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Functional
Ecology
2007
21
, 387–393
387
© 2007 The Authors.
Journal compilation
© 2007 British
Ecological Society
Blackwell Publishing Ltd
EDITORIAL
Evolution on ecological time-scales
S. P. C ARRO LL,* † A. P. H ENDRY,‡ D. N. REZ NICK § and C. W. FOX¶
*
Department of Entomology and Center for Population Biology, University of California, Davis, CA 95616 USA
,
‡
Redpath Museum and Department of Biology, McGill University, Montréal, Québec, Canada H3A 2K6
,
§
Department of Biology, University of California, Riverside, Riverside, CA 92521, USA
, ¶
Department of
Entomology, University of Kentucky, Lexington, KY 40546, USA
Summary
1.
Ecologically significant evolutionary change, occurring over tens of generations or
fewer, is now widely documented in nature. These findings counter the long-standing
assumption that ecological and evolutionary processes occur on different time-scales,
and thus that the study of ecological processes can safely assume evolutionary stasis.
Recognition that substantial evolution occurs on ecological time-scales dissolves this
dichotomy and provides new opportunities for integrative approaches to pressing ques-
tions in many fields of biology.
2.
The goals of this special feature are twofold: to consider the factors that influence
evolution on ecological time-scales – phenotypic plasticity, maternal effects, sexual
selection, and gene flow – and to assess the consequences of such evolution – for
population persistence, speciation, community dynamics, and ecosystem function.
3.
The role of evolution in ecological processes is expected to be largest for traits that
change most quickly and for traits that most strongly influence ecological interactions.
Understanding this fine-scale interplay of ecological and evolutionary factors will
require a new class of eco-evolutionary dynamic modelling.
4.
Contemporary evolution occurs in a wide diversity of ecological contexts, but
appears to be especially common in response to anthropogenic changes in selection
and population structure. Evolutionary biology may thus offer substantial insight to
many conservation issues arising from global change.
5.
Recent studies suggest that fluctuating selection and associated periods of contemporary
evolution are the norm rather than exception throughout the history of life on earth.
The consequences of contemporary evolution for population dynamics and ecological
interactions are likely ubiquitous in time and space.
Key-words
: contemporary evolution, anthropogenic change, modern syntesis, natural selection, rapid evolution
Functional Ecology
(2007)
21
, 387–393
doi: 10.1111/j.1365-2435.2007.01289.x
Introduction
Clearly, our thinking must not exclude the possibi-
lity of animals attaining to extremely rapid rates of
evolution ... (Johnston & Selander 1964)
A wave of recent studies has shown that many organisms
can undergo adaptive phenotypic evolution over just
a few generations. Across a wide range of taxa, this
contemporary evolution has been observed in many
biological contexts, on fine spatial scales, and in the
presence of gene flow (e.g. Hendry & Kinnison 1999;
Schluter 2000; Kinnison & Hendry 2001; Palumbi 2001;
Gilchrist
et al.
2004; Hargeby, Johansson & Ahnesjo
2004; Garant
et al.
2005; Strauss, Lau & Carroll 2006;
Hendry
et al.
2006; Phillips
et al.
2006). Moreover, strong
selection on observable time frames has caused sub-
stantial differentiation among populations within species,
in some cases enough to represent the early stages of
speciation (Hendry
et al.
2000; Filchak, Roethele &
Feder 2000; Schwarz
et al.
2005; Hendry, Nosil &
Rieseberg 2007). Evolution may also occur quickly
enough to alter the outcomes of ecological interactions
and ecosystem-level processes (Thompson 1998; Agrawal,
Lau & Hambäck 2006; Fussmann, Loreau & Abrams
2007), including those that generate selection in the first
place (Yoshida
et al.
2003; Hairston
et al.
2005). Given
all of these effects, ignoring contemporary evolution
will likely limit success in many biological disciplines.
†Autho r to whom corresp ondence shou ld be addressed.
E-mail: spcarroll@ucdavis.edu
388
S. P. Carroll
et al.
© 2007 The Authors.
Journal compilation
© 2007 British
Ecological Society,
Functional Ecology
,
21
, 387–393
Recognizing the potentially broad influences of
contemporary evolution leads to many new ques-
tions. Are the high frequencies and rates of evolution
observed in modern times unusual, perhaps the result
our own increasing impact on the selective environments
of other taxa? Or, has evolution always occurred on
ecological time-scales, but been largely overlooked
until recently? How do rates of evolution measured in
ecological time relate to those assessed by macroevo-
lutionary and palaeontological studies? What are the
genetic, developmental, behavioural and demographic
factors that influence rates of contemporary evolution?
And, what are the implications of contemporary
evolution for general principles and problems in
evolutionary biology, ecology, conservation biology,
medicine and agriculture? Empirically, the reality of
contemporary evolution means that many evolutionary
processes thought approachable only by inferential
study, such as speciation, can now be studied directly.
This special feature of
Functional Ecology
starts
from the premise that contemporary evolution is a
common phenomenon. This recognition allows us to
move forward beyond simply documenting the process
to examining the factors that influence it and that are
influenced by it. We specifically emphasize evolution
on ecological time-scales; that is, ecologically significant
evolution occurring on a time frame over which ecolog-
ical studies typically consider populations to be evolu-
tionarily static. Our goal is to synthesize and broadcast
what is known about the biology of contemporary
evolution, to highlight the opportunities it presents,
and to identify current gaps in our understanding.
In this introductory essay, we review the history of
thought on the pace of evolution and its importance in
ecological pattern and process. This introduction is
followed by seven papers – each a general review of
conceptual ideas and empirical support for them –
examining emerging topics in this field. Four of these
papers treat factors that influence evolution on ecological
time-scales – phenotypic plasticity, maternal effects,
sexual selection, and gene flow. The remaining papers
examine the ecological consequences of contemporary
evolution for population persistence, speciation, and
community/ecosystem dynamics.
Natural selection and the pace of evolution
Darwin’s impeccable logic allowed him to infer the
primary mechanism of evolution without being able
to observe it directly. Despite knowing that artificial
selection could produce phenotypic change very
quickly, his intuition about the rate of evolution in
nature appears conservative: ‘We see nothing of these
slow changes in progress, until the hand of time has
marked the long lapse of ages’ (1859, p. 84). Elsewhere,
however, he seems to have considered a contemporary
and reciprocal interplay between ecology and evolution:
‘But it is difficult to tell ... whether habitats generally
change first and structure [morphology] afterwards;
or whether slight modifications of structure lead to
changed habitats; both probably often change almost
simultaneously’ (1859, p. 183). His experiences in the
Southern Hemisphere as naturalist on the
Beagle
sensitized him to the fact that environments do in fact
sometimes change swiftly and suddenly (1860, e.g. pp. 120,
311). Yet, the perception that evolutionary change can
really only be documented through palaeontological
study remained dominant for more than a century.
With the exception of a few examples documenting
that natural selection occurs in nature (e.g. Bumpus
1899), little progress was made towards understanding
the causes and consequences of natural selection
until the rise of population genetics in the 1920s. Bridg-
ing the gap, Turreson (1922, 1930) proposed the study of
‘genecology’ – adaptive properties of populations in
relation to the environment, which include the concept
of ‘ecotypes’. The first thorough experimental field
studies of selection appeared during the Modern
Synthesis. Clausen, Keck & Heisey (1940) conducted
transplantation/common garden experiments that
documented the genetic basis of adaptive morphological
differences across an elevational gradient in the western
North American herb,
Potentilla glandulosa
. During
the same period, Dobzhansky (1948) conducted
myriad studies of adaptation in
Drosophila
, including
the demonstration of cyclical annual evolution in
relation to habitat seasonality. These studies showed
that natural selection is a decipherable, quantifiable
phenomenon, subject to hypothesis testing. Under the
influence of these pioneers, biologists came to recognize
the value of spatial and temporal comparisons for
inferring the role of selection in nature.
The 1950s marked the emergence of the British
School of Ecological Genetics, and Ford’s first book
with that title was published in 1964. Like Clausen’s
work, these were not studies of ‘evolution in action’
per se, but rather inferential analyses of colour variation
in relation to predation (e.g. Brower 1958; Sheppard
1959; Clarke & Murray 1962). Of all such examples,
that of ‘industrial melanism’ in the peppered moth
(
Biston betularia
Linnaeus), became most well known
(e.g. Kettlewell 1956). Along with studies of heavy
metal tolerance evolving in plants on mine tailings
(Antonovics, Bradshaw & Turner 1971), and adaptation
to fertilizer treatment in the Park Grass Experiment
(Snaydon 1970, reviewed in Silvertown
et al
. 2006), this
‘smoking gun’ of the industrial revolution was widely
taught in biology courses as the principal heuristic
exposition of evolution by natural selection. What set
these last examples apart from earlier work was that
adaptation had occurred in contemporary time.
Despite this burgeoning literature on natural selection,
it proved difficult to shake the perception that evolution
and ecology happened on very different time-scales.
Slobodkin (1961) formalized this perspective by
arguing that ‘ecological time’ was the ten-or-so genera-
tion period over which a population might, on average,
maintain a steady state. ‘Evolutionary time’, in contrast,
389
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ecological
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© 2007 The Authors.
Journal compilation
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Ecological Society,
Functional Ecology
,
21
, 387–393
was considered the approximately half-a-million-year
period sufficient for evolutionary change to disrupt
ecological steady states. Bucking this inertia, Johnston
& Selander (1964) and Berry (1964) found convincing
evidence of something quite different. In the first case,
house sparrows (
Passer domesticus
) introduced to North
America diversified phenotypically as they colonized
different habitats. In the second, house mice introduced
to the Welsh island of Skokholm changed dramatically
in skeletal measurements over only 70 years. This was
soon followed by other examples, such as the evolution
in < 100 generations of host races of
Rhagoletis pomonella
(apple maggot fly) in response to introduced apples
(Bush 1969). A few scattered studies in the 1970s
added to the pantheon of contemporary evolution.
The 1980s, however, ushered in an important
innovation in the study of contemporary evolution:
the use of common garden or experimental methods to
determine the genetic basis for phenotypic differences
that have evolved in contemporary time. Chief examples
include Endler’s (1980) and Reznick & Bryga’s (1987;
Reznick, Bryga & Endler 1990) work on colour patterns
and life-history traits in guppies introduced to new
predator environments, Stearns’ (1983) work on life-
history traits in mosquitofish introduced to Hawaii,
and Williams & Moore’s (1989) work on phenotypic
traits in rabbits introduced to New Zealand. These and
other studies confirmed that substantial genetically
based adaptive genetic change can indeed occur in
contemporary time.
Drawing on examples such at these, Endler’s (1986)
book
Natural Selection in the Wild
clearly established
for the first time that evolution by natural selection was
a potent ongoing phenomenon. He reviewed enough
cases to show that measurable responses to con-
temporary natural selection, if not common, were at
least not exceedingly rare or aberrant. Simultaneously,
modern ‘selection thinking’ (e.g. Charnov 1982) was
making broad incursions into ecology, yielding pro-
gressively more evidence of precise specialization and
fine-scale adaptive population differentiation in nature.
The field seemed to explode in the late 1990s,
probably spurred by the publication of high-profile
papers that compared rates of phenotypic change in
the fossil record to those observed in experimental
introductions of guppies (Reznick
et al
. 1997) and
Anolis
lizards (Losos, Warheit & Schoener 1997). These
empirical examples were followed by the influential
reviews of Thompson (1998) and Hendry & Kinnison
(1999), both showing that many studies have found
substantive phenotypic changes occurring on ecological
(contemporary) time-scales: ‘... claims of rapid
microevolution should not necessarily be considered
exceptional, and perhaps represent typical rates of
microevolution in contemporary populations facing
environmental change’ (Hendry & Kinnison 1999,
p. 1650).
This emerging shift in mindset brought into sharper
relief a paradox earlier elucidated by Gingerich (1983):
short-term studies often find dramatic and rapid
rates of phenotypic change, whereas long-term studies
seem to manifest much slower rates of change (see also
Estes & Arnold 2007). Fortunately, long-term studies
of selection and evolution had, by the late 1990s, con-
tinued for long enough to provide some resolution.
Specifically, selection and evolution often fluctuate
dramatically in direction through time, presumably
tracking fluctuating environments, so that rapid short
term changes rarely accumulate into long-term direc-
tional trends (Gibbs & Grant 1987; Hairston & Dillon
1990; Ellner
et al.
1999; Grant & Grant 2002, 2006).
A related concern was the realization that many
examples of contemporary evolution result from
anthropogenic disturbances (Palumbi 2001; Rice &
Emery 2003, Carroll
et al.
2005; Strauss
et al
. 2006;
Hendry
et al.
2006). Since humans are reshaping
selective environments across the globe, contemporary
evolution may be more frequent, and directional
selection may be less ephemeral, in the present than
under more ‘natural’ conditions prevailing in the past
(Strauss
et al
. 2006). Although the global scale of
environment change caused by human disturbance is
largely unprecedented in evolutionary history, the
magnitude of the resulting genetic change does not
seem abnormally high. Instead, observed rates of con-
temporary evolution may mirror historical processes,
and what currently appears ‘rapid’ may have been
common throughout the history of life, though less
contemporaneous among such a wide diversity of
organisms.
If contemporary evolution is often reversible and
rarely accumulates into long-term changes, how do
larger changes in taxa ultimately arise. Comparing
various genetic models, Estes & Arnold (2007) found
that the distribution of observed evolutionary rates
is best explained by a model that assumes a fitness
optimum that can move but within fixed limits (i.e. an
‘adaptive zone’). Under this scenario, most of evolu-
tionary time is characterized by stabilizing selection at
the level of an adaptive zone, within which populations
can move to new local adaptive peaks. Indeed, Darwin’s
finches, which show such dramatic change on short
time-scales (Grant & Grant 2002, 2006), continue to
appear much as they did millions of years ago (Carroll
1997). Macroevolution may thus require progressive
change in the environment that takes evolution con-
sistently in one direction (such as increased body size).
It is important to note, however, that this may some-
times be a gradual process, interspersed with many
temporary reversals of direction. Macroevolution may
thus be nothing more than an aggregate of many small
events, and is entirely explainable by events that we can
observe and quantify over the course of our lives
as investigators. The frequency with which evolution
stemming from ongoing human disturbance will
likewise show a highly iterated or reversible nature,
versus consistent and enduring shifts, remains to be
discovered.
390
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et al.
© 2007 The Authors.
Journal compilation
© 2007 British
Ecological Society,
Functional Ecology
,
21
, 387–393
Historically, macroevolution has been equated with
substantive adaptive change, and rapid and fluctuating
evolution has been regarded as evolutionary noise.
Thus, debate continues over what constitutes trivial
versus important evolution. We argue that the real
distinction between macro- and microevolution may
lie only in the degree to which the factors causing evolu-
tion are fluctuating or are gradually and persistently
directional, and not in the ecological significance of
that evolution.
The ecological significance of evolution
Recognition of the rate and ecological significance of
adaptive evolution is heralding a new epoch where the
dichotomy between ecological and evolutionary time
and, more importantly, between ecological and
evolutionary processes is breaking down. Here, again,
Thompson (1998) was formative in noting that high
rates of evolution, even when transient, are ‘constantly
reshaping populations and interspecific interactions
through frequency-dependent selection, seasonal
fluctuations in allele frequencies, selective death of
genotypes within and among cohorts, and the con-
stantly shifting genetic configuration of populations
and interspecific interactions across broad geographic
landscapes’ (p. 331). In other words, the continuously
evolving face of the biotic environment, including
simultaneously co-evolving relationships among
(and likely within) populations, drives dynamic
variation in selective environments that are linked to
adaptation.
A general prediction is that the importance of
contemporary evolution to ecological interactions will
increase with (i) increasing changes in traits under
selection, and (ii) increasing links between that trait
variation and ecological interactions (Hairston
et al.
2005). Hairston
et al.
2005 propose a quantitative
approach to assessing concurrent rates of evolution-
ary and ecological change in populations, and to mod-
elling the direct contribution of evolution to ecological
change. Using comprehensive and long-term data for
Darwin’s finches and freshwater copepods, they con-
clude that population growth rate can sometimes be at
least twice as sensitive to evolutionary changes as to
important ecological changes in abiotic factors, such
as rainfall. In other words, evolutionary change can have
substantial effects on ecological dynamics.
Thus, while it has long been recognized that adaptive
evolution occurs in an ecological context, it is now
clear that evolutionary change feeds back directly and
indirectly to demographic and community processes,
and that ecological studies should consider this
reciprocal interplay between ecology and evolution.
Because we are now in a period of accelerated environ-
mental change, evolutionary biologists have an unpre-
cedented opportunity to study evolutionary processes
and their influence on ecological processes. Likewise,
evolutionary biology has an unprecedented opportunity
to contribute to other areas of basic and applied biology
(e.g. Palumbi 2001; Stockwell, Hendry & Kinnison
2003; Strauss
et al.
2006; Carroll in press).
What is explored in this special feature?
The foregoing sections illustrate that contemporary
evolution is common and is likely important to ecological
pattern and process. We can thus turn to the main goal
of this special feature, which is to move beyond simple
demonstrations of evolution on ecological time-scales
to a more complete and nuanced consideration of
(i) factors influencing contemporary evolution, and
(ii) the effects of contemporary evolution on specific
aspects of ecology. Here we provide an introduction to
what the reader will encounter in the other contribu-
tions to this special feature.
Genetic and phenotypic interactions within and
among populations and species may strongly influence
the interplay of ecological and evolutionary change,
and thereby determine our ability to predict the impact
of environmental alteration on individual and popula-
tion characters of interest (‘eco-evolutionary dynamics’,
e.g. Kinnison & Hairston 2007; Fussmann
et al.
2007).
To broaden the applicability of eco-evolutionary models,
a next step will be incorporating even more complex
variables and scenarios during simultaneous ecological
and evolutionary change. For example, in determining
the phenotypic and fitness values of traits under selec-
tion, the interaction between genes, development and
ongoing evolution itself should be a fundamental
source of variation (
sensu
Coulson
et al.
2006, Carroll
in press). Two such interactions include phenotypic
plasticity and maternal effects.
Phenotypic plasticity results from the developmental
and behavioral responses of genotypes to environmental
influences, and operates both within and across gen-
erations (Ghalambor
et al.
2007). Plasticity may have
a variety of influences on contemporary evolution,
ranging from being a constraint on adaptive evolution
(owing to reduced selection) all the way to being a pro-
moter of adaptive evolution (by bringing populations
into the domain of attraction of new fitness peaks)
(Price, Qvarnström & Irwin 2003). Plasticity may also
serve to buffer populations from environmental change,
thus reducing the risk of extinction and making
adaptive evolution possible. Maternal effects are like
phenotypic plasticity in that they can change fitness
in the absence of genetic change, and can thereby alter
the frequencies of phenotypes (Rasanen & Kruuk 2007).
Moreover, reaction norms (patterns of plasticity)
and maternal effects may both evolve substantially
across just a few generations (e.g. Hairston
et al.
2001;
Ghalambor
et al.
2007; Rasanen & Kruuk 2007), thus
changing the response of organisms to environmental
heterogeneity.
Sexual selection is another factor that may influence
contemporary evolution, and this process can be
thought of as co-evolutionary dynamics within and
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© 2007 The Authors.
Journal compilation
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Ecological Society,
Functional Ecology
,
21
, 387–393
between the sexes within a species. The tight cycling of
sexual selection in models of mate choice, for example,
suggests that contemporary evolution should be
especially apparent in such systems. Surprisingly,
despite the high volume of scientific activity aimed at
sexual selection in general, few studies have considered
its role in contemporary evolution, although the few
examples include some classics (e.g. Endler 1980).
Svennson & Gosden (2007) conclude that the apparent
rarity of contemporary evolution in secondary sexual
traits may simply result from an absence of long-term
studies of sexual selection within a population biology
framework. It is also possible that inherent peculiarities
of the traits themselves may somehow limit rapid
responses to altered selection pressures (Svennson &
Gosden 2007; Karim
et al
. in press).
Contemporary evolution will be strongly influenced
by rates and patterns of gene flow and dispersal. For
example, gene flow may hinder adaptive differentiation
by reducing the genetic independence of populations
inhabiting different selective environments, but may
sometimes promote adaptive differentiation by sup-
plementing the genetic variation on which selection
can act. Gene flow can thus variously promote or retard
adaptive divergence between populations, and may
thus act to structure adaptation across the geographic
range of a species. Garant, Ford & Hendry (2007)
highlight the important differences between traits
adapting to relatively static environmental conditions
versus those caught up in co-evolutionary dynamics,
with the latter likely benefiting more from the genetic
variation brought by gene flow. One of the main
implications of this perspective is that environmental
change will alter the complex, dynamic relationships
between adaptation and gene flow, and will therefore
influence adaptive divergence and speciation (Hendry
et al.
2007). Gene flow may have similar complicated
and dynamic influences on population persistence
(Kinnison & Hairston 2007).
Lastly, the eco-evolutionary dynamics of inter-
specific interactions may strongly influence community
dynamics, food web structure and ecosystem function.
Fussmann
et al.
(2007) review a series of simple
theoretical models that have long shown important
feedbacks from evolution to ecology – if only empiri-
cists had paid more attention! The main theoretical
hurdle is moving beyond the current simple models to
ones that more accurately assess the complexities of
ecological communities and ecosystems. Empirical work
in natural populations has lagged behind theoretical
developments, but we now at least have a number of
intriguing examples of how genetic variation in
populations can influence community dynamics and
ecosystems function (Whitham
et al
. 2006). The next
step is to document how contemporary
changes
in
genetic variation influence these same dynamics and
functions. Some of the best examples here come from
interactions between hosts and pathogens in introduced
species (Fussmann
et al.
2007).
The frequency, magnitude and importance of
evolution on ecological time-scales are likely to grow
as human environmental impacts increase and their
effects ramify. The dialogue in evolutionary biology
will soon be dominated by discussions of systems in
which evolution is ongoing, directional and in many
cases non-reversing because environments are changed
so substantially. Community and conservation ecology
will necessarily become microevolutionary disciplines,
and other disciplines will follow. As part of this trans-
formation, discussions of evolutionary rates and their
significance will broaden as we progress from more
unified perspectives about the challenges and prospects
to be addressed. One such perspective has given birth
to the nascent field of evolutionary conservation man-
agement, in which genotypes and population processes
may be manipulated to promote the resilience of
threatened populations, combat invasive species, or
promote coexistence (e.g. Ashley
et al.
2003; Stockwell
et al.
2003; Carroll & Watters in press). Evolution on
ecological time-scales makes conservation efforts more
complex, as taxa become ecological moving targets.
Yet many conservation goals will only be met with the
assistance of adapting taxa, even if prior phenotypic
and genotypic states disappear as a result (e.g. Carroll
et al.
2005). Theoretical advances are bringing us
closer to applied fields of evolutionary demography
and community ecology (Kinnison & Hairston 2007;
Fussmann
et al.
2007, respectively). To the extent that
contemporary evolution is inevitable, predictable, and
central to the generation of biodiversity, we must har-
ness its power to ameliorate damage and reconstitute
species and ecosystem functions. Doing so may be the
chief path towards sustainable conservation practice.
Acknowledgements
N. Hairston Jr., C. Ghalambor, and P. Abrams contributed
directly to this review, and G. Fussmann, L. Rieseberg,
E. Svensson, and S. Forde also made helpful comments.
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Received 17 April 2007; accepted 18 April 2007
Editor: Liz Baker
