the nature of climate change - the rspb wildlife charity...
TRANSCRIPT
The nature of climate changeEurope’s wildlife at risk
Contents
Foreword from the RSPB Chief Executive Mike Clarke.................................. 3
Sections: 1) The impacts of climate change on wildlife .................................................. 4
2) Shifting ranges........................................................................................... 6
3) New arrivals ............................................................................................. 10
4) Changing populations and communities .................................................. 13
5) Changing interactions between species ................................................... 17
6) Extreme weather ..................................................................................... 21
7) The future ................................................................................................ 24
8) Protected areas and climate change ....................................................... 28
9) Managing sites and landscapes for adaptation ........................................ 31
10) Creating new sites and expanding site networks ................................... 35
11) Helping people to adapt ......................................................................... 38
12) Implications for conservation ................................................................. 42
13) References ............................................................................................ 43
The RSPB is a registered charity in England & Wales 207076, in Scotland SC037654. 272-0092-15-16
3
Foreword
We are at a point in recent geological history where the rate of human-induced
climate change will far outstrip the ability of species to adapt successfully,
especially when the resilience of nature has been reduced by habitat loss,
non-native species introductions and over-exploitation. The disruption to the web
of life is a threat not just to wildlife, but to the lives of people around the world.
This report presents the evidence that wildlife in the UK and beyond is already facing a more challenging
time due to the climate change that has occurred; and that things are, for the most part, only likely
to get worse. Higher rainfall will likely have a detrimental effect on bearded tits, capercaillie and shags for
example, and warmer temperatures in southern Europe will result in habitat loss for Dartford warblers.
Moreover, the evidence points to global scale patterns of change, such as the collapse in kittiwake
populations linked to sea surface temperatures and the timing of plankton blooms. Of course it’s not all
bad news, and we’ve already seen exciting new species colonise the UK and begin breeding here, some
of them on our very own RSPB reserves. Birds like little egrets, black-winged stilts and little bitterns are
spectacular and enticing new arrivals.
But we have to face up to the fact that in general the changes to our climate will be challenging for wildlife.
One of the most important conclusions of this report is that protected areas and nature reserves will be
vital in the future for helping wildlife to cope with climate change. This makes it even more important
that we ensure EU decision makers leave the Birds and Habitats Directives intact and that they are
better implemented.
Protecting wildlife from these threats means each country, in Europe and in the UK, playing an active role
in mitigating carbon emissions. We need a clear policy direction to keep rolling out renewable energy that
doesn’t harm nature, and to invest more in ways that benefit the climate and help nature and people to
adapt, such as by restoring our upland peatlands.
Nature is demonstrating that we’re already into the world of change. This report has a clear message – we
need to act, and fast, to reduce our greenhouse gas pollution and to face up to the impacts we’re already
living with. Then we can make better homes for nature, and ourselves.
Dr Mike Clarke
Chief Executive
The RSPB
4
1) The impacts of climate change on wildlife
Background and purpose of the report
Human actions are causing the climate to change extremely rapidly. Since the Industrial Revolution, global
temperatures have risen, driven by rising greenhouse gas concentrations in the atmosphere1. Comparison
with historical records suggests that temperatures are rising at an unprecedented rate2. It is very likely that
temperatures will continue to rise throughout the 21st century: we are committed to further climate change3, 4
.
Climate change is already changing the natural world. Plants and animals are adapted to certain climatic
conditions that they require to survive and thrive. As the climate changes, becoming more or less favourable,
species will be impacted by, and will have to respond to, the new conditions. There is abundant evidence of
this already occurring. This report aims to document some of the impacts of climate change on wildlife that
we’ve already seen, changes we might expect in the future, and some of the possible conservation
responses that could benefit both wildlife and people.
The focus of this report is on areas where the RSPB has taken a closer role in understanding,
limiting or managing the impacts on wildlife. This may have been via on-the-ground conservation
action, leading scientific research, or collaboration with other researchers.
The scale of climate change means we must think beyond national boundaries, so this report considers
wildlife throughout Europe. Because entire ecological communities are experiencing the effects of climate
change, it considers species from plankton and plants, through to insects and other invertebrates, up to
birds and mammals.
Structure of the report
The report is structured in themed sections. In each section, examples are presented from the wider
scientific literature to illustrate the impacts already occurring and those expected in the future.
Each section also provides more detailed case studies focusing on the science and conservation
work to which the RSPB has contributed. Together, these examples show how climate change is driving
fundamental changes to wildlife, and how conservation action can continue to help wildlife to adapt.
Within individual plants and animals, genetic, physical and functional changes have been detected;
within populations of individual species, changing abundances and changes in geographic distribution
are increasingly observed; within entire ecological communities, changes in the species present and in
food web functioning have been documented5. Responses to the warming climate are complex: if one
species in an ecological community changes, there will be knock-on impacts for others, leading to
ecosystem-scale impacts6.
5
It is essential that we understand these impacts of climate change on wildlife, so that appropriate
conservation measures can be taken. As the area with suitable climate for species changes and shifts,
species may move into new areas (section 2 from page 6, and section 3 from page 10) to track
suitable climatic conditions. In a given area, abundances of some species will increase whilst others will
decline, and species new to the area will establish populations, leading to big changes in ecological
communities (section 4 from page 13)6. As interactions between species (section 5 from page 17)
are altered, reduced prey abundance, increased predation risk, or new diseases and pathogens could prove
to be substantial threats7. As the intensity and frequency of extreme weather (section 6 from page 21)
increases, wildlife populations could suffer severe mortality or reproductive failure8. Climate change therefore
poses a threat to biodiversity, and will change the biological communities with which we are familiar.
Future projections (section 7 from page 24) suggest that further, substantial changes will occur,
potentially bringing increased risk of extinction9, so nature conservation also needs a longer term outlook.
We need to consider how vulnerable, sensitive and exposed species are and will become and assess
how different conservation methods might help10
. Existing conservation methods will be vital. Protected
area networks (section 8 from page 28) are essential to help species survive and track suitable climate.
Site management (section 9 from page 31) and creation of new sites (section 10 from page 35), to
aid population resilience and accommodate changes, will also help species to adapt. Landscape-scale
approaches will help link sites, and encourage ecosystem-based adaptation (section 11 from page 38)
that benefits both people and wildlife. Finally, the report concludes with the implications for conservation
(section 12 from page 42).
6
2) Shifting ranges
Many species are already moving, predominantly northwards and to higher elevations.
This is exactly what we would expect as a response to a warming climate.
Climate is important in determining where a species can live – its range. Species are adapted to particular
climatic conditions, but as global temperatures rise, the conditions they experience are changing. Some
species may tolerate the change at a location, by rapid adaptation. However, given the unprecedented pace
of current climate change, many species will need to move to track the conditions suitable for them.
The climate shows natural gradients: broadly-speaking, it is warmer near the equator, cooler near the
poles, and cooler at higher elevations. A species has a natural range, often bounded at higher latitudes or
elevations where it becomes too cold for the species, and at lower latitudes or elevations beyond which it
becomes too warm. Even if a thermal limit is not reached, a species’ range may be bounded by other factors
such as moisture availability, habitat suitability, or availability of land. As temperatures rise, lower latitudes
and elevations of a species’ range could become less climatically suitable, while higher latitudes and
elevations (if available) could become more suitable. If species track these changes, their ranges will
shift toward the poles (ie northwards in Europe) and uphill.
One way to study this is to examine changes in the most extreme northerly points of a species’ range – the
northern range margin. As the climate warms, areas further north are being colonised, shifting the
northern range margin. Across 59 British bird species, this margin shifted north by an average of 19 km
between 1968– 72 and 1988– 9111
. Further shifts happened between 1988– 91 and 2008–11, with the
northern range margin of 77 species moving north by an average of 13.5 km12
.
Even larger shifts have been observed. Between the 1960s–70s and 1980s–90s, the northern range margins
of many British animals, including mammals, grasshoppers, beetles and spiders, shifted north by an average
of 12.5–19 km per decade13
. An updated analysis indicated that across 1,573 British species from 21 groups,
including birds, bees, dragonflies and moths, the northern range margin shifted 23.2 km per decade on
average between the 1960s–70s and 1980s–90s, and a further 18.0 km per decade between the 1980s–90s
and the 2000s14
. Similarly, another analysis found that 24 British dragonfly and damselfly species moved an
average of 88 km further north between 1960–70 and 1985–9515
. In France, the pine processionary moth
– a species that causes severe damage to pine trees – advanced 87 km between 1972 and 200416
.
Conversely, populations at the southern range margin face the risk of local extinction. For instance,
two butterfly species of northern Britain, the northern brown argus and scotch argus, showed southern range
margin retractions of 70–100 km between the 1970s and late 1990s, driven by local extinctions in southerly
sites17
. Likewise, between 1960–70 and 1985–95, southern range margins of northern British dragonflies
and damselflies shifted 44 km north on average15
. However, overall, fewer southern range margin shifts have
been observed so far. This is perhaps because complete disappearance from an area is harder to confirm or
7
because other factors, such as species interactions or habitat availability, are currently more important
determinants of population persistence at southern margins15
.
As a result of these expansions and retractions, species’ entire ranges are shifting. The most dramatic
examples of this are in the sea. Plankton species found in the Bay of Biscay and waters around the
south-western UK in the 1960s were, by the 1990s, found approximately 1,100 km further north, around
northern Scotland18
. Fifteen North Sea fish species, including commercially-fished species such as cod,
moved an average of 170 km north (ranging 48–403 km) between 1977 and 200119
. On land, between
1981 and 2000, wintering distributions of grey plover and curlew in north-west Europe shifted 115 km
and 119 km north-east, respectively20
, whilst for 122 British breeding bird species, the central point of
their distributions shifted an average 20.4 km north between 1988–91 and 2008–1112
.
There is also good evidence for changes in the elevations over which species occur. Four northern
British butterfly species moved an average of 41 m (metres) uphill between 1970 and the late 1990s,
because of local extinctions at lower sites and colonisation of higher sites21
. In the Alps, dung beetles
moved uphill 65 m on average between 1992–93 and 2007, mainly by colonising higher locations, whilst in
the Spanish Sierra Nevada mountains, they shifted 321 m uphill between 1981–82 and 2006–07, driven by
both colonising higher elevations and loss from lower elevations22
. In the Sierra de Guadarrama mountains in
Spain, the lower limit of 16 butterfly species moved uphill by an average of over 200 m between 1967–73
and 2004, but the upper limit was unchanged, so the overall range shrank23
. In the sea, however,
temperatures are higher closer to the surface: consequently, 28 bottom-dwelling fish in the North Sea
moved an average of 3.6 m per decade deeper between 1980 and 200424
.
In some cases, rapid changes in species’ behaviour or form appear to have facilitated range shifts
linked to climate change. For instance, in Britain, northern expansion of the brown argus butterfly has been
driven by greater use of a widespread geranium species for egg-laying, so the extent of suitable breeding
habitat has increased25
. Some changes are even more fundamental. Northward shifts of the garden tiger
moth26
and a species of bush cricket27
in Britain have been associated with changes to their wing shapes,
which are linked to improved dispersal capability.
Species are moving as we would expect in response to climate change, showing that, by moving to new
areas, they can adapt to changing conditions. However, whilst shifting ranges could aid species’ persistence
in the longer term, not all species are responding adequately to the changing climate. In Finland, 48
butterfly species shifted their northern range margin an average of 60 km between 1992–96 and 2000–04,
but the shifts were smaller for species that were threatened or less mobile, with these traits likely to mean
that the species’ required habitats were too rare and patchy, or too hard to reach28
. In the North Sea, larger
fish species and those that reached reproductive age later were less likely to shift ranges in response to
warming19
. In the Alps, breeding birds shifted uphill between 1992–94 and 2003–05, but the changes were
small and less than expected from observed temperature increases29
. In Sweden, of 101 breeding bird
species, around 20% showed range shifts in the direction expected, but many did not fully compensate for
the extent to which temperatures had increased30
. While some variation in ability to respond is due to
variation between the species themselves, for instance in specialism or dispersal ability, a key factor is
8
availability of suitable habitats: a species can only shift range if there is suitable habitat to move into
(see sections: on Protected areas, from page 28; Site management, from page 31; and Creating new sites,
from page 35).
Case study: Dartford warbler range change (Wotton et al., 2009; Bradbury et al., 2011)31, 32
The Dartford warbler is a bird species of dense, low scrub in Western Europe and North Africa. At the
northern extent of its global range in southern Britain, where it is a heathland specialist, the climate is only
marginally suitable. In 1963, following two very cold winters, the population may have declined to as few
as 11 pairs.
National surveys for the species from the 1970s to the 2000s showed major changes. Protection afforded by
the 1979 EU Birds Directive enabled the population in the species’ core area to increase, and new areas
were colonised. These areas included sites at higher elevations and further north, suggesting winters were
no longer so strongly limiting: higher temperatures have aided the Dartford warbler’s northward expansion.
Severe winters in the late 2000s led to some population declines33
, but the Dartford warbler has recovered
well and maintained the range extension, with further expansion likely in the coming years.
Dartford warbler distributions from UK national surveys. White spots show population of 1–4 territories,
yellow shows 5–14 territories, orange shows 15–49 territories and red shows over 50 territories. Over time,
the population has grown and spread north and west.
Adapted from Wotton et al. (2009), British Birds 102(5), 230–246.
9
However, the UK range expansion as temperatures rise tells only part of the story. In Spain, the core of its
range, the Dartford warbler is declining, likely through habitat loss34
. Models of future climate change suggest
that during the 21st century, the climate will become unsuitable for them across large parts of their southern
range35
. Therefore, whilst climate change could lead to the Dartford warbler increasing in Britain, its
global future may be less positive. This highlights the importance of coordinated, continent-scale action to
support wildlife adaptation.
10
3) New arrivals
Species are starting to successfully colonise new territories as they track suitable climate.
Conservation strategies will need to support this.
As the climate changes, populations of species are establishing in areas where they are not known to
have occurred previously. Many range shifts discussed in the previous section considered how species
moved within a territory: British birds, Finnish butterflies, North Sea fish. However, species moving tens or
hundreds of kilometres in a decade will inevitably arrive in areas where they are not “native” species: they
will have colonised a new region or country. Observing these colonisations, understanding them and
providing suitable habitat for range expansion are key parts of the conservation response to climate change.
The UK provides good opportunities to study colonisation: as a series of islands supporting well-studied
wildlife, incoming species are often well-recorded. These records have been used to examine the occurrence
of over 7,000 species from a range of groups, including plants, spiders, butterflies and moths, freshwater
fish, birds and mammals: since 1900, at least 120 species have colonised Britain36
. Over time, the number
of species colonising Britain has increased, as we would expect if the warming climate was
responsible36
, although other factors, such as global trade, are also important.
Some of the best-known recent colonists of the UK are waterbirds. Since 2008, there have been first
breeding records for cattle egrets, purple herons and great white egrets, the commencement of
regular breeding by little bitterns, and breeding attempts by black-winged stilts37
. These species
all breed in western continental Europe, where several of them have shown increasing populations over
recent years; combined with predictions that the British climate will become increasingly suitable, this could
make colonisation of Britain more likely37
. Another waterbird, the little egret, used to be only an occasional
springtime visitor to Britain but from the late 1980s, the frequency and duration of visits increased, leading
to confirmed breeding by 199638
. It is now well-established. Little egrets are sensitive to periods of cold
weather, so climate change appears to be linked to the species’ establishment38
.
Not only birds are moving: there are several examples of colonists from other groups. The small
red-eyed damselfly has spread north through Europe since the 1980s, and was found on the east coast
of England in 1999, thereafter spreading further; this colonisation is believed to be linked to the warming
climate39
. The wasp spider, which was first observed on the south coast of England in 1922, has spread
throughout south-east England36
; its northward spread in Europe may be associated both with a warming
climate and recent adaptations that allow the species to tolerate slightly cooler climates40
. Climate change
might also have played a role in Nathusius’ pipistrelle bats starting to breed in the UK in the 1990s41
.
Successful colonists appear, typically, to be more mobile species36
: if species are more mobile, they will be
better able to track the changing climate. However, this reinforces two issues. First, even mobile species
need sufficient, suitable habitat in the new territories. For example, waterbird colonisations of the UK
would be facilitated by wetland protection and the re-creation of large wetlands37
.
11
Second, less mobile species may not be able to naturally colonise new territories. It has therefore
been argued that conservationists could help these species to establish populations in newly suitable areas
– so-called “assisted colonisation”42, 43
. This has been shown to be feasible for marbled white and small
skipper butterflies within the UK44
, and has been suggested for the critically endangered Iberian lynx within
Spain45
. However, moving species between countries could be associated with substantial and unpredictable
impacts, so other conservation measures may provide safer options46
. The potential risks and benefits of
assisted colonisation, and its potential applications, are therefore important parts of the debate around
conservation under climate change.
The wildlife we typically accept as being part of our “native” flora and fauna is moving, and new species
are arriving as colonists, partly driven by climate change. The assemblage of species we consider “native”
is therefore in a state of flux. We cannot arrest the changes, so to aid adaptation it will be important to
enable species to colonise new areas via provision of sufficient, suitably-protected habitat, in areas
that will become more climatically suitable over time (see sections on Protected areas, from page 28;
and Creating new sites, from page 35).
Case study: breeding birds colonise the UK via protected areas (Hiley et al., 2013)47
Between 1960 and 2013, 20 waterbird species bred in the UK for the first time. Of these, six species
established regular breeding populations: little egret, common crane, whooper swan, Cetti’s warbler,
goldeneye and Mediterranean gull. Several of these colonisations appear to be associated with the
warming climate.
To understand how these species colonised the UK, the location of breeding records was studied for the
six colonists that now breed regularly. For all six, the first breeding records were in protected areas – British
Sites of Special Scientific Interest (SSSIs) and Northern Irish Areas of Special Scientific Interest (ASSIs).
For all except goldeneye, subsequent breeding records increasingly occurred outside of protected areas as
populations grew: the species spread into the wider landscape. Therefore, protected areas appear to have
aided successful natural colonisations of waterbirds in the UK.
12
The proportion of colonists’ breeding pairs (little egrets) or singing males (Cetti’s warblers) in protected areas
(dots), and total population size (lines). For both species, the proportion of breeding records in protected
areas decreased as the population grew: the species used protected areas as a platform for colonisation,
before spreading into the wider landscape.
Adapted from Hiley et al. (2013), Proceedings of the Royal Society B 280 (1760), 20122310.
0
500
1000
1500
2000
2500
0
10
20
30
40
50
60
70
80
90
100
1970 1980 1990 2000 2010
Nu
mb
er
of
bre
ed
ing
pa
irs
% s
ing
ing
ma
les
in
pro
tec
ted
are
as
Year
Cetti's warbler
0
100
200
300
400
500
600
700
800
0
10
20
30
40
50
60
70
80
90
100
1995 2000 2005 2010N
um
be
r o
f b
ree
din
g p
air
s
% b
ree
din
g i
n p
rote
cte
d a
rea
s
Year
Little egret
13
4) Changing populations and communities
Changes to the species present in an area, and the abundances of these species, show that
climate change is already affecting ecological communities.
As the climate warms, the species present in a given area, and their respective abundances, are changing.
New species are arriving, whilst others are disappearing; some populations are increasing, whilst others are
declining. These changes do not occur uniformly across species: mobile species, those that can use a wider
range of habitats, and those adapted to warmer conditions can respond better than others.
Together, this means that ecological communities – the combination of species present in a given area
at a given time – are changing. This is important, as community composition influences how an ecosystem
functions. However, community composition data also provide useful information on climate change impacts.
The population monitoring of different groups can give an impression of the overall impacts of climate
change, whilst indicator statistics can be calculated to show the changing balance of warm-adapted and
cold-adapted species.
Large-scale surveys and long-term monitoring programmes give widespread evidence of community
changes in line with those expected under climate change. Data from the UK’s Environmental Change
Network (ECN) show that at northern and upland sites, between 1993 and 2007, butterflies and moths
typically found in the south increased in abundance, whilst ground beetles typically found in the north
declined48
. In the plant communities of Scottish alpine habitats – those found above the tree line on
mountains – specialist alpine and northern species declined in abundance between 1963–87 and 2004–06,
whilst generalist lowland species increased49
. Whilst some of these changes to Scottish alpine plants could
be linked to nitrogen pollution, many align with expectations of climate change impacts49
.
British breeding birds show similar patterns. Between 1994 and 2006, the abundance and distribution of
generalist species increased as temperatures rose, whilst those of specialist species declined, creating more
homogeneous communities50
. Even communities of relatively immobile groups have shown changes: in the
Netherlands, between 1979 and 2001, southerly or warm-adapted lichen species increased in abundance
whilst northerly or cold-adapted species decreased51
.
Changes have also been observed in marine communities. The plankton community of the North Sea
changed substantially in the 1980s, linked with a strong increase in water temperature: abundances of
cool-adapted plankton species declined, abundances of warm-adapted plankton species increased, and
overall species diversity increased52
. The plankton community changes (and associated changes to fish
populations) were so large that the period of change has been termed a “regime shift”52
.
Information about species composition and abundances can be used to calculate metrics, known as
“indicators”, that tell us more about how climate change is affecting communities. The Community
Temperature Index (CTI) combines data on species composition, population abundance, and the average
14
temperature with which species are associated53
. The CTI has been calculated for breeding bird
communities in France53
and Sweden54
, plant, butterfly and bird communities in Switzerland55
, and for
the breeding bird and butterfly communities across the whole of Europe56
. In all cases, the CTI increased
between the late 20th century and the 2000s, showing that community composition shifted towards
warm-adapted species. This could be caused by abundances of warm-adapted species increasing,
abundances of cold-adapted species decreasing, new warm-adapted species arriving, or cold-adapted
species becoming locally extinct. However, in most cases the temperature shifted more than community
composition: communities are lagging behind climate change53, 54, 56
.
Case study: a climate change indictor for birds in Europe (Gregory et al, 2009)57
The Climatic Impacts Indicator (CII) indicates the overall impact of climate change on the European bird
community, by examining the changing abundances of those species predicted to benefit from climate
change, and those predicted to be adversely affected. To construct the indicator, Gregory et al modelled the
impact of future climate change on European bird species’ distributions. Species predicted to increase their
range (30 species, including bee-eater, hoopoe and Cetti’s warbler) were grouped together, as were species
predicted to experience range decreases (92 species, including lapwing, wood warbler and willow tit).
They then examined how the different groups actually fared over recent years, using data from large-scale
bird surveys across Europe. These indices reflect observed population trends and the climatic sensitivity
of the species: higher values mean the species were doing better; lower values mean the species were
doing worse.
Birds predicted to do well under climate change, often species associated with warmer conditions,
declined between 1980 and 1990, but since then have shown a big increase. Species predicted to do
poorly under climate change, often those associated with cooler conditions, declined from 1980 onwards.
Overall, this shows that European bird populations appear to be responding to climate change,
with warm-adapted species increasing in abundance and cool-adapted species decreasing.
15
Weighted population trends of European birds expected to benefit from climate change (red line) and those
expected to be adversely affected by climatic change (blue line). Species population trends were weighted
by modelled climatic sensitivity. Until 1990, both groups show declining populations. After 1990, the trends
diverge: species predicted to benefit show strong population increases, whilst species predicted to suffer
continue to decline.
Figure produced following the methods of Gregory et al. (2009), PLoS ONE 4(3), e4678:
RSPB/EBCC/University of Durham/University of Cambridge.
60
65
70
75
80
85
90
95
100
105
1980 1985 1990 1995 2000 2005 2010
CL
IM g
rou
p i
nd
ex
Year
16
The final CII was constructed by comparing the two population trends. Simply, the more the trends diverge,
the more the CII rises, giving a measure of the total impact of climate change on European bird populations.
Between 1980 and 1990, there was a slight decrease in the CII, coinciding with a period of colder winters
and with land-use changes. However, from 1990 onwards, the CII showed a strong increase, coinciding with
strong temperature rises: climate change is already having an impact on bird communities across
Europe, and the impacts are becoming stronger.
The final CII indicator, showing the overall impact of climate change on European bird populations. After an
initial slight decrease through the 1980s, the CII increases strongly through the 1990s and 2000s: the impact
of climate change on European bird populations is increasing.
Figure produced following the methods of Gregory et al. (2009), PLoS ONE 4(3), e4678:
RSPB/EBCC/University of Durham/University of Cambridge.
60
70
80
90
100
110
120
130
140
1980 1985 1990 1995 2000 2005 2010
Clim
ati
c Im
pac
t In
de
x
Year
Decreasing climatic impact on bird populations
Increasing climatic impact on bird populations
17
5) Changing interactions between species
Some of the strongest impacts of climate change on wildlife come from changes in
interactions between species, in particular the relationships between predators and prey.
An ecological community comprises a number of different species interacting with each other: predators
and prey, herbivores and food plants, parasites and hosts. As shown in the previous section, climate change
is altering the make-up of ecological communities. This can dramatically affect the nature and dynamics of
interactions between the species within communities. Indeed, a recent review suggested that changed
interactions among species could be the single most important mechanism behind climate change
impacts on wildlife, particularly for predatory species7, and is potentially the biggest driver of local
extinctions58
. Understanding changing species interactions could therefore be vital for understanding
climate change threats to wildlife.
One important way in which the dynamics of ecological communities is changing is via shifts in
the timing, or phenology, of seasonal events. This is particularly notable in spring, when temperature
dependent events, such as commencing breeding, eggs hatching or tree buds opening, are occurring
earlier. Across 726 UK species, covering marine, freshwater and terrestrial environments, spring and
summer events occurred 3.9 days earlier per decade on average between 1976 – 200559
. However, not
all species responded in the same way: primary producers (plants and plankton) and primary consumers
(herbivores) responded faster than secondary consumers (predators). This “phenological mismatch”
could disrupt interactions between species within the food web, resulting in food being available
at the wrong time59
.
Some of the best examples of phenological mismatch come from woodland food chains in the Netherlands.
Between 1988 and 2005, oak leaves emerged 1.7 days earlier per decade, caterpillar abundance peaked
7.5 days earlier per decade, egg hatching of great tits, blue tits, coal tits and pied flycatchers became
3.6–5.0 days earlier, while egg hatching of sparrowhawks did not advance60
. Hence, over the course of the
study, the matching between different levels of the food web became poorer. Pied flycatcher populations
declined most strongly in those areas where the mismatch between the caterpillar peak and timing of
breeding was greatest, suggesting that earlier caterpillar emergence was detrimental to the birds61
.
For great tits, the number of chicks fledged and fledgling weight was influenced by how well egg laying
matched the caterpillar peak62
.
Phenological mismatch is also a risk in marine environments. In the North Sea, different groups, from
plankton, through fish, up to seabirds, are shifting timings at different rates63
. When North Sea plankton
groups were studied between 1958 and 2002, diatoms, a group of algae, did not bloom any earlier, but
plankton species higher up the food chain, including fish larvae, peaked substantially earlier, possibly
reducing efficiency of energy transfer through the food chain64
.
18
However, phenological mismatch may not have severe consequences in all habitats and for all
species. For the marbled white butterfly in the UK, impacts of timing mismatches with its main nectar
source, greater knapweed, might be minimised by using plants in different microhabitats – such as warmer
south-facing slopes and cooler north-facing slopes – that produce variety in the plant’s flowering time65
.
In Dutch long-distance migratory birds, although forest-dwelling species declined strongly between 1984
and 2005, marsh-dwelling species did not; this is possibly because forests are highly seasonal systems
with short food peaks, raising the risk of phenological mismatch, whereas marshes are less seasonal66
.
In the British uplands, phenological mismatch between golden plover egg laying and emergence of their
cranefly prey appears to be less important than cranefly abundance (itself influenced by climate) in
influencing plover populations67
.
Evidence of impacts on predator-prey relationships from a range of systems suggests that changing prey
availability or quality, or abundances of predators, could have substantial impacts on wildlife. In the
Czech Republic, higher temperatures have allowed populations of the edible dormouse to increase, leading
to increased nest predation on great tits and collared flycatchers68
. In Norway, willow grouse chick production
is lower in years following hot Augusts: after hot summers, bilberry plants provide lower-quality food for the
grouse, and moths that are important prey for grouse chicks are less abundant69
. In Welsh upland streams,
high temperatures in two years in the early 1990s were associated with very low abundances of freshwater
invertebrates such as caddisfly larvae, in turn leading to local extinction of one of their predators, a flatworm
species associated with cooler habitats70
.
As seems to be the case with phenological change, the risk is magnified up the food chain. In German
grasslands, more than twenty years of data on plants and insects showed that community composition of
producers (the plants) was less sensitive to climate change than was that of herbivores (some grasshoppers,
true bugs), which in turn was less sensitive than that of carnivores (beetles, spiders)71
.
Case study: food web disruption and North Sea kittiwakes (Carroll et al, 2015)72
The UK black-legged kittiwake population has declined by around 70% since 1986. Both breeding success
(the number of chicks fledged) and adult survival rates are falling. Work on the Isle of May population in
Scotland indicates that breeding success and survival could be affected by two key things73
. First, North Sea
kittiwakes rely heavily on sandeels for food during the breeding season, so sandeel fisheries could remove
vital prey. Second, when the sea was warmer, survival and breeding success were lower. So, human activity
and changing ocean conditions – driven by a warming climate – could be acting together to threaten
kittiwakes. However, it is likely that the impacts of rising temperatures are not direct, instead reflecting shifts
in the food web.
Rising temperatures have been associated with a dramatic change in the North Sea plankton community52
,
as cool-adapted plankton species are replaced by those adapted to warmer waters, but which seem less
able to support the sandeels that feed on them74
. However, temperature is not the only important
environmental factor. In the spring, changes to seawater temperature and salinity lead to sharp density
differences between deeper and shallower waters – this is called stratification. Earlier stratification could
19
cause a mismatch between plankton availability, sandeel development, and kittiwake food requirements75
,
while stronger stratification can favour less nutritious plankton species and make the seabed less suitable
for sandeels76, 77
. Driven by rising temperatures and changes to stratification conditions, changed
species interactions could combine to reduce food availability for kittiwakes78, 79
.
Recently, large-scale tracking studies using lightweight GPS tags have yielded unprecedented amounts of
information about where kittiwakes feed during the breeding season. Tracking data from 11 colonies (from
northern Scotland, to eastern England and Wales) were used to model the effects of changing sea
conditions in feeding areas on kittiwake breeding success.
Foraging areas of kittiwakes (coloured patches), identified by attaching lightweight GPS tags to a number of
birds at eleven colonies (white diamonds) during the breeding season. The foraging area of each colony is
identified with a different colour. Data on ocean conditions were extracted for these areas, to see how ocean
conditions in feeding areas affected breeding success.
Figure reproduced courtesy of Matthew Carroll.
20
Results indicated that higher sea temperatures led to lower kittiwake breeding success, supporting previous
findings that warming seas are a threat. Further to this, earlier, stronger ocean stratification did indeed
reduce breeding success, confirming suggestions that climate change could affect kittiwakes via more than
temperatures alone. Based on these relationships, projections for the late 21st century suggested that the
breeding success of the eleven colonies could fall by 21–43%.
Modelled kittiwake breeding success for the eleven colonies for a 20th century “baseline” and the late
21st century. Driven by warming seas and stronger, earlier stratification, the model suggested that
kittiwake breeding success could fall 21–43%.
Adapted from Carroll et al. (2015), Climate Research 66, 75–89.
21
6) Extreme weather
Increases in the frequency and intensity of extreme weather events can be expected to have
an increasing impact on wildlife populations.
Wildlife already deals with a variable climate, but since 1950 there have been increases in the length and
frequency of heatwaves, the frequency and intensity of heavy rainfall events, and the frequency and
intensity of droughts in some areas1. Since 2000, numerous weather events have broken records, with,
for instance, the hottest European summer in 500 years occurring in 2003, thus appearing to show that the
climate is becoming more extreme80
. IPCC (The Intergovernmental Panel on Climate Change) projections
show that, in the future, the frequency and intensity of such events will increase3, 4
, potentially
representing a major source of impacts on wildlife.
From the perspective of wildlife impacts, extreme weather means events of large magnitude and low
frequency relative to the lifespan of the animals or plants being considered8. As these are infrequent
events, to see how wildlife populations could react to increasingly extreme weather, we can examine
how they already respond to severe weather events and climatic variability.
One of the most obvious impacts of severe weather on wildlife is high mortality. Because their
plumage is not fully waterproof, shags in Scotland survive less well in years with strong onshore winds and
high rainfall in February, sometimes leading to mass mortality events known as “wrecks”81
. Projections show
that more variable weather could lead to increased risk of extinction for shag populations81
. Temperature
extremes can also affect survival. Swiss barn owls monitored between 1934 and 2002 had lower survival
rates in winters when snow-cover lasted longer82
. Conversely, lambs of mouflon (wild sheep) in southern
France survived much less well during the severe summer drought of 2003 (caused by the hottest summer
in over 500 years80
), probably because of food shortages83
.
Extreme weather can also affect reproductive success. Slavonian grebes in Scotland had higher
breeding success when temperatures during chick rearing were higher, this relationship being driven
primarily by poor chick production if there were very cold spells during breeding84
. Slavonian grebe
populations were also sensitive to rainfall, with heavy rain during the breeding season leading to smaller
populations84
. In the Netherlands, oystercatcher chick fledging success was always low in years with high
summer flood risk85
. This could be increasingly problematic, as floods increased in frequency and intensity
from 1971–2008 and may increase further as sea levels rise85
. Plants also suffer impacts of severe weather
on reproduction: in the Apennines in Italy, foxtail grass and a vetch species produced fewer flowering stems
and fewer flowers per stem in the 2003 summer heat wave86
.
The ability of animals and plants to cope with extreme weather impacts varies within and between
species, and can be influenced by the way humans manage the landscape. In the 2003 heat wave,
French bird species with a wider temperature tolerance (the difference between the hottest and coldest
temperatures found in the species’ European range) were most resilient to the extremely high
22
temperatures87
. Following a severe drought in the UK in 1995, ringlet butterfly populations crashed; those in
drier areas experienced the biggest crashes, whilst those in areas with more woodland, and less fragmented
woodland, declined less and recovered faster88
. Similarly, following a severe drought in autumn 2002,
common frogs in Finnish farmland showed much reduced egg production and strong population declines,
but populations were more resilient in landscapes with more varied habitats89
. Finally, in an extreme drought
in Sweden in 1992, bush cricket populations were more resilient in tall rather than short grassland, likely due
to increased humidity in taller vegetation90
. Further, the crickets moved into forests during the drought; this
habitat would usually be avoided, but was presumably favoured during the drought due to increased
humidity90
. These examples show that it is important to consider how sensitive species are to extreme
weather, and how habitats and landscapes can be managed to improve population resilience (see
section on “Managing sites”, from page 31).
Case study: impacts of extreme rain and floods on UK birds
Capercaillie breeding success and rainfall (Summers et al, 2004; Summers et al, 2010)91, 92
The capercaillie is a large woodland grouse that is an iconic species of Scottish native pine forests. However,
its population has declined rapidly since the 1970s, linked to low breeding success and collision mortality in
adult birds. The capercaillie population in Abernethy Forest was studied from 1989–99 to examine why the
species suffers from low breeding success. Over this period, in years with heavy rainfall, productivity was
very low: when June rainfall exceeded 80–100 mm, virtually no chicks were raised. Breeding success
was also affected by predation, but rainfall was the overriding factor: breeding success was low in wet years,
even when predation was low.
High June rainfall causes high chick mortality, likely due to reduced food supplies and increased chilling
of chicks93
. It was thought possible to mitigate some of the impacts of high June rainfall with vegetation
management, but trials involving cutting pathways through tall heather failed to increase breeding success,
even with a predator control programme in place. Historical weather data show that very wet Junes have
increased in frequency, so in the longer term, this could have an increasing impact on capercaillie breeding
success: even if Highland summers become drier overall, increasingly intense June rainfall events could still
be detrimental to the capercaillie.
Bearded tit survival and flooding (Wilson & Peach, 2006)94
With a distribution restricted to reedbeds, the bearded tit is a species of conservation concern in the UK.
At one breeding site, Leighton Moss in north-west England, the adult population increased from 60 to 180
between 1992 and 2000. However, in October to December 2000, exceptionally high rainfall occurred
in England – the highest total for over 300 years – leading to a severe winter flood and a bearded tit
population crash of 94%. Survival was very low, with the flood appearing to be the main cause. The flood
prevented the birds foraging in the layer of reed litter in the reed bed, severely reducing food availability.
Adults weighed in December 2000 were 20% lighter than in preceding years. As climate change projections
suggest that autumn and winter rainfall could increase over time, the risk posed by extreme floods to
bearded tit populations could increase in the future.
23
Estimates of bearded tit survival and population size at Leighton Moss. Bars show the number of adults,
circles and the solid red line show adult survival, and diamonds and the dashed dark blue line show survival
of birds in their first winter. Extreme rainfall and a resulting flood between the 2000 and 2001 surveys led to
much-reduced survival and a 94% population crash. After this, survival increased again, but population levels
recovered slowly.
Adapted from Wilson & Peach (2006), Animal Conservation 9 (4), 463–473.
0
0.2
0.4
0.6
0.8
1
0
50
100
150
200
Su
rviv
al ra
te
Nu
mb
er
of
ad
ults
Year
24
7) The future
Further, large shifts could occur in species’ ranges in the future, with many species losing
large areas of suitable climate, creating a substantial threat to their survival.
Examples so far have shown that species distributions, species interactions, reproduction and survival
have already been affected by the changing climate. IPCC projections show that by the end of the 21st
century, global temperatures are likely to have increased by at least 0.9°C and as much as 5.4°C, relative
to 1850–19004. The impacts on wildlife are therefore likely to become more severe
95, so it is important to
use our understanding of ecology to examine what might happen in the future.
Much work to understand future impacts of climate change on wildlife involves the use of species distribution
models, which establish statistical relationships between species’ current distributions and important climate
variables, including moisture availability, minimum temperatures and summer warmth. Once these
associations are established, projections of future climatic conditions can be used to see where might be
suitable for a species in future. Understanding how these potential distributions change in location
and size is vital to understand and prepare for longer-term impacts of climate change on wildlife.
The future potential geographic ranges of a wide range of species are projected to shrink and shift
northwards under climate change. For some groups of animals, assessments of future potential ranges
have been made for all species in Europe. Many European butterfly species are projected to shift north over
the 21st century and the climatically-suitable area will decrease for many96
. If populations can’t disperse well,
and under a high climate change scenario, nearly a quarter of butterfly species could lose over 95% of their
current range by 2080, and nearly four-fifths would lose over 50%. Even if climate change proceeds at an
intermediate rate, nearly a tenth of species would lose over 95% of their current range, and two-thirds would
lose more than 50%96
. Similarly, for European bumblebees, around a third of species could lose over 80% of
their current range by 2100 under a severe climate change scenario, whilst around two-fifths of species could
lose over 50%97
. The greater the loss of suitable climate, the greater the threat to the species’ survival.
Projections have also been made for a variety of other groups. One study modelled the distributions of
1,350 European plant species – just over 10% of all plant species in Europe98
. Assuming a high rate of
climate change and no migration of species, over a fifth of plant species modelled could lose over 80% of
their range by 2080, and 2% could go extinct altogether; even allowing for migration and with limited
climate change, many species face losing large proportions of their current ranges98
. Among 1,648
European freshwater species, including plants, fishes, molluscs and amphibians, many were projected to
shift north-east by the 2050s under an intermediate climate change scenario; nearly three-fifths of the
species were projected to lose over 50% of their current range; rare species were most threatened, with
nearly four-fifths losing more than 90% of their current range99
. For fish populations in the North Atlantic,
rising sea temperatures were projected to drive substantial distribution changes in a range of fish species
by the 2090s: southerly-distributed species such as anchovy and horse mackerel were projected to occur
25
throughout the North Sea, whilst northerly-distributed species like haddock and saithe were projected to
be lost from the southern North Sea100
.
Although many models consider how distributions might change, conservationists also need
to consider population abundances. In a model of British breeding birds, populations of two
northerly-distributed British breeding bird species, the Eurasian curlew and meadow pipit, were projected
to decline over the 21st century, with the curlew decline particularly sharp101
. Conversely, the same study
suggested that two southerly-distributed species, the Eurasian nuthatch and green woodpecker, could
increase substantially in both abundance and range101
.
Habitat availability is another key influence on future distributions and conservation strategies.
Modelling studies of the Austrian Alps suggest that, even with only 1.8°C warming, many plant and
invertebrate species found only there – endemic species – could lose their mountain-top habitat as the tree
line moves uphill in response to warming102
. Large European mammals are also projected to be threatened,
with changing land use interacting with the changing climate to drive substantial habitat loss: on average,
10–25% of habitat could be lost by 2050, with high-latitude or high-altitude species such as reindeer and
chamois likely to fare particularly badly103
.
It is also important to consider how populations might behave and species interact in landscapes
changed by climate change. A model of Iberian lynx populations combined projections of climatic suitability
with population models of both the lynx and of rabbits, the lynx’s primary prey. Results indicated that, without
significant conservation interventions, climate change could drive the Iberian lynx to extinction by 205045
.
Ecological modelling can use relationships from the present to make valuable projections of the future.
It is important to note that projections are not firm forecasts or predictions: they show what might happen
based on a set of possible conditions, and there are many factors the models do not yet incorporate.
However, although projections might seem detached from real impacts, populations are already showing the
responses projected (see section on “Community change”, from page 13). Therefore, climate change looks
set to increasingly affect wildlife populations, and whilst some species could do well in some places,
many populations, and perhaps even entire species, could be at risk of extinction.
Case study: climate change and breeding bird distributions (Huntley et al, 2007)35
Huntley et al developed models of the distributions of 453 European breeding bird species using information
on breeding distribution between 1961–1990 and three important climate variables; one indicating the
degree of winter cold, another indicating available summer warmth, and the last indicating moisture
availability. These models were then combined with projections of future climatic conditions for 2070–99,
to give an indication of potential future distributions.
These projections indicate the potential range: the areas where climatic suitability suggests a species could
occur. However, for a species to occupy its climatically-suitable area, there must be appropriate habitat and
sufficient other resources, and it must be able to disperse across the landscape. On average, by 2070–99,
26
species’ potential ranges were projected to be nearly 550 km further north and north-east. However, there
was variation in the directions that species were projected to move in: ecological communities will change as
species respond differently to the changing climate.
Present and future potential distribution maps for Dartford warbler, a southerly-distributed species, and
willow (red) grouse, a northerly distributed species, in 1961–90 and the late 21st century. Blue squares show
where climate is suitable for the species to occur; yellow squares show where the species are predicted to
be absent. For both species, the area with suitable climate will decrease, and will show limited overlap with
the existing range. For Dartford warbler, Britain could become more important under climate change; for
willow grouse, much of the existing British range could become unsuitable.
Figures reproduced with permission from Huntley et al. (2007),
“A Climatic Atlas of European Breeding Birds”, Durham University, RSPB and Lynx Edicions.
Simulated distribution for 1961-1990 Projected distribution for late 21st Century
Dartford warbler
Future range: 86% size of current range, 39% overlap with current range
Simulated distribution for 1961-1990 Projected distribution for late 21st Century
Willow grouse
Future range: 50% size of current range, 39% overlap with current range
27
Under all climate change scenarios, future breeding distributions were only around 80% of the size of
current distributions. These shifting, shrinking distributions led to limited overlap (40% on average)
with recent distributions – 27 species had no overlap between recent and future distributions, while
51 species had less than 10% overlap. The smaller the overlap, the greater the extinction risk. Endemic
species – those found only in Europe – appeared to be at greater risk, with an average overlap of only
around 15%. Overall, the changes to the European bird community could be substantial.
These shifting distributions could lead to more species in some areas, and fewer in others. The UK
could become important for some species currently found further south, reinforcing the need for regional
co-operation in adaptation, but species currently found in the north and uplands could face substantial
loss of suitable locations.
28
8) Protected areas and climate change
As species’ ranges shift, both existing and new protected areas will play an important role
in helping wildlife cope with the effects of climate change.
Protected areas are crucial in conservation. In the much-altered landscapes of Europe, they provide
plants and animals with the conditions and habitat they need to survive. They are not solely nature
reserves, but represent “a clearly defined geographical space, recognised, dedicated and managed, through
legal or other effective means, to achieve the long-term conservation of nature with associated ecosystem
services and cultural values”104
. Around 21% of EU land or water is protected in some way, covering around
1,200,000 km2 104
. It is important to examine the role protected areas will play in a world increasingly affected
by climate change.
Empirical evidence shows that protected areas are already proving important as the climate changes.
Protected areas can aid threatened wildlife by allowing larger, more resilient populations to become
established. Across 415 European breeding bird species, long-term (1980–2012) population trends were
in the direction expected under climate change, confirming that climate change is already influencing the
species105
. However, those species protected under Annex I of the EU Birds Directive, of which a key
element is the establishment of Special Protection Areas (SPAs) for important habitats and populations,
had more positive population trends. Provision of protected areas, along with other protection
measures, aids populations despite the underlying effects of climate change105
.
Similarly, across 90 bird species in Finland, between the 1970s and 2000s, some communities showed
declining diversity and others showed increasing diversity (both in line with predictions of climate change
impacts)106
. However, regardless of the trend, protected areas maintained higher numbers of species than
unprotected areas106
.
As well as helping species to establish larger, more resilient populations within their existing range,
protected areas also help species colonise and establish in new areas, even though the protected
areas there were not designated for those species. The silver-spotted skipper butterfly has expanded its UK
range in recent years: protected areas actively managed for conservation were over three times more likely
to have been colonised than unmanaged protected sites or unprotected sites, and population survival was
higher in protected sites107
. Of 57 butterfly species and 42 dragonfly and damselfly species in the UK, 61
species were more abundant inside protected areas than outside, when looking at areas colonised since
the 1960s–80s: protected areas appear to have helped species to establish larger populations during range
expansion108
. Likewise, in Europe, as the wintering smew distribution has shifted north-east, population
growth in the north-east has been nearly twice as fast inside SPAs than it has been outside.109
But what about the future? A modelling analysis, looking at plants, mammals, birds, amphibians and
reptiles, considered the importance of protected areas across Europe: climatic suitability in protected areas
was projected to decrease by 2080 for 58–63% of species modelled, based on their current distributions
(ie, assuming species cannot move in response to climate change)110
. Although some species could see
29
climatic conditions improving, a greater proportion of the species modelled would see climatic conditions
deteriorating (ie “losers” were projected to outnumber “winners” across most groups)110
. However, for most
groups, protected areas would retain suitable climate as well as, or even better than, areas outside of the
network110
. More northerly areas, such as parts of the UK and Scandinavia, and higher-elevation areas, such
as the Alps, were projected to have more “winner” species, whilst more southerly countries were projected to
have more “loser” species110
.
However, even with some reductions in climate suitability, protected area networks should continue
to function effectively. A study of SPAs in the UK found that although climate change could cause the
abundance of breeding seabirds and wintering waterbirds to decline over the next 70 years, most SPAs
would keep large enough populations of their key species to warrant continued legal protection111
.
Further, although some SPAs may lose some qualifying species, new species are expected to arrive:
sites that are currently important should remain important and the site network as a whole should
remain resilient111
.
A similar modelling exercise considered how four bird of prey species would be protected by Natura 2000
sites under future conditions: although some sites could become less important, the network as a whole
should maintain current performance112
. Increasing connectivity between southern and northern protected
sites would benefit lesser spotted eagles, whilst adaptation for griffon vultures, golden eagles and Egyptian
vultures can be achieved by maintaining and expanding existing protected areas112
. Finally, 2,500 plant
species in the French Alps were modelled, and the effectiveness of existing protected areas was assessed:
by 2080, although plant communities could experience substantial changes, the protected area network
would remain effective, with rare species experiencing smaller impacts, and threatened species protected
just as well in the future as the present113
.
It is clear, however, that additional conservation efforts will be needed to supplement the existing protected
area network. A study of 1,200 European plant species generated a theoretical reserve network based on
current distributions of species; future distributions under climate change were then projected, and the
effectiveness of the simulated reserve network assessed114
. Depending on assumptions used, 6–11% of
species lost suitable climatic conditions from the “current” reserve network114
. Another analysis considered
plant species in protected areas in Western Europe: under future climates, the current protected area
network could need to expand by 30–40%115
. Given that further protected areas are likely to be required,
it will be important to identify areas that would be important under both current and future climates115
(see section on “Creating new sites”, from page 35). In some cases, other conservation measures
such as translocation (or even assisted colonisation outside the current range) might need to
be considered.114
.
30
Case study: protected areas, survival and colonisation
Protected areas and shifting range margins (Gillingham et al, 2015)116
The role of protected areas at expanding northern range margins and retracting southern range margins has
been examined for seven butterfly species and eleven bird species in Britain. Some are found in southern
Britain, so might spread north into new areas, and some are found in northern Britain, so might lose the
southernmost parts of their range. Distributions were compared between an early time period (ranging from
the 1970s to the 1990s, dependent upon data availability) and a later time period (ranging from the 1990s to
the 2000s). Protected areas were defined as sites with Site of Special Scientific Interest (SSSI) designation.
Among the northern species considered, survival at lower latitudes and altitudes was more likely in areas
with a higher coverage of SSSIs – species were more likely to persist at their southern range margin in
areas with greater protection. Among the southern species considered, colonisation was more likely in
areas with higher coverage of SSSIs – as their range expanded, species were more likely to colonise
areas with greater protection. Overall, therefore, protected areas appeared to facilitate both population
persistence and colonisation as species’ ranges shifted.
Protected areas and range expansions (Thomas et al, 2012)117
Seven British species – two butterflies and five birds – were examined in detail to see whether their
northwards range expansion was influenced by protected areas. Species distributions were compared
between the 1970s–80s and the 2000s, to identify sites that had been colonised. Protected areas were
colonised 4.2 times more frequently than expected given the available coverage of protected areas. Across
256 invertebrate species with less detailed survey data, 251 were recorded more frequently than expected
in protected areas in newly-colonised parts of their ranges. This suggests that protected areas are already
facilitating range expansions in response to the warming climate.
Map of Dartford warbler breeding distribution (left figure), showing sites where breeding commenced before
1991 (blue squares) and where breeding commenced after 1991 (red squares). Expanded view of the black
box (right figure) shows SSSI coverage in green, and locations where Dartford warblers were observed: sites
colonised after 1991 (red crosses) are clustered in and around protected areas.
Reproduced with permission from Thomas et al. (2012),
Proceedings of the National Academy of Sciences 109 (35), 14063–14068.
31
9) Managing sites and landscapes for adaptation
Different site management techniques will become more important to help wildlife to adapt
to climate change.
The way sites are managed – both protected areas and other sites – influences the role they play
in helping wildlife adapt to climate change. It is already clear that areas with active conservation
management are more beneficial than those without107
. However, site management can also interact
with wider landscape management to influence how species respond to climate change.
Three common aims of site management for climate change adaptation include: increasing resistance
to climate-driven change; helping populations to be more resilient, to bounce back from impacts of
climate-driven change; and accommodating climate-driven change118
. Much site management for adaptation
targets the first two aims: counteracting climate change effects with action to reduce the severity of the
impacts, and tackling non-climatic stressors to compensate for climate change effects119, 120
.
A key action is to provide a range of microhabitats and microclimates – small-scale patches where
habitat or climate differ from prevailing local conditions. Increased variation creates a range of options
for species that might need to adapt. For Glanville fritillary butterflies in the UK, egg-laying occurs on plants
that are warmer than the ambient air temperature, and abundances are higher where the habitat contains
bare ground and short grass121
. Creating a mosaic of short turf and taller vegetation would provide variation
in thermal conditions, allowing flexibility in egg-laying locations to counteract variable temperatures under
climate change121
. In European deciduous forests, managing forest canopies to retain denser cover could
help cool-adapted plants on the forest floor, with cooler microclimates limiting some of the effects of
climate change122
.
Such thinking is already being put into practice. At the Winterbourne Downs RSPB nature reserve, a lowland
grassland site in southern England, S-shaped chalk banks have been created and planted with butterfly food
plants. This creates a range of different microclimates, providing warmer and cooler areas so that insects
can find suitable conditions under a range of climatic conditions.
Hydrological management is also often important. Shallow drainage channels on farmland, which can
produce small surface floods, are associated with greater abundances of breeding lapwings, and more nests
nearby123
. With climate change expected to increase risk of both flooding and droughts, these shallow drains
could help retain water throughout the breeding season and prevent larger floods following heavy rain123
.
Black-tailed godwits in the UK nest in wet meadows used for floodwater storage; so have much reduced
breeding success during floods124
. Increased flood risk under climate change could threaten the populations,
so flood mitigation measures in the meadows and provision of better habitat in adjacent farmland is needed
to safeguard the population124
.
32
The area and quality of habitat patches, along with characteristics of the wider landscape, are
key considerations. The silver-spotted skipper butterfly is expanding its range in the UK; between 2000
and 2009, higher survival in colonising populations was associated with larger habitat patches, increased
availability of sheep’s fescue grass, and intermediate bare ground cover125
. Providing such habitat features
is important to aid expansion of the species’ range125
. Similarly, woodland specialist bird species in the
UK, such as nuthatch and green woodpecker, recovered better from small population sizes when in
larger woodlands126
. Landscape characteristics can also influence the extent of climate impacts:
generalist woodland species are less sensitive to very cold winters in landscapes with less fragmented
woodland cover126
.
Climate change could also interact with changing management of the wider landscape. In the
Netherlands, the black-tailed godwit population has declined: this appears to be linked to a combination of
land management and climate change, with rising temperatures and agricultural intensification combining to
reduce the amount and quality of the habitats that chicks need for foraging127
. Agri-environment schemes
could help to address some of the issues, but need to be designed and implemented to take into account
the effects of both agricultural impacts and rising temperatures127
.
In the French Alps, a modelling study considered how pasture management and climate change could
interact in the future: intensification of pasture use would cause rapid loss of plant diversity, with climate
change gradually leading to further losses; abandonment of pasture could initially increase diversity, but
climate change could then decrease diversity further as the tree line rapidly expands128
. Maintaining low
intensity pasture may therefore be the best option under climate change128
.
Another modelling study considered the effects of landscape management on breeding wading birds in the
Netherlands: conversion of pasture land to bioenergy crops, which could be an element of human climate
change adaptation, would reduce habitat suitability for the birds by reducing habitat openness and raising
groundwater levels129
. Climate change will influence the way humans manage land, and these
changes will be an important determinant of how wildlife adapts to climate change.
Protected areas may be specifically managed for conservation purposes; evaluating and altering this
management is an important part of improving the resilience of wildlife populations under climate change,
helping species move through the landscape effectively, and ameliorating negative impacts. However,
large areas of Europe are under intensive agricultural or forestry management. The management of this
wider landscape must therefore also be considered, with larger-scale management changes such as land
abandonment or shifts to new crops potentially affecting species’ responses to climate change.
33
Case study: climate change resilience in upland birds (Carroll et al, 2011; Carroll et al, 2015)130, 131
The European golden plover breeds in the UK uplands, but these breeding populations are among the most
southerly in its range, potentially making them susceptible to climate change impacts. During the breeding
season, golden plovers rely on craneflies for food, but cranefly larvae are sensitive to drought, meaning
that climate change could threaten the birds’ food supply132
. Modelling suggests that the abundance and
persistence of golden plover populations in the Peak District of northern England could decline as warmer,
drier summers reduce cranefly abundances.
However, management could increase resilience of these populations. Throughout the 20th century, many
British upland peatlands were drained with the intention of improving agriculture, but drainage could
exacerbate cranefly declines and have further impacts on ecosystem functioning. Experimental examination
of three drained peatlands has shown that blocking drains as part of restoration programmes leads to wetter
peat and higher cranefly abundances.
Modelled golden plover abundances in the Peak District under recent conditions and for a future climate
change scenario: darker red indicates higher abundances, greys and lighter reds indicate lower abundances.
Abundances are projected to decline overall and populations to retract into the wettest areas where cranefly
abundances remain high.
Reproduced with permission from Carroll et al. (2015), Nature Communications 6, 7851.
34
Cranefly abundance at drains that have been blocked during restoration and those that remain open. Higher
abundances were found at blocked drains. Sampling occurred immediately at the drain edge (near) and
10 m away (far), showing that the benefits of drain blocking extend away from the drains.
Adapted from Carroll et al. (2011), Global Change Biology 17 (9), 2991–3001.
Blocking drainage ditches could therefore provide more food for golden plovers in drained peatlands,
aiding populations under climate change. Other ecosystem services benefits could also be achieved
by drain blocking, such as improved carbon storage, improved water quality and reduced flood risk
(eg Wilson et al, 2011)133
.
Blanket bog restoration and drain blocking have already been carried out by the RSPB and partners in
projects such as the Sustainable Catchment Management Programme (SCaMP) in the Peak District and the
Active Blanket Bog Wales EU LIFE project. These projects show how, by acting on site management now,
we can provide immediate benefits as well as longer-term climate change resilience.
35
10) Creating new sites and expanding site networks
Creating, re-creating and protecting new sites for wildlife conservation will play important
roles, from helping populations to disperse through inhospitable landscapes, to increasing
the size of existing protected areas to enhance their resilience, to providing entirely new
locations for populations to inhabit.
Creating new areas of habitat and new protected areas may be necessary to compensate for areas where
existing habitat is under threat, in areas that are likely to become more important under climate change, and
where populations need to move across the landscape to track suitable climate. It is therefore important to
consider how the creation of new sites could aid wildlife.
One key adaptation action is to increase the ability of populations to track suitable climate. Habitat
fragmentation, where formerly continuous areas of habitat are broken up into smaller,
unconnected pieces, could worsen the impacts of climate change by limiting species’ ability to
colonise newly-suitable areas134
. However, many habitats are already fragmented, and protected areas
may be surrounded by a matrix of less suitable habitat, meaning that a core function of site creation could be
to improve the connectivity between areas of suitable habitat. Connectivity could be achieved by making the
wider landscape more wildlife-friendly, but could also be done using “wildlife corridors”, whereby smaller
areas of habitat are created to join together larger main patches135
. Some empirical evidence shows that
species use corridors or scattered habitat patches to move across landscapes136
. Further, if populations in
core areas decline, due to extreme weather for example, then improved connectivity might allow “rescue”
of that population as individuals from nearby areas recolonise119
.
A model of habitat creation in Yorkshire suggested that linking existing good-quality habitat patches to
prevent “bottlenecks” (where species cannot reach good quality habitat) would be the most effective way
of promoting movement across the landscape137
. However, adding more habitat near to good-quality,
well-connected patches might be better at increasing population persistence, suggesting that promoting
dispersal and increasing persistence might require two complementary, but different, habitat
creation strategies137
. On balance, larger, better quality habitats might aid wildlife populations even
more than smaller, well-connected habitats138
.
In some instances, creating new sites for wildlife might not seek to simply link together existing
areas, but to provide entirely new opportunities in areas likely to become climatically suitable. In
Italy, a modelling study suggested that both nationally-designated protected areas and Natura 2000 sites
are crucial to achieve good protection of amphibians under climate change139
. However, areas that would
become more important under climate change were also discovered, suggesting that the creation of further
reserves in parts of Sardinia, Sicily and the north-east mainland would enhance the network further139
.
Similarly, modelling of otter distributions in Europe identified areas that are likely to become more suitable
under climate change, where otters don’t currently occur, such as areas of central-northern Italy, southern
36
France and eastern France140
. Habitat creation and protection could be put in place in these
climatically-suitable areas to promote otter expansion under climate change140
.
Observations of ongoing changes can inform approaches to creating new sites. For example, the
RSPB has re-created rare lowland heathland further north than the core range of many associated species,
so that as the climate warms, there will be habitat available for the populations to colonise118
. Similarly, it
has been suggested that large wetlands should be created in Britain to accommodate incoming waterbird
species that require large areas of habitat to establish breeding populations: existing wetlands in Somerset,
Cambridgeshire, Norfolk and Suffolk could be the focus of such habitat creation37
. Habitat creation can also
counter some impacts of sea-level rise: managed coastal realignment has been carried out at sites such as
the RSPB’s Wallasea Island reserve, to re-create important intertidal habitats such as salt marshes, thus
helping to counteract the loss of intertidal habitats elsewhere through development and “coastal squeeze”118
.
Case study: creating wetlands for bitterns (Ausden, 2014; Gilbert et al, 2010; Wotton et al, 2009)118, 141, 142
Bitterns became extinct as a breeding species in Britain in the 19th century following persecution and
habitat loss, but re-colonised in the early 20th century and grew to a peak of 7080 males by the 1950s.
However, the population declined again as the species' freshwater reedbed habitat degraded in some
areas, reaching a minimum of 11 males in 1997. The majority of these were at coastal reedbeds in Suffolk,
which were vulnerable to coastal flooding. Flooding with seawater reduces bitterns' food supply, in turn
affecting their breeding success and survival. Climate change-driven sea-level rise therefore threatened
these populations, as saltwater incursion was expected to become more frequent; mathematical modelling
suggested that the UK population would be even more vulnerable if these core sites were flooded
more often.
An ambitious habitat creation initiative began to tackle this problem, underpinned by the bittern’s legal
protection under the 1979 EU Birds Directive. Reedbeds were restored and new reedbeds created at sites
further inland, away from areas vulnerable to coastal flooding. Population monitoring indicates that bittern
populations have rapidly increased in the new reedbeds, so that as the total national population has grown,
the risks posed by sea-level rise have decreased. Other species have benefited too, with the first records of
breeding by purple heron and great egret, and the first regular breeding of little bittern occurring in newly
created reedbeds. Creating new reedbeds therefore appears to have reduced climate change risk for
bitterns, whilst providing opportunities for colonising species.
37
Number of male bitterns on RSPB reserves under different degrees of coastal flood risk. As the population
has grown, the proportion of the population at risk from seawater flooding has reduced, as populations
establish in newly created reedbeds further inland.
Figure adapted and updated from Ausden et al. (2014), Environmental Management 54, 685–698.
0
10
20
30
40
50
60
70
Nu
mb
er
of
bo
om
ing
ma
le b
itte
rns
Year
<1 in 75 annual risk of coastal flooding (reedbeds created since the mid-1990s)
<1 in 75 annual risk of coastal flooding (reedbeds present before the mid-1990s)
>1 in 75 annual risk of coastal flooding (reedbeds present before the mid-1990s)
38
11) Helping people to adapt
Humans and wildlife both need to adapt to climate change. Restoring and conserving
natural ecosystems could, in some cases, provide benefits to nature and humans at the
same time.
Like wildlife, human society also needs to adapt to climate change. A previously-discussed example
showed how land management to aid human adaptation, if not done carefully, could reduce habitat quality
for breeding wading birds129
. Whilst such conflicts are possible, they are not inevitable and adaptation for
humans can even be achieved via measures that benefit ecosystems and wildlife.
“Ecosystem-based adaptation” is defined by the Convention on Biological Diversity as “the use of
biodiversity and ecosystem services as part of an overall adaptation strategy to help people to adapt to the
adverse effects of climate change”143
. Adaptation of this kind includes managing uplands, water courses and
flood plains to store water and reduce flood risks, managing coasts to prevent flooding and erosion, and
managing forests to stabilise land and maintain water flow144
.
These “soft”, ecosystem-based approaches can be contrasted with “hard” approaches based on engineered
solutions. Although they can be effective, “hard” approaches have drawbacks. For example, sea walls can
damage or alter intertidal habitats such as salt marshes, which in turn leads to a reduction in natural flood
protection, as well as biodiversity loss. In a “soft” approach, combining natural defence from more extensive
intertidal habitats with smaller sea walls, flood protection can be more cost-effective, provide longer-term
protection under rising sea-levels and benefit biodiversity145
. Further, “hard” interventions often only provide
the service for which they were built, and are inflexible in a world that is increasingly defined by change,
whereas ecosystem-based approaches provide multiple services and are very flexible146
.
Many examples of ecosystem-based adaptation already exist, addressing adaptation to current climatic
variability, rather than long-term change147
. They are concerned, for example, with how to manage water
scarcity or flooding, or how to maintain agricultural productivity, whilst the hazards addressed include
changing rainfall patterns, high temperatures and sea-level rise; encouragingly, most examples report
positive outcomes147
.
Controlling flooding is a key concern under climate change. Sustainable floodplain management can
be an important way to reduce flood risks148
. Floodplain restoration around the Skjern river in Denmark
commenced in the late 1990s to counteract a range of “engineered” changes made in the 1960s, such as
river straightening and meadow drainage149
. As well as benefiting wildlife, the restoration has saved water
pumping costs, reduced nutrient concentrations in the water, reduced carbon dioxide emissions, provided
recreation opportunities and reduced flood risk; the benefits outweigh the costs of restoration149
. Similarly,
areas of floodplain around the lower Danube have recently been restored after agricultural conversion and
dike building led to increased flood peaks: in 2005, a single flood cost €396 million in damages, and flood
frequency is expected to increase under climate change150
. Whilst some “hard” management was retained,
39
the floodplain has been restored by decommissioning dikes; restoration is intended to benefit wetland
wildlife, provide new recreation and economic opportunities, and improve flood protection151
.
Impacts of rising temperatures can, in some cases, also be combated with ecosystem-based
approaches. In the UK, rising river temperatures could harm freshwater wildlife, notably including brown
trout and salmon152
. To counteract this, trees and other vegetation can be replanted alongside rivers to
reduce temperatures by providing shade; further benefits for human adaptation could come from lowered
flood risk and reduced erosion152
. Similarly, in cities, increasing green space can help to reduce
temperatures whilst improving urban biodiversity; modelling suggests that adding only 10% of green cover
to city centres could keep maximum surface temperatures similar to or lower than 20th century levels even
under high climate change scenarios153
.
Ecosystem management could help to reduce overall greenhouse gas emissions (climate change
mitigation). Peatland restoration is an example of both climate change adaptation and mitigation, with
improved flood management and carbon storage both resulting from better peatland management. Peatland
restoration in the UK has already been shown to benefit breeding birds, carbon storage and hydrology130, 133
.
Another example comes from Belarus, where a partnership co-ordinated by the RSPB was established to
restore degraded peatlands. Between 2009 and 2011, over 17,000 ha (hectares) of drained and degraded
peatland was re-wetted154
. The restoration could reduce carbon emissions by an amount equivalent to
30,400 tonnes of CO2 each year, providing substantial climate change mitigation benefits154
. Threatened bird
species such as the aquatic warbler and greater spotted eagle should benefit from the restored habitat, and
local human communities should benefit from increased fish populations and cranberry production154
.
Successful application of ecosystem-based adaptation principles is increasing, but there is still more to
learn about how and where such approaches can be applied, and how they compare in cost and efficacy
to alternative adaptation measures. Integrating wildlife into climate change adaptation and mitigation
measures could provide much wider benefits than considering humans separately from the rest of
the natural world.
Case study: Wallasea Island Wild Coast (Ausden, 2014; Ausden, 2015)118, 155
Wallasea Island on England's Essex coast is the subject of a major project to provide valuable habitat for
wildlife. This started with a scheme to compensate, under the EU Birds Directives, for loss of intertidal habitat
to port development in the region. It has subsequently expanded to compensate more broadly for the loss of
coastal and intertidal habitats in the area and nationally, and to aid adaptation to sea-level rise.
There have been large, historical losses of saltmarsh in Essex through land claim, with the county now
retaining only around 2,000 ha of saltmarsh, from an estimated 40,000 ha in the 14th century. Until recently,
the island was low-lying farmland protected by sea walls. However, the sea walls were of low flood defence
standard, raising the risk of them being breached during a storm surge and to upgrade them would be highly
costly. Uncontrolled breaching would have big consequences for the rest of the estuary, with the associated
increased volume of water flowing into and out of the rest of the estuary impacting navigation and the
40
estuary’s existing conservation interest, as well as increasing pressures on flood defences in the
surrounding area.
Wallasea Island before controlled breaching of the seawall, showing landscaping to create creeks, islands
and pools that will allow important intertidal habitats to form.
Photo credit: BAM Nuttall.
The RSPB, in partnership with Crossrail, Defra, the Environment Agency and Natural England, has
re-created intertidal habitat through managed realignment on part of the island, which will provide benefits
for wildlife together with a range of benefits for people. Crossrail, Europe’s largest infrastructure project, is
the development of a high frequency, high capacity railway for London and the South East of England.
Using clean soil excavated during the Crossrail project, the level of the island has been raised so that
following flooding, the increase in volume of water flowing into and out of the rest of the estuary will be kept
within acceptable limits. Landscaping to create creeks, pools, islands and lagoons has been combined with
the controlled breaching of sea walls to introduce tidal flooding, leading to the re-creation of various intertidal
habitats. These will provide valuable habitat for a range of species, including wading birds, rare plants and
invertebrates. The design of the site is also intended to provide suitable breeding habitat for spoonbills, and
so facilitate their re-colonisation of the UK. Furthermore, the project benefits people by storing water in the
event of storm surges and thereby reducing pressure on flood defences in the surrounding area,
sequestering carbon in buried sediment, and providing recreation opportunities.
41
Wallasea Island shows how, with careful planning, big benefits can be provided for both people and wildlife adapting to climate change.
Map of the current and intended habitats on Wallasea Island. The top map shows the whole island,
indicating the range of habitats, while the lower map shows greater detail for Jubilee Marsh on the
east of the island.
Figure reproduced courtesy of Malcolm Ausden.
Map of Wallasea Island Wild Coast
Detail of Jubilee Marsh
Water Vole
mitigation
area
Salt pan
Saline
lagoon
Regulated tidal exchange
area containing islands
and lagoons
Artificial
saline creek
network
0 1 km
N
Visitor hub Rotationally
dried-out
lagoons Currently farmland, but
intended eventually to
become wetland habitat
Grazing
marsh and
seasonal
floods
Wild bird
cover Managed realignment
with islands & lagoons
(Jubilee Marsh)
Reptile
mitigation area
0
Islands in
a saline
lagoon
Sub-tidal lagoon
Breach 1
Shingle ridge for
nesting terns &
ringed plovers
Breach 2
Spoonbill
nesting island
Breach 3
Sea wall and
sea-level-rise
adaptation zone
Permanent water
Temporary water
Mudflat
Shingle, sand and cockle shells
Saltmarsh
Saltmarsh/grassland transition
Grassland
Grassland (reptile mitigation area)
Farmland
Wild bird cover
Scrub
Key
1 km
42
12) Implications for conservation
This report provides widespread evidence that climate change is already affecting wildlife across Europe,
and that impacts are likely to escalate in the not-too-distant future. This will add to, and interact with, many
other pressures faced by wildlife. How should we respond?
Greenhouse gas emissions should be reduced to limit the degree of warming: although wildlife
is already being affected, future climate change is likely to lead to more severe impacts. Reducing
emissions is a key step in reducing impacts.
Species’ ranges are changing as they track suitable climate: providing sufficient, suitable habitat
is key in ensuring species can track the conditions to which they are adapted.
Some species could benefit from climate change, at least at certain geographical scales, whilst many
could be adversely affected: resulting changes in ecological communities should be monitored
to ensure efficient targeting of conservation action, and identification of mechanisms behind impacts.
Extreme weather will increasingly affect populations, and wider climatic conditions could become
less suitable for many species: under these circumstances it is important to identify site and
landscape management actions to aid population resilience and adaptation.
Non-climatic pressures on wildlife should be reduced. As climate change will have an increasing
impact on wildlife, it will be important to ensure other threats are reduced.
Robust protected area networks are essential now, and will continue to be essential under
climate change. Protected areas support larger populations and more diverse communities, and
aid colonisation of new areas and survival in existing ranges.
The existing protected area network could be enhanced: “more, bigger, better and joined”
sites156
will aid population persistence and species’ movement across the landscape.
Ecosystem-based adaptation could aid human climate change adaptation and benefit
biodiversity at the same time. Working with nature could provide much wider benefits than
considering humans separately from the environment.
43
13) References
1. Hartmann, DL et al. (2013) Observations: atmosphere and surface, in Climate Change 2013: The Physical Science Basis. Contribution of Working Group I to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change, Stocker, TF, et al (Editors). Cambridge University Press, Cambridge, UK and New York, NY, USA.
2. Diffenbaugh, NS and Field, CB. (2013) Changes in ecologically critical terrestrial climate conditions. Science 341, 486–492.
3. Kirtman, B et al. (2013) Near-term climate change: projections and predictability, in Climate Change 2013: The Physical Science Basis. Contribution of Working Group I to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change, Stocker, TF, et al (Editors). Cambridge University Press, Cambridge, UK and New York, NY, USA. 953–1028.
4. Collins, M et al. (2013) Long-term climate change: projections, commitments and irreversibility, in Climate Change 2013: The Physical Science Basis. Contribution of Working Group I to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change, Stocker, TF, et al (Editors). Cambridge University Press, Cambridge, UK and New York, NY, USA. 1029–1136.
5. Peñuelas, J et al. (2013) Evidence of current impact of climate change on life: a walk from genes to the biosphere. Global Change Biology 19(8), 2303–2338.
6. Walther, GR. (2010) Community and ecosystem responses to recent climate change. Philosophical Transactions of the Royal Society B-Biological Sciences 365, 2019–2024.
7. Ockendon, N et al. (2014) Mechanisms underpinning climatic impacts on natural populations: altered species interactions are more important than direct effects. Global Change Biology 20(7), 2221–2229.
8. Jentsch, A, Kreyling, J and Beierkuhnlein, C. (2007) A new generation of climate-change experiments: events, not trends. Frontiers in Ecology and the Environment 5(7), 365–374.
9. Thomas, CD et al. (2004) Extinction risk from climate change. Nature 427, 145–148. 10. Dawson, TP et al. (2011) Beyond predictions: biodiversity conservation in a changing climate.
Science 332, 53–58. 11. Thomas, CD and Lennon, JJ. (1999) Birds extend their ranges northwards. Nature 399, 213–213. 12. Gillings, S, Balmer, DE and Fuller RJ. (2015) Directionality of recent bird distribution shifts and
climate change in Great Britain. Global Change Biology 21(6), 2155–2168. 13. Hickling, R et al. (2006) The distributions of a wide range of taxonomic groups are expanding
polewards. Global Change Biology 12(3), 450–455. 14. Mason, SC et al. (2015) Geographical range margins of many taxonomic groups continue to shift
polewards. Biological Journal of the Linnean Society 115(3), 586–597. 15. Hickling, R et al. (2005) A northward shift of range margins in British Odonata. Global Change
Biology 11(3), 502–506. 16. Battisti, A et al. (2005) Expansion of geographic range in the pine processionary moth caused by
increased winter temperatures. Ecological Applications 15(6), 2084–2096. 17. Franco, AMA et al. (2006) Impacts of climate warming and habitat loss on extinctions at species'
low–latitude range boundaries. Global Change Biology 12(8), 1545–1553. 18. Beaugrand, G et al. (2002) Reorganization of North Atlantic marine copepod biodiversity and
climate. Science 296, 1692–1694. 19. Perry, AL et al. (2005) Climate change and distribution shifts in marine fishes. Science 308,
1912–1915. 20. Maclean, IMD et al. (2008) Climate change causes rapid changes in the distribution and site
abundance of birds in winter. Global Change Biology 14(11), 2489–2500. 21. Hill, JK et al. (2002) Responses of butterflies to twentieth century climate warming: implications for
future ranges. Proceedings of the Royal Society B 269, 2163–2171. 22. Menéndez, R et al. (2014) Climate change and elevational range shifts: evidence from dung beetles
in two European mountain ranges. Global Ecology and Biogeography 23(6), 646–657. 23. Wilson, RJ et al. (2005) Changes to the elevational limits and extent of species ranges associated
with climate change. Ecology Letters 8(11), 1138–1146. 24. Dulvy, NK et al. (2008) Climate change and deepening of the North Sea fish assemblage: a biotic
indicator of warming seas. Journal of Applied Ecology 45(4), 1029–1039. 25. Pateman, RM et al. (2012) Temperature-dependent alterations in host use drive rapid range
expansion in a butterfly. Science 336, 1028–1030. 26. Anderson, SJ et al. (2008) Phenotypic changes and reduced genetic diversity have accompanied the
rapid decline of the garden tiger moth (Arctia caja) in the U.K. Ecological Entomology 33(5), 638–645.
44
27. Simmons, AD and Thomas, CD. (2004) Changes in dispersal during species’ range expansions. The American Naturalist 164(3), 378–395.
28. Pöyry, J et al. (2009) Species traits explain recent range shifts of Finnish butterflies. Global Change Biology 15(3), 732–743.
29. Popy, S, Bordignon, L and Prodon, R. (2010) A weak upward elevational shift in the distributions of breeding birds in the Italian Alps. Journal of Biogeography 37(1), 57–67.
30. Tayleur, C et al. (2015) Swedish birds are tracking temperature but not rainfall: evidence from a decade of abundance changes. Global Ecology and Biogeography 24(7), 859–872.
31. Wotton, S et al. (2009) The status of the Dartford Warbler in the UK and the Channel Islands in 2006. British Birds 102(5), 230–246.
32. Bradbury, RB et al. (2011) The influence of climate and topography in patterns of territory establishment in a range-expanding bird. Ibis 153(2), 336–344.
33. Holling, M and The Rare Breeding Birds Panel. (2013) Rare breeding birds in the United Kingdom in 2011. British Birds 106, 496–554.
34. BirdLife International. Sylvia undata. The IUCN Red List of Threatened Species. Version 2015.2. 2012. Information correct as of 3rd August 2015. Available from iucnredlist.org.
35. Huntley, B et al. (2007) A climatic atlas of European breeding birds. Lynx Barcelona. 36. Gurney, M. (2015) Gains and losses: extinctions and colonisations in Britain since 1900. Biological
Journal of the Linnean Society 115(3), 573–585. 37. Ausden, M et al. (2014) Managing and re-creating wetlands in Britain for potential colonists. British
Birds 107, 726–755. 38. Lock, L and Cook, K. (1998) The little egret in Britain: a successful colonist. British Birds 91(7),
273–280. 39. Watts, PC, Keat, S and Thompson, DJ. (2010) Patterns of spatial genetic structure and diversity at
the onset of a rapid range expansion: colonisation of the UK by the small red-eyed damselfly Erythromma viridulum. Biological Invasions 12(11), 3887–3903.
40. Krehenwinkel, H and Tautz, D. (2013) Northern range expansion of European populations of the wasp spider Argiope bruennichi is associated with global warming – correlated genetic admixture and population-specific temperature adaptations. Molecular Ecology 22(8), 2232–2248.
41. Lundy, M, Montgomery, I and Russ, J .(2010) Climate change-linked range expansion of Nathusius’ pipistrelle bat, Pipistrellus nathusii (Keyserling & Blasius, 1839). Journal of Biogeography 37(12), 2232–2242.
42. Thomas, CD. (2011) Translocation of species, climate change, and the end of trying to recreate past ecological communities. Trends in Ecology & Evolution 26(5), 216–221.
43. Hoegh-Guldberg, O et al. (2008) Assisted colonization and rapid climate change. Science 321, 345–346.
44. Willis, SG et al. (2009) Assisted colonization in a changing climate: a test-study using two UK butterflies. Conservation Letters 2(1), 45–51.
45. Fordham, DA et al. (2013) Adapted conservation measures are required to save the Iberian lynx in a changing climate. Nature Climate Change 3(10), 899–903.
46. Ricciardi, A and Simberloff, D. (2009) Assisted colonization is not a viable conservation strategy. Trends in Ecology and Evolution 24(5), 248–253.
47. Hiley, JR et al. (2013) Protected areas act as establishment centres for species colonizing the UK. Proceedings of the Royal Society B 280, 20122310.
48. Morecroft, MD et al. (2009) The UK Environmental Change Network: emerging trends in the composition of plant and animal communities and the physical environment. Biological Conservation 142(12), 2814–2832.
49. Britton, AJ et al. (2009) Biodiversity gains and losses: evidence for homogenisation of Scottish alpine vegetation. Biological Conservation 142(8), 1728–1739.
50. Davey, CM et al. (2012) Rise of the generalists: evidence for climate driven homogenization in avian communities. Global Ecology and Biogeography 21(5), 568–578.
51. van Herk, CM, Aptroot, A and van Dobben, HF. (2002) Long-term monitoring in the Netherlands suggests that lichens respond to global warming. The Lichenologist 34(2), 141–154.
52. Beaugrand, G. (2004) The North Sea regime shift: evidence, causes, mechanisms and consequences. Progress in Oceanography 60(2–4), 245–262.
53. Devictor, V et al. (2008) Birds are tracking climate warming, but not fast enough. Proceedings of the Royal Society B 275, 2743–2748.
54. Lindström, Å et al. (2013) Rapid changes in bird community composition at multiple temporal and spatial scales in response to recent climate change. Ecography 36(3), 313–322.
55. Roth, T, Plattner, M and Amrhein, V. (2014) Plants, birds and butterflies: short-term responses of species communities to climate warming vary by taxon and with altitude. PLoS ONE 9(1), e82490.
56. Devictor, V et al. (2012) Differences in the climatic debts of birds and butterflies at a continental scale. Nature Climate Change 2(2), 121–124.
45
57. Gregory, RD et al. (2009) An indicator of the impact of climatic change on European bird populations. PLoS ONE 4(3), e4678.
58. Cahill, AE et al. (2013) How does climate change cause extinction? Proceedings of the Royal Society B 280, 20121890.
59. Thackeray, SJ et al. (2010) Trophic level asynchrony in rates of phenological change for marine, freshwater and terrestrial environments. Global Change Biology 16(12), 3304–3313.
60. Both, C et al. (2009) Climate change and unequal phenological changes across four trophic levels: constraints or adaptations? Journal of Animal Ecology 78(1), 73–83.
61. Both, C et al. (2006) Climate change and population declines in a long-distance migratory bird. Nature 441, 81–83.
62. Visser, ME, Holleman, LJM, and Gienapp, P. (2006) Shifts in caterpillar biomass phenology due to climate change and its impact on the breeding biology of an insectivorous bird. Oecologia 147(1), 164–172.
63. Burthe, S et al. (2012) Phenological trends and trophic mismatch across multiple levels of a North Sea pelagic food web. Marine Ecology Progress Series 454, 119–133.
64. Edwards, M and Richardson, AJ. (2004) Impact of climate change on marine pelagic phenology and trophic mismatch. Nature 430, 881–884.
65. Hindle, BJ et al. (2015) Topographical variation reduces phenological mismatch between a butterfly and its nectar source. Journal of Insect Conservation 19(2), 227–236.
66. Both, C et al. (2009) Avian population consequences of climate change are most severe for long-distance migrants in seasonal habitats. Proceedings of the Royal Society B 271, 1657–1662.
67. Pearce-Higgins, JW et al. (2010) Impacts of climate on prey abundance account for fluctuations in a population of a northern wader at the southern edge of its range. Global Change Biology 16(1), 12–23.
68. Adamík, P and Král, M. (2008) Climate- and resource-driven long-term changes in dormice populations negatively affect hole-nesting songbirds. Journal of Zoology 275(3), 209–215.
69. Selås, V et al. (2011) Climate change in Norway: warm summers limit grouse reproduction. Population Ecology 53(2), 361–371.
70. Durance, I and Ormerod, SJ. (2010) Evidence for the role of climate in the local extinction of a cool-water triclad. Journal of the North American Benthological Society 29(4), 1367–1378.
71. Voigt, W et al. (2003) Trophic levels are differentially sensitive to climate. Ecology 84(9), 2444–2453. 72. Carroll, MJ et al. (2015) Effects of sea temperature and stratification changes on seabird breeding
success. Climate Research 66, 75–89. 73. Frederiksen, M et al. (2004) The role of industrial fisheries and oceanographic change in the decline
of North Sea black-legged kittiwakes. Journal of Applied Ecology 41(6), 1129–1139. 74. Frederiksen, M et al. (2013) Climate, copepods and seabirds in the boreal Northeast Atlantic –
current state and future outlook. Global Change Biology 19(2), 364–372. 75. Scott, BE et al. (2006) The use of biologically meaningful oceanographic indices to separate the
effects of climate and fisheries on seabird breeding success, in Top Predators in Marine Ecosystems: Their Role in Monitoring and Management, Boyd, IL, Wanless, S and Camphuysen, CJ (Editors). Cambridge University Press, Cambridge, UK. 46–62.
76. Beare, DJ et al. (2002) Prevalence of boreal Atlantic, temperate Atlantic and neritic zooplankton in the North Sea between 1958 and 1998 in relation to temperature, salinity, stratification intensity and Atlantic inflow. Journal of Sea Research 48(1), 29–49.
77. Behrens, JW et al. (2009) Oxygen deficiency impacts on burying habitats for lesser sandeel, Ammodytes tobianus, in the inner Danish waters. Canadian Journal of Fisheries and Aquatic Sciences 66(6), 883–895.
78. Wanless, S et al. (2005) Low energy values of fish as a probable cause of a major seabird breeding failure in the North Sea. Marine Ecology Progress Series 294, 1–8.
79. Frederiksen, M et al. (2006) From plankton to top predators: bottom-up control of a marine food web across four trophic levels. Journal of Animal Ecology 75(6), 1259–1268.
80. Coumou, D and Rahmstorf, S. (2012) A decade of weather extremes. Nature Climate Change 2(7), 491–496.
81. Frederiksen, M et al. (2008) The demographic impact of extreme events: stochastic weather drives survival and population dynamics in a long-lived seabird. Journal of Animal Ecology 77(5), 1020–1029.
82. Altwegg, R et al. (2006) Demographic effects of extreme winter weather in the barn owl. Oecologia 149(1), 44–51.
83. Garel, M et al. (2004) The effects of a severe drought on mouflon lamb survival. Proceedings of the Royal Society B 271(Supplement 6), S471–S473.
84. Ewing, SR et al. (2013) Effects of weather variation on a declining population of Slavonian Grebes Podiceps auritus. Journal of Ornithology 154(4), 995–1006.
46
85. van de Pol, M et al. (2010) Do changes in the frequency, magnitude and timing of extreme climatic events threaten the population viability of coastal birds? Journal of Applied Ecology 47(4), 720–730.
86. Abeli, T et al. (2012) Effect of the extreme summer heat waves on isolated populations of two orophitic plants in the north Apennines (Italy). Nordic Journal of Botany 30(1), 109–115.
87. Jiguet, F et al. (2006) Thermal range predicts bird population resilience to extreme high temperatures. Ecology Letters 9(12), 1321–1330.
88. Oliver, TH, Brereton, T and Roy, DB. (2013) Population resilience to an extreme drought is influenced by habitat area and fragmentation in the local landscape. Ecography 36(5), 579–586.
89. Piha, H et al. (2007) Anuran abundance and persistence in agricultural landscapes during a climatic extreme. Global Change Biology 13(1), 300–311.
90. Kindvall, O. (1995) The impact of extreme weather on habitat preference and survival in a metapopulation of the bush-cricket Metrioptera bicolor on Sweden. Biological Conservation 73(1), 51–58.
91. Summers, RW et al. (2004) An experimental study of the effects of predation on the breeding productivity of capercaillie and black grouse. Journal of Applied Ecology 41(3), 513–525.
92. Summers, RW, Dugan, D and Proctor, R. (2010) Numbers and breeding success of Capercaillies Tetrao urogallus and Black Grouse T. tetrix at Abernethy Forest, Scotland. Bird Study 57(4), 437–446.
93. Moss, R. (1986) Rain, breeding success and distribution of Capercaillie Tetrao urogallus and Black Grouse Tetrao tetrix in Scotland. Ibis 128(1), 65–72.
94. Wilson, J and Peach, W. (2006) Impact of an exceptional winter flood on the population dynamics of bearded tits (Panurus biarmicus). Animal Conservation 9(4), 463–473.
95. Leadley, P et al. (2010) Biodiversity scenarios: projections of 21st century change in biodiversity and associated ecosystem services. Montreal, Canada: Secretariat of the Convention on Biological Diversity.
96. Settele, J et al. (2008) Climatic risk atlas of European butterflies. BioRisk 1, 710pp. 97. Rasmont, P et al. (2015) Climatic Risk and Distribution Atlas of European Bumblebees. BioRisk 10,
246pp. 98. Thuiller, W et al. (2005) Climate change threats to plant diversity in Europe. Proceedings of the
National Academy of Sciences of the United States of America 102(23), 8245–8250. 99. Markovic, D et al. (2014) Europe's freshwater biodiversity under climate change: distribution shifts
and conservation needs. Diversity and Distributions 20(9), 1097–1107. 100. Lenoir, S, Beaugrand, G and Lecuyer, É. (2011) Modelled spatial distribution of marine fish and
projected modifications in the North Atlantic Ocean. Global Change Biology 17(1), 115–129. 101. Renwick, AR et al. (2012) Modelling changes in species’ abundance in response to projected climate
change. Diversity and Distributions 18(2), 121–132. 102. Dirnböck, T, Essl, F and Rabitsch, W. (2011) Disproportional risk for habitat loss of high-altitude
endemic species under climate change. Global Change Biology 17(2), 990–996. 103. Rondinini, C and Visconti, P. (2015) Scenarios of large mammal loss in Europe for the 21st century.
Conservation Biology 29(4), 1028–1036.
104. European Environment Agency (2012) Protected areas in Europe an overview: EEA Report No 5/2012. Copenhagen, Denmark.
105. Sanderson, FJ et al. (2015) Assessing the performance of EU nature legislation in protecting target bird species in an era of climate change. Conservation Letters. (Online early.)
106. Virkkala, R et al. (2014) Protected areas alleviate climate change effects on northern bird species of conservation concern. Ecology and Evolution 4(15), 2991–3003.
107. Lawson, CR et al. (2014) Active management of protected areas enhances metapopulation expansion under climate change. Conservation Letters 7(2), 111–118.
108. Gillingham, PK et al. (2015) High Abundances of Species in Protected Areas in Parts of their Geographic Distributions Colonized during a Recent Period of Climatic Change. Conservation Letters 8(2), 97–106.
109. Pavon-Jordan, D et al. (2015) Climate-driven changes in winter abundance of a migratory waterbird in relation to EU protected areas. Diversity and Distributions 21(5), 571–582.
110. Araújo, MB et al. (2011) Climate change threatens European conservation areas. Ecology Letters 14(5), 484–492.
111. Johnston, A et al. (2013) Observed and predicted effects of climate change on species abundance in protected areas. Nature Climate Change 3(12), 1055–1061.
112. Mazaris, AD et al. (2013) Evaluating the connectivity of a protected areas' network under the prism of global change: the efficiency of the European Natura 2000 network for four birds of prey. Plos One 8(3).
113. Thuiller, W et al. (2014) Are different facets of plant diversity well protected against climate and land cover changes? A test study in the French Alps. Ecography 37(12), 1254–1266.
47
114. Araújo, MB et al. (2004) Would climate change drive species out of reserves? An assessment of existing reserve-selection methods. Global Change Biology 10(9), 1618–1626.
115. Hannah, L et al. (2007) Protected area needs in a changing climate. Frontiers in Ecology and the Environment 5(3), 131–138.
116. Gillingham, PK et al. (2015) The effectiveness of protected areas in the conservation of species with changing geographical ranges. Biological Journal of the Linnean Society 115(3), 707–717.
117. Thomas, CD et al. (2012) Protected areas facilitate species’ range expansions. Proceedings of the National Academy of Sciences 109(35), 14063–14068.
118. Ausden, M. (2014) Climate change adaptation: putting principles into practice. Environmental Management 54(4), 685–698.
119. Green, RE and Pearce-Higgins, JW. (2010) Species management in the face of a changing climate., in Species Management: Challenges and Solutions for the 21st Century, Baxter, JM and Galbraith, CA (Editors). HMSO: Edinburgh. 517–536.
120. Pearce-Higgins, JW. (2011) Modelling conservation management options for a southern range-margin population of Golden Plover Pluvialis apricaria vulnerable to climate change. Ibis 153(2), 345–356.
121. Curtis, RJ and Isaac, NJB.(2015) The effect of temperature and habitat quality on abundance of the Glanville fritillary on the Isle of Wight: implications for conservation management in a warming climate. Journal of Insect Conservation 19(2), 217–225.
122. De Frenne, P et al. (2013) Microclimate moderates plant responses to macroclimate warming. Proceedings of the National Academy of Sciences 110(46), 18561–18565.
123. Eglington, SM et al. (2008) Restoration of wet features for breeding waders on lowland grassland. Journal of Applied Ecology 45(1), 305–314.
124. Ratcliffe, N, Schmitt, S and Whiffin, M. (2005) Sink or swim? Viability of a black-tailed godwit population in relation to flooding. Journal of Applied Ecology 42(5), 834–843.
125. Lawson, CR, et al. (2012) Local and landscape management of an expanding range margin under climate change. Journal of Applied Ecology 49(3), 552–561.
126. Newson, SE et al. (2014) Can site and landscape-scale environmental attributes buffer bird populations against weather events? Ecography 37(9), 872–882.
127. Kleijn, D et al. (2010) Adverse effects of agricultural intensification and climate change on breeding habitat quality of Black-tailed Godwits Limosa l. limosa in the Netherlands. Ibis 152(3), 475–486.
128. Boulangeat, I et al. (2014) Anticipating the spatio-temporal response of plant diversity and vegetation structure to climate and land use change in a protected area. Ecography 37(12), 1230–1239.
129. van Dijk, J et al. (2015) Modeling direct and indirect climate change impacts on ecological networks: a case study on breeding habitat of Dutch meadow birds. Landscape Ecology 30(5), 805–816.
130. Carroll, MJ et al. (2011) Maintaining northern peatland ecosystems in a changing climate: effects of soil moisture, drainage and drain blocking on craneflies. Global Change Biology 17(9), 2991–3001.
131. Carroll, MJ et al. (2015) Hydrologically driven ecosystem processes determine the distribution and persistence of ecosystem-specialist predators under climate change. Nature Communications 6, 7851.
132. Pearce-Higgins, JW. (2010) Using diet to assess the sensitivity of northern and upland birds to climate change. Climate Research 45, 119–130.
133. Wilson, L et al. (2011) The impact of drain blocking on an upland blanket bog during storm and drought events, and the importance of sampling-scale. Journal of Hydrology 404(3–4), 198–208.
134. Opdam, P and Wascher, D. (2004) Climate change meets habitat fragmentation: linking landscape and biogeographical scale levels in research and conservation. Biological Conservation 117(3), 285–297.
135. Heller, NE and Zavaleta, ES. (2009) Biodiversity management in the face of climate change: a review of 22 years of recommendations. Biological Conservation 142(1), 14–32.
136. Doerr, VAJ, Barrett, T and Doerr, ED. (2011) Connectivity, dispersal behaviour and conservation under climate change: a response to Hodgson et al. Journal of Applied Ecology 48(1), 143–147.
137. Hodgson, JA et al. (2011) Habitat re-creation strategies for promoting adaptation of species to climate change. Conservation Letters 4(4), 289–297.
138. Hodgson, JA et al. (2011) Habitat area, quality and connectivity: striking the balance for efficient conservation. Journal of Applied Ecology 48(1), 148–152.
139. D’Amen, M et al. (2011) Will climate change reduce the efficacy of protected areas for amphibian conservation in Italy? Biological Conservation 144(3), 989–997.
140. Cianfrani, C et al. (2011) Adapting global conservation strategies to climate change at the European scale: the otter as a flagship species. Biological Conservation 144(8), 2068–2080.
141. Gilbert, G, Brown, AF and Wotton SR. (2010) Current dynamics and predicted vulnerability to sea-level rise of a threatened Bittern Botaurus stellaris population. Ibis 152(3), 580–589.
142. Wotton, S et al. (2009) Boom or bust – a sustainable future for reedbeds and bitterns? British Wildlife 20(5), 305–315.
48
143. Secretariat of the Convention on Biological Diversity (2009) Connecting biodiversity and climate change mitigation and adaptation: report of the second ad hoc technical expert group on biodiversity and climate change: Montreal, Canada. 126 pages.
144. Munroe, R et al. (2012) Review of the evidence base for ecosystem-based approaches for adaptation to climate change. Environmental Evidence 1(1), 13.
145. Campbell, A et al. (2009) The linkages between biodiversity and climate change mitigation. UNEP World Conservation Monitoring Centre.
146. Jones, HP, Hole, DG and Zavaleta, ES. (2012) Harnessing nature to help people adapt to climate change. Nature Climate Change 2(7), 504–509.
147. Doswald, N et al. (2014) Effectiveness of ecosystem-based approaches for adaptation: review of the evidence-base. Climate and Development 6(2), 185–201.
148. Doswald, N and Osti, M. (2011) Ecosystem-based approaches to adaptation and mitigation: good practice examples and lessons learned in Europe. Bundesamt für Naturschutz, Federal Agency for Nature Conservation: Bonn, Germany.
149. Dubgaard, A. (2004) Cost-benefit analysis of wetland restoration. Water and Land Development 8, 87–102.
150. Hulea, O, Ebert, S and Strobel, D. (2009) Floodplain restoration along the Lower Danube: A climate change adaptation case study. IOP Conference Series: Earth and Environmental Science 6(40), 402002.
151. Ebert, S, Hulea, O and Strobel, D. (2009) Floodplain restoration along the lower Danube: A climate change adaptation case study. Climate and Development 1(3), 212–219.
152. Lenane, R. (2012) Keeping rivers cool: getting ready for climate change by creating riparian shade. Environment Agency: Bristol, UK. 41 pages.
153. Gill, SE et al. (2007) Adapting cities for climate change: the role of the green infrastructure. Built Environment 33(1), 115–133.
154. Tanneberger, F and Wichtmann. W (Editors). (2011) Carbon credits from peatland rewetting: climate, biodiversity, land use. Schweizerbart Science Publishers, Stuttgart, Germany.
155. Ausden, M et al. (2015) Wallasea: a wetland designed for the future. British Wildlife 26(6), 382–389. 156. Lawton, JH et al. (2010) Making space for nature: a review of England’s wildlife sites and ecological
network. Report to DEFRA (Department for Environment Food & Rural Affairs).