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Transcript
Jeffrey S. Dukes, Lorena Torres Martinez and Michael Schuster
Distribution of Terrestrial Ecosystems and Changes in Plant Community Composition
SpringerReference
Distribution of Terrestrial Ecosystems and Changes in Plant
Community Composition
Abstract
Plant communities have been transformed by global changes such as land-use change and biological invasion in recent
decades, and climate change will drive further transformation in the coming decades. Plant species can respond to
changing climates by shifting their ranges to new areas in order to track optimal conditions and/or by adapting to these
changes in situ. Future climates have the potential to alter species’ productivity, phenology and biotic interactions. In
addition, species’ ranges are expected to shift towards higher latitudes, tracking favorable climate conditions as the planet
warms. The same trend is expected along altitudinal gradients, where species are expected to move uphill. However,
distributions of communities and ecosystems depend on their individual species’ responses to climate change, and are
therefore more complicated to predict. Future plant community assemblages will differ from the ones that have existed
historically, and these changes could themselves influence the rate of global climate change.
Keywords
Terrestrial ecosystem distribution; plant community composition; distributional shifts; niche envelope models; biotic
interactions; niche evolution; adaptive potential
Definition
Climatic variables strongly influence the distributions and growth rates of plant species. As climates change, plant
responses will alter the composition, distribution, and functioning of terrestrial ecosystems.
Key Information
Altered temperature, precipitation, and resource availability will lead to changes in ecosystem distributions and changes in
the composition of plant communities. When large-scale changes in the environment occur, especially in the climate,
plants and other organisms can respond in only a few ways. One possibility is that species tolerate or adapt to the new
conditions. Alternatively, species' distributions can shift into new areas, such that populations continue to inhabit suitable
climates. These range shifts contribute to a shift in the greater ecosystem’s distribution and composition. Species unable
to tolerate, adapt to, or shift their ranges in response to the new climate will go locally extinct.
Climate change is only one of many factors of global change. Land use change, exotic species introductions,
anthropogenic nitrogen deposition, and elevated CO2 (although closely tied to climate change) also affect terrestrial
ecosystems. Land use change and climate change are expected to be the two most important factors driving changes in
biodiversity over the course of this century; however the relative importance of each differs among biomes. Climate is
expected to be most important in biomes with relatively less human disturbance, such as artic, alpine and boreal systems;
whereas land use is already an important factor in areas closely tied to human activity, such as grassland, Mediterranean
and tropical systems (Sala et al. 2000). Therefore, while climate change is an increasingly important driver of global
change, it should be considered within a broader context.
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Jeffrey S. Dukes, Lorena Torres Martinez and Michael Schuster
Distribution of Terrestrial Ecosystems and Changes in Plant Community Composition
SpringerReference
Species Responses to Global Change and the Re-structuring of Plant Communities
Climate affects many aspects of plant growth and function. In response to a changing climate, species may modify
phenology, thermal tolerance, and dispersal traits. Phenological responses to warming can prolong growing periods and
enhance growth in many species. Warming commonly alters plant flowering time and advances bud break. Differences in
phenological responses across species can alter inter-specific competitive ability by changing the timing of light,
temperature, and water availability at the extremes of the growing season. Phenological changes can also alter plant
competition for pollinators or cause life cycle asynchrony between plants and mutualist animals by affecting flowering time
and plant distribution. Life cycle asynchrony can disrupt pollination cycles, potentially reducing plant reproductive output
and population density.
Rising temperature, altered precipitation, elevated atmospheric CO2 concentration, and increasing anthropogenic
nitrogen deposition can affect species diversity patterns within communities, affect competitive relationships among
species, and influence trophic interactions. Rising CO2 levels have been found to facilitate invasion of C4 grasslands by C
woody species, although warming is expected to favor warm season C 4 grasses. Morgan et al. (2011) found that
combined CO2 enrichment and warming favored higher productivity in C4 grasses, suggesting future grasslands may be
3
more resilient to C3 invasion. Furthermore, nitrogen deposition and other global environmental changes are thought to
have opened some ecosystems to invasion by non-native plant species (Hellmann et al. 2008). These changes result in
novel interactions of species that lack a common evolutionary history, making impacts on community composition difficult
to predict.
At the microevolutionary scale, the high level of local adaptation to climate and the great amount of genetic
variation for traits defining species’ ecological niches suggests a high potential for species to adapt to new climatic
conditions. For instance, several studies have reported significant genetic variation of functional traits related to
phenology, growth or gas exchange and drought tolerance. However, niche evolutionary potential will require more than
just genetic variation in defining traits. It will also strongly depend on constraints imposed by the species’ realized niche,
such as interspecific competition, demographic dynamics, and trait plasticity (Lavergne et al. 2009).
Projected Species Range Shifts and Biome Distributions
As climate change shifts species ranges, it will also alter the distribution of ecosystems and biomes. As the planet warms,
species are expected to shift to higher latitudes, tracking favorable climate conditions. Species are largely influenced by
their tolerance to temperature and precipitation extremes. For many species, low winter temperatures prevent expansion
into higher latitudes and competition with better climatically-adapted species limits expansion into lower latitudes. A
similar trend exists along altitudinal gradients. The commonly adopted altitude-for-latitude temperature model suggests
that distributional shifts resulting from temperature changes will occur proportionally along altitudinal and latitudinal
gradients. Because temperature decreases by 1°C over a ~167m increase in altitude or a ~145km increase in latitude,
each 1°C increase in surface temperature should cause ranges to shift ~167m higher and ~145km poleward (Walther,
2003). Communities previously relegated to lower elevations have already started moving uphill, replacing communities of
higher elevations at their distributional margins. Over the past 50 years, mountain treelines have commonly shifted
upwards by up to 130m (Jump et al. 2009, Walther 2003). This trend will reduce the extent of mountaintop communities,
which are not adapted for warmer conditions. Given sufficient warming, many mountain communities may be
outcompeted by communities from lower altitudes as mountaintops become more hospitable to more competitive
species. Precipitation is another critical component influencing species distributions, and as such, altered precipitation
regimes can determine which plant communities occur in a given region (Wu et al. 2011). Expected changes in
precipitation depend on the region being considered, and sometimes on the climate model, especially in the temperate
zone. Differences among model projections add to the uncertainty in predictions of future ecosystem distributions,
although general patterns can be observed across models. Inter- and intra-annual variability in precipitation are expected
to become more extreme, exposing ecosystems to greater risk of prolonged drought and heat waves (Knapp et al. 2008).
This will have consequences for community composition, as described previously, but will also make otherwise suitable
areas unfit for some ecosystems, thus modifying the poleward and uphill trend expected when only considering changes
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Jeffrey S. Dukes, Lorena Torres Martinez and Michael Schuster
Distribution of Terrestrial Ecosystems and Changes in Plant Community Composition
SpringerReference
in temperature. Topography also determines the diversity of microclimates experienced by any ecosystem, and can thus
influence distributional shifts. Furthermore, changes in species distributions may be significantly influenced by human
habitat modification or a combination of both climate and human disturbances. For example, Hättenschwiler and Körner
(1995) have suggested that the high level of nitrogen deposition and the cessation of cattle grazing in the Swiss Central
Alps may have enhanced the replacement of Pinus sylvestris seedling populations by Pinus cembra, a species previously
unreported in at that altitude.
Modeling Impacts of Climate Change
Projections of future ecosystem distributions can be based on species distribution models (SDM) of representative
species of that ecosystem. SDMs consider the contemporary climate of regions associated with a given species and map
future distributions of those species based on where similar climates are expected to exist in the future. Compiling the
model projections for several species can suggest possible future distributions of their respective ecosystems. However,
SDMs are static models and are limited by their assumptions. This limitation is based on whether the model considers a
species’ fundamental niche (all possible locations where that species could potentially exist considering abiotic factors) or
a species’ realized niche (only locations where that species exists in reality, which is determined by abiotic and biotic
factors). While SDMs utilize the observed range, and therefore the realized niche, of the considered species in order to
define that species niche, they often only consider the fundamental niche of a species when making projections. Thus,
SDMs partially omit the importance of the biotic interactions that can define the realized niche, and therefore the
distribution, of the species. For instance, temporary downhill species range shifts in the French mountain forests might be
facilitated by competitive release from other alpine species at the range margins brought on by climate-induced
disturbance (Lenoir et al. 2010). Furthermore, most models do not necessarily include the dispersal patterns and
evolutionary potential of species (see Elith & Leathwick 2009) although some attempts have been made to include these
parameters. Despite their limitations, species distribution models are a useful tool in predicting the impacts of climate
change.
A variety of abiotic and biotic factors will limit or alter the response of each species to changes in climate. These
limitations will put some species at risk of being unable to establish in a region with a suitable climate. For instance, soil
characteristics can hamper the otherwise successful establishment of plant species in new regions. Barriers to dispersal,
such as habitat fragmentation or a lack of land, can also limit the ability of species to track suitable climates. A lack of
natural area will be particularly detrimental to many species’ mobility in regions with high human populations, while
natural barriers will be of greater concern in high latitudes and mountainous regions. In some cases the time required for
a species to establish in a new community and spread to others may be too great compared to the rate of climate change.
As a result, some species may decline or become extinct.
The idiosyncratic responses of individual species to global change can lead to complex and unexpected
responses at the community level. Consequently, predicting the composition of future plant communities is difficult.
Species-level responses to climate change are likely to result in assemblages of species that have not historically
interacted. These new assemblages, known as “no-analog communities,” could support novel species interactions.
No-analog communities may function differently than contemporary communities, and consequently provide different
ecological services. Thus, while models can provide an estimate of individual species responses, the communities of the
future may be very different from the ones that have been observed historically.
Changes in ecosystem distribution and plant community composition feed back to climate through several
pathways. Forests store large amounts of carbon; consequently, alterations to their community composition, productivity,
or distribution will influence global carbon dynamics and climate. In addition, ecosystems influence climate by influencing
the fraction of radiation absorbed by the surface, the rate of evapotranspiration, which contributes to cloud creation and
other atmospheric processes and the physical structure of the land surface.
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Jeffrey S. Dukes, Lorena Torres Martinez and Michael Schuster
Distribution of Terrestrial Ecosystems and Changes in Plant Community Composition
SpringerReference
Figure 1. Relationship between individual species responses, community changes and ecosystem modifications with climate change.
Idiosyncratic responses of individual species will create no-analog communities. Feedbacks from terrestrial ecosystems further
complicate predictions of future climate.
References
Elith J, and Leathwick JR (2009) Species distribution models: ecological explanation and prediction across space and
time. Annual Review of Ecology Evolution and Systematics 40: 677-697
Hättenschwiler S, and Körner C (1995) Responses to recent climate warming of Pinus sylvestris and Pinus cembra within
their montane transition zone in the Swiss Alps. Journal of Vegetation Science 6: 357-368
Hellman JJ, Byers JE, Bierwagen BG, Dukes JS (2008) Five potential consequences of climate change for invasive
species. Conservation Biology 22(3): 534-543
Knapp AK, Beier C, Briske DD, Classen AT, Luo Y, Reichstein M, Smith MD, Smith SD, Bell JE, Fay PA, Heisler JL,
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ecosystems. Bioscience 58(9): 811-821
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Jeffrey S. Dukes, Lorena Torres Martinez and Michael Schuster
Distribution of Terrestrial Ecosystems and Changes in Plant Community Composition
SpringerReference
Additional Recommended Reading
Franklin J (2009) Mapping Species Distributions: Spatial Inference and Prediction. Cambridge University Press, New York
Loarie SR, Duffy PB, Hamilton H, Asner GP, Field CB, Ackerly DD (2009) The velocity of climate change. Nature 462:
1052-1055
Sala OE, Chapin FS III, Armesto JJ, Berlow E, Bloomfield J, Dirzo R, Huber-Sanwald, E, Huenneke LF, Jackson RB,
Kinzig A, Leemans R, Lodge DM, Mooney HA, Oesterheld M, Poff NL, Sykes MT, Walker BH, Walker M, Wall DH (2000)
Global biodiversity scenarios for the year 2100. Science 287: 1770-1774
Schweiger O, Biesmeijer JC, Bommarco R, Hickler T, Hulme PE, Klotz S, Kühn I, Moora M, Nielsen A, Ohlemüller R,
Petanidou T, Potts SG, Pyšek P, Stout JC, Sykes MT, Tscheulin T, Vilà M, Walther GR, Westphal C, Winter M, Zobel M,
Settele J (2010) Multiple stressors on biotic interactions: how climate change and alien species interact to affect
pollination. Biological Reviews 85: 777-795
Thuiller W, Albert C, Araújo MB, Berry PM, Cabeza M, Guisan A, Hickler T, Midgley GF, Paterson J, Schurr FM, Sykes
MT, Zimmermann NE (2008) Predicting global change impacts on plant species’ distributions: future challenges.
Perspectives in Plant Ecology, Evolution and Systematics 9: 137-152
Williams JW, Jackson ST (2007) Novel climates, no-analog communities, and ecological surprises. Frontiers in Ecology
and the Environment 5: 475-482
Distribution of Terrestrial Ecosystems and Changes in Plant Community Composition
Michael
Schuster
Purdue University, West Lafayette, USA
Lorena Torres
Martinez
Department of Biological Sciences, Purdue University, West Lafayette, USA
Jeffrey S.
Dukes
Department of Forestry and Natural Resources, Department of Biological
Sciences, Purdue University, West Lafayette, USA
DOI:
10.1007/SpringerReference_300096
URL:
http://www.springerreference.com/index/chapterdbid/300096
Part of:
Global Environmental Change
Editor:
Dr. Bill Freedman
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