, 250– 252 consumption efficiency, 253–256 ecosystem variations, 247–248 energy flow, controls on, 246–250, 254–255 in low-resource environments, 248–249 nutrient transfers, 259–261 plant defenses against consumption, 248–250 production efficiency, 256– 257 seasonal patterns, 258–259 and succession, 298–299 trophic cascades, 257–258 Trophic transfers, process of, 244–245 Tropical forests biomass distribution of, 138 climate of, 41–42 decomposition and uptake of nutrients, 159 diversity in, 42–43 dry forests, 41 and mineralization, 206 net primary production (NPP), 130, 131–132 nitrogen fixation, 200 wet forests, 41–43 Tropopause, 23, 26 Troposheric ozone, 22 Troposphere, 22 Tundra See Arctic tundra Turn over, of carbon, 335 U Uplift, plates of Earth, 48, 51 Upwelling, ocean nutrients, 233 Upwelling areas, oceans, 29 Urban areas, planetary boundary layer (PBL), 23 Urease, 204 Utisols, 61 V Valleys, and soil deposition, 49 Vapor pressure, meaning of, 73 Vapor pressure deficit (VPD), meaning of, 73 Vapor pressure gradient, 83 Vegetation albedo of, 32–34 carbon dioxide in, 337 and chemical weathering, 54–55 and climate, 32–34 deserts, 77 and evapotranspiration, 91– 92 normalized difference vegetation index (NDVI), 118–119 rooting depths by biome type, 82 and runoff, 95 and soil formation, 50 and soil transfers, 57 and water table, 93 See also Plants Vertisols, 61 Volatile emissions, plant respiration, 124, 127 Volcanic eruptions See Disturbances to ecosystem Vostok ice core, 35–36 435 W Water and carbon inputs See Photosynthesis and energy budget, 72–73 energy status of, 78–79 global sources of, 350–351 inputs to ecosystem, 77–78 leaving landscape, pathways for, 52–53 relationship to energy budgets, 72–73 soil, water contents of, 63–64 and soil loss, 51–53 soil storage of, 78 and soil transfers, 56–57 Water cycle, 350–354 and human activities, 351– 352 hydrologic problems related to, 352, 354 Water flow, and soil loss, 52–53 Water-holding capacity, 63–64 Water inputs, and water losses, 89, 92–93 Water losses, 89–95 evaporation, 89–90 evapotranspiration, 90–92 runoff, 93–95 and water inputs, 89, 92–93 water storage changes, 92–93 Water movements, 78–88 day versus night, 85 and nutrient uptake, 179–180 through plants, 83–89 principles of, 78–80 within soil, 80–81 from soil to roots, 81–82 Water potential, 63, 79, 81 Water purification, New York City, 368 Watershed study of, vegetation, effects of, 95 Water sources, isotopic signatures of, 84–85 Water use efficiency (WUE), photosynthesis, 113–114 Water vapor atmospheric, 22 and precipitation, 32–33 Water vapor density, meaning of, 72 Water vapor feedback, 71 PTEIndex 10/9/2002 436 7:40 PM Page 436 Index Wavelength and aquatic systems, 229 solar radiation, 18–19 Weather and currents, 31 tropospheric origination of, 23 Weathering See Rock weathering White-rot fungi, 153 Winds atmospheric circulation, 24–28 and soil loss, 53 wind-flow patterns, map of, 28 X Xanthophyll cycle, 100–101 Xylem, water movement through, 86 Z Zebra mussel, 238 PTE(Color Plate) 10/9/2002 7:41 PM Page Color Plate I Plate The global distribution of Earth’s major biomes and the seasonal patterns of monthly average temperature and precipitation at one representative site for each biome (Bailey 1998) Climate data are monthly averages of the entire period of record for selected sites through the year 2000 (http://www.ncdc.noaa.gov/ol/climite/ research/ghcn/ghcn.hlml) PTE(Color Plate) 10/9/2002 7:41 PM Page Color Plate II Plate The global pattern of mean annual temperature and total annual precipitation (New et al 1999) Temperature is highest at the equator and lowest at the poles and at high elevations (Reproduced with permission from the Atlas of the Biosphere .) PTE(Color Plate) 10/9/2002 7:41 PM Page Color Plate III Plate The global pattern of net primary productiviy (Foley et al 1996, Kucharik et al 2000) The patterns of productivity correlates more closely with precipitation than with temperature, including a strong role of moisture in regulating the productivity of the biosphere (Reproduced with permission from the Atlas of the Biosphere .) PTE(Color Plate) 10/9/2002 7:41 PM Page Color Plate IV -90 -60 -30 30 60 90 -180 ice