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The distributions of many terrestrial organisms are currently shifting in latitude or elevation in response to changing climate. Using a meta-analysis, we estimated that the distributions of species have recently shifted to higher elevations at a median rate of 11.0 meters per decade, and to higher latitudes at a median rate of 16.9 kilometers per decade. These rates are approximately two and three times faster than previously reported. The distances moved by species are greatest in studies showing the highest levels of warming, with average latitudinal shifts being generally sufficient to track temperature changes. However, individual species vary greatly in their rates of change, suggesting that the range shift of each species depends on multiple internal species traits and external drivers of change. Rapid average shifts derive from a wide diversity of responses by individual species.
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Acknowledgments: This work was supported by the Howard
Hughes Medical Institute (C.B.L., S.R.S., D.M.K., D.H.),
the NSF (CAREER-0644282 to M.K., DBI-0644111 to
A.S.), the NIH (R01-HG004037 to M.K., P50- HG02568
to D.M.K., U54-HG003067 to K.L-T., 1U01-HG004695
to C.B.L., 5P41-HG002371to B.J.R.), the Sloan
Foundation (M.K.), and the European Science Foundation
(EURYI to K.L-T.).
Supporting Online Material
www.sciencemag.org/cgi/content/full/333/6045/1019/DC1
Materials and Methods
Figs. S1 to S9
Tables S1 to S12
References (2949)
10 January 2011; accepted 24 June 2011
10.1126/science.1202702
Rapid Range Shifts of Species
Associated with High Levels
of Climate Warming
I-Ching Chen,
1,2
Jane K. Hill,
1
Ralf Ohlemüller,
3
David B. Roy,
4
Chris D. Thomas
1
*
The distributions of many terrestrial organisms are currently shifting in latitude or elevation in response
to changing climate. Using a meta-analysis, we estimated that the distributions of species have
recently shifted to higher elevations at a median rate of 11.0 meters per decade, and to higher latitudes
at a median rate of 16.9 kilometers per decade. These rates are approximately two and three times
faster than previously reported. The distances moved by species are greatest in studies showing the
highest levels of warming, with average latitudinal shifts being generally sufficient to track temperature
changes. However, individual species vary greatly in their rates of change, suggesting that the
range shift of each species depends on multiple internal species traits and external drivers of change.
Rapid average shifts derive from a wide diversity of responses by individual species.
Threats to global biodiversity from climate
change (1-8) make it important to identify
the rates at which species have already
responded to recent warming. There is strong evi-
dence that species have changed the timing of
their life cycles during the year and that this is
linked to annual and longer-term variations in
temperature (912). Many species have also
shifted their geographic distributions toward
higher latitudes and elevations (1317), but this
evidence has previously fallen short of demon-
strating a direct link between temperature change
and range shifts; that is, greater range shifts have
not been demonstrated for regions with the high-
est levels of warming.
We undertook a meta-analysis of available
studies of latitudinal (Europe, North America,
and Chile) and elevational (Europe, North Amer-
ica, Malaysia, and Marion Island) range shifts for
a range of taxonomic groups (18)(tableS1).We
considered N= 23 taxonomic group × geographic
region combinations for latitude, incorporating
764 individual species responses, and N=31
taxonomic group × region combinations for ele-
vation, representing 1367 species responses. For
the purpose of analysis, the mean shift across all
species of a given taxonomic group, in a given
region, was taken to represent a single value (for
example, plants in Switzerland or birds in New
York State; table S1) (18).
The latitudinal analysis revealed that spe-
cies have moved away from the Equator at a
median rate of 16.9 km decade
1
(mean = 17.6
km decade
1
, SE = 2.9, N= 22 species group ×
region combinations, one-sample ttest versus
zero shift, t= 6.10, P< 0.0001). Weighting each
study by the (numberofspecies)inthegrou
region combination gave a mean rate of 16.6 km
decade
1
. For elevation, there was a median shift
to higher elevations of 11.0 m uphill decade
1
(mean = 12.2 m decade
1
, SE = 1.8, N=30spe-
cies groups × regions, one-sample ttest versus
zero shift, t= 7.04, P< 0.0001). Weighting ele-
vation studies by (number of species) gave a
mean rate of uphill movement of 11.1 m decade
1
.
A previous meta-analysis (14)ofdistribu-
tion changes analyzed individual species, rather
than the averages of taxonomic groups × regions
that we used, and also included data on latitu-
dinal and elevational shifts in the same analysis
(18). It concluded that ranges had shifted toward
higher latitudes at 6.1 km decade
1
andtohigh-
er elevations at 6.1 m decade
1
(14), whereas
the rates of range shift that we found were sig-
nificantly greater [N= 22 species groups × regions,
one-sample ttest versus 6.1 km decade
1
,t=
3.99, P= 0.0007 for latitude; N= 30 groups ×
regions, one-sample ttest versus 6.1 m decade
1
,
t= 3.49, P= 0.002 for elevation (18)]. Our
estimated mean rates are approximately three
and two times higher than those in (14), for
1
Department of Biology, University of York, Wentworth Way,
York YO10 5DD, UK.
2
Biodiversity Research Center, Academia
Sinica, 128 Academia Road, Section 2, Nankang Taipei 115,
Taiwan.
3
School of Biological and Biomedical Sciences, and
Institute of Hazard, Risk and Resilience, Durham University,
South Road, Durham DH1 3LE, UK.
4
Centre for Ecology &
Hydrology, Crowmarsh Gifford, Wallingford, Oxfordshire,
OX10 8BB, UK.
*To whom correspondence should be addressed. E-mail:
chris.thomas@york.ac.uk
Fig. 1. Relationship between observed and expected range shifts in response to climate change, for (A)
latitude and (B) elevation. Points represent the mean responses (TSE)ofspeciesinaparticulartax-
onomic group, in a given region. Positive values indicate shifts toward the pole and to higher ele-
vations. Diagonals represent 1:1 lines, where expected and observed responses are equal. Open circles,
birds; open triangles, mammals; solid circles, arthropods; solid inverted triangles, plants; solid square,
herptiles; solid diamond, fish; solid triangle, mollusks.
19 AUGUST 2011 VOL 333 SCIENCE www.sciencemag.org
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latitude and elevation respectively, implying much
greater responses of species to climate warming
than previously reported (18). Most of the data
we analyzed are from the temperate zone and
from tropical mountains (table S1), where eco-
systems are at least partly temperature-limited;
different rates of change might be observed in
moisture-limited ecosystems (19).
Published studies have shown nonrandom
latitudinal and elevational changes (1, 7, 1317)
but have not previously demonstrated a statis-
tical linkage between range shifts and levels
of warming. We found that observed latitudinal
and elevational shifts (the latter more weakly)
have been significantly greater in studies with
higher levels of warming (mean latitudinal shift
versus average temperature increase; N=23spe-
cies groups × regions, Pearson correlation coef-
ficient (r) = 0.59, P= 0.003; mean elevational
shift versus temperature increase; N= 31, r=
0.37, P= 0.042). Temperature gradients differ
across the world, so a given level of warming
leads to different expected range shifts of spe-
cies in different regions (20), assuming that spe-
cies track climate changes. To estimate the
expected shifts, we calculated the distances in
latitude (kilometers) and elevation (meters) that
species in a given region would have been re-
quired to move to track temperature changes
and thus to experience the same average tem-
perature at the end of the recording period as
encountered at the start (18) (table S1). We
found that both observed latitudinal and ele-
vation range shifts were correlated with predicted
distances (Fig. 1A, N= 20 species groups ×
regions, r= 0.65, P= 0.002 for latitude; Fig.
1B, N= 30 groups × regions, r= 0.39, P=
0.035 for elevation), so our analyses directly
link terrestrial range shifts to regional and study
differences in the warming experienced.
Despite reports that many species lag behind
climate change (2123), nearly as many studies
of observed latitudinal changes fall above as
below the observed = expected line in Fig. 1A
(9 points above, 11 below; c
2
= 0.20, 1 df, P=
0.65), suggesting that mean latitudinal shifts are
not consistently lagging behind the climate. The
lag in elevation response (Fig. 1B; 2 points above
the 1:1 line, 28 below; c
2
= 22.53, 1 df, P<
0.001) is equally surprising because the required
distances to track climate are much shorter than
for latitudinal shifts (20). Real and apparent ele-
vation lags may arise if suitable new conditions
at higher elevations occur only in locations that
cannot be reached easily (for example, on other
mountain peaks), or they may reflect the topo-
graphic and microclimatic complexity of moun-
tainous terrain [for example, cooler locations
may be on poleward-facing slopes rather than
higher (24)]; the need for finer-resolution analy-
ses (25); and additional topographic, climatic, ge-
ological, and ecological constraints [for example,
causing declines in cloud forest species (2628)].
Taxonomic differences are not consistent pre-
dictors of recent response rates. For example,
birds seem to have responded least in terms of
elevational shifts but had a slightly greater than
expected latitudinal shift (Fig. 1). Much greater
variation is associated with differences among
species within a taxonomic group than between
taxonomic groups (Fig. 2 and table S2). For lat-
itudinal studies, on average 22% (average of
N= 23 species groups × regions) of the species
actually shifted in the opposite direction to that
expected. Similarly, 25% of species shifted down-
hill rather than to higher elevations (average of
N= 29 species groups × regions). Thus, despite
an overall significant shift toward higher lati-
tudes and elevations, which is greatest where
the climate has warmed the most, and despite
around three-quarters of species shifting pole-
ward and to higher elevations, we found that
species have exhibited a high diversity of range
shifts in recent decades.
At least three processes are likely to generate
the high diversity of range shifts among species:
time delays in speciesresponses, individualistic
physiological constraints, and alternative and in-
teracting drivers of change. Species may lag be-
hind climate change if they are habitat specialists
or immobile species that cannot colonize across
fragmented landscapes (17,2123), or if they
possess other traits associated with low extinc-
tion or colonization rates (29). Species may also
show individualistic physiological responses to
different aspects of the climate, such as different
sensitivities to maximum and minimum temper-
atures at critical times of their life cycles. These
sensitivities will combine with variable wait times
for different novel climatic extremes to take
place (30). Species are also affected to dif-
ferent extents by nonclimatic factors and by
multispecies interactions, which themselves de-
pend on a diversity of environmental drivers
(21,28). For example, a species might retreat
toward the Equator at its poleward margin if it
contracts with habitat loss faster than it expands
through climate warming; whereas the poleward
range margin of a species that thrives in novel ag-
ricultural landscapes may spread at a rate exceed-
ing that expected, were warming the sole driver.
We found that rates of latitudinal and eleva-
tional shifts are substantially greater than reported
Fig. 2. Observed latitudinal shifts of the northern range boundaries of species within four exemplar
taxonomic groups, studied over 25 years in Britain. (A) Spiders (85 species), (B)groundbeetles
(59 species), (C) butterflies (29 species), and (D) grasshoppers and allies (22 species). Positive
latitudinal shifts indicate movement toward the north (pole); negative values indicate shifts toward the
south (Equator). The solid line shows zero shift, the short-dashed line indicates the median observed
shift, and the long-dashed line indicates the predicted range shift.
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in a previous meta-analysis, and increase with
the level of warming. We conclude that average
rates of latitudinal distribution change match
those expected on the basis of average temper-
ature change, but that variation is so great within
taxonomic groups that more detailed physio-
logical, ecological and environmental data are
required to provide specific prognoses for indi-
vidual species.
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Acknowledgments: We thank A. Bergamini, R. Hickling,
R. Wilson, and B. Zuckerberg for data; H.-J. Shiu for
statistical assistance; S.-F. Shen, the Ministry of Education
in Taiwan, a UK Overseas Research Scholarship Award,
and the Natural Environment Research Council for
support; and anonymous referees for comments on the
manuscript. We are particularly grateful to the many
thousands of volunteers responsible for collecting most of
the original records of species. All data sources are listed
in the supporting online material.
Supporting Online Material
www.sciencemag.org/cgi/content/full/333/6045/1024/DC1
Materials and Methods
Tables S1 and S2
References (3151)
1 April 2011; accepted 6 July 2011
10.1126/science.1206432
Aneuploidy Drives Genomic
Instability in Yeast
Jason M. Sheltzer,
1
Heidi M. Blank,
1
Sarah J. Pfau,
1
Yoshie Tange,
2
Benson M. George,
1
Timothy J. Humpton,
1
Ilana L. Brito,
3
Yasushi Hiraoka,
2,4
Osami Niwa,
5
Angelika Amon
1
*
Aneuploidy decreases cellular fitness, yet it is also associated with cancer, a disease of enhanced
proliferative capacity. To investigate one mechanism by which aneuploidy could contribute to
tumorigenesis, we examined the effects of aneuploidy on genomic stability. We analyzed 13 budding
yeast strains that carry extra copies of single chromosomes and found that all aneuploid strains
exhibited one or more forms of genomic instability. Most strains displayed increased chromosome loss
and mitotic recombination, as well as defective DNA damage repair. Aneuploid fission yeast strains
also exhibited defects in mitotic recombination. Aneuploidy-induced genomic instability could facilitate
the development of genetic alterations that drive malignant growth in cancer.
Whole-chromosome aneuploidyor a
karyotype that is not a multiple of the
haploid complementis found in great-
er than 90% of human tumors and may contrib-
ute to cancer development (1,2). It has been
suggested that aneuploidy increases genomic
instability, which could accelerate the acquisition
of growth-promoting genetic alterations (1,3).
However, whereas aneuploidy is a result of ge-
nomic instability, there is at present limited evi-
dence as to whether genomic instability can be a
consequence of aneuploidy itself. To test this
possibility directly, we assayed chromosome seg-
regation fidelity in 13 haploid strains of Saccha-
romyces cerevisiae that carry additional copies
of single yeast chromosomes (4). These aneu-
ploid strains (henceforth disomes) display im-
paired proliferation and sensitivity to conditions
that interfere with protein homeostasis (4,5).
We measured the segregation fidelity of a yeast
artificial chromosome (YAC) containing human
DNA and found that the rate of chromosome
missegregation was increased in 9 out of 13 di-
somic strains relative to a euploid control (Fig.
1A). The increase ranged from 1.7-fold to 3.3-
fold, comparable to the fold increase observed
in strains lacking the kinetochore components
Chl4 or Mcm21. Consistent with chromosome
segregation defects, 8 out of 13 disomic strains
displayed impaired proliferation on plates con-
taining the microtubule poison benomyl, includ-
ing a majority of the strains that had increased
rates of YAC loss (Fig. 1B).
Chromosome missegregation can result from
defects in chromosome attachment to the mitotic
spindle or from problems in DNA replication or
repair. Defects in any of these processes delay
mitosis by stabilizing the anaphase inhibitor
Pds1 (securin) (6). Five out of five disomes (di-
somes V, VIII, XI, XV, and XVI) exhibited de-
layed degradation of Pds1 relative to wild type
after release from a pheromone-induced G
1
arrest
(Fig. 1C and fig. S1). Defective chromosome bi-
orientation delays anaphase through the mitotic
checkpoint component Mad2 (6). Deletion of
MAD2 had no effect on Pds1 persistence in four
disomes, but eliminated this persistence in disome
V cells (fig. S1). Disome V also delayed Pds1 deg-
radation after release from a mitotic arrest in-
duced by the microtubule poison nocodazole,
which demonstrated that this strain exhibits a bi-
orientation defect. Disome XVI, which displayed
Mad2-independent stabilization of Pds1, recov-
ered from nocodazole with wild-type kinetics (fig.
S2). Thus, Pds1 persistence results predominant-
ly from Mad2-independent defects in genome
replication and/or repair (see below).
We next investigated whether aneuploidy
could affect the rate of forward mutation. Di-
somes V, VIII, X, and XIV displayed an in-
creased mutation rate at two independent loci,
whereas disome IV displayed an increased
mutation rate at CAN1 but not at URA3 (Fig.
2A). The fold increase ranged from 2.2-fold to
7.1-fold, less than the 9.5-fold and 12-fold in-
creases observed in a recombination-deficient
rad51Dmutant and a mismatch repairdeficient
msh2Dmutant, respectively. Additionally, in an as-
say for microsatellite instability, we found that di-
somes VIII and XVI displayed increased instability
in a poly(GT) tract (fig. S3), which demonstrated
that aneuploidy can enhance both simple se-
quence instability and forward mutagenesis.
To define the mechanism underlying the
increased mutation rate in aneuploid cells, we
1
David H. Koch Institute for Integrative Cancer Research and
Howard Hughes Medical Institute (HHMI), Massachusetts In-
stitute of Technology, Cambridge, MA 02139, USA.
2
Graduate
School of Frontier Biosciences, Osaka University 1-3 Yamadaoka,
Suita 565-0871, Japan.
3
Department of Ecology, Evolution and
Environmental Biology, Columbia University, New York, NY
10027, USA.
4
Kobe Advanced ICT Research Center, National Insti-
tute of Information and Communications Technology 588-2 Iwaoka,
Iwaoka-cho, Nishi-ku, Kobe 651-2492, Japan.
5
The Rockefeller
University, 1230 York Avenue, New York, NY 10065, USA.
*To whom correspondence should be addressed. E-mail:
angelika@mit.edu
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... This ideal temperature range is one of the main drivers limiting the biogeographic range suitable to animals (Du Plessis et al. 2012). Climate change causing warmer temperatures globally alters the distribution of ideal temperature ranges, forcing species to shift their distribution and phenology accordingly (Parmesan 2006;Chen et al. 2011). Long-term upslope range shifts driven by temperature increases have been recorded in many montane species but, in numerous cases, range shifts are lagging behind climate change (Chen et al. 2011). ...
... Climate change causing warmer temperatures globally alters the distribution of ideal temperature ranges, forcing species to shift their distribution and phenology accordingly (Parmesan 2006;Chen et al. 2011). Long-term upslope range shifts driven by temperature increases have been recorded in many montane species but, in numerous cases, range shifts are lagging behind climate change (Chen et al. 2011). Upslope range shift among montane species is not a viable long-term tactic to avoid thermal stress in a warming world. ...
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Anthropogenic climate change is having measurable effects on the phenology and distribution of organisms, with many species moving polewards or to higher elevations. Temperature-driven elevational range shifts can vary between species and therefore interactions within multitrophic systems are expected to alter along elevational gradients, however there is limited empirical evidence to test this hypothesis. The tritrophic interactions between the seed-feeding moth Coleophora alticolella , its larval ectoparasitoids and its host plant, the rush Juncus squarrosus , were surveyed along the same elevational gradient in northern England in 1977 and 1978, and again in 2019 and 2021. Spatio-temporal changes in these trophic levels over the intervening forty-year period were more complex than a simple synchronous upslope shift of the whole system. Seed production by J. squarrosus greatly increased at equivalent elevations since 1977/1978. Floret production by inflorescences was significantly higher than expected from recent temperature changes. Seed capsule ripening was also greater at higher elevations in the warmer summers of 2019 and 2021. The elevational limit of C. alticolella distribution and the elevation of peak larval densities both rose more than the change in the isotherm recorded for this area since 1977. The rate of larval parasitisation was greater at equivalent sites in 2019 and 2021 than previously and the structure of the ectoparasitoid community on C. alticolella larvae had changed. The ectoparasitoids also occurred at higher elevations but their elevational gains were less than for their host. Implications for insect conservation: Understanding the impact of climate-related changes on insects is currently of major importance for insect conservation. In addition to assessing the effect of these changes on an individual species, this study shows that measuring longer-term changes within a multitrophic system across a spatial dimension can reveal additional insights for conservation and management strategies.
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The changes in range margins of birds breeding in Finland was analysed from published atlas data for a 12-year period (1974-1979 to 1986-1989). The change in range margin was statistically corrected for changes in species' distribution using linear regression. For species predominantly occurring in southern Finland (n = 116), the expected range margin shift, if their distribution would not have changed, was 18.8 km northwards. Northerly species (n = 34) showed no such significant range margin shift. A similar result was found earlier for UK birds. Recent range margin shifts in birds therefore seem to be a general phenomenon, which may be related to climate change.
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Changes in geographic distributions of 5 bird species endemic to the Great Plains of North America were examined over the last few decades based on the United States Breeding Bird Survey. Examining the mean latitude of individuals of each species, 3 species showed significant or near-significant northward shifts, and 1 a significant shift southward. Over all 5 species examined, colonization events were concentrated in the northern part of the distributions of the species; in 3 species, extinctions were concentrated in the southern part of the distributions of the species. The conclusion is that significant distributional changes have taken place, but they have been subtle, and might be associated with global climate change.
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While poleward species migration in response to recent climatic warming is widely documented, few studies have examined entire range responses of broadly distributed sessile organisms, including changes on both the trailing (equatorward) and the leading (poleward) range edges. From a detailed population census throughout the entire geographical range of Aloe dichotoma Masson, a long-lived Namib Desert tree, together with data from repeat photographs, we present strong evidence that a developing range shift in this species is a ‘fingerprint’ of anthropogenic climate change. This is explained at a high level of statistical significance by population level impacts of observed regional warming and resulting water balance constraints. Generalized linear models suggest that greater mortalities and population declines in equatorward populations are virtually certainly the result, due to anthropogenic climate change, of the progressive exceedance of critical climate thresholds that are relatively closer to the species’ tolerance limits in equatorward sites. Equatorward population declines are also broadly consistent with bioclimatically modelled projections under anticipated anthropogenic climate change but, as yet, there is no evidence of poleward range expansion into the area predicted to become suitable in future, despite good evidence for positive population growth trends in poleward populations. This study is among the first to show a marked lag between trailing edge population extinction and leading edge range expansion in a species experiencing anthropogenic climate change impacts, a pattern likely to apply to most sessile and poorly dispersed organisms. This provides support for conservative assumptions of species’ migration rates when modelling climate change impacts for such species. Aloe dichotoma's response to climate change suggests that desert ecosystems may be more sensitive to climate change than previously suspected.
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The conservation status of 845 zooxanthellate reef-building coral species was assessed by using International Union for Conservation of Nature Red List Criteria. Of the 704 species that could be assigned conservation status, 32.8% are in categories with elevated risk of extinction. Declines in abundance are associated with bleaching and diseases driven by elevated sea surface temperatures, with extinction risk further exacerbated by local-scale anthropogenic disturbances. The proportion of corals threatened with extinction has increased dramatically in recent decades and exceeds that of most terrestrial groups. The Caribbean has the largest proportion of corals in high extinction risk categories, whereas the Coral Triangle (western Pacific) has the highest proportion of species in all categories of elevated extinction risk. Our results emphasize the widespread plight of coral reefs and the urgent need to enact conservation measures.
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It is predicted that climate change will cause species extinctions and distributional shifts in coming decades, but data to validate these predictions are relatively scarce. Here, we compare recent and historical surveys for 48 Mexican lizard species at 200 sites. Since 1975, 12% of local populations have gone extinct. We verified physiological models of extinction risk with observed local extinctions and extended projections worldwide. Since 1975, we estimate that 4% of local populations have gone extinct worldwide, but by 2080 local extinctions are projected to reach 39% worldwide, and species extinctions may reach 20%. Global extinction projections were validated with local extinctions observed from 1975 to 2009 for regional biotas on four other continents, suggesting that lizards have already crossed a threshold for extinctions caused by climate change.
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Significant changes in physical and biological systems are occurring on all continents and in most oceans, with a concentration of available data in Europe and North America. Most of these changes are in the direction expected with warming temperature. Here we show that these changes in natural systems since at least 1970 are occurring in regions of observed temperature increases, and that these temperature increases at continental scales cannot be explained by natural climate variations alone. Given the conclusions from the Intergovernmental Panel on Climate Change (IPCC) Fourth Assessment Report that most of the observed increase in global average temperatures since the mid-twentieth century is very likely to be due to the observed increase in anthropogenic greenhouse gas concentrations, and furthermore that it is likely that there has been significant anthropogenic warming over the past 50 years averaged over each continent except Antarctica, we conclude that anthropogenic climate change is having a significant impact on physical and biological systems globally and in some continents.
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Mean global temperatures have risen this century, and further warming is predicted to continue for the next 50-100 years. Some migratory species can respond rapidly to yearly climate variation by altering the timing or destination of migration, but most wildlife is sedentary and so is incapable of such a rapid response. For these species, responses to the warming trend should be slower, reflected in poleward shifts of the range. Such changes in distribution would occur at the level of the population, stemming not from changes in the pattern of indivduals' movements, but from changes in the ratios of extinctions to colonizations at the northern and southern boundaries of the range. A northward range shift therefore occurs when there is net extinction at teh southern boundary or net colonization at the northern boundary. However, previous evidence has been limited to a single species or to only a portion of the species' range. Here we provide the first large-scale evidence of poleward shifts in entire species' ranges. In a sample of 35 non-migratory European butterflies, 63% have ranges that have shifted to the north by 35-240 km during this century, and only 3% have shifted to the south.
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Aim To estimate whether species have shifted at equal rates at their leading edges (cool boundaries) and trailing edges (warm boundaries) in response to climate change. We provide the first such evidence for tropical insects, here examining elevation shifts for the upper and lower boundaries shifts of montane moths. Threats to species on tropical mountains are considered. Location Mount Kinabalu, Sabah, Malaysia. Methods We surveyed Lepidoptera (Geometridae) on Mount Kinabalu in 2007, 42 years after the previous surveys in 1965. Changes in species upper and lower boundaries, elevational extents and range areas were assessed. We randomly subsampled the data to ensure comparable datasets between years. Estimated shifts were compared for endemic versus more widespread species, and for species that reached their range limits at different elevations. Results Species that reached their upper limits at 2500–2700 m (n= 28 species, 20% of those considered) retreated at both their lower and upper boundaries, and hence showed substantial average range contractions (−300 m in elevational extent and −45 km2 in estimated range area). These declines may be associated with changes in cloud cover and the presence of ecological barriers (geological and vegetation transitions) which impede uphill movement. Other than this group, most species (n= 109, 80% of the species considered) expanded their upper boundaries upwards (by an average of 152 m) more than they retreated at their lower boundaries (77 m). Main conclusions Without constraints, leading margins shifted uphill faster than trailing margins retreated, such that many species increased their elevational extents. However, this did not result in increases in range area because the area of land available declines with increasing elevation. Species close to a major ecological/geological transition zone on the mountain flank declined in their range areas. Extinction risk may increase long before species reach the summit, even when undisturbed habitats are available.
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Recent warming has caused changes in species distribution and abundance, but the extent of the effects is unclear. Here we investigate whether such changes in highland forests at Monteverde, Costa Rica, are related to the increase in air temperatures that followed a step-like warming of tropical oceans in 1976 (refs4, 5). Twenty of 50 species of anurans (frogs and toads) in a 30-km2 study area, including the locally endemic golden toad (Bufo periglenes), disappeared following synchronous population crashes in 1987 (refs 6-8). Our results indicate that these crashes probably belong to a constellation of demographic changes that have altered communities of birds, reptiles and amphibians in the area and are linked to recent warming. The changes are all associated with patterns of dry-season mist frequency, which is negatively correlated with sea surface temperatures in the equatorial Pacific and has declined dramatically since the mid-1970s. The biological and climatic patterns suggest that atmospheric warming has raised the average altitude at the base of the orographic cloud bank, as predicted by the lifting-cloud-base hypothesis,.
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Aim To investigate altitudinal range shifts of bryophytes in Switzerland by comparing recent altitudinal distributions with historical distributions derived from herbarium specimens. Location Switzerland, covering 41,285 km2 in Central Europe. Methods We used a dataset of 8520 herbarium specimens of 61 bryophyte species and compared altitudinal data between the two periods 1880–1920 and 1980–2005. The records we used were not specifically sampled for climatological analyses, but originate from non-systematic fieldwork by various collectors. Historical and recent records were distributed all over Switzerland with occurrences in all major biogeographical areas. To account for different sampling efforts in the two time periods, different subsampling procedures were applied. Results Overall, we found a significant mean increase in altitude of 89 ± 29 m which was mainly driven by the cryophilous species (+222 ± 50 m). The mean increase in altitude of cryophilous species corresponds to a decadal upward shift of 24 m. The upper range limit of cryophilous species also increased by 189 ± 55 m, but there was no effect on the lower range limit. For intermediate and thermophilous species neither mean, nor upper or lower range limits changed. However, the proportion of records of thermophilous to cryophilous species increased considerably at lower altitudes, but levelled off above approximately 1800 m. Main conclusions We conclude that cryophilous bryophytes are expanding their range to higher elevations in Switzerland and that at lower elevations, a slow extinction process is going on, probably as a result of climate warming trends. The observed decadal upward shifts of cryophilous species closely match those reported from vascular plants in Europe and those expected, given recent estimates of climate warming trends. We emphasize that herbaria provide valuable data that can be used to detect ongoing changes in the distribution of species.