Download Impacts of climate change on diversity in forested ecosystems: Some

Impacts of climate change on diversity in forested ecosystems:
Some examples
by Paul A. Gray1
ABSTRACT
Ecological diversity (the product of ecosystem, species, and genetic diversity) will change significantly in the 21st Century
in response to the combined influence of climate, human activities, the movement of indigenous and non-indigenous
species, and natural disturbances like fire (also modified by climate). Many species will acclimate (phenotypic variation)
and/or adapt (genotypic variation) to changing conditions. Many will not. Species with a high rate of reproduction that
are able to move long distances, rapidly colonize new habitats, tolerate humans, and survive within a broad range of biophysical conditions will be most successful in finding new niches. Large changes in ecosystem composition, structure, and
function are expected to occur at northern latitudes and higher altitudes. In some areas novel ecosystems likely will replace
existing subalpine, alpine, boreal forest, and tundra ecosystems.
Key words: climate change, ecodiversity, forest, ecosystem diversity, species diversity, genetic diversity
RÉSUMÉ
La diversité écologique (le produit de la diversité des écosystèmes, des espèces et de la diversité génétique) sera significativement modifiée au cours du XXIe siècle en réaction à l’influence combinée du climat, des activités humaines, du
déplacement des espèces indigènes et non –indigènes et des perturbations naturelles comme les feux de forêts (également
modifiés par le climat). Plusieurs espèces s’acclimateront (variation phénotypique) ou encore s’adapteront (variation
génotypique) aux conditions changeantes. Mais ce ne sera pas le cas pour plusieurs espèces. Les espèces avec un fort taux
de reproduction qui sont capables de se déplacer sur de grandes distances, de coloniser de nouveaux habitats, de tolérer
les humains et de survivre sous une grande variété de conditions bio-physiques seront celles qui connaîtront le plus de
succès dans la recherche d’une nouvelle niche. D’importants changements dans la composition des écosystèmes, dans la
structure et les fonctions devraient s’opérer sous des latitudes plus nordiques et à des élévations supérieures. Dans certaines régions, de nouveaux écosystèmes devraient vraisemblablement remplacer les écosystèmes subalpins, alpins,
boréaux et de la toundra.
Mots clés: changements climatiques, écodiversité, écosystème forestier, diversités des espèces, diversité génétique
Introduction
The remaining half of Earth’s
pre-industrial forest covers
approximately 3500 million
hectares (IPCC 2001a), provides habitat for most terrestrial species, and is as important to the maintenance and
health of the ecosphere as it is
central to human cultures and
their economies. These forested ecosystems will change significantly in coming decades
Paul A. Gray
in response to the continuing
variety of pressures exerted by an expanding human population, including human-induced climate change, which is now
a leading agent of ecospheric change. In fact, since 1900 the
Earth’s surface has warmed by 0.6 ± 0.2oC, and most of this
warming is attributable to human activities since the end of
World War II (IPCC 2001b). The Earth’s surface is projected
to warm by 1.4 to 5.8oC over the next 100 years, with land
areas warming more than the oceans, and with the high lati-
1Ontario
tudes warming more than the tropics (IPCC 2001a). This
increasing atmospheric heat (energy) will continue to change
moisture regimes, wind patterns, the frequency of extreme
events, and Earth’s ecosystems and their constituent organisms. This paper reviews some of the known and potential
impacts of climate change on forest ecosystem diversity in the
northern hemisphere.
Ecological Diversity
Ecological diversity or ecodiversity includes ecosystems
(ecosystem diversity) and the organisms (species and genetic
diversity) that live and die in them. It is in this context that an
ecosystem can be described as a recognizable piece of Earth
space in which the flow of energy and the transformation of
matter in-space-in-time creates networks of organisms,
atmosphere, rock, soil, and water, interacting with each other
and with other ecosystems. Each ecosystem can be described
according to its composition (the elements such as water and
plants), structure (the chemical and physical organization of
the system such as wildlife habitat), and function (the flow
and transfer of energy and the creation and dissolution of
matter). Species diversity is the variety of organisms and
Ministry of Natural Resources, 300 Water Street, Peterborough, Ontario K9J 8M5. E-mail: [email protected]
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655
genetic diversity is the variation of individual genes (polymorphism) that enables populations to adapt to constantly
changing ecological conditions through successive generations.
Describing and Delineating Ecosystems in a Changing
Climate
Ecosystems have horizontal (longitudinal width and latitudinal length) and vertical (above- and below-ground depth)
dimensions, and although every one is unique, the concept of
ecosystem as an entity comprising interacting parts of matter
and energy permits us to describe and organize them in various ways, including into a hierarchy where smaller ecosystems
are nested in larger ecosystems (Fig. 1). We define and characterize ecosystems of a certain size and level in the hierarchy
on the basis of common features, which set them apart from
other ecosystems in other levels. For example, the boundaries
of large, sub-continental ecosystems are a function of northsouth variations in temperature, east-west variations in
humidity (Hills 1961, Bailey 1996), altitudinal variation in
temperature (Bailey 1996), and topography (Wiken 1986),
while the boundaries of smaller ecosystems can be delineated
and classified according to vegetation and soil types (e.g.,
Bergeron et al. 1992, Corns 1992, MacKinnon et al. 1992).
Forces of Ecological Change in the 21st Century
The ecosphere is being altered by the combined effects of
human-induced primary and secondary impacts, climate,
and natural disturbances. Pollution, modification of land and
water bodies, the unprecedented intentional and accidental
global redistribution of species, and unsustainable use of
some natural resources are rapidly affecting ecospheric composition, structure, and function, and in some cases causing
the elimination (and subsequent novel creation) or the reconfiguration of entire ecosystems.
Climate is “average weather” described statistically in
terms of the mean and variability of temperature, precipitation, and wind over time that can range from a few months to
millions of years (IPCC 2001b). Climate is primarily fuelled
by energy (heat) from the sun and created by dynamic and
complex interactions between the atmosphere, the hydrosphere, the cryosphere, land, and organisms. Human-induced
climate change is a secondary (cumulative) impact resulting
from the extraction and burning of fossil fuels, the emission
of manufactured chemicals, the drainage of wetlands, and the
conversion of forests and grasslands to other uses such as
urban development. Current climate change results from an
increase in the amount of energy trapped in the atmosphere
by increased concentrations of carbon dioxide (CO2), nitrous
oxide (N2O), methane (CH4), and other greenhouse gases.
For example, atmospheric CO2 has increased 31% since preindustrial times (IPCC 2001b).
Climate is a significant force at all scales in the hierarchy of
ecosystems where temperature, precipitation, and wind
directly affect isothermal boundaries and the distribution and
abundance of organisms through life cycle events based on
physiological and morphological tolerance limits (Cossins
and Bowler 1987) and phenological cues (Root and Hughes
2005). Climate change indirectly affects ecosystems and their
organisms through its influence on disturbance events such
as fire (Weber and Flannigan 1997, Stocks et al. 1998,
656
Flannigan et al. 2000, Li et al. 2000), insects and pathogens
(Fleming et al. 2002a, b; Hogg et al. 2002), extreme events
such as ice storms (Irland 2000; Smith 2000; Dale et al. 2001;
Hopkin et al. 2001, 2003), and invasive species (Ayres and
Lombardero 2000, Simberloff 2000, Torchin et al. 2003).
Ecosystems and species respond differently to the combined variety of human-caused and natural forces, and in
many ecosystems global warming may not necessarily be the
dominant or the only force of change at the current time.
Land use change in Carolinian forest types of southern
Ontario may exert greater influence on ecosystem composition, structure, and function than natural disturbance and/or
climate change. For example, the southern flying squirrel
(Glaucomys volans) migrated north through the remaining
contiguous forests of southeastern Ontario and southwestern
Quebec, and not the severely fragmented forests of southwestern Ontario (Bowman et al. in review). In the Great
Lakes – St. Lawrence forest on the other hand, the northward
migration of the squirrel has been influenced more by interannual variations in the minimum January temperature and
food availability (Bowman et al. in review). In the Boreal forest, climate-driven fire regimes may be more important to
ecosystem function than the direct effects of climate change
(Weber and Flannigan 1997), while the direct effects of
warmer temperatures at mountain tops will be a primary
force of change to ecosystem composition, structure, and
function (Beniston 2003).
Species Diversity in a Changing Climate
Although climate change impact studies are relatively recent,
compelling evidence provided in the literature indicates that
plants, animals, and other organisms have been and will continue to respond to climate change in a variety of ways (see
Walther et al. 2002, Parmesan and Yohe 2003, Root et al. 2003,
Crick 2004, Malcolm et al. 2005, Parmesan 2005, and many
others). Most species live on other species as parasites, commensals, or mutualists (Thompson 1999), and given that it is
unlikely that all species will respond in the same way to climate change, existing species assemblages will be disrupted
through phenological miscues (Inouye et al. 2000), phenological disjunction between species (Inkley et al. 2004), and differential physiological responses.
In the presence of new and emerging ecological constraints, animal species with a high rate of reproduction that
can move long distances, rapidly colonize new habitats, that
can readily use new forage or prey species, tolerate humans,
and survive in a broad range of physical conditions
(Rejmánek and Richardson 1996, Inkley et al. 2004) will be
most successful in finding and using new niches. Generally,
the survival, distribution, and abundance of plant species will
depend on good health and access to appropriate soil types,
migratory pathways, pollinator species, and asexual and sexual reproduction capabilities (Cherry 1998, Thompson et al.
1998).
To survive in a new habitat or persist in its current habitat,
an organism must be able to complete its life cycle (Fleming
et al. 2002a), and through combinations of physiological and
behavioural responses, individuals will need to successfully
meet emergent ecological challenges (often simultaneously)
in the form of weather, predation, parasites, diseases, food
supply, and shelter. For example, a combination of higher
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Fig. 1. Using the ecological land classification terminology described by Wiken (1986) and Crins (2002), this graphic depicts how
ecosystems of varying size and shape are related to each other in ladder-like levels. Use of this type of classification system allows natural asset managers to design and deliver programs within an ecologically meaningful spatial framework.
summer temperatures, reduced availability of optimal habitat, the northward expansion of white-tailed deer (Odocoileus
virginianus), corresponding elevated predation by wolves
(Canis lupus), and increased mortality from the brain worm
(Parelaphostrongylus tenuis) carried by white-tailed deer suggest to Thompson et al. (1998) that moose (Alces alces) will be
relegated to more northerly areas.
Genetic Diversity in a Changing Climate
Species capable of responding to climate change through
acclimation and/or natural selection of genotypes produced
by mutation and recombination, and spread by gene flow
(Bawa et al. 1991, Hewitt and Nichols 2005) stand a better
chance of survival in the new, emerging climate than those
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that do not. Species with short generation times, such as
insects and annual plants, are more likely to evolve rapidly
and adapt to ecological change (Brubaker 1986, RodríguezTrelles et al. 1998) than long-lived species such as trees
(Mátyás 1997). Notwithstanding climate variability and
short-term disruptions, long-term and consistent warming
will promote successive generations of directed natural selection (Fleming et al. 2002a), which may be enhanced in species
(e.g., insects) that produce multiple generations each year. As
a result, genotypes best suited for survival in warmer habitats
will become increasingly common, resulting in genetically
adapted populations (Fleming et al. 2002a).
Although there is a significant positive correlation
between temperature and species diversity, the rate of warm-
657
ing in the 21st Century will potentially erode this relationship. While many species evolved in response to climate
change in pre-human times (Harris 1993), and some species
can adaptively evolve over short periods of time ranging
from days to decades (Ashley et al. 2003, Stockwell and
Ashley 2004), many will not have time to adapt to the rate
and magnitude of changing thermal habitats. In addition,
high species diversity is related to stable conditions
(Tambussi et al. 1993), and the faster that climate changes
and continually disrupts habitat structure and function, the
likelihood of adaptation and high species diversity is
reduced.
Species capable of phenotypic and genotypic responses
at a rate dictated by the changing climate reduce their risk of
extinction in a changed climate. For example, Réale et al.
(2003) reported that the timing of breeding in a red squirrel
(Tamiasciurus hudsonicus) population in the southwestern
Yukon advanced as a result of the combination of phenotypic changes within generations (phenotypic plasticity)
and genetic changes (micro-evolutionary response) among
generations in response to rapidly changing habitat. It is
important to remember, however, that as temperatures continue to rise there is a point on the thermal continuum at
which this, and any other, species will loose the phenotypic
and/or genotypic ability to respond.
Ecosystem Diversity in a Changing Climate
Ecosystem boundaries based in whole or in part on climate
and vegetation will change in response to temperature and
humidity regimes and the different metabolic response of
species to changing thermal regimes (Root and Hughes
2005). For example, large sub-continental ecosystem
boundaries based on north-south variations in temperature
and east-west variations in humidity such the site regions
(ecoregions) described by Hills (1961) and Crins (2002) for
Ontario will shift in response to a warmer climate. And the
boundaries of smaller ecosystems based on vegetation will
be re-defined by species from new configurations in
response to the combined impacts of climate change,
human activities, and natural disturbance (Peters 1992,
Walter and Patterson 1994, Davis et al. 1998).
The most significant changes in ecosystem composition,
structure, and function are expected to occur at northern
latitudes and higher altitudes such as boreal forest ecosystems, where changes in weather-related disturbance regimes
(e.g., fire) and nutrient cycling are primary controls on productivity (IPCC 2001a). Malcolm et al. (2002a) estimated
migration rates of vegetation biomes in response to climate
change using several General Climate Models (GCMs) and
two vegetation models (MAPSS and BIOME3), and determined that northern ecosystem species require very high
migration rates to keep up with the projected rate of climatic change. In a related study, Malcolm et al. (2002b) used
MAPSS and BIOME3 to model equilibrium distribution of
generalized plant types for the present and future climates
and determined that high latitude boreal forest ecosystems
and arctic ecosystems are particularly vulnerable. In fact,
some subalpine, alpine, and boreal forest ecosystems are
expected to disappear completely and be replaced by novel
configurations. Examples of known and potential climate
induced ecosystem change include:
658
Loss of Alpine Forest
Climate, vegetation, and hydrology change rapidly with altitude over relatively short distances in mountainous ecosystems creating high biodiversity, sharp vegetation ecotones,
and equally rapid changes from vegetation and soil to snow
and ice (Beniston 2003). Mountains are susceptible to rapid
climate change because many ecosystems, their habitats and
species, are endemic as a result of isolation at the higher elevations. Some lower altitude ecosystems will move upwards
(Woodward et al. 1995) while upper altitude systems will disappear (Cumming and Burton 1996). In the United States,
several forest ecosystems, including alpine and subalpine
spruce/fir forest types, will decline. In the western United
States, for example, habitats for several subalpine conifers,
including Engelmann spruce (Picea engelmannii), mountain
hemlock (Tsuga mertensiana), and several fir (Abies) species
are projected to contract (Hansen and Dale 2001). Grabherr
et al. (1994), Theurillat and Guisan (2001) and others suggest
that species restricted to low mountain tops or whose range is
limited by soil and other factors are particularly susceptible to
loss.
Subalpine Birch Forests to Subalpine Heath Communities
in Finland
As a result of multiple ecosystem changes, resilience of longestablished disturbance recovery regimes can be disrupted
resulting in an ecosystem flip. For example, as a result of climate-modified defoliation patterns of the autumn moth
(Epirrita autumnata) and reindeer (Rangifer tarandus tarandus) grazing patterns in northern Finland, subalpine birch
forest was replaced by subalpine heath communities with little potential for re-establishment of forest in the future
(Chapin et al. 2004).
Boreal Forest to Wetland
A warmer climate will eliminate large areas of continuous
and discontinuous permafrost in circumpolar ecosystems.
GCM experiments by Smith and Burgess (1999) indicate that
under a 2 CO2 scenario permafrost could disappear from
half of the present-day Canadian permafrost region, because
higher ground temperatures and deeper seasonal thawing will
stimulate thermokarsting. This warming will cause some
ecosystems to flip. For example, some boreal forest ecosystems in central Alaska were transformed into extensive wetlands during the last few decades of the 20th Century as a
result of thermokarsting (Oosterkamp et al. 2000).
Boreal Forest to Great Lakes – St. Lawrence Forest
Paleoecological evidence indicates that ecosystems have successfully responded to climate change in the past. For example, at the end of the last ice age Boreal and Great Lakes – St.
Lawrence forest ecosystems followed the retreating glaciers,
and during a warmer period from 7000 to 3000 B.P., thermal
habitats were suitable for Great Lakes-St. Lawrence forest as
far north as Timmins before receding to a line south of
Gogama (Liu 1990). It is anticipated that that Great Lakes –
St. Lawrence forest ecosystems will move north in response to
temperature, and their characteristics will be similar to current forest composition and structure in areas where fire
regimes remain the same or are less severe than today
(Thompson et al. 1998).
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Treeline to Boreal Grassland-Steppe
A number of northern hemisphere treeline migration studies
demonstrate northward shifts during warming in the early
part of the 20th Century. Migration was less pronounced during the warming trends of the latter part of the 20th Century,
perhaps in response to some combination of water stress,
insect infestation (Kullman 1986, Lescop-Sinclair and Payette
1995, Briffa et al. 1998), and other factors. This response by
treeline species could potentially lead to novel (potentially
non-forested) ecosystems in these areas over the next century. For example, Chapin and Starfield (1997) simulated the
advance of the Alaskan treeline and estimated a 150- to 250year time lag in the forestation of Alaskan tundra and suggested that with rapid warming under dry conditions a novel
ecosystem type, a boreal grassland-steppe, could emerge.
Evidence for this scenario comes from lack of response of
trees to warming in Alaska (Barber et al. 2000) and
Fennoscandia (Linderholm et al. 2003).
Boreal Forest to Aspen Parkland
In boreal ecosystems with warmer, drier climates and a higher fire severity index, a temporal disequilibrium will occur
because the rotation time exceeds the development of a new
climate regime (Suffling 1995, Thompson et al. 1998). Fire
frequency influences post-fire succession, and with sufficiently frequent fires, forests can become shrublands, followed by
prairie (Clark 1990). For example, Schindler (1998) reports
that due to increased evaporation and a decline in precipitation, several large fires burned in the Experimental Lakes
Study area of northwestern Ontario in 1974, followed by
rapid vegetation recovery that was subsequently impacted by
drought, creating more fuel and a second burn in 1980, which
resulted in a denuded landscape followed by a slow recovery
of trees. In many areas, bedrock remained exposed after 17
years. Even at maturity this forest will be less dense and dominated by deciduous trees like trembling aspen (Populus
tremuloides) and balsam poplar, (Populus balsamifera) resembling the arid aspen parkland forests of western Canada, as
predicted by Rizzo and Wiken (1992) and Hogg and Hurdle
(1995).
Summary Remarks
Greenhouse gas emissions will contribute to ecospheric
change throughout the 21st Century. While it is widely recognized that stabilization of greenhouse gas emissions is a critical part of any serious attempt to mitigate the impacts of
global warming, and initiatives under the auspices of the
Kyoto Protocol will lead to reductions, the realized size of the
reductions is unknown at this time. Accordingly, forest management agencies, industry, and forest-dependent communities must begin preparations to adapt to a range of climatic
conditions and emerging ecosystems (Spittlehouse 2005). Key
management strategies include:
1. Understanding the impacts of climate change through science, including research, inventory, monitoring, and
assessment (IPCC 2001a, b).
2. Mitigating the impacts of climate change using coping
strategies comprised of an integrated suite of tools that
include institutional commitment and support, partnership, policy initiatives based on the principles of adaptive
management, strategic planning, and effective on-site
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management programs (Hansen et al. 2001, Gray 2004,
Scott and Lemieux 2005).
3. Helping people adapt through economic diversification,
and on-going education, extension, and training programs
(Williamson et al. 2005).
Acknowledgements
I thank S. Colombo, D. Spittlehouse, M. Taylor and
T. Williamson for providing comments on an earlier version
of this paper.
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