Diversity and Distributions - 2010 - Thomas - Climate Climate Change and Range Boundaries

Diversity and Distributions, (Diversity Distrib.
) (2010) 16, 488–495

A Journal of Conservation Biogeography
BIODIVERSITY Climate, climate change and range
REVIEW
boundaries
Chris D. Thomas*
Department of Biology, University of York, ABSTRACT

Heslington, York YO10 5YW, UK
Aim A major issue in ecology, biogeography, conservation biology and invasion
biology is the extent to which climate, and hence climate change, contributes to
the positions of species’ range boundaries. Thirty years of rapid climate warming
provides an excellent opportunity to test the hypothesis that climate acts as a
major constraint on range boundaries, treating anthropogenic climate change as a
large-scale experiment.
Location UK and global data, and literature.
Methods This article analyses the frequencies with which species have responded
to climate change by shifting their range boundaries. It does not consider
abundance or other changes.
Results For the majority of species, boundaries shifted in a direction that is
concordant with being a response to climate change; 84% of all species have
expanded in a polewards direction as the climate has warmed (for the best data
available), which represents an excess of 68% of species after taking account of the
fact that some species may shift in this direction for non-climatic reasons. Other
data sets also show an excess of animal range boundaries expanding in the
expected direction.
Main conclusions Climate is likely to contribute to the majority of terrestrial and
Diversity and Distributions
freshwater range boundaries. This generalization excludes species that are

endemic to specific islands, lakes, rivers and geological outcrops, although these
local endemics are not immune from the effects of climate change. The observed
shifts associated with recent climate change are likely to have been brought about
through both direct and indirect (changes to species’ interactions) effects of
climate; indirect effects are discussed in relation to laboratory experiments and
invasive species. Recent observations of range boundary shifts are consistent with
the hypothesis that climate contributes to, but is not the sole determinant of, the
position of the range boundaries of the majority of terrestrial animal species.
*Correspondence: Chris D. Thomas,
Keywords
Department of Biology, University of York,
Heslington, York YO10 5YW, UK. Adaptation, biological invasions, climate warming, distributions, extinction,
E-mail: cdt2@york.ac.uk range margins, thermal ecology.
range expansions and range contractions, potentially threat-

INTRODUCTION
ening large numbers of species with extinction (e.g. Thomas
Understanding the factors determining the distributions and et al., 2004; McClean et al., 2005; Malcolm et al., 2006).
abundances of species has been a major focus throughout the Research on the factors that determine range boundaries has
history of ecology (e.g. Andrewartha & Birch, 1954; MacAr- often resorted to the interpretation of unintended, large-scale
thur, 1972), an interest that remains undiminished as we face ‘experiments’. For example, the success of introduced species
practical issues that relate to both shrinking (extinction) and provides numerous examples of how geographical barriers to
expanding (invasives) ranges. If climate is, inter alia, one of the dispersal can limit species’ ranges and also of how biological
important determinants of species’ range boundaries, then we interactions determine large-scale distribution patterns (native
can expect anthropogenic climate change to generate both species that are reduced or eliminated by invasives). Until
DOI: 10.1111/j.1472-4642.2010.00642.x
488 www.blackwellpublishing.com/ddi ª 2010 Blackwell Publishing Ltd
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Climate and range boundaries
recent decades, the impact of climate on species’ contemporary

Climate Other constraints
distributions has been much harder to assess because the temperature, rainfall geology, barrier
climate had been thought of as moderately stable since 1800,
the period for which most distributional data are available.
Palaeoecological studies have shown that changing climates
affect species’ distributions (e.g. Pitelka et al., 1997; Hewitt,
1999), as have studies of biological responses to the Little Ice
Age (Maunder Minimum) between 1645 and 1715 (Grove,
Other species &
1988). But fossil data and pre-1800 documented records are habitat structure
inevitably sparse and rarely of sufficient resolution to identify
the precise locations of range boundaries. Given the nature of
the data, most such data document the past distribution
responses of relatively common and widespread species.
Anthropogenic climate change provides the ideal opportunity
to test the hypothesis that climate is an important determi- Boundary
nants of species’ range boundaries more generally. location
The death of an individual near the edge of a species’ range
might be caused directly by the climate; by extreme cold, heat Figure 1 Schematic diagram of the effects of climate on the
or drought. However, most deaths in most species appear to be location of the range boundary of a species. Climate may affect the
caused by natural enemies or through a failure to compete boundary directly (e.g. through physiological and life history
successfully for resources, rather than by climate per se (e.g. Sih responses; left-hand arrow), as may other constraints (right-hand
arrow). Climate and other constraints also combine to determine
et al., 1985; Cornell & Hawkins, 1995). Similarly, birth rates
which other species are present at a given location, and their
are affected by other species as well as by the physical
abundances, as well as to determine the structure (e.g. vegetation
environment. The abundance and diversity of natural enemies, height and complexity) of habitat; these indirect species/habitat
of competitors, of mutualists and of other species that effects act together to affect the location of the range boundary.
constitute the resources used by an organism may nevertheless ‘Other species’ effects include positive (mutualists, resources)
be affected by climate. In addition, the ability of an individual and negative (competitors, enemies) impacts. Note that human
to avoid or resist natural enemies or compete with other activities are combined with those of other species.
species can be affected by climate. Therefore, climate may
affect range boundaries indirectly through changes to species
interactions and through climate-driven changes to the but which are too distant to have been colonized naturally (e.g.
physical structure of habitats, as well as having direct impacts Williamson, 1996; Peterson, 2003; Svenning & Skov, 2007;
on birth and death rates (Fig. 1). Duncan et al., 2009). A combination of population/species
This article uses recent climate change to assess the history and inadequate time to colonize represent important
frequency with which climate contributes to the position of constraints on species’ distributions at a global scale. This
range boundaries, analysing the frequencies with which species article concentrates, instead, on evaluating whether climate
have responded to climate change by shifting their distribu- plays a role in affecting the edges of existing ranges (i.e. within
tions in the ‘expected’ direction. In addition, it discusses some relatively easy colonization range).
alternative means of assessing the impact of climate on the
positions of range boundaries.
CLIMATE-DISTRIBUTION CORRELATIONS
Aside from the palaeo-record, which is beyond the scope of
TERRESTRIAL RANGE LIMITS
this article, deductions that climate is an important determi-
Most species are highly localized, with a small minority being nant of species’ current distribution margins have mainly been
geographically widespread (Gaston, 1996, 2003). Thus, most based on indirect information. The most frequent approach is
species have at least some range boundaries that are not to match (correlate) the existing distribution of species to the
coastlines; continental centres of endemism contain many such spatial distribution of climatic variation; variously termed
species. Why do all of these species not spread from their distribution, climate envelope or niche modelling (Elith &
existing boundaries into adjacent land? Some kind of limit Leathwick, 2009). Correlations between climate variables and
(climatic or otherwise) must logically exist or have existed until distributions are widely observed.
quite recently, at their existing boundaries that prevents them One interesting approach is to compare the match between
from spreading. climate variables and the distributions of real species with the
The fact that at least part of the existing/recent range match of climate variables with the distributions of fictional
boundary of most terrestrial species is/was limited by climate is ‘null’ species that have the same spatial attributes (range size
compatible with the idea that it may be possible to find other, and level of distributional aggregation) as the real species.
disjunct parts of the world that are suitable for these species, Taking this approach, Beale et al. (2008) conclude that 68% of
Diversity and Distributions, 16, 488–495, ª 2010 Blackwell Publishing Ltd 489
C. D. Thomas
bird species have distributions that are no more closely support for climate as an important determinant of range
associated with spatial variation in the climate than expected at boundaries.
random; alternatively, 32% are statistically associated with
spatial variation in the climate. However, 32% may be an
OBSERVED SHIFTS IN RANGE BOUNDARIES
underestimate (Araújo et al., 2009) for several reasons,
including: (1) if the spatial characteristics (range size and Recent climate change represents a large-scale experiment to
aggregation) of real species are genuinely generated by spatial test directly whether species’ boundaries are shifting in the
patterns of climatic variation, then ‘null’ species are not truly ‘expected’ direction. Here, I examine the frequency with which
‘null’ but can be thought of as potential climatic niches that the positions of range boundaries have moved along thermal
species could have, were there to be an infinite number of gradients. The reason for concentrating on thermal gradients is
species on Earth; and (2) it is not clear how the null because it is possible to make clear predictions for the direction
distribution analysis performs when some but not all range of range shifts, whereas shifts along moisture gradients are less
boundaries of a species are set by climate and when many of easy to interpret. I only consider studies in which all species
the species are not restricted (endemic) to the region analysed within a specified group/region are considered (given specified
(Peterson et al., 2009). This is a useful approach, but it is data quality inclusion criteria). The reason for concentrating
ultimately difficult or impossible to deduce causation from on multi-species studies is because single-species studies are
correlation. prone to publication bias and could therefore bias estimates of
A potentially stronger approach is to evaluate whether the the proportion of species showing distributional responses to
distributions of species in their native ranges can be used to climate change.
predict the ranges of the same species in parts of the world to Parmesan & Yohe (2003) reviewed then-available multi-
which they have been introduced (and vice versa). This species studies, the majority from the temperate zone. Of
approach has had mixed success, identifying that both climate studies classified by these authors as regional/continental (as
and other factors (species’ traits, human activities, propagule opposed to local, which I do not consider), and where multiple
pressure, time since arrival) contribute to species’ distributions (‡10 spp. in each study) terrestrial species were examined, 106
within their introduced ranges (e.g. Beerling et al., 1995; range boundaries were classified as shifting in the direction
Peterson, 2003; Roura-Pascual et al., 2004; Thuiller et al., 2006; expected on the basis of climate warming (shifting northwards
Richardson & Thuiller, 2007; Duncan et al., 2009). These in the northern hemisphere), 84 as stable, and 36 as shifting in
studies provide strong support to the view that climate the opposite direction. Significantly, more range shifts were in
contributes to the success and distribution of a species, the direction expected from climate change than expected by
following introduction, but (1) time since arrival is a predictor chance (Parmesan & Yohe, 2003). These numbers provide an
of introduced range size (Wilson et al., 2007; Williamson et al., estimate of 47% of range boundaries shifting in the direction
2009), which implies that the range boundaries of many species expected. This may exaggerate the percentage responding
have not yet have come to ‘equilibrium’ in the introduced because some of the 106 boundaries shifting polewards might
range, and (2) introduced species interact with new continental have done so for other reasons (e.g. land use changes, changes
biotas after translocation, such that their realized climatic in persecution), just as the 36 boundaries that moved towards
niches may differ between their native and introduced ranges the equator are likely to have been responding to other
(Broennimann et al., 2007). For both of these reasons, this pressures. Simplistically, we might conclude that an excess of
approach is likely to provide a minimum estimate of the role of 31% of boundaries of these 226 species’ boundaries shifted
climate in determining stable or native range boundaries. towards higher latitudes; i.e. 100*(106)36)/226. Conversely,
Another approach is to evaluate whether climate envelopes most of the studies included by Parmesan & Yohe (2003) were
can be used to predict changes to the distributions of species relatively early within the recent phase of anthropogenic
that have already been observed (e.g. Walther et al., 2005; warming, so they might under-estimate the climate sensitivity
Berger et al., 2007). Araújo et al. (2005) analysed bird distri- of range boundaries; the 84 ‘stable’ boundaries may hide small
bution changes in Britain and found rather inconsistent changes in one direction or another. If we ignore the ‘stable’
matches between predicted and observed changes, depending species, 75% of species shifted in the direction expected, an
on the species and modelling approach used. Green et al. excess of 49%.
(2008) concluded that bird population changes did tend to The other compilation of regional range shifts was provided
track predicted changes in climatic suitability at a European by Hickling et al. (2006), who considered data for the northern
scale, although this study did not specifically examine changes (high latitude, or poleward) range boundaries of many
to the locations of range boundaries. Because realized rates of different taxonomic groups in Britain (millipedes; woodlice;
response (which determine the statistical match between harvestmen; spiders; aquatic bugs; butterflies; carabid, long-
projected and observed changes) depend not only on climate horn & soldier beetles; dragonflies & damselflies; grasshopper
but also on the traits of the species and the landscapes through relatives; lacewings; fish; herptiles; birds; mammals). These are
which the distributions are shifting (e.g. Warren et al., 2001; arguably the most reliable data in the world available to assess
Willis et al., 2009; Wilson et al., 2009), it is not clear whether such changes. Of 329 species meeting data quality criteria, 275
these results should be regarded as providing strong or weak (84%) boundaries expanded northwards, two remain exactly
490 Diversity and Distributions, 16, 488–495, ª 2010 Blackwell Publishing Ltd
0.35 capacity to spread into new regions (Warren et al., 2001). In

contrast, the most rapidly retreating species are likely to be
0.3
those that experience a combination of deteriorating climate
Proportion of species
0.25 and other pressures, and declining species that are rarely
recorded may tend to be excluded on the basis of low data
0.2
quality. Therefore, it is conceivable that data selection results in
0.15 an over-estimate of the fraction of range boundaries showing
expansion at potentially leading range edges and an under-
0.1 estimate of retreat rates at trailing edges. However, it is also
0.05 possible that many additional (rarer) species are responding at
more local scales, such as expanding into new habitats (Davies
0 et al., 2006), which have not yet been detected in geographical-
00
00
00
00
0
scale analyses.
10
20
30
40
–3
–4
–2
–1
Range margin change (km) Another complication is whether land use changes or other
environmental drivers might themselves show latitudinal
Figure 2 Shifts at the northern range boundaries of southerly gradients of intensity and hence mimic the effects of climate
distributed animal species in Britain. Northwards shifts are posi- change. For most of the British taxa considered by Hickling
tive values, and southwards shifts are negative, with distances et al. (2006), this seems unlikely, given that most of the
moved over approximately 25 years. Data are for 329 species from
southern species are spreading northwards across human-
the following taxa: millipedes; woodlice; harvestmen; spiders;
dominated landscapes; land use appears to be responsible for
aquatic bugs; butterflies; carabid, longhorn & soldier beetles;
dragonflies & damselflies; grasshopper relatives; lacewings; fish; limiting the level of response, in this case, rather than
herptiles; birds; mammals. Data and further details from Hickling accelerating it.
et al. (2006). Comparable analyses of the distribution boundaries of
individual plant species require further development; Harsch
et al. (2009) report advances of the tree line since 1900 at 52%
the same and 52 retreated southwards (Fig. 2). Again, some of sites, whereas retreat was observed at only 1% of sites, an
northwards shifts are likely to be related to non-climatic excess of 51%.
factors, so the excess northwards shift is approximately 68%; The direction of response is relevant simply to ask whether
i.e. 100*(275)52)/329. The data in Hickling et al. (2006) are climate contributes to the location of range boundaries. On the
only up to 2000, so these estimates may already be exceeded. above evidence, I conclude that over half, and perhaps around
Evaluating whether low latitude range boundaries are two-thirds, of observed animal range boundaries have already
retreating with equal frequency is harder to assess, mainly shown a response to 1970–2000 anthropogenic warming.
because the data are inadequate (Thomas et al., 2006).
Parmesan & Yohe (2003) found that expansions at leading
ARE TROPICAL SPECIES SIMILARLY LIMITED?
edges was potentially more frequent than contractions at
trailing edges. However, expansions are typified by increases in The above studies were from the temperate zone, and there is a
abundance and expansion as a ‘front’, which is reasonably easy dearth of studies of geographical-scale range boundary shifts
to detect, whereas retreats more usually proceed by increasing for tropical species. More local studies provide evidence of
fragmentation of populations (Wilson et al., 2004; Hampe & upwards shifts on tropical mountains, in Costa Rica, Mada-
Petit, 2005), making it harder to identify when the last local gascar and Borneo (Pounds et al., 1999; Raxworthy et al., 2008;
population in a region has disappeared, and whether climate Chen et al., 2009), but they are insufficient to draw strong
change is a possible cause. Thomas et al. (2006) estimated that conclusions about the frequencies with which species have
17 out of 21 (81%, an excess of 62%) butterfly species for shown geographical-scale boundary shifts. Combining data for
which there had been detailed surveys of their trailing edges Costa Rican birds (Pounds et al., 1999) and Madagascar
had shown elevational or latitudinal retreats – comparable to herptiles (Raxworthy et al., 2008), 91 (75%) upper boundaries
the expansion figures for leading edges. moved upwards, 18 moved downwards and 12 remained
It is possible that the species that have been assessed could stable, an excess of 60% shifting upwards. For the lower
represent a biased sample, even in multi-species studies. boundaries of montane species, however, no such pattern was
Species for which there are sufficient records for comparison apparent; 48 (38%) lower boundaries shifted upwards, 40
across time may disproportionately be relatively common downwards and 38 remained stable, an excess of only 8%
species (i.e. they meet ‘data quality’ criteria for inclusion more moving upwards. However, both studies failed to detect some
frequently than do rarer species because they have many high elevation species in the more recent surveys (and hence
records in two or more time periods). Habitat generalists that they were excluded from comparisons of boundaries between
are relatively common and widespread within their distribu- time periods), and if these species are extinct, then the true
tions will normally (1) meet data quality criteria, and (2) not proportion retreating upwards is higher than the above figures
find barriers to dispersal, and hence they will have the greatest suggest. I-C Chen et al. (unpublished) found quite comparable
C. D. Thomas
upwards shifts of the lower and upper boundaries when (10–25C and 15–30C). Experiments considered three Dro-
considering all geometrid moth species on Mt. Kinabalu, in sophila species on their own, and in two- and three-species
Borneo, although this overall result hid differences at different mixtures, with and without a shared parasitoid (natural
elevations on the mountain (greater expansion upwards than enemy), and with and without dispersal (by opening and
retreat at some elevations, but the opposite elsewhere). blocking the connecting tubes). The Drosophila species and the
Over large parts of the tropics, moisture availability is parasitoid affected the thermal ranges and abundances of each
probably a more important determinant of range boundaries other in these experiments, showing very clearly that distribu-
than temperature. The tree Aloe, Aloe dichotoma, appears to be tions arise from a combination of biotic and physical
showing population decreases in the driest parts of its (temperature) factors in these simplified environments. A
geographical range, an observation that is consistent with major message that has been taken up in the literature citing
climate change (Foden et al., 2007). However, multi-species this work (over 300 and 100 citations of Davis et al., 1998a,b,
geographical studies of distribution responses to drying and respectively; Web of Science, December 2009) is that responses
wetting trends are not yet available. Part of the difficulty is that to climate change will be unpredictable because of complex
predictions are far from straight-forward; would species be species interactions.
expected to expand into drier areas because of increased water The conclusion that multi-species boundaries are unpre-
use efficiency (atmospheric CO2 enrichment enables plants to dictable seems premature. In Davis et al.’s results, the ‘winner’
keep stomata closed more of the time), or retract because of of pairwise competition experiments between Drosophila
increased desiccation arising from higher temperatures or species was entirely predictable at each temperature (but the
reduced precipitation? winner was different at different temperatures). The ‘optimum’
Whilst further data would be desirable, these preliminary temperature of each species was also largely unaffected by
analyses suggest that the responses of the range boundaries of which other species were in the experiment (they always lined
tropical montane species are not obviously different from those up with D. subobscura being most abundant in the coolest
observed at larger geographical scales in the temperate zone; cages, D. simulans in intermediate cages and D. melanogaster in
the majority of upper boundaries have already shifted upwards the hottest). And, the standard errors of the mean abundances
in response to climate warming. Responses of lower bound- (in chambers of a given temperature) in multi-species clines
aries on tropical mountains, and especially distributional were just as small as in single-species replicates; densities in
responses to changing moisture gradients, are poorly docu- multi-species experiments were just as predictable as in single-
mented. The importance of climate to species range bound- species chambers. In other words, the outcomes of species
aries’ in the tropics receives some further support from the interactions were highly predictable throughout the sequence
observation that centres of endemism tend to be located in of experiments, for a given temperature, and given a particular
climatically unusual regions and in regions with steep climatic set of species, etc. These effects might be indirect (Fig. 1), but
gradients (Ohlemüller et al., 2008). temperature strongly determined the observed patterns. When
the experimenters applied climate change to the thermal clines
(cline of 15–30C vs. 10–25C), the differences in abundance
THE CLIMATE–SPECIES INTERACTION
were largely predictable on the basis of the previous patterns.
The main indirect means by which climate affects the locations My own interpretation is that thermally driven changes to
of species’ range boundaries is likely to be through its impact multi-species interactions will in most cases not produce major
on the interactions between species (Fig. 1) (MacArthur, surprises, although this will happen sometimes; most of the
1972). Evidence that ‘other species’ are major determinants surprises will arise when entirely different species or functional
of distribution boundaries comes from studies of invasive groups, usually from other continents (invasive species), join
species (e.g. van Riper et al., 1986; Channell & Lomolino, the set of interacting species.
2000a,b; Short & Turner, 2000) and from competition/ Invasive species often show variation in their propensity to
predation experiments (Davis et al., 1998a,b; Pople et al., impact negatively on native species along environmental
2000). From Gause onwards, researchers have commonly gradients; threatened species commonly survive (last) at the
observed that it is difficult to maintain two species in culture margins of their former distributions (e.g. Clout & Craig, 1995;
together in a single controlled environment but that the Channell & Lomolino, 2000a,b; Short & Turner, 2000). For all
outcome (which species ‘wins’) of the interaction can be species that eventually co-exist with an invader in a subset of
changed by altering the environment. If this holds in general their previous distributions, the locations of new boundaries
with respect to climatic environments, then climate is expected are likely to be set by factors that limit the occurrence or
to be a critical determinant of the locations of species virulence of the invader or increase the capacity of the native
transitions and so indirectly affects range boundaries. species to resist it; such as climate, geology or dispersal failure
Two microcosm articles by Andrew Davis and colleagues (e.g. failure to reach offshore islands). The pervasive impacts of
(Davis et al., 1998a,b) are particularly relevant. Davis et al. some non-native species on others do not in any way disprove
kept local Drosophila populations (in different incubators) at the hypothesis that climate contributes to the locations of
different temperatures, and linked the populations by existing range boundaries; it reminds us, rather, that climatic
connecting tubes, forming a thermal cline spanning 15C limits must be seen in the context of multi-species interactions.
If climate change drives malaria-resistant introduced birds, 2008). When the climate changes, almost all species are likely
mosquitoes and bird malaria to higher elevations in the to be affected indirectly through the responses of species that
Hawaiian Islands, we might expect them to make further are affected directly (Fig. 1). Such species are potentially
inroads into the susceptible (now largely montane) native bird susceptible to changes in ecosystem productivity and species
fauna (van Riper et al., 1986; Freed et al., 2005). interactions wrought by climate change (O’Reilly et al., 2003).
In conclusion, laboratory experiments and observations of
invasive species commonly show that the outcome of interac-
CONCLUSION
tions between species depends on the environment. In as much
as that ‘climate’ is part of the environment, this implies that Climate change has now provided a preliminary answer to the
climate does contribute to range boundaries. When the climate old question of whether climate is an important determinant of
shifts, so do those range boundaries that are set by the the recent range boundaries of species. Over half of species’
outcomes of interspecific interactions. boundaries that have been examined have already responded to
the quite modest level of global warming already experienced
between 1970 and 2000. It seems likely that climate contributes
NON-CLIMATIC LIMITS
to (but is not the sole determinant of) the locations of
Some species have range boundaries set by factors that are distribution boundaries for the majority of terrestrial species in
completely unrelated to climate. A land species that is endemic continental regions; even higher fractions of range boundaries
to and occurs throughout a particular oceanic island or an are likely to respond to further warming. The contribution of
aquatic species that occurs throughout the waters of a single climate to range boundaries is in many cases likely to be
lake are obvious examples. There are also many other island- indirect, through alterations to species’ interactions (Fig. 1).
like environments, particularly geologies (e.g. Serpentine Because responses to climate change are already so common,
outcrops), which may constrain range boundaries irrespective virtually every species is already experiencing changes to these
of the climate (this does not include cold mountain tops where interspecific interactions, and hence feeling at least the indirect
the island-like nature of the environment is determined largely impacts of climate change.
by climate). Narrowly distributed endemics associated with
these localized environments may exhibit realized climatic
ACKNOWLEDGEMENTS
niches that are a small fraction of their potential climatic niche,
such that their distributions are not immediately affected by I thank I-Ching Chen, Rachael Hickling, Jane Hill, Richard Fox
climate change (see below). Even so, quite a high proportion of and David Roy, all of the amateur natural history recorders
local endemics still show range limits within these areas, such responsible for documenting range changes, and NERC for
as species that are restricted by elevation on an island, by depth financial support. Janet Franklin, Dave Richardson, Mark
in a lake, or by aspect or soil moisture on an unusual geological Schwartz and an anonymous referee provided helpful com-
outcrop (e.g. van Riper et al., 1986; Daniel & Fox, 1999; ments on the manuscript.
Mackay et al., 2006). These species are expected to shift their
local distribution boundaries higher, deeper, or to a shadier
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BIOSKETCH
Sih, A., Crowley, P., McPeek, M., Petranka, J. & Strohmeier, K.
(1985) Predation, competition, and prey communities: a Chris Thomas is Professor of Conservation Biology at the
review of field experiments. Annual Review of Ecology & University of York. His research concentrates on the impacts of
Systematics, 16, 269–311. climate change and habitat fragmentation on species’ distri-
Svenning, J.C. & Skov, F. (2007) Ice age legacies in the geo- butions. He is also interested in the implications of these
graphical distribution of tree species richness in Europe. impacts for conservation.
Global Ecology and Biogeography, 16, 234–245.
Thomas, C.D., Cameron, A., Green, R.E., Bakkenes, M.,
Beaumont, L.J., Collingham, Y.C., Erasmus, B.F.N., de Editor: David Richardson

Diversity and Distributions - 2010 - Thomas - Climate Climate Change and Range Boundaries

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Diversity and Distributions, (Diversity Distrib.

) (2010) 16, 488–495

Department of Biology, University of York, ABSTRACT

freshwater range boundaries. This generalization excludes species that are

range expansions and range contractions, potentially threat-

recent decades, the impact of climate on species’ contemporary

0.35 capacity to spread into new regions (Warren et al., 2001). In

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