Annu. Rev. Ecol. Syst. 1992. 23:1-14 Copyright © 1992 by Annual Reviews Inc. All rights reserved GLOBAL ENVIRONMENTAL CHANGE: An Introduction Peter M. Vitousek Department of Biological Sciences, Stanford University, Stanford, California 94305 KEYWORDS: land use, human population, greenhouse gas, climate change, biological diversity INTRODUCTION This issue of the Annual Review of Ecology and Systematics includes a set of papers on the ecological causes and consequences of global environmental change, and on the methods by which these can be studied. Global studies are likely to be among the most important concerns of ecological research and teaching for some decades. While the Annual Review of Ecology and Systematics has published a number of papers in this area (cf 4, 7, 22, 41, 44), the Editorial Committee believed that it would be useful to bring together a Set of reviews selected to illustrate the breadth of ecologists’ contributions to research on global environmental change. This collection is designed to demonstrate that a very wide array of ecological research is crucial to the analysis of global change—and that both our basic work as ecologists and our contribution to the understanding of Earth as a system will be enhanced if we keep that relevance in mind. Earth is a dynamic system; global environmental change has always been a part of its functioning. A recent and prominent example is the glacial/inter- glacial cycles of the past 2 million years (12, 13, 16). The current interest in global change arises from the fact that some components of human-caused global change have reached a magnitude at least equal to that of natural changes—and the human-caused changes are often more rapid than and beyond the bounds of natural change, at least for the past millions of years. COMPONENTS OF GLOBAL ENVIRONMENTAL CHANGE Global changes are defined as those that alter the well-mixed fluid envelopes of the Earth system (the atmosphere and oceans) and hence are experienced 1 0066-4 162/92/1120-0001$02.002 VITOUSEK globally, and those that occur in discrete sites but are so widespread as to constitute a global change. Examples of the first category include: change in the composition of the atmosphere, climate change, decreased stratospheric ozone concentrations and increased ultraviolet input. The second type of global change is exemplified by land use change, loss of biological diversity, biological invasions, and changes in atmospheric chemistry. Each of these is discussed below. Change in the Composition of the Atmosphere Increases in global concentrations of carbon dioxide have been documented very clearly. Careful atmospheric measurements have been carried out since 1957; carbon dioxide concentrations increased from 315 ppm then to over 350 ppm in 1988 (29), and the rate of increase itself increased over that interval. Could this relatively short record have happened to coincide with a natural fluctuation in carbon dioxide concentrations? No—a record of earlier concentrations is preserved in polar ice, and analyses of cores taken from this ice demonstrate that concentrations were quite stable near 280 ppm for at least 1000 years, then began increasing around the year 1750 (63). A longer-term record covering the past 160,000 years (3) shows that there are natural fluctuations in carbon dioxide concentrations through glacial-intergla- cial cycles—but the recent increase began at a time when concentrations already were high and took them outside the bounds of known Pleistocene variation. The recent increase is now similar in magnitude to, and much more rapid than, the fluctuations occurring over glacial-interglacial time (Figure 1). There is no doubt that current increases in carbon dioxide are caused primarily by fossil fuel combustion and secondarily by changes in land use. C02 (ppm) Human population (b \o~COSSC«* 1000 years before present Figure 1 Natural variation in concentrations of carbon dioxide for the past 160,000 years from the Vostok ice core (3), together with recent anthropogenic increases in carbon dioxide concentrations and the growth of the human population.GLOBAL ENVIRONMENTAL CHANGE = 3 The timing and magnitude of fossil fuel combustion alone are more than sufficient to account for the global increase (51). Moreover, the atmospheric abundances of the carbon isotopes '°C and '*C relative to '7C have decreased over time in a pattern that reflects dilution of the atmospheric pool by carbon released from fossil fuel combustion (which is '*C-free and '*C-depleted) and loss of terrestrial biomass ('*C-depleted) (56, 57). The increased concentra- tions of carbon dioxide are expected to have substantial direct effects on plants, herbivores, and ecosystems (4, 38). A number of other gases are also increasing globally (40, 63). The wholly anthropogenic chlorofluorocarbons (CFCs) have been increasing most rapidly (66). Concentrations of methane have more than doubled since 1750; this increase is believed to be caused by a combination of agricultural and industrial activities (11). Nitrous oxide is increasing more slowly; the reasons for its increase are less certain, but they are probably primarily related to tropical land use change and agricultural activity (34). The history of increase in tropospheric concentrations of these four relatively stable gases is summarized in Figure 2; an excellent review of their sources and dynamics can be found in a recent Intergovernmental Panel on Climate Change (IPCC) report (63). z 3 E 360 1800 B = A B = ~ 340h 5 § 1400. = sz0h. S = = & 3007 i000 § 8 8 5 200] 5 ede 8 ¢ cS 2601+ 600 = oO oO S S 5 a 3 2 10 0.3 B ER 2 < § § 300 doz 2 S £ £ 5 = © 290] for 2 3 5 2 8 § 8 Q, 2001 : 7 0.0 2" 1750 1800 1850 1900 1950 2000 ¥e00 1850 1900 1950 2000 © Year Figure 2 Increases since 1750 in the concentrations of relatively stable greenhouse gases: carbon dioxide (A), methane (B), nitrous oxide (C), and chlorofluorocarbon-11 (D). From (27)4 VITOUSEK Climate Change The four gases whose status is summarized in Figure 2 are all greenhouse gases, in that they are transparent to incoming solar radiation but absorb outgoing infrared radiation (IR) emitted by Earth’s surface and atmosphere. The continuing increase in their concentrations therefore will enhance the naturally occurring greenhouse effect, which has maintained the average temperature of Earth’s surface more than 30°C warmer than it would have been in the absence of atmospheric absorption of infrared radiation (24). That the greenhouse effect will be enhanced by anthropogenic increases in these gases is wholly uncontroversial, and that this enhancement will cause global warming is nearly so. A recent “best estimate” for the temperature increase likely to result from a doubling of carbon dioxide (and equivalent forcing from the other gases) is 2.5°C globally, with greater changes towards the poles (27). This corresponds to about half of the average temperature change from full-glacial to interglacial conditions; its effect on the distribution and dynamics of natural and agricultural ecosystems would be substantial (6). However, more detailed predictions of the rate, magnitude, and distributions of changes in temperature and (particularly) precipitation contain substantial uncertainties. These reflect: (i) an imperfect understanding of some funda- mental processes in the climate system; (ii) the inherent complexity of the climate system and the difficulty of representing it in general circulation models, no matter how detailed and demanding these are; (iii) the existence of both positive and negative feedbacks in the response of the climate system to altered conditions—even where all of the feedbacks can be identified, it is difficult to determine their net effects; and (iv) the influence of offsetting human activities such as the generation of aerosols and cloud condensation nuclei from anthropogenic sulfur dioxide emissions (10). A number of current summaries of the certainties and uncertainties of climate change and the likely effects of climate change on ecosystems are available; these range from moderately technical articles (53) and books (52) to the IPCC Scientific Assessment discussed above (15, 35, 36). Decreased Stratospheric Ozone Concentrations and Increased Ultraviolet Input Ozone in the stratosphere absorbs incoming ultraviolet radiation (UV), thereby reducing the amount reaching Earth’s surface. Since 1974, it has been known that chlorofluorocarbons break down in the stratosphere, liberating chlorine and thereby catalyzing the breakdown of stratospheric ozone (37). These findings, and models based on them, sparked substantial discussion and debate—but the discovery in the mid-1980s of a growing springtime Antarctic ozone hole (20) came as a nasty surprise to all parties. Subsequent proof that CFCs cause Antarctic ozone destruction through a previously unsuspectedGLOBAL ENVIRONMENTAL CHANGE 5 series of processes (cf 45, 47) illustrates that global environmental change is not always a gradual, predictable process. More recently, significant stratospheric ozone depletion has been docu- mented over a much wider area of Earth. The consequent increase in UV radiation is likely to affect plants, ecosystems, and human health (9, 58), although to date experimental studies of these effects at realistic levels of increased UV are relatively sparse. This component of global environmental change is unique in that widespread concern over the effects of increased UV has led to an international agreement to eliminate CFC emissions globally by early in the next century (5). Another type of global environmental change occurs in enough discrete sites or regions to constitute a global change. Examples of this type are land use change, loss of biological diversity, biological invasions, and changes in atmospheric chemistry. Land Use Change Changes in land use are reviewed in this volume by Meyer & Turner; they are discussed only briefly here. Unlike the atmosphere or oceans, terrestrial ecosystems do not mix on any time scale relevant to global change—and hence any global effect of land use change must be a consequence of many local changes. In fact, human-caused land use change alters enough local ecosys- tems substantially enough to affect global budgets of greenhouse gases (as discussed above; also see 26), and to alter regional climate (17, 55), regional atmospheric chemistry (2, 30), and the dynamics of major river systems (60). Nevertheless, its most important global effect may be simply the alteration of so many local ecosystems (59); some major ecosystem-types (such as tall grass prairie and tropical deciduous forests) have virtually disappeared (cf 28), and many more have been significantly altered and/or fragmented. Loss of Biological Diversity Human activity is now causing the extinction of species and genetically distinct populations at a rate far above background (65). Most species are ephemeral features of the Earth system; under normal conditions (between rare episodes of mass extinction), an average species lasts perhaps 10 million years (19). Human activity is now making many species much more ephemeral; observations of well-studied groups such as birds, together with calculations based on species/area curves, suggest that current rates of extinction are several orders of magnitude above background rates. So far the increase is due primarily to land use change (secondarily to biological invasions, below), but other components of global environmental change are likely to contribute increasingly in the future. If these increased rates of extinction continue, the next few centuries will6 VITOUSEK witness a mass extinction of a magnitude not seen since the Cretaceous-Ter- tiary boundary—and this time, there will be no need to argue about the cause of the episode. The causes, rates, and consequences of the loss of biological diversity at the species and population levels are discussed briefly in Ehrlich & Wilson (19), and at greater length in Wilson & Peter (65). Biological Invasions One consequence of humanity’s extraordinary mobility is an increase in the mobility of numerous species that we transport either deliberately or inadvertently. The resultant biological invasions reduce the biological distinc- tiveness of the various continents and islands, leading toward a homogenization of the biota of Earth (18, 39). This component of global environmental change, and its consequences for the composition, structure, and functioning of terrestrial ecosystems, is reviewed in this issue by D’ Antonio & Vitousek. Changes in Atmospheric Chemistry A syndrome of elevated tropospheric ozone concentrations, acidic precipita- tion, and elevated nitrogen deposition occurs over most economically devel- oped regions of Earth. Similar changes are now being observed seasonally in developing tropical regions (21). Increases in tropospheric ozone are driven by emissions of reactive nitrogen oxide gases from internal combustion engines, agriculture, and other human activities (43); biomass burning is the most important source in developing tropical regions (2, 30). These nitrogen oxides interact with anthropogenic or natural hydrocarbons and carbon monoxide to lead to the photochemical production of tropospheric ozone (14, 32). Acid precipitation (and dry deposition) results from the oxidation of nitrogen and sulfur oxides to nitric and sulfuric acid, and their subsequent rain-out or deposition downwind. This deposition, and that of ammonia emitted from agricultural systems, can increase atmospheric inputs of nitrogen several-fold for hundreds to thousands of kilometers downwind of highly developed regions. These alterations to regional atmospheric chemistry occur more-or-less independently in a number of large regions; they can therefore be considered a global environmental change. The effects of altered atmospheric chemistry include threats to human health (23), observed reductions in agricultural productivity (46), and changes in the dynamics of terrestrial and aquatic ecosystems (1, 41, 50 54). IMPORTANCE AND INTERACTIONS OF THE COMPONENTS The relative importance of the various components of global environmental change is a matter for debate. A number of factors must go into any suchGLOBAL ENVIRONMENTAL CHANGE = 7 weighting, including the relative impact of plausible changes in each component, the relative certainty of that change, and the reversibility of change. Of these, the relative certainty of change can be addressed straight- forwardly. Changes in the atmosphere, especially increased carbon dioxide, methane, and nitrous oxide concentrations in the troposphere and decreased ozone concentrations in the stratosphere, already are well documented; globally significant land use change clearly has occurred; and an ongoing loss of biological diversity can be inferred with confidence. On the other hand, human-caused global climate change has not been detected unequivocally as yet (64). Our understanding of the climate system is sufficient to say that it will occur, but the complexity of that system is such that the magnitude, timing, and geographic distribution of change all remain uncertain. The reversibility of components of global environmental change—the rate at which they would return to previous levels if human forcing were reduced or eliminated—differs widely. The greenhouse gas methane has an atmospheric lifetime of about a decade, and so its concentrations could return to background levels relatively rapidly. The other major greenhouse gases have effective atmospheric lifetimes in excess of a century (31). Reversing the effects of land use change is a successional process (decades to centuries)—although where soils have been degraded substantially, some millennia may be required for recovery (assuming, of course, that it is feasible to take land out of food production). Extinction, however, is much less reversible than the other components. While overall levels of diversity might be reestablished in a very few million years following an episode of extinction, the loss of particular species and the information they contain is wholly irreversible. In any case, except for CFCs and ozone depletion, reversibility is a moot point. Most components of global environmental change are becoming more important rather than less so, and some will continue to do so for the foreseeable future. The probable impact of the various components of global change is more difficult to assess, in part because it is dependent on the time scale of concern. Ultimately, climate change probably has the greatest potential to alter the functioning of the Earth system; its direct effects on natural and managed systems ultimately could become overwhelming, and it interacts strongly with most of the other components of global change, as discussed below. It is particularly worrisome that climate change may not be a gradual phenomenon; rather, the Earth system may switch rapidly from one climate mode to another (8). Nevertheless, the major effects of climate change are mostly in the future, while most of the others are already with us. There is a reasonably strong consensus among terrestrial ecologists that for the next several decades, land use change is likely to be the most certain and the most significant component of global change, followed by changes in the composition of the atmosphere. Analyses of the causes and significance of global environmental change are complicated greatly by the fact that many of the components of change8 VITOUSEK interact with each other directly and are modified further by feedbacks from their effects on ecosystems. For example, the increasing concentrations of carbon dioxide will drive climatic change by enhancing the greenhouse effect, which in turn is likely to alter the balance between rates of net primary production and decomposition in terrestrial ecosystems (42, 48, 49). This change can feed back to alter carbon dioxide concentrations in the atmo- sphere—if this process is not offset by direct effects of elevated carbon dioxide on carbon storage (4), or the indirect effects of climatic warming on the biological availability of soil nitrogen (42, 49). Similarly, land use change is the major process responsible for the extinction of species and genetically distinct populations (so far), but the loss of genetic diversity is itself a global change that could have significant effects on land use (19). The complexity of causal interactions among components of global change and their consequences, together with our uncertainty concerning some of these effects, makes simple, universal statements concerning chains of causation difficult. Nevertheless, there is no uncertainty concerning the ultimate cause of most current global change—it occurs because the scale of human activity has become large relative not just to that of other species, but to the flow of energy and materials on a global scale. For example, humanity consumes less than 1% of the terrestrial net primary productivity of Earth but in doing so dominates or destroys nearly 40% of the total (61). All of the other tens of millions of species must either adapt to us or subsist on the remainder. Our activity more than doubles rates of nitrogen fixation globally (62) and alters aspects of the global sulfur cycle to a greater extent (10). Certain components of the global influence of humanity are amenable to technological fixes—for example, we can control chlorofluorocarbon emis- sions, and hence ultimately stratospheric ozone depletion, without altering our way of life fundamentally. Nevertheless, the human population will continue to cause—but not control—global environmental change as long as our population and level of activity loom so large on the planet. ECOLOGY AND GLOBAL CHANGE The reviews in this volume cover diverse aspects of global environmental change. Webb & Bartlein examine the rich record of past global change—a record that helps place the ongoing human-driven changes in context and also provides a basis for testing the validity of modern climate models. Much of this record is a biological history, one that is read through an understanding of the ecology of marine foraminifera or the pollen of terrestrial plants. D’Antonio & Vitousek consider an often-neglected component of global environmental change—the loss of the regional distinctiveness of Earth’s biota brought about by human-caused breakdown of biogeographic barriers toGLOBAL ENVIRONMENTAL CHANGE 9 species dispersal. They use the widespread introduction of exotic grasses, often from Africa, as an illustration. Invasions by these grasses are driven in part by physiological and demographic characteristics of the grasses themselves and in part by biotic interactions in the invaded communities. However, grass invasions have substantial, interactive effects on fire regimes and other aspects of ecosystem function; they thereby represent a significant threat to the maintenance of biological diversity and to regional biogeochemistry. The next three reviews, by Field et al, Carpenter et al, and Smith & Buddemeier, examine the interactive effects of global change on terrestrial ecosystems, freshwater ecosystems, and coral reefs respectively. All three demonstrate that ecosystems are far from passive participants in global environmental change. Rather, changes to ecosystems caused by one process can alter other components of global change. For example, land use change in both upland and wetland systems alters microbial processes that cause emissions of radiatively active (greenhouse) and chemically reactive trace gases. These in turn can drive changes in the composition of the atmosphere and ultimately in global climate. In addition, Smith & Buddemeier suggest that while substantial attention has been paid to the effects of climate change on coral reefs, the effects of land use change (sedimentation and nutrient loading) and elevated carbon dioxide may be more significant for the next several decades. This is not to downplay climate change, which is a real concern, but to suggest that the other components of global environmental change may be even more important. The similarity of their ranking to that of terrestrial ecologists (above) is striking. Wessman discusses a problem that is common to all scientific efforts to evaluate global phenomena—the disparity between the spatial and temporal scale of most ecological measurements and that of the Earth system that we seek to analyze. How can we “scale up” from local measurements to global dynamics? Wessman illustrates how this disparity can be addressed in ecological theory and through a variety of technological developments that include satellite- and aircraft-based remote sensing, coarse-scale measure- ments of gas fluxes, and global tracer models. These new technologies have made a true science of the biosphere into an attainable goal. Shugart et al discuss a set of models of plant community dynamics that is based on the replacement dynamics of individual trees. They consider how physiological and ecosystem processes have been and are being incorporated into these models. The resultant syntheses can be used to evaluate the effects of interacting components of global change on the distribution and dynamics of terrestrial ecosystems. The models are also useful in the analysis of regional and global feedbacks resulting from alterations to ecosystems. Because these models are based on the mechanisms that control species replacement and10 VITOUSEK consequent changes in ecosystem function, they are more useful for dealing with wholly novel conditions of land use, climate, and atmospheric composi- tion than are correlative models. Finally, Meyer & Turner evaluate global changes in land use and land cover, and demonstrate the importance of understanding the human dimension of global environmental change. They point to the distinction between land conversion (often the focus of ecologists) and agricultural intensification; the latter may be more significant in its global consequences. They also discuss the mixed evidence for the role that human population growth per se plays in driving land use change. Most importantly, they show that an understanding of the overall causes of land use change, and hence any rational attempt to affect this most important component of global environmental change, must be based on an understanding of the reasons for the human actions that drive it. Social science, and social scientists, therefore must be integral parts of any global change research program. A few other reviews were commissioned but for various reasons not received; as with any collection, the ones that got away are particularly missed. These included reviews about the use of globally distributed measurements and models to determine the global sources and sinks of carbon dioxide, the continental-scale analysis of ecosystem function, the current rate of loss of biological diversity, and pelagic marine ecosystems. Even if these reviews had been included, the papers in this volume would be far from a complete introduction to global environmental change. Further information can be obtained from the many references cited herein. Readers interested in the directions taken by global research programs concerned with ecological aspects of global change should consult the planning and opera- tional documents of the International Geosphere-Biosphere Program (IGBP). IGBP is the coordinating body for a number of global projects that include Global Change in Terrestrial Ecosystems (GCTE), International Global Atmospheric Chemistry project (IGAC), Joint Global Ocean Flux Study (JGOFS), Global Changes of the Past (PAGES), Biospheric Aspects of the Hydrologic Cycle (BAHC), and Land-Ocean Interactions in the Coastal Zones (LOICZ). All of these include greater or lesser amounts of ecological research in their core program. The addresses of the project offices are appended to this introduction. In addition, the Ecological Society of America has suggested a set of research priorities that includes a focus on ecological causes and conse- quences of global environmental change, on biological diversity, and on the development of sustainable human uses of the biosphere. Their document (33) includes a rich array of suggestions for ecological research that will further the understanding, and perhaps ultimately the management, of global environ- mental change.GLOBAL ENVIRONMENTAL CHANGE i ACKNOWLEDGMENTS I thank J. M. Melillo for his contribution to putting together the set of reviews, and C. B. Field, H. A. Mooney, and A. R. Townsend for comments on an earlier draft. C. Nakashima prepared the manuscript for publication. Research was supported by a Pew Foundation Fellowship. Appendix: Addresses Of International Geosphere-Biosphere Program Core Project Offices Global Change in Terrestrial Ecosystems Dr. Brian Walker CSIRO Division of Wildlife and Ecology P. O. Box 84 Lyneham, Canberra, ACT 2602 Australia Joint Global Ocean Flux Study Dr. Geoffrey Evans Institut fiir Meereskunde Diisternbrooker Weg 20 D-2300 Kiel, Germany International Global Atmospheric Chemistry Project Professor Ronald Prinn Department of Earth, Atmosphere, and Planetary Sciences Room 54-1312 Massachusetts Institute of Technology Cambridge, MA 02139 USA Global Changes of the Past Dr. Hans Oeschger Institute of Physics University of Bern Silderstrasse 5 CH-3012 Bern, Switzerland Land-Ocean Interactions in the Coastal Zone Chair, Dr. Patrick Holligan Natural Environmental Research Council Plymouth Marine Laboratory Prospect Place West-Hoe, Plymouth PL1 3DH, United Kingdom12 Biospheric Aspects of the Hydrologic Cycle VITOUSEK Institut fiir Meteorologie Freie Universitat Berlin Dietrich-Schafer-Weg 6-10 D-1000 Berlin 41, Germany Literature Cited 10 u B Aber, J.D.,Nadelhoffer, K.J., Steudler, P.. Melillo, J. M. 1989. 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