10.

Rising atmospheric CO2 and global climate change:

responses and management implications for grazing lands

Jack A. Morgan1

Abstract

Concentrations of the greenhouse gases CO2, CH4 and N2O are increasing in the
atmosphere, and evidence suggests that this is already causing significant changes in
the Earths climate. Results from numerous investigations indicate that increased
atmospheric CO2 often leads to increased plant production, greater water use
efficiency and higher soil water content, but may also result in reduced forage
quality, with consequent lowered digestibility, particularly in nutrient-limited
systems. Predicted future increases in temperature will have varied effects on plants
and animals through alterations in growing season length, metabolism, energy
dynamics and system water relations. The potential for increased plant productivity
due to CO2 fertilization as temperatures increase will often be greater in present-day
mesic, temperate and cold ecosystems, but may be offset to the extent temperatures
increase and conditions become more desiccating in world regions that are predicted
to become more drought prone. Rainfall is predicted to increase in some areas, but
may not result in more production in cases where increased storm intensity will lead
to more runoff and erosion. These basic responses will be modulated by changes in
plant and animal species composition that result from competitive species
interactions in native ecosystem responses to global climate change, or are the
consequence of management strategies to select better adapted plant and animal
species or genotypes. Management options for adapting to global climate change are
discussed for intensively managed improved pastures as well as for native
rangelands.

Introduction

We live in a time when the imprint of human activities can be detected in virtually every corner of
the globe. A rapidly expanding human population is the main driver behind this, causing dramatic
changes in the Earths ecosystems. Much of this transformation can be viewed as a success story,
like the green revolution that combined advances in plant breeding, irrigation and fertilizer
technology to significantly increase agricultural productivity in developing nations. Yet these new
technologies have often created or been accompanied by new problems that have required
attention and new innovation.

Most such development-related environmental problems (e.g. surface and groundwater pollution
by N fertilization, salinization, erosion) were observed primarily in the local neighbourhood. In
more recent times, the environmental consequences of human development have been more far-
reaching. Such is the nature of the increasing concentrations of trace gases in Earths atmosphere,
and associated global climate change. We cannot see these gases that are being emitted by
human activities in greater quantities now than in any recent time, yet there is increasing
evidence that they may have already caused significant changes in Earths climate and
ecosystems, and are likely to induce even greater changes in the twenty-first century. The
invisibility of greenhouse gas emissions and their global reach create unique problems: the issue is
unappreciated by many, who, not seeing the problem, are sceptical; trace gas emissions from one
country mix in the atmosphere and affect the entire world, thereby complicating control
strategies; and the complex interactions among greenhouse gases and their varied effects on
earths climate and ecosystems confound simple predictions of future consequences.

In this chapter, some of the evidence and explanations of trace greenhouse gas emissions and
consequences for climate change and grassland responses are reviewed. The ecological and
management implications for grazing lands will also be discussed, focusing primarily on altered
CO2 concentrations, increased temperature and altered precipitation patterns. The discussion
includes both native rangeland systems and more productive grasslands. Rangelands are natural
or semi-natural areas of typically low productivity and are grazed by wild and domesticated
animals (Polley et al., 2000). In contrast, productive grasslands are largely created by human
activities (Nsberger, Blum and Fuhrer, 2000), and so are more intensely managed than
rangelands. They range from intensively managed monocultures to species-rich natural or semi-
natural grasslands. In general they are considerably less limited by precipitation than rangelands,
accounting for their greater productivity.

The case for global climate change

The anthropogenic release of significant quantities of greenhouse gases (CO2, CH4 and N2O) into
the atmosphere began in the Industrial Age, and continues today at increasingly higher rates
(Albritton et al., 2001). Greenhouse gases reduce the efficiency with which the earth radiates its
energy into space, and their increased atmospheric concentrations warm the lower atmosphere
and surface of the planet. The warming potential of these gases can be expressed as radiative
forcing. Figure 10.1 illustrates how increases in atmospheric CO2, CH4 and N2O concentrations
over the past 200 years have increased the respective radiative forcings of these greenhouse
gases. Increases in atmospheric SO2 (Figure 10.1), also a result of industrialization, cause the
opposite effect, since they are precursors to aerosols, and result in cooling through cloud
formation and reflectance of solar radiation back into space. Aerosols have a much shorter lifetime
(days to weeks) compared with most greenhouse gases (decades to centuries), but can have
important effects on radiative transfer. Despite the cooling due to aerosols, the net effect of gas
emissions is one of warming.

Greenhouse gas emissions resulting from human activities occur against a background of variable
solar radiation cycles, natural emissions of greenhouse gases (volcanic activity), and other natural
changes in climate forcing unrelated to greenhouse gases, like the El Nio-Southern Oscillation
(ENSO) phenomenon (ocean-atmospheric interactions in the Pacific tropics, which affect world
climates). Evaluating the influence of anthropogenic gas emissions on past and projected future
climates must account for these other natural anomalies. A modelling exercise by Mitchell et al.
(2001) included both anthropogenic and natural forcings to evaluate the performance of a coupled
ocean-atmosphere climate model; the model performed well when compared with measured
temperature anomalies from the late 1800s until 2000 (Figure 10.2). The results suggest we may
have a good degree of confidence in our ability to predict future scenarios climate change induced
by greenhouse gases.

a)
 b

Figure 10.1 Records of changes in atmospheric gases. (a) Atmospheric concentrations

of CO2, CH4 and N2O over the past 1 000 years. Ice core data for several sites in
Antarctica and Greenland (shown by different symbols) are supplemented with data from
 atmospheric samples taken over the past few decades (shown by line for CO2 or
incorporated into the curve for CH4). The estimated radiative forcing is indicated on
right-hand scale. (b) Sulphate concentration in several Greenland ice cores and total SO2
emissions from sources in the USA and Europe (crosses).

Source: Figure reprinted with permission from the Intergovernmental Panel on Climate Change (Albritton
et al., 2001; Figure 8).

Figure 10.2 Global mean surface temperature anomalies relative to the 1880 and
1920 mean from the instrumental record (red line) compared with ensembles of four
simulations with a coupled ocean-atmosphere climate model (black line). Simulations
include both anthropogenic and natural forcings.
Source: Figure reprinted with permission from the Intergovernmental Panel on Climate Change (Albritton
et al., 2001; Figure 15c).

Table 10.1 Estimates of confidence in observed and projected changes in extreme weather and
climate events. The table depicts an assessment of confidence in observed changes in extremes of
weather and climate during the latter half of the twentieth century (left column) and in projected
changes during the twenty-first century (right column). This assessment relies on observational
and modelling studies, as well as physical plausibility of future projections across all commonly
used scenarios and is based on expert judgment. Table reprinted with permission from the
Intergovernmental Panel on Climate Change.

Confidence in
Confidence in projected
observed changes
Changes in phenomenon changes
(latter half of the
(during the 21st century)
20th century)
Likely Higher maximum Very likely
temperatures and more hot
days over nearly all land
areas
Very likely Higher minimum Very likely
temperatures, fewer cold
days and frost days over
nearly all land areas

Very likely Reduced diurnal temperature Very likely

 Very likely Reduced diurnal temperature Very likely
range over most land areas
Likely, over many areas Increase of heat index over Very likely, over most areas
land areas(1)
Likely, over many More intense precipitation Very likely, over many areas
Northern Hemisphere events(2)
mid- to high-latitude
land areas
Likely, in a few areas Increased summer Likely, over most mid-latitude
continental drying and continental interiors (lack of
associated risk of drought consistent projections in other
areas)
Not observed in the few Increase in tropical cyclone Likely, over some areas
analyses available peak wind intensities(3)
Insufficient data for Increase in tropical cyclone Likely, over some areas
assessment mean and peak precipitation
intensities(3)

NOTES: (1) Heat index: A combination of temperature and humidity that measures effects on
human comfort.
(2) For other areas there are either insufficient data or conflicting analyses.
(3) Past and future changes in tropical cyclone location and frequency are uncertain.

Source: Albritton et al., 2001; Table 4.

The construction of future scenarios of climate change relies on projected patterns in

demographic, economic and technical forces that affect greenhouse gas and sulphur emissions.
Since it is impossible to develop one or even several scenarios that capture the many possibilities
for Earths continued human development, the Intergovernmental Panel on Climate Change
utilizes up to 40 different scenarios as evaluation tools to predict a range of possible future
climates. In their latest report, Albritton et al. (2001) present results from numerous models,
which were run for 35 different scenarios, and concluded that globally averaged surface
temperatures would increase by between 1.4 and 5.8C from 1990 to 2100, with a wider range
and warmer temperatures for terrestrial surfaces, especially in winter and in the Northern
Hemisphere, where warming is predicted to be maximal. A similar exercise was done for
precipitation patterns, and the results suggest increased summer and winter precipitation at
higher latitudes. Increased winter precipitation is also predicted over northern mid-latitudes,
tropical Africa and Antarctica, and in the summer for southern and eastern Asia. Decreases in
winter precipitation are predicted for Australia, Central America and southern Africa.

A summary of some of the major predictions and confidences in them are given in Table 10.1. The
models predict warmer global temperatures, more hot days over most land areas, increased heat
index, and more intense precipitation events. Since more intense storms will lead to increased
surface runoff and erosion (Campbell, Stafford Smith and McKeon, 1997), increased rainfall in
some areas may not translate into more mesic conditions. In fact, with warming added to the
picture, there will probably be increased continental drying and associated risk of drought
(Table 10.1).

Primary Production Responses

Carbon dioxide

Carbon dioxide (CO2) is a substrate for photosynthesis. Increases in its concentration above
present atmospheric levels result in higher photosynthesis rates in plants and plant communities,
and to a lesser extent, lead to greater aboveground primary production (Drake, Gonzlez-Meler
and Long, 1997; Jackson et al., 1994; Lawlor and Mitchell, 1991; Mielnick et al., 2001; Morgan et
al., 2001, 2004; Owensby et al., 1996). However, the degree of plant response to CO2 can vary
between species or environmental conditions (Porter, 1993; Wand et al., 1999), so information on
species or functional group responses is important in predicting responses of plant communities
and ecosystems to rising atmospheric CO2 concentrations.

Differences in photosynthetic pathway may be a useful indicator of a plants potential to respond

to CO2. Plants with the C3 photosynthetic pathway are the most abundant and widely distributed
class of plants, and have a photosynthetic metabolism that is unsaturated at present atmospheric
CO2 concentrations. In contrast, photosynthesis of the second most abundant class of plants,
those with the C4 pathway, is saturated near present atmospheric CO2 concentrations of near

370 mlitre litre-1 air. Because of these differing photosynthetic sensitivities to CO2, early
predictions were that increasing CO2 would tend to give C3 plants a photosynthetic advantage
over C4 plants in future CO2-enriched atmospheres. (Bazzaz, 1990; Pearcy and Ehleringer, 1984).
However, subsequent experimentation revealed that, while the relative photosynthetic and growth
responses are on average less in C4 than C3 species, the differences are often not as great as
expected based on their underlying biochemistry (Bowes, 1993; Porter, 1993; Wand et al., 1999).

There are a couple of explanations of why elevated CO2 does not induce large differences in
photosynthetic and growth responses between C3 and C4 plants. First and foremost, raising CO2
induces stomatal closure in most herbaceous species, regardless of photosynthetic class (Field,
Jackson and Mooney, 1995; Kimball and Idso, 1983; Wand et al., 1999). This reduces
transpiration, resulting in higher plant and soil water potentials, and results in enhanced plant
biomass yields through improved water use efficiency (Drake, Gonzlez-Meler and Long, 1997;
Knapp, Hamerlynck and Owensby, 1993; Morgan et al., 2001; Niklaus, Spinnler and Krner, 1998;
Owensby et al., 1997; Sindhj et al., 2000; Volk, Niklaus and Krner, 2000). Such water relations
responses may be most important in native grassland and rangelands, where water scarcity is an
important limiting factor in production. Second, photosynthetic acclimation, a downward
adjustment of photosynthetic capacity in C3 plants exposed long-term to elevated CO2, can reduce
or even eliminate the photosynthetic enhancement achieved under elevated CO2 (Lee et al., 2001;
Sage, Sharkey and Seeman, 1989; Sage, 1994). Acclimation lessens CO2 response differences
between C3 and C4 species.

Some plants may respond more to CO2 due to different strategies in balancing C assimilation and
its utilization for growth and maintenance. Porter (1993) noted that herbaceous C3 crop plants
were more responsive than wild C3 species to CO2 enrichment, and also that CO2-induced growth
responses were greater in fast-growing compared with slow-growing wild species. He suggested
that plants with stronger C sinks may transport reduced C compounds more effectively away from
leaves and reduce metabolic feedbacks that lower photosynthetic activity, and, ultimately, reduce
growth response to CO2.

Legumes are one of the most responsive functional groups to CO2, in part because of their ability
to fix atmospheric N (Hebeisen et al., 1997; Jablonski, Wang and Curtis, 2002; Lscher et al.,
2000; Porter, 1993; Reich et al., 2001; Wand et al., 1999). Insufficient N appears to be one of the
major causes behind photosynthetic acclimation and reduced growth responses of plants to
elevated CO2 (Lee et al., 2001; Morgan et al., 2001; Sage, Sharkey and Seeman, 1989), so
legumes may have an advantage over non-legumes by virtue of their ability to fix atmospheric N
into organic compounds. Not only does the N fixation benefit the plant by providing a strong sink
for carbohydrates, but it also provides the N needed for biosynthesis of proteins, nucleic acids and
other vital plant compounds. However, legumes may not be as responsive to CO2 enrichment in
soils with low available phosphorus (Stcklin, Schweizer and Krner, 1998; Warwick, Taylor and
Blum, 1998), or when soil N is in abundant supply (Geeske et al., 2001).

As a group, forbs tend to be more responsive than grasses (Leadley et al., 1999; Morgan et al.,
2001; Owensby et al., 1999; Porter, 1993; Potvin and Vasseur, 1997; Reich et al., 2001;
Teyssonneyre et al., 2002). However, no satisfactory explanation has yet been proposed to
explain apparent high CO2 sensitivity in this diverse group of plants.

Growth of plants at elevated CO2 may also affect their respiration. This has important
consequences for the atmosphere as well as for plants, since as much as 50 percent of C fixed in
C3 photosynthesis may be respired back into the atmosphere (Farrar, 1985). There is some
evidence that plant respiration may be reduced in CO2-enriched environments (Bunce, 1990;
Drake, Gonzlez-Meler and Long, 1997; Rogers et al., 1997). However, since CO2 often increases
the root : shoot ratio in plants, and root systems have higher specific respiration rates than
shoots, whole plant respiration may tend to increase as CO2 rises (Rogers et al., 1997).
Unfortunately, much plant respiration research has not been conducted in a manner to evaluate
the particular mechanism behind the plant responses (Rogers et al., 1997), or to evaluate yet
whether interspecific differences may be an important aspect of variable species responses to CO2.

Temperature and CO2

Temperature is a significant factor in setting the boundaries on growing seasons and plant
communities, as well as determining growth rates, plant developmental responses and the
availability and utilization of resources like water, radiation and nutrients. Warmer climates will
lengthen growing seasons and will accelerate plant development; growth rate will increase up to a
point, and then decline as super-optimal temperatures are reached. A meta-analysis by Rustad et
al. (2001) suggests that global warming may enhance primary production in this century by an
average of 19 percent in tundra, grassland and forested biomes, and the response is likely to be
greater in the colder ecosystems, particularly in the tundra.

There is considerable evidence that CO2 and temperature responses are not additive, so it is
useful to understand warming in the context of rising CO2. For instance, the stimulatory effect of
CO2 enrichment on plant growth is often enhanced at warmer temperatures (Cure, 1985; Greer et
al., 2000). For C3 plants this stimulation may be related to CO2-induced changes in
photorespiration (Long, 1991) and activity of RuBisco, the enzyme responsible for fixation of CO2
into organic compounds (Bowes, 1993); changes in both tend to increase photosynthesis more at
high than low temperatures (Greer et al., 2000; Sage, Sharkey and Seeman, 1989). For many
plants, warmer temperatures enhance carbohydrate sink demand (Arp, 1991; Farrar and Williams,
1991), which may stimulate the CO2 growth response (Greer et al., 2000; Newton et al., 1994).
For these reasons, the effect of elevated CO2 on primary production in temperate pastures is likely
to be highly seasonal (Newton et al., 1994; Nsberger, Blum and Fuhrer, 2000).

Many of the present predictions for combined CO2 and temperature responses of rangelands,
mostly based on simulation models, suggest increasing CO2 and warming will lead to greater
primary production of most grasslands (Baker et al., 1993; Coughenour and Chen, 1997; Neilson
et al., 1998; Parton et al., 1995; Riedo, Gyalistras and Fuhrer, 2000, 2001), although the degree
of response varies considerably depending on the particular ecosystem and other environment
conditions. Warmer temperatures may be especially important at enhancing relative CO2
production responses at high altitudes and at high- and mid-latitude regions where low
temperatures currently limit CO2 production responses (Krner, 1996; Riedo, Gyalistras and
Fuhrer, 2001; Rounsevell, Brignall and Siddons, 1996).

Precipitation, temperature and CO2

Ninety percent of the variability in rangeland primary production is accounted for by variation in
annual precipitation (Campbell, Stafford Smith and McKeon, 1997). Production is therefore likely
to follow any changes in precipitation patterns to the extent they impact upon seasonal soil water
dynamics. In more mesic and productive grasslands, the consequences of altered rainfall will
probably be less, but will still be important since seasonal productivity is commonly limited by
distribution and amounts of precipitation (Knapp, Briggs and Koelliker, 2001).

While increased precipitation in many parts of the world may lead to more production, these
changes must be considered in the context of global warming, which will tend to offset the
precipitation responses by desiccation (Polley et al., 2000; Rounsevell, Brignall and Siddons,
1996). Changes in the seasonal distribution and intensity of rainfall will affect seasonal soil water
dynamics and the efficiency of plant use, and may have a larger impact on rangelands than
changes in precipitation amounts (Giorgi et al., 1998). More intense storms are likely to lead to
increased surface runoff and erosion and concentration of water in smaller portions of the
landscape (Campbell, Stafford Smith and McKeon, 1997). The negative impact of increased global
temperatures through desiccation may be especially important in present arid and semi-arid
regions of the world, especially in Central America, Australia, the Near East, Southeast Asia and
southern Africa, where precipitation may be little affected or may decline (Albritton et al., 2001).
However, as mentioned previously, rising atmospheric CO2 will lead to increased water use
efficiency and improved water relations. Thus, the overall hydraulic and plant productivity
response to changes in precipitation will depend on complicated interactions with temperature and
CO2.

Plant reproductive and recruitment responses

In addition to growth responses, reproductive and plant recruitment response to CO2 enrichment
and associated climate change can have important impacts on productivity, species composition
and, ultimately, biodiversity (Edwards, Clark and Newton, 2001; Grnzweig and Krner, 2001).

There are a number of ways in which CO2 may effect recruitment of new individuals. In terms of
direct plant responses, elevated CO2 is often, but not always (Farnsworth and Bazzaz, 1995;
Smith et al., 2000), associated with increases in seed number (Edwards, Clark and Newton, 2001;
Grnzweig and Krner 2001; Jablonski, Wang and Curtis, 2002; Lawlor and Mitchell, 1991; Smith
et al., 2000), size or weight (Edwards, Clark and Newton, 2001; Jablonski, Wang and Curtis,
2002; Lawlor and Mitchell, 1991; Newton, 1991; Steinger, Gall and Schmid, 2000). These
responses, however, vary with both species and environmental conditions (Jablonski, Wang and
Curtis, 2002; Grnzweig and Krner, 2001); often responses in one reproductive trait (seed
number or size) offset or cancel out another. Further, CO2-induced changes in seed number, size
and dispersal become important only to the extent that seedling recruitment is limited by seed
number, which may often be the case for arid and semi-arid grasslands, but less so for mesic
grasslands (Turnbull, Crawley and Rees, 2000). Seed size may not necessarily be a good indicator
of reproductive success, especially in CO2-enriched atmospheres where larger seeds produced
may sometimes be depleted in nutrients (Jablonski, Wang and Curtis, 2002; Steinger, Gall and
Schmid, 2000). Other factors, such as competitive plant interactions (Bazzaz and McConnaughay,
1992) and seed predation (Arnone et al., 1995), can prevent the establishment of simple
relationships between seed number, seed size and recruitment. Nevertheless, CO2-induced
changes and species differences in seed dispersal and seed size can be important in affecting
recruitment of new individuals (Edwards, Clark and Newton, 2001). Ultimately, the outcome will
depend on interactions between reproduction and the microsite conditions that determine
germination and establishment of the seedling.

Proper soil temperature and moisture for germination and establishment of seedlings is an
important factor in the establishment of grassland species (Defosse, Bertiller and Robberecht,
1997; Minnick and Coffin, 1999; Wester, 1995). Thus, any changes in temperature or precipitation
patterns that affect soil water dynamics have the potential to affect germination and recruitment
processes. Higher CO2 also has the potential to enhance recruitment indirectly through its
improvement of soil water content resulting from reduced plant transpiration.

Below-ground responses

In native grasslands, the majority of plant production occurs below ground, and root biomass
pools often exceed above-ground pools by a factor of two to five (Arnone et al., 2000; Stanton,
1988). In more productive grassland systems, growth responses to CO2 are often greater and
more consistent in below-ground than above-ground plant organs (Newton, 1991; Rogers, Runion
and Krupa, 1994; Rogers et al., 1997). Further, the long-term cycling of nutrients, their
availability for plant growth, and the dependence of soil processes on dynamic root responses may
control, in large part, the long-term response of ecosystems to CO2 enrichment and climate
change (Arnone et al., 2000; Curtis et al., 1994; Norby and Jackson, 2000; Thornley and Cannell,
2000). An understanding of these complex but important interactions is required in considering
ecosystem responses and possible management options for grazing lands.

Root responses
Increasing global air temperatures will lead to higher soil temperatures (Schlesinger and Andrews,
2000). As a result, root production and mortality should increase, assuming soil water and
nutrients remain at sufficient levels (Norby and Jackson, 2000). In natural systems, conditions of
sufficient water and nutrients are often not met, making the prediction of root dynamic responses
to increased temperature difficult. The seasonality of root activity will also be affected by climate
change (Reeder, Franks and Milchunas, 2001), with root activity commencing earlier in the
growing season, and continuing later as growing seasons extend.

In rangelands where plant production is limited mostly by water, any change in soil water
dynamics, whether resulting directly from altered precipitation patterns or indirectly from warmer
temperatures or increased CO2, will alter root activity. In arid and semi-arid rangelands, increased
water will enhance root activity. In more mesic grasslands, increased precipitation may have both
positive and negative consequences for root activities. Under dry conditions, increased nutrient
mineralization and root activity will be important features of belowground system responses to
increased water (Mosier, 1998; Rogers et al., 1999) leading to enhanced plant growth. Under
wetter conditions, negative consequences for roots may occur with water additions, such as
leaching of mobile nutrients, loss of N through de-nitrification, or soil erosion (Rounsevell, Evans
and Bullock, 1999).

Although elevated CO2 can significantly enhance production of belowground organs in productive
grasslands (Newton, 1991; Nsberger, Blum and Fuhrer, 2000; Rogers, Runion and Krupa, 1994),
the response of less productive native grasslands is less certain. About half of native grassland
studies exhibit little or no response in root biomass to growth at elevated CO2, while the other half
exhibit substantial increases in root biomass under elevated CO2 (Arnone et al., 2000). These
variable responses will probably result from the differing soil and environmental conditions of the
experiments, and the consequences of those conditions on plant growth and on root-shoot
partitioning (Larigauderie, Hilbert and Oechel, 1988).

Nutrient cycling and soil feedbacks

Soil feedbacks may ultimately determine ecosystem responses to CO2 and climate change.
Perhaps the strongest evidence for this has been the almost universal decline in shoot N
concentration that occurs in CO2-enrichment studies, especially those in which CO2-induced
production changes are evident (Drake, Gonzlez-Meler and Long, 1997). While part of this
response may be due to the enhanced resource-use efficiency that plants experience under
elevated CO2, there is good evidence that soils cannot release N quickly enough to keep pace with
CO2-induced production increases (Zak et al., 2000). Most research has focused on the N cycle,
but nutrient immobilization and release processes also involve other essential nutrients.

Briefly, CO2-enhanced production will lead to increased carbon substrates entering the soil, which
will fuel microbial growth and initially immobilize soil nutrients (Figure 10.3). However, the
additional microbial biomass will eventually cycle and release more nutrients back into the soil
through mineralization (Morgan, 2002). Thus, the balance between these and other competing
processes can result in increased, decreased or no change in plant available nutrients (Zak et al.,
2000). In the long term, soil nutrient cycles may adjust to changes in organic inputs, and nutrient
availability will increase to meet new demands in CO2-enriched atmospheres (Thornley and
Cannell, 2000); the time this takes to occur will differ among ecosystems due to differences in
species composition and soil type (Reeder, Franks and Milchunas, 2001), as well as to differences
in climate.

The degree to which the N, or any other nutrient, cycle is altered due to CO2-enhanced plant
production changes may account for much of the variability among studies and ecosystems in
their responses to CO2. For instance, the increase in root-shoot ratios in many CO2-enrichment
studies has been interpreted as an indirect response to CO2-induced plant N deficits, which causes
plants to shift more substrates below ground to acquire more N (Larigauderie, Hilbert and Oechel,
1988; Newton, 1991; Rogers, Runion and Krupa, 1994; Rogers et al., 1997, 1999; Stulen and den
Hertog, 1993). The acclimation of plants that often occurs under long-term CO2 enrichment,
whereby photosynthetic and growth responses become less responsive over time, appears to be
related to the build up of carbohydrates that occurs more readily under limited soil N (Arp, 1991;
related to the build up of carbohydrates that occurs more readily under limited soil N (Arp, 1991;
Sage, 1994).

Warmer temperatures may lead to increased organic matter decomposition rates in mesic
grasslands, but the same may not hold for dry rangelands. Globally, warmer regions are known to
exhibit higher rates of soil respiration and organic matter decomposition compared with colder
regions (Epstein, Burke and Lauenroth, 2002; Rustad et al., 2001). However, decomposition is
typically modelled by an interaction of temperature and soil water (Esser, 1992; Parton et al.,
1987, 1995; Rastetter et al., 1991). As water becomes limiting, decomposition and respiration are
predicated less by temperature and more by soil water content (Epstein, Burke and Lauenroth,
2002), with lower soil water resulting in reduced decomposition rates. To the extent it affects soil
water dynamics, rising atmospheric CO2 will also affect decomposition processes (Mosier, 1998;
Rogers et al., 1999).

Figure 10.3 Nutrient Cycling Feedbacks. While CO2 enrichment may lead to increased
photosynthesis and enhanced plant growth, the long-term response will depend on nutrient
cycling feedbacks. Litter from decaying plants and root exudates enters a large soil nutrient
pool that is unavailable to plants until they are broken down and released by microbial
activity. Soil microbes may also fix available nutrients into new microbial biomass, thereby
temporarily immobilizing them. The balance between these and other nutrient release and
immobilization processes determines available nutrients and ultimate plant response.

Source: Figure reprinted with permission from Science (Morgan, 2002).

Plant Community Changes

While responses of plant community productivity to CO2 and climate change are predicted to be
important in many ecosystems, long-term changes in plant community composition may be even
more important with regard to ecological impacts and effects on ecosystem goods and services
(Krner, Bazzaz and Field, 1996; Leadley and Krner, 1996; Polley et al., 2000; VEMAP Members,
1995).
Climatic changes

The distribution of plant types is affected strongly by spatial and temporal patterns of precipitation
and temperature (Epstein et al., 1997; Paruelo and Lauenroth, 1996). Water is the most important
determinant of geographical plant distribution (Stephenson, 1990) and productivity (Sala et al.,
1988; Webb et al., 1978). Any changes in climate that affect soil water dynamics will have an
equally important effect on plant communities. Although precipitation is predicted to increase over
much of the globe, the concentration of more rainfall into fewer rainfall events will lead to more
uneven spatial distribution of water across the landscape, and more drought in some regions
(Campbell, Stafford Smith and McKeon, 1997). In rangelands, where plant community responses
are commonly event-driven, changes in timing and amount of precipitation will have especially
important consequences for plant community responses (Polley et al., 2000). Patterns of
precipitation that significantly alter the vertical distribution of water will correspondingly alter the
proportion of shallow-rooted versus deep-rooted species (Ehleringer et al., 1991).

Temperature is also important in the distribution and functioning of plant communities (Epstein et
al., 1997; Field and Forbe, 1990; Pyankov et al., 2000; Terri and Stowe, 1976). The increased
presence of C4 grasses toward tropical latitudes is one of the simplest and yet most successful
models of plant distribution (Pyankov et al., 2000: Terri and Stowe, 1976). Modelling exercises
that consider both temperature and precipitation aspects of climate change predict an increase in
relative abundance of C4 grasses and a decrease in C3 grasses in North and South American
grasslands and shrublands (Coffin and Lauenroth, 1996; Epstein et al., 1997; Epstein, Burke and
Lauenroth, 2002; VEMAP Members, 1995). In Europe, latitudinal vegetation gradients in
calcareous grasslands exhibit good correlations between species composition and summer
temperature (Duckworth, Bunce and Malloch, 2000), suggesting that present-day transitions in
grassland communities from north to south may be good indicators of future plant community
changes to warming.

Predictions of plant community changes for shrublands are less certain than for grasslands. There
is some evidence that warming or precipitation increases, or a combination, may lead to an
increase in subtropical shrublands, but a decrease in temperate shrublands (Epstein, Burke and
Lauenroth, 2002; VEMAP, 1995). However, these predictions are uncertain given our incomplete
understanding of shrubland ecology. The encroachment of shrubs into former semi-arid and arid
grasslands has been attributed to livestock grazing, fire suppression, increased atmospheric CO2,
and climate change (Archer, Schimel and Holland, 1995; Bond and Midgley, 2000; Moore et al.,
2001; Polley et al., 1997; Roques, OConnor and Watkinson, 2001). Unfortunately, sorting out the
relative importance of these competing forces is difficult at best, and predictions based on them
may be unreliable.

The long-term responses of grazing land plant communities to warming and precipitation changes
will be determined to a large extent by soil nutrient cycling and water dynamic feedbacks, which
affect plant productive, reproductive and recruitment responses, and alter the competitive balance
in ecosystems. The rate at which these climate changes occur will be an important factor in
determining the ability of plant species to expand their distribution range into new regions.

Rising atmospheric CO2

Knowledge of individual plant traits like photosynthetic pathway, domesticated compared with wild
species, ability to fix atmospheric N, reproductive traits, and morphological considerations such as
rooting morphology and depth, have been examined to determine adaptability and responses to
future CO2-enriched atmospheres (Bazzaz, 1990; Bowes, 1993; Cure, 1985; Gitay et al., 2001;
Krner, 1996; Long, 1991; Porter, 1993; Wand et al., 1999). However, studies involving field
experiments with complex plant communities have revealed that single species traits and
responses are often not good predictors of field performance and competitive interactions (Ackerly
and Bazzaz, 1995; He, Bazzaz and Schmid, 2002; Joel et al., 2001; Leadley et al., 1999; Morgan
et al., 2001; Owensby et al., 1996, 1999; Reich et al., 2001). For instance, while legumes may be
particularly responsive to CO2, He, Bazzaz and Schmid (2002) observed reduced growth of N-
fixers under elevated CO2 in communities with high species richness. Competitive interactions in
plant communities can modify the dynamics of soil resources and affect the ability of species to
plant communities can modify the dynamics of soil resources and affect the ability of species to

respond to CO2 (Geeske et al., 2001). There is some evidence that more diverse ecosystems may
exhibit increased capacity to adapt and respond to rising CO2 (He, Bazzaz and Schmid, 2002;
Niklaus et al., 2001). Like climatic changes, the long-term consequences of CO2 enrichment on
water and nutrient cycling will probably determine major species and ecosystem responses (Joel
et al., 2001; Thornley and Cannell, 2000).

Multiple global changes

Present knowledge would suggest that plants with the C3 photosynthetic pathway, forbs and
legumes might be among the more responsive plant types to rising atmospheric CO2
concentrations, and thus may expand their distribution in future CO2-enriched environments.
However, significant increases in temperature and altered precipitation patterns will tend to offset
these functional group responses. Warmer, drier temperatures will tend to favour C4 metabolisms;
so significant increases in temperature may cancel out some of the competitive advantage
afforded C3 plants at elevated CO2 levels. Further, other global changes, such as increased ozone
and atmospheric N deposition, may be important modifiers of these responses, especially near
urbanized centres. Increased atmospheric ozone, a pollutant in rural areas of industrial countries,
can reduce the contribution of legumes in grass-legume mixtures (Nsberger, Blum and Fuhrer,
2000). Increased N deposition, a function of animal production as well as industrial activity, will
add N to grazing lands (Mosier, 1998; Vitousek et al., 1997), thereby countering the tendency of
rising CO2 to result in reduced available soil N. Both of these will tend to reduce the importance of
legumes in maintaining sufficient N in systems to promote CO2-induced growth responses.

Livestock responses

Since animals attempt to maintain an optimal temperature (Rtter and Van De Geijn, 1999), an
understanding of the changing climate and how that affects the thermal balance of animals is
critical to understanding how livestock production systems respond to climate change (Parsons et
al., 2001).

In general, heat stress reduces feed intake, feed efficiency, animal gain, milk production, and
reproduction (Adams et al., 1998; Baker et al., 1993; Bianca, 1965; Hanson, Baker and Bourdon,
1993; Klinedinst et al., 1993), and extreme conditions ultimately lead to animal disease and death
(Gitay et al., 2001; Rtter and Van De Geijn, 1999). These negative impacts of global warming
will be especially pertinent in regions where summer temperatures are already warm and further
increases will significantly affect animal performance (Gitay et al., 2001; Rtter and Van De Geijn,
1999). However, livestock operations in cooler regions of the earth may be little affected by
increasing temperature, or may even benefit through reduced feed requirements (Thompson,
1973), increased growth, increased survival and decreased energy costs (Rtter and Van De Geijn,
1999). Further, the particular effect of any change in temperature will also depend on the breed.

In addition to the direct impacts of altered climates, livestock performance will also be influenced
by changes in the amount and quality of forage produced. While most research indicates greater
primary production under elevated CO2, some research has also indicated a change in forage
quality. Plant N concentrations tend to decline under elevated CO2, especially when production
responses are large (Krner, 2002). However, elevated CO2 can increase total non-structural
carbohydrates (TNC), which tends to enhance forage quality (Lilley et al., 2001). Where N is not
limiting for animal performance (e.g., in improved pastures), elevated CO2 may enhance forage
quality (Lilley et al., 2001). Conversely, in native rangelands and semi-natural grasslands where N
is typically limiting, the constraints of reduced N and increased fibre in CO2-enriched forage may
reduce digestibility and have a negative impact on ruminant performance (Krner, 2002; Morgan
et al., 2004; Owensby, Cochran and Auen, 1996). A change in species composition due to CO2
enrichment or to climatic change may also affect forage quality (Krner, 2002; Morgan et al.,
2004), and the results can be either positive or negative for quality, depending on the particular
species involved.

Finally, the range of many animal diseases and pests will also increase at warmer temperatures
(Stem et al., 1988). Animal management problems resulting from these expansions will be
(Stem et al., 1988). Animal management problems resulting from these expansions will be

experienced most acutely in regions just outside of present-day boundaries of particular diseases
and pests.
Management of Grazing lands for the Future

The rate and character of future grazing land changes will incorporate responses to land use
changes and management, as well as to climatic change and rising CO2 concentrations.
Management of these systems will ultimately determine their ecological and economic trajectories.
Thus, an understanding of management options for future grazing lands is important not only as a
practical matter, but also to predict the future character of grazing lands. In a general sense, land
managers will need to be adaptive and proactive in considering how to match their management
practices with the changing environment, and should not be satisfied with management that has
worked in the past. Decision-support systems that can assess multiple environmental changes will
become increasingly important as managers build risk assessment more explicitly into their
enterprises.

A list of the more important grazing land factors and responses to increased atmospheric CO2 and
climate change discussed previously are presented in Table 10.2, followed by some management
options. The following discussion will elaborate on most of the options listed, and will separately
consider rangelands and productive pastures.

Table 10.2 CO2 and climate change responses and management options for grazing land factors.

RESPONSES TO RISING CO2 MANAGEMENT

AND CLIMATE CHANGE OPTIONS

Primary production
Increase or little change with rising CO2: Applies to most systems, Adjust
especially water-limited rangelands. N may limit CO2 response in some Forage
systems. harvesting:
Stocking rates.
Increases or little change with temperature: Applies to most temperate and Grazing systems.
wet systems. Mowing practices
(productive
Decreases with temperature: Applies to arid and semi-arid systems that grasslands).
experience significantly enhanced evapotranspiration and drought,
particularly where precipitation is not expected to increase. Develop and utilize
adapted forage
Variable responses with precipitation: Depends on present climate, and species (e.g.
nature of precipitation change. Increases in production in regions where legumes, C4
water is limiting, but increasing temperatures and more intense grasses where
precipitation events will reduce this. appropriate, more
drought-resistant
species and
cultivars).

Enterprise change
(e.g. movement to
more or to less
intensive
agricultural
practices).
Plant community species composition
Global changes will drive competitive responses that alter plant All of the above.
communities: In some systems, legumes and C3 species may be favoured in
Weed control:
future CO2-enriched environments, but community reactions will be variable
Fire management
and highly site specific. Warmer environments will favour C4 metabolisms. and/or grazing
Both productive and reproductive responses will be featured in community practices to
changes. Ultimate plant community responses will probably reflect convert woody
alterations in soil nutrients and water, and involve complex interactions lands to
alterations in soil nutrients and water, and involve complex interactions lands to
between changes in CO2, temperature and precipitation. Weed invasions grasslands.
may already be underway, due to rising atmospheric CO2. Proximity to Herbicides where
urban areas will add complex interactions with ozone and N deposition. appropriate to
control
undesirables.

Enterprise change
or emphasis:
Change between
intensive/extensive
practices.
C storage
strategy.
Tourism,
hunting, wildlife.
Biodiversity.
Forage quality
Increasing CO2 will alter forage quality. In productive grasslands with ample Utilize or interseed
N, forage quality may increase due to more TNC. In N-limited native legumes where N
systems, CO2-induced reduction in N and increased fibre may lower quality. is limiting and
practice is feasible.

Fertilize where
feasible.

Alter supplemental
feeding practices.
Animal performance to altered climate
Increased temperature, warm regions: Reduced feed intake, feed efficiency, Animal usage:
animal gain, milk production and reproduction. Increased disease Select adapted
susceptibility, and death. animal breeds from
different world
Increased temperature, cold regions: Enhanced animal performance, regions to match
lowered energy costs. new climate.
Improve animal
genetics.
Select different
animal species (i.e.
camels, sheep and
goats for more
drought-prone
areas).

Adjust forage
harvesting (above)

Alter management
(e.g., timing of
breeding, calving,
weaning)

Enterprise change
(above)

Rangelands

Stocking rate and stocking system decisions are based on seasonal variability in forage
production, compositional changes in the plant community, forage quality, and the particular
animal breed. Thus, rising CO2 and climate change will lead to a variety of altered livestock
stocking rates and systems to adapt to new environments of the future (Ojima and Lackett, 2002;
Polley et al., 2000; Campbell et al., 2000).

Increases or decrease in stocking rates will follow changes in primary productivity, and may not be
great in many ecosystems due to compensating changes in production resulting from multiple
global changes (Campbell et al., 2000; Smith and Lazo, 2001). Significant changes in seasonality
of production and plant community composition will alter the timing of animal operations like
calving, lambing or weaning (Ojima et al., 2002), as well as grazing season duration, and may
affect the necessity and timing of supplemental livestock feeding. In warmer, drier regions, even
small reductions in stocking rates, resulting from lowered forage production, may drive some
producers out of business or subsistence people into famine (Campbell et al., 2000).

Changes in plant community composition will be among the most important changes for
rangelands. One particular community change that has been underway for the past 100 years, and
that might involve both rising atmospheric CO2 concentrations and climate change, is the
incursion of woody plants into former semi-arid and arid grasslands (Polley et al., 1997; Scholes
and Archer, 1997). Unfortunately, management considerations for converting some shrublands
back to native grasslands are constrained by the creation of new, stable states that resist further
transition (Schlesinger et al., 1990; Whitford, Martinez-Turanzas and Martinez-Meza, 1995).
Nevertheless, in systems where change is still feasible, the re-institution of fire and light grazing
may be used to reverse brush encroachment (Roques, OConnor and Watkinson, 2001). These
types of management will be even more important in regions where increased precipitation will
tend to push systems further towards woody plant invasion. At the same time, maintaining
shrublands of low economic value may be a cost-effective carbon offset for reducing greenhouse
gas emissions (Moore et al., 2001), in which case continued fire suppression may be desired.

In contrast to the above story, invasion of thousands of hectares of sagebrush steppe rangelands
in western North America by the exotic Bromus tectorum may be linked to increased atmospheric
CO2 concentrations (Smith et al., 2000). This is converting native sagebrush steppe vegetation
into a fire-controlled annual grassland. Numerous other weed invasions are underway around the
world, and while much of these are the result of human transportation and movement of exotics
around the globe, and are often linked to domestic livestock grazing, the changing climate and
increased atmospheric CO2 are also implicated in some of these invasions (Watkinson and
Ormerod, 2001). Management research that traditionally has focused on livestock performance
must increasingly evaluate the ecological responses of rangelands to grazing systems, and in
particular, to alterations in nutrient and water cycles that may be important in the growth and
recruitment of exotics. Differentiating between responses due to CO2 or climatic changes and
responses due to management practices will be fundamental in developing management practices
that can cope with the rapidly changing environment.

Some potentially useful plant community changes for livestock production may occur in response
to rising CO2 concentrations, like increasing the competitiveness or presence of legumes, thereby
providing additional N to support CO2-enhanced plant growth. An increase in legumes may be
especially important for enhancing forage quality that otherwise may decline due to lowered plant
N concentrations (Owensby, Cochran and Auen, 1996). It may be useful to consider interseeding
legumes into rangelands as a means to facilitate this process, as well as a means to sequester C in
the soil, thereby abating greenhouse gas emissions (Mortenson, Schuman and Ingram, 2003).
However, research needs to determine whether such increases in legumes will have effects on the
exchange of other trace gases like N2O or methane, which contribute to greenhouse gas warming.
Selection of appropriate legume species and the development of improved germplasm and
management practices may be required to maximize the benefits of such legume usage for
rangelands.

Climate change will no doubt lead to altered animal usage. Camels, sheep and goats are better
adapted to dry regions, and will probably become more important in regions that experience
increased drought (Squires and Sidahmed, 1997). Goats may become economically more practical
in subtropical rangelands experiencing more climatic variability (Souza Neto et al., 1998). A high
degree of genetic diversity already exists within livestock species, so warming in temperate
regions may be accommodated by selecting livestock from more tropical climates (Squires and
Sidahmed, 1997). Attention has been given in the past to adapting the genetics of livestock to
targeted environments, like the tropics (Bonsma, 1949; Rtter and Van De Geijn, 1999). However,
targeted environments, like the tropics (Bonsma, 1949; Rtter and Van De Geijn, 1999). However,
there is little information on how these adapted animals would perform with the further warming
predicted for this century, and virtually no information on how such selected or altered livestock
will perform in future rangelands that will undoubtedly undergo significant plant compositional
changes (Squires and Sidahmed, 1997).

Implications for rangeland livestock management systems

Two types of grazing operations prevail on rangelands: pastoralism and ranching (Squires and
Sidahmed, 1997). Pastoralism is a practice dating to ancient times, and traditionally centred on
herding domestic livestock, but may include some cropping where feasible. Pastoralists typically
maintain herds as insurance against extreme drought or hardship. It is a system that in modern
times has tended to exist on marginal lands, and is currently in crisis due to a complex interaction
of ecological, socio-economic and population pressure problems (Squires and Sidahmed, 1997).
Commercial ranching, a more recently developed enterprise, tends to be more economically
motivated (Squires and Sidahmed, 1997). Private ownership of land is common, operational scale
is large, and improvements in the way of fencing, improved pastures and access to water and
other resources are common.

The impacts of climate change are likely to differ between pastoralists and commercial ranchers.
For instance, Smith and Lazo (2001) concluded that climate change will have less impact on
livestock yields than on forage yields, in part because there currently exists excess capacity for
livestock, or that some expansion of rangelands is possible. However, this may not apply to some
pastoral groups that are already constrained by human population growth and decreasing grazing
lands (Squires and Sidahmed, 1997). Further, pastoral grazing often involves complex social
arrangements in the seasonal movement of animals across the landscape (Parish and Funnell,
1999). Although pastoralists are characterized by an ability to adapt to extreme climatic events
(Mortimore and Adams, 2001), their complex socio-economic structures and dwindling land
resources in some regions may make them more susceptible than ranchers to changes brought
about by climate change (Squires and Sidahmed, 1997; Turner, 1993). Climate changes that
affect production and plant community composition may challenge social arrangements that have
developed over hundreds of years, and further complicate modern-day attempts to protect the
landscape while preserving important cultural and societal arrangements as human populations
expand.

Other uses for rangelands

In addition to livestock grazing, rangelands provide many other goods and services, including
biodiversity, tourism, fuelwood (Gitay et al., 2001; Watkinson and Ormerod, 2001), and more
recently, C stores (Meeting et al., 2001; Moore et al., 2001; Schuman, Herrick and Janzen, 2001).
Undoubtedly, climate change, government policy and economics will alter the balance among
these uses, and may shift management from one enterprise to another. Commercial ranching is
already moving away from intensive livestock grazing and towards urbanization, game ranching
and transfer of homelands back to indigenous peoples (Polley et al., 2000). Some rangelands will
be taken out of production and converted to cropping, as human populations increase and crop
production on marginal lands becomes economically more feasible. The contribution livestock
make to climate change through methane production is an important problem that will no doubt
be considered in future research and management options (Gitay et al., 2001; McCrabb and
Hunter, 1999). However, less intensive livestock practices may be encouraged due to the lower
net contribution of rangeland to methane production, and thus to greenhouse gas warming
compared with more intensive livestock operations (Subak, 1999).

Productive grasslands

Production in many temperate humid grasslands is likely to remain unchanged (Rounsevell,

Brignall and Siddons, 1996), or may increase due to increased CO2 and warmer temperatures,
which will extend growing seasons, and in some cases increase primary production. Because water
limitations are generally less severe in productive grasslands, the negative impact of warmer
temperatures on desiccation will be less of a problem than in semi-arid and arid rangelands.
However, temperature increases in excess of 4C are likely to have negative consequences
However, temperature increases in excess of 4C are likely to have negative consequences

because of increased drought stress and reduced growing season lengths (Baethgen, 1997;
Rounsevell, Brignall and Siddons, 1996). Subtropical and tropical grasslands may be more
vulnerable to climate change than cooler, temperate grasslands. A major risk in subtropics and
tropics would be a decline in forage quality (Polley et al., 2000), increased heat stress in livestock,
and in dry zones or regions with periodic droughts increased incidences of drought or more
severe droughts (Baethgen, 1997). However, predicted increased precipitation in some regions
may counter the negative response to temperature. As for rangelands, management options may
have much to do with altered species composition, changed stocking rates and systems, and, in
some situations, different grazing animals or breeds. The options in productive grasslands will
tend to be greater since manipulations are economically more practicable than in rangelands.

In semi-natural grasslands with high species diversity, species composition is expected to

gradually change due to rising CO2 (Teyssonneyre et al., 2002) and changes in climate
(Duckworth, Bunce and Malloch, 2000). Defoliation can affect the balance of functional group and
species composition, so management practices developed to deal with rising CO2 and climate
change will have to balance animal productivity responses with management that maintains or
enhances biodiversity (Teyssonneyre et al., 2002). As with rangelands, an understanding of rising
CO2 and climate change impacts on grassland invasions will be required to develop viable
management practices to combat those invasions (Buckland et al., 2001; Davis, Grime and
Thompson, 2000; Watkinson and Ormerod, 2001).

Contrasting rangelands and productive grasslands

While many of the same issues brought on by rising atmospheric CO2 and climate change are
likely to occur in rangeland and productive grasslands, differences between these two types of
systems are likely to affect how they are manifested and addressed. For instance, livestock
grazing practices for improved pastures rely on the introduction of adapted species, as well as on
ecological considerations of maintaining the optimal balance of desired species that is important
for both rangelands and productive grasslands. Sowing legumes into productive grasslands is
already an accepted practice for improving N nutrition of pastures. Thus, the increased
development and use of legumes to combat N limitations in a future CO2-enriched world should be
a relatively easy technology to adapt (Gitay et al., 2001; Schenk, Jger and Weigel, 1997). The
same may not be true for rangelands, where establishment and subsequent control of introduced
species is more problematic. In general, much of the species changes in improved pastures will be
due to conscious management decisions to sow or nurture particular adapted species. In
rangelands, species issues will involve management of the ecological changes and states resulting
from combined climate change and past management to ensure that future desired plant
communities result.

Conclusions

Our present understanding of atmospheric CO2 and climate change and their impact on grazing
lands is modest. We know that atmospheric trace gas concentrations are rising and that significant
changes in the Earths climate are already under way, and we are beginning to understand how
these global changes will manifest in different world regions. They will have important
consequences for grazing lands, leading ultimately to management modifications, and in some
instances, changes in enterprise. In addition to the ecological effects, significant socio-economic
consequences will surely follow. But our ability to predict and prepare for possible future scenarios
of ecological and management responses is poor.

Our challenge as we look to the future of world grazing lands is to acknowledge that these
complex ecosystem responses to CO2, climate change and other aspects of global change like
increased N deposition or higher levels of ozone are under way, and to conduct research to
understand what has already occurred, as well as develop appropriate technologies and decision-
support systems to deal with future changes. The success of global change research programmes
will depend in large part on our ability to involve ranchers, pastoralists and other land managers in
the design of new research projects to develop proactive strategies and technologies to prepare
for the future (Campbell et al., 2000). Where past research has focused on the responses of
ecosystems to changes in one or two environmental factors, future research must consider the
ecosystems to changes in one or two environmental factors, future research must consider the

multi-factorial nature of global changes. More work is needed to understand the complex below-
ground processes that will ultimately govern long-term responses. Animal responses and their
effects on grazing systems need more attention if we are to realistically simulate combined global
change and management responses. And we must advance our capability to respond by
identifying low-risk, realistic strategies that allow adaptation, but make sense in the face of
uncertainty concerning the degree and rate of global changes and ecosystem responses. Grazing
land agriculture has always been characterized by change and adaptability despite our
increasing knowledge about the Earths environment, there is little reason to indicate that it will be
any different in the future.

Acknowledgements

The author would like to thank Dana Blumenthal, Dan LeCain, Jean Reeder, Robin Kelly and Justin
Derner, who made many useful contributions that helped greatly in developing and clarifying this
manuscript.

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