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Resurrection ecology and global climate change research in freshwater ecosystems
Author(s): David G. Angeler
Source: Journal of the North American Benthological Society, 26(1):12-22. 2007.
Published By: The Society for Freshwater Science
DOI: http://dx.doi.org/10.1899/0887-3593(2007)26[12:REAGCC]2.0.CO;2
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%5D2.0.CO%3B2
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J. N. Am. Benthol. Soc., 2007, 26(1):12–22
Ó 2007 by The North American Benthological Society
PERSPECTIVES
This section of the journal is for the expression of new ideas, points of view, and comments on topics of interest to
benthologists. The editorial board invites new and original papers as well as comments on items already published in
J-NABS. Format and style may be less formal than conventional research papers; massive data sets are not
appropriate. Speculation is welcome if it is likely to stimulate worthwhile discussion. Alternative points of view should
be instructive rather than merely contradictory or argumentative. All submissions will receive the usual reviews and
editorial assessments.
Resurrection ecology and global climate change research in
freshwater ecosystems
David G. Angeler1
Institute of Environmental Sciences (ICAM), University of Castilla–La Mancha, Avda. Carlos III s/n,
E-45071 Toledo, Spain
Abstract. The complex effects of global climate change on freshwater ecosystems limit our ability to
predict biological responses in a standard way and may compromise ecosystem management with respect
to potential changes. I present a theoretical framework that shows the usefulness of resurrection ecology for
standardizing cross-system comparisons of ecological responses to global climate change. Resurrection
ecology makes use of plant seed and animal resting-egg (propagule) banks that integrate past
environmental histories in the gene pools of their organisms. Resurrected organisms that have undergone
different periods of dormancy can be studied comparatively using evolutionary/genetic and experimental
approaches. Both approaches combined can provide insights into how the dimensions of species’ ecological
niches have shifted over time and could help reveal whether direct effects of climate change (increased
temperatures and CO2 concentrations and hydrological alterations) or other anthropogenic stressors (e.g.,
contamination, landuse change) have caused microevolution. Insights gained from resurrection ecology
could be used to manage gene flow between populations and to help prevent extinctions of threatened
populations. These insights could be used to help manage ecosystem structure and function and maintain
ecological sustainability. However, our ability to apply results from resurrection ecology to organisms that
do not have long-term dormancy stages in their life cycles may be limited, and the usefulness of
resurrection ecology will have to be evaluated along gradients of hydroperiod and flood frequency, which
may determine rates of microevolution in aquatic ecosystems.
Key words: anthropogenic stress, contemporary evolution, dormancy, seed and resting egg banks, paleolimnology.
‘‘The contribution of climate change to future extinctions
depends on how quickly species can respond to change.’’
McCarty (2001)
(life-cycle events triggered by environmental cues),
geographic range, and physiology have been attributed to climate change. Worst-case scenarios depict
increasing rates of extinction and loss of biodiversity
resulting from physiological stress, interactions with
invading species, habitat loss, and habitat conversion
(Vitousek 1994, Hughes 2000, Root et al. 2003, Thomas
et al. 2004).
The strong consensus in the scientific community is
that freshwater ecosystems are particularly vulnerable
to projected climate changes (Carpenter et al. 1992,
The hypothesis that the Earth currently is undergoing anthropogenic global climate change has accumulated considerable supporting evidence. Global climate
change already is generating ecological responses
(reviewed by Hughes 2000, McCarty 2001, Walther et
al. 2002, Parmesan and Yohe 2003). Shifts in phenology
1
E-mail address: [email protected]
12
2007]
GLOBAL CLIMATE CHANGE
Firth and Fisher 1992, Magnuson et al. 1997, Lake et al.
2000, Lodge 2001, McCarthy et al. 2001, Schindler
2001, Poff et al. 2002, Alvarez-Cobelas et al. 2005a).
Aquatic systems are generally resilient to short- and
long-term anthropogenic disturbances, but the pace of
climate change is probably too rapid to allow many
aquatic organisms to adapt to future environmental
conditions. Many species are thought to be at risk
because their ability to adapt through natural selection
is limited and because anthropogenic alteration of the
environment reduces the dispersal and migration
potential of organisms, thereby limiting their ability
to find suitable habitat (Poff et al. 2002). Moreover, the
synchronization between life-cycle events and seasonal
changes in habitats may be disrupted (Winder and
Schindler 2004, Harper and Peckarsky 2006), limiting
the ability of organisms to reproduce.
Cumulative effects and complex interactions of
climate-driven environmental changes (chiefly alterations of natural hydrological cycles) and other
anthropogenic disturbances (e.g., contamination, habitat fragmentation, introductions of exotic species)
limit our ability to forecast with certainty how aquatic
ecosystems and their biota will change in response to
novel environmental conditions (e.g., Dale 1997,
Magnuson et al. 1997, Schindler 2001), thereby leading
to ecological surprises (Paine et al. 1998). This problem
limits our ability to manage aquatic ecosystems in a
way that will be sustainable and will conserve
ecosystems services for human populations that are
projected to increase over the next several decades
(Magnuson et al. 1997, IPCC 2001).
Mechanistic relationships between environmental
changes and biological responses must be understood
if we are to adopt sound mitigation and conservation
strategies in the future. I provide a theoretical
framework that shows how resurrection ecology (Kerfoot et al. 1999) could be used to predict evolutionary
responses of organisms to environmental changes
caused by global warming. Resurrection ecology has
the potential to enable us to determine whether higher
temperatures, higher CO2 concentrations, and hydrological alterations (direct effects of climate change) or
other anthropogenic stressors (contamination, exotic
species, landuse change) could exert strong selection
pressure on biota in aquatic ecosystems. Comparisons
of results obtained from local studies of lake and
wetland sediments could enable freshwater ecologists
to assess the consistency of evolutionary responses
across regional and climatic domains. Remedial
actions intended to mitigate the impacts of global
climate change could include artificial management of
gene flow between isolated populations that have
different abilities to adapt to environmental change (as
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RESURRECTION ECOLOGY
13
revealed by resurrection ecology), which could help
counteract effects that might lead to extinction.
Back to the Future: What Can We Learn from Past
Adaptations to Rapid Climatic Change? The
Pleistocene–Holocene Transition
‘‘Paleolimnological reconstructions . . . can provide evidence of the past environmental and climatic conditions that
impacted [aquatic] communities.’’ Porinchu et al. (2003)
Current rates of global warming are unprecedented
based on the last 10,000 y of paleoclimatological
evidence. Global mean surface temperature increased
by ;0.68C during the 20th century, and modeling
approaches using different CO2 emission scenarios
suggest that temperature will increase another 1.4 to
5.88C by 2100 (IPCC 2001), depending on the
willingness of humans to reduce current CO2 emissions. Independent of which scenario will have
happened in 100 y, managing ecosystems to guarantee
sustainability will be challenging.
Paleoclimatologists advocate learning from the past
to estimate future shifts in ecological systems related to
the projected range of temperature increases. We can
use the paleoecological record to trace other relatively
abrupt changes in Earth’s climate history and determine biological responses to those events. Those
records might enable us to generalize about the rate
and degree of biological adaptation to global warming
(but see Ammann et al. 2000 for methodological
difficulties).
The abrupt change in climate between the Pleistocene and the Holocene is an instructive example.
According to paleoclimatological data (e.g., Brauer et
al. 1999), temperatures (derived from O2 isotopic
concentrations) appear to have increased ;78C (Johnsen et al. 1992, Alley et al. 1993) in ;50 y during the
transition from the Younger Dryas cold period to the
Preboreal warm period ;11,600 calibrated y (paleoclimatological reconstructions calibrated to calendar
years) ago. Reconstructions of terrestrial habitats using
remains of plants (summary in Hoek 2001) and beetles
(Atkinson et al. 1987) and aquatic studies using
remains of chironomids (Ammann et al. 2000, Porinchu
et al. 2003, Massaferro et al. 2005), cladocerans
(Ammann et al. 2000), algae, or plant seeds (Brauer et
al. 1999) suggest that many ecosystems responded very
quickly to the transition. Many systems showed no lag
or responded in ,20 y, whereas other systems had
slower response times (hundreds of years). However,
most of the biological shifts must be understood as
catastrophic collapses of ecosystem structure or as
switches between different stable states. Only a few
systems showed gradual biological change in response
14
D. G. ANGELER
to abrupt warming. Abrupt shifts in community
structure probably disrupted functional interactions
among species and destabilized ecosystems (Ammann
et al. 2000). Paleoecological evidence also suggests that
the biological changes varied from site to site and were
influenced by local and regional environmental characteristics. Hoek (2001) suggested that the magnitude
of responses to temperature change might also have
been strongly influenced by nonclimatic variables
including migration lags, edaphic characteristics, landscape settings, dryness, and windiness.
Global Climate Change in Freshwater Ecosystems:
Projected Impacts and Biological Responses
‘‘Investigations into climate change have begun to
highlight the importance of strongly nonlinear, complex,
and discontinuous responses.’’ Schneider and Sarukhan
(2001)
Examples from the Pleistocene–Holocene transition
show that biological responses to climate change were
highly variable, even within similar ecosystem types.
Highly variable responses also are expected as a result
of present-day anthropogenic interference with the
climate, despite differences in the spatiotemporal
pattern (abrupt in the past vs creeping in the present)
and nature (shifting from cold to warm in the past vs
warm to warmer in the present/future) of the 2
transitions. Moreover, humans have inflicted other
forms of stress that were not a factor for ecosystems in
the past and that will make the biological responses of
freshwater ecosystems to climate change even more
complex and unpredictable (IPCC 2001).
Many modeling studies indicate that slight increases
in temperature will have profound effects on the
amount and frequency of precipitation and that these
effects will alter runoff patterns and hydrological
functioning of aquatic ecosystems (IPCC 2001, Poff et
al. 2002). A detailed review of these effects is beyond
the scope of my paper, but the hydrological alterations
(precipitation patterns, evapotranspiration, hydroperiods, flood frequency, lake mixis, etc.) may vary
strongly between different regions and climatic domains (IPCC 2001, Poff et al. 2002). Anthropogenic
stressors will continue to degrade aquatic ecosystems
through unsustainable land uses (e.g., habitat fragmentation, contamination) and other practices that
alter hydrology (e.g., overexploitation of ground
water, dam construction). New combinations of
anthropogenic stress and climate change will affect
ecosystems at the local scale, and these combinations
could be more deleterious than the effects arising from
climate change alone (Przeslawski et al. 2005, Xenopoulos et al. 2005, Franco et al. 2006). These effects
[Volume 26
will depend on local, regional, and climatic conditions
and will affect ecological responses across the biological hierarchy from organisms to communities (Fig. 1).
Several mechanisms are thought to influence responses of aquatic organisms to climate change and
other anthropogenic stressors (Fig. 2). Migration and
dispersal are likely to be especially important because
climate change will pose significant challenges to
aquatic organisms seeking suitable new habitat (e.g.,
Poff et al. 2002). Natural genetic adaptation also is
likely to be a key mechanism because organisms may
be able to adapt rapidly to environmental change
through contemporary evolution (i.e., adaptation
occurring within time periods of years and decades;
reviewed by Ashley et al. 2003, Stockwell et al. 2003).
The potential for rapid adaptation to climate change is
encouraging because reduced gene flow between
populations caused by habitat destruction and fragmentation may force populations to rely more on
genetic adaptation than on migration/dispersal to new
habitats (Etterson and Shaw 2001).
Contemporary evolution in plants and animals has
occurred in response to a wide range of anthropogenic
disturbances, including exotic species introduction,
contamination, and overharvesting (Thompson 1998,
Ashley et al. 2003, Stockwell et al. 2003). Most of our
understanding of rapid evolutionary responses to
climatic change relates to adaptation to temperature
at the molecular level for some cellular processes
(Clarke 2003). How these molecular adaptations to
changes in temperature will be reflected in organism
performance and fitness and how these adaptations
will affect population, community, and ecosystem
processes are not well understood (Clarke 2003).
A few recent studies have shed light on how
ecological systems may adapt evolutionarily to climate
change. Studies of evolutionary responses of model
organisms (e.g., Drosophila, Arabidopsis) to increases in
temperature or CO2 over multiple generations in the
laboratory (e.g., Cavichi et al. 1995, Ward and Kelly
2004) indicate that adaptation to selective agents can
be rapid (Ward et al. 2000). Rates of natural genetic
change in annual plants (Etterson and Shaw 2001),
Drosophila (Rodrı́guez-Trelles and Rodrı́guez 1998,
Levitan 2003, Levitan and Etges 2005), other dipterans
(Bradshaw and Holzapfel 2001), or mammals (Berteaux et al. 2004) have been measured in field studies.
A detailed review of these studies is beyond the scope
of my paper, but the available evidence highlights
several important features:
1. As in the Pleistocene–Holocene transition, biological
responses may be complex, hierarchical, and multivariate, making generalizations across ecological
2007]
GLOBAL CLIMATE CHANGE
AND
RESURRECTION ECOLOGY
15
FIG. 1. Conceptual model summarizing impacts in freshwater ecosystems that will arise as a result of disruption of natural
hydrological functioning mediated by global climate change and other anthropogenic stressors as influenced by local, regional, and
biome characteristics.
systems difficult. The type of response will vary
among populations and communities depending on
dispersal strategies and natural (latitudinal and
altitudinal boundaries) and anthropogenic constraints (e.g., dams, land use) on dispersal, migration, and natural selection (Fig. 2). The type of
response and the constraints on dispersal will affect
population genetics, source–sink dynamics, metapopulations and metacommunities, and other community-structuring concepts.
2. Adaptation to climate change is taxon and context
specific and difficult to predict (Holt 1990, Gomulkiewicz and Holt 1995, Grant and Grant 1995,
Abrams 1996, Stockwell et al. 2003). Some species
may adapt to change, but with negative consequences. For example, microevolutionary changes
could result in depauperate genomes and loss of
fitness at the population level (Rodrı́guez-Trelles
and Rodrı́guez 1998, Stockwell et al. 2003), leading
to reduced evolutionary potential because of reduced gene flow or genetic drift. In other cases, rates
of adaptive evolution may be unable to keep pace
with the predicted rate of climate change (Etterson
and Shaw 2001).
3. Our inability to assess mutation rates in relation to
climate change (a technical limitation) will influence
our ability to measure the degree to which natural
selection will determine responses to climate change
(Fig. 2; Berteaux et al. 2004).
4. Methods problems associated with field studies
limit our ability to attribute observed changes
unambiguously to global warming. Variables other
than climate can easily confound the effects of
climate change (Hughes 2000).
The degree of certainty with which we can predict how
biological systems in nature will respond directly or
indirectly to climate change depends on how well we
16
D. G. ANGELER
[Volume 26
FIG. 2. Conceptual model showing biological responses to global climate change across different spatial scales and the potential
uses of resurrection ecology for assessing those responses. The model emphasizes contemporary evolution and adaptation to novel
ecosystem characteristics that may arise from complex biotic and abiotic changes across different spatial scales.
can integrate disturbance histories, biological interactions, and evolution in experiments. I provide arguments below that will show the usefulness of
resurrection ecology for this task.
Finding the Cause of Change: The Potential of
Resurrection Ecology
‘‘Lake sediments store genotypes and species from the past
and so can provide insight into the rates and trajectories of
past ecological and evolutionary changes.’’ Hairston and
Kearns (2002)
The complex impact–response patterns caused by
global climate change in aquatic ecosystems limit our
understanding of biological responses to climate
change. However, this knowledge will be essential
for deriving sound management options in the future.
Determining the forces that drive natural selection in
ecosystems in response to change could be especially
helpful. However, Conner (2003) pointed out the
difficulty of assigning specific causes of natural
selection in the wild. These difficulties arise because
the present-day adaptations of many organisms
cannot be attributed unambiguously to one or more
selective forces acting within the multidimensional
selection regimes in nature. Our inability to benchmark autecological traits of present-day organisms
against those of their evolutionary ancestors further
complicates the issue. Resurrection ecology has strong
potential to overcome these limitations.
Resurrection ecology is based on the notion that
plant seed and animal resting-egg (propagule) banks
are spatiotemporal integrators of genetic-, phenotypic-,
species-, and community-level variation, which can
serve as important biodiversity surrogates in aquatic
ecosystems. Traditionally, the record of organism
remains in the propagule bank has been used
descriptively to reconstruct paleoenvironments and
2007]
GLOBAL CLIMATE CHANGE
historical environmental changes. However, propagule
banks contain resting stages that can remain viable for
decades to centuries (Hairston et al. 1995). Therefore,
resurrection of organisms that have been dormant for
known lengths of time, extraction and analysis of their
DNA, and comparisons of autecological traits along
different niche axes offer great promise for experimental determination of microevolution. This combination
of paleoecological, genetic, and experimental disciplines has given rise to the promising research area of
resurrection ecology (Kerfoot et al. 1999). The techniques have been used successfully with Daphnia to
assess evolutionary responses to metal contamination
(Kerfoot et al. 1999), morphological (Kerfoot and
Weider 2004) and phototactic behavioral responses to
predation (Cousyn et al. 2001), and responses to toxic
cyanobacteria during eutrophication (Hairston et al.
1999, 2001).
When, Where, and How Will Resurrection Ecology
Work in Climate-Change Research?
Constraints on resurrection ecology
Life-history traits.—Use of resurrection ecology is
limited to organisms that remain dormant in sediments for long periods, e.g., protists (Fryxell 1983,
McQuoid et al. 2002) and microinvertebrates (Hairston
et al. 1995, Belk 1998, Thorp and Covich 2001).
Macrophytes (Baskin and Baskin 1998) and many
aquatic insects (Williams 2006) also have dormant
stages (e.g., seeds, turions, eggs, and pupae) in their
life cycles. However, we do not know how long many
of the species in these groups can survive in resting
stages, rendering the application of resurrection
ecology to them uncertain. Many other important
biota in aquatic ecosystems (reptiles, amphibians,
birds, and fish) do not produce long-lived resting
stages, and resurrection ecology cannot be applied to
these groups (Fig. 3A).
Abiotic.—Propagule banks are present in many types
of aquatic ecosystems, including lakes of all depths,
permanent and temporary ponds, riverine floodplains,
swamps, and coastal marshes. Riverine floodplains are
integral parts of many fluvial systems. If the signatures
of anthropogenic stress in fluvial systems can be read
in floodplain propagule banks, resurrection ecology
may offer the potential for determining responses to
climate change in fluvial systems as well as in lentic
systems. Thus, resurrection ecology could provide
useful and easily standardized tools for making crosssystem comparisons of evolutionary change and could
be useful for comparing responses to environmental
stress across regions and climatic domains.
However, potential abiotic constraints to contempo-
AND
RESURRECTION ECOLOGY
17
rary evolution must be considered if we are to evaluate
further the usefulness of resurrection ecology for crosssystem comparisons. Organisms must be able to
undergo full life cycles constantly for microevolution
to take place. Thus, the ability to respond evolutionarily to environmental change is expected to diminish
along gradients of habitat permanence and flood
frequency (Fig. 3B). The prerequisite of constant
reproduction is met in permanently flooded systems
such as lakes (left end of the gradient in Fig. 3B).
However, climate change could outpace the rate of
adaptive genetic responses in ephemeral wetlands that
flood only sporadically on a temporal scale of decades
(right end of the gradient in Fig. 3B). In such habitats,
resurrection ecology is unlikely to be useful for
detecting evolutionary responses to change.
Temporary wetlands have been traditional choices
for conversion to agricultural use, a practice that will
pose an increased threat as global climate changes
push aquatic ecosystems toward shorter hydroperiods
and reduced flood frequencies (Alvarez-Cobelas et al.
2005b). Once the structure of the propagule bank is
destroyed by agricultural practices, potential responses to environmental change can no longer be studied
using resurrection ecology. Moreover, some evidence
suggests that disruption of natural hydroperiods and
flood frequencies can degrade propagule banks over
time (M. Brock, University of New England, Armidale,
Australia, personal communication). This evidence
suggests that propagule banks can deteriorate to a
point that could reduce the usefulness of resurrection
ecology unless adaptive management helps to counteract excessive ecological degradation (Hulme 2005).
Because resurrection ecology can be applied only to
a limited set of organisms in specific kinds of
environments, the inferences that can be drawn from
resurrection studies may be limited. For example,
many aquatic vertebrates have longer life spans and
generation times than organisms with long-term
dormancy stages in their life cycles. Therefore,
evolutionary responses of aquatic vertebrates to
environmental stress could be decoupled from the
responses of organisms banked in lake and wetland
sediments, i.e., genetic adaptation to stress is expected
to be slower in many vertebrates than in most
invertebrates. As a further example, inferring ecosystem responses (arguably the sum of responses of all
biotic [benthic and pelagic] constituents of the system)
directly from the responses of a small subset of taxa
with specific life-history traits will be problematic.
Nevertheless, the responses of propagules banked in
wetland and lake sediments may be useful surrogates
for ecosystem responses to global climate change (Fig.
3C), and resurrection ecology offers a number of
18
D. G. ANGELER
[Volume 26
FIG. 3. Conceptual model showing the influences of life-history traits of organisms (A) and hydrological characteristics of
aquatic ecosystems (B) on potential uses of resurrection ecology and the potential contributions of resurrection ecology to global
climate change research (C). The dotted lines indicate speculative applications of resurrection ecology.
important advantages over conventional methods of
investigating responses to environmental change
(Table 1).
Advantages
Rapid response times.—Many of the organisms banked
in the sediments of aquatic ecosystems grow quickly
and have short generation times, so their evolutionary
responses to environmental change should be detectable within years or decades (Kerfoot et al. 1999,
Hairston et al. 1999, 2001, Cousyn et al. 2001, Kerfoot
and Weider 2004), and trends in responses could be
identified relatively quickly. The impacts of future
climate change will depend on the specific combination
of direct effects of climate change and other anthropogenic stressors imposed on an ecosystem (Fig. 1).
Organism responses to these specific combinations
should be registered over time in the genotypes stored
in the historical archives of the propagule banks, and
this biological memory could help us to determine the
specific forces driving natural selection. More specifically, propagule banks of aquatic ecosystems could act
as useful indicators of Red Queen dynamics (the idea
from Alice in Wonderland that organisms must run just
to remain in place; Van Valen 1973) and help to
GLOBAL CLIMATE CHANGE
2007]
TABLE 1.
AND
RESURRECTION ECOLOGY
19
Advantages and limitations of resurrection ecology in global climate change research.
Advantage
Allows cross-system comparisons in ecosystems that
contain propagule banks
Serves as a standardized indicator for environmental
change across local, regional, and biome scales
Probably could be used to manage gene flow between
populations and to help prevent extinction
Could help inform management of ecosystem services
Integrates paleoecological, evolutionary/genetic, and
experimental disciplines, allowing
straightforward determination of ecological responses
determine the impacts in aquatic ecosystems to which
organisms must adapt evolutionarily.
Potential for experimental manipulation.—Autecological studies of organisms obtained from past and
contemporary resting eggs are probably the most
powerful tools for identifying evolutionary responses
to climate change. Manipulations of present-day and
past niche dimensions could be useful for determining
evolutionary responses of organisms to direct or
indirect effects of climate change (Fig. 2). Specifically
designed manipulative experiments could be used to
address questions such as: 1) Do species in natural
environments adapt to higher temperatures or CO2
levels? 2) Do species evolve to cope with the
hydrological alterations of the ecosystems they inhabit? 3) How will species respond to anthropogenic
stressors such as introduced species, contamination, or
change in land uses? 4) Will species have evolutionary
priorities if different stressors act in concert, i.e., will
evolutionary responses to subtle and slower changes
in temperature be similar to their responses to other
anthropogenic and more-incisive perturbations? 5) If
yes, will these priorities be the same across ecosystems,
regions, and climatic domains? Additional paleolimnological studies based on radiometric dating, pollen,
diatoms, and invertebrate remains could provide
information on the nature and magnitude of environmental change that have acted on genotypes.
Resurrection Ecology: Managing Gene Flow and
Mitigating Impacts of Climate Change
‘‘We may not be able to prevent accidents, but we may be
able to diminish their impact.’’ Rosenzweig (1995)
A long history of ecological experimentation and
theory supports the hypothesis that ecosystem goods
and services and the ecosystem properties from which
they are derived depend on biodiversity (Hooper et al.
2005). The available evidence suggests that biodiversity will decrease because of the interactive effects of
Limitation
Gradients of hydroperiod and flood frequency could affect the
rate of contemporary evolution
Limited to organisms that have long-term dormancy stages
in their life cycles (not applicable to many aquatic vertebrates)
Probably high budgetary and personnel demands
climate change and other disturbances (Thomas et al.
2004, Lovejoy and Hannah 2005). Decreased biodiversity, in turn, will affect ecosystem services (Metzger et
al. 2005, Schröter et al. 2005) and could compromise
human welfare.
Resurrection ecology has the potential to be an
indicator of possible organismal responses to climate
change and other anthropogenic stressors across
ecosystem types. However, its contributions to climate-change research could go beyond simple detection of evolutionary responses to mitigation of the
effects of global climate change through artificial
management of gene flow between populations (Fig.
3C). Research related to natural genetic selection and
artificial management of gene flow in natural systems
(Conner 2003, Stockwell et al. 2003) is still preliminary.
Thus, the ideas presented here must be considered
speculative (reflected by the dotted arrows in Fig. 3C).
However, results obtained from resurrection ecology
studies of the rates and magnitudes of evolutionary
responses to selection forces have some potential to
inform artificial management of gene flow between
populations. If we were able to manage gene flow
among populations with different degrees of evolutionary adaptation to the selective forces accompanying climate change, we could help increase
evolutionary potential, help species resist environmental change, and reduce the risk of extinction. Of course,
artificial management of gene flow within the context
of the results of resurrection ecology studies would be
limited to species with long-term dormancy in their
life cycles. Nevertheless, if the goal of artificial
management of gene flow were to help preserve the
ability of ecosystems to provide ecosystem services,
resurrection ecology could target those services that
are provided directly by organisms with long-term
dormancy as a life-history trait. Application of
resurrection ecology to the management of services
that are provided by the sum of aquatic biota is less
straightforward.
20
D. G. ANGELER
Global Climate Change Research and Resurrection
Ecology: A Synthesis
The nonlinear, complex, and discontinuous nature of
global climate change and biological response patterns
limits our understanding of how ecosystems will
change under projected climate-change scenarios and
how these changes will be distributed across geographical regions and biomes. This lack of knowledge
currently limits our ability to manage ecosystems and
implement strategies that could help mitigate the
effects of global climate change. The strongest application of resurrection ecology lies in the use of
comparative studies of propagule banks that can be
standardized across systems at local, regional, and
biome scales. Resurrection ecology also has some
potential to provide useful insights into ways to
manage gene flow between populations to help
counteract factors that might lead to extinction.
However, the use of resurrection ecology is inherently
limited to organisms with long-term dormancy in their
life cycles, so that results obtained from resurrection
ecology probably cannot be applied to other aquatic
biota (chiefly vertebrates). Thus, resurrection ecology
is less likely to be helpful in efforts to determine
ecosystem responses to global climate change or to
manage ecosystem services derived from the sum of all
biotic constituents of aquatic ecosystems. Given that
future scenarios of the effects of global climate change
are pessimistic regarding loss of biodiversity and
sustainability of ecosystem services, politicians, scientists, and managers should exploit every tool available
that could help conserve our environment (Osmond et
al. 2004). Resurrection ecology could be very useful for
some of these tasks.
Acknowledgements
The input of 2 anonymous referees, W. H. Clements,
and P. Silver helped to strengthen the message of this
research and is gratefully acknowledged. Financial
support was provided by EU and JCCM/FEDER
research grants (EVG1-2002-00019 and PAI-05-020).
This paper is dedicated to the memory of my
grandfather, Joseph Camilleri (1920–2005).
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Received: 2 February 2006
Accepted: 12 September 2006