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ORIGINAL PAPER
Recent evidence for the climate change threat to Lepidoptera
and other insects
Robert J. Wilson •Ilya M. D. Maclean
Received: 19 May 2010 / Accepted: 3 September 2010 / Published online: 19 September 2010
ÓSpringer Science+Business Media B.V. 2010
Abstract Climate change is now estimated by some biol-
ogists to be the main threat to biodiversity, but doubts persist
regarding which species are most at risk, and how best to
adapt conservation management. Insects are expected to be
highly responsive to climate change, because they have short
life cycles which are strongly influenced by temperature.
Insects also constitute the most diverse taxonomic group,
carrying out biotic interactions of importance for ecological
functioning and ecosystem services, so their responses to
climate change are likely to be of considerable wider eco-
logical significance. However, a review of recent published
evidence of observed and modelled effects of climate change
in ten high-ranking journals shows that comparatively few
such studies have focused on insects. The majority of these
studies are on Lepidoptera, because of the existence of
detailed contemporary and historical datasets. These biases
in published information may influence conclusions
regarding the threat of climate change to insect biodiversity.
Assessment of the vulnerability of insect species protected
by the Bern Convention on the Conservation of European
Wildlife and Natural Habitats also emphasises that most
information is available for the Lepidoptera. In the absence
of the necessary data to carry out detailed assessments of the
likely effects of climate change on most threatened insects,
we consider how autecological studies may help to illumi-
nate the potential vulnerability of species, and draw
preliminary conclusions about the priorities for insect con-
servation and research in a changing climate.
Keywords Bioclimate models Conservation
priorities Flagship species Global change Range shifts
Introduction
Insects are expected to show rapid responses to climate
change, because they are ectotherms with short life cycles
and wide variation in population size over space and time
(Bale et al. 2002). The insects are also a highly diverse tax-
onomic group, and their biotic interactions as natural ene-
mies of plants and other animals, decomposers, and
pollinators are critical for ecological functioning and eco-
system services (Ayres and Lombardero 2000; Bradford
et al. 2002; Harrington 2002; Kremen et al. 2007). Therefore
the effects of climate change on insect biodiversity have
considerable applied ecological importance.
Some of the first clear evidence for ecological responses
to climate change was drawn from studies on insects.
Poleward shifts in the distributions of butterflies in Europe
and North America represented clear signals of recent
warming (Parmesan 1996; Parmesan et al. 1999), as did
phenological advances towards earlier dates in the spring
activity of temperate Lepidoptera (Roy and Sparks 2000;
Pen
˜uelas et al. 2002; Stefanescu et al. 2003). Subsequently,
evidence has also been presented for range shifts (Hickling
et al. 2005,2006) and phenological shifts (Gordo and Sanz
2005,2006; Hassall et al. 2007) in other insect groups.
Now that the ecological effects of climate change have
been widely documented (Walther et al. 2002; Parmesan
and Yohe 2003), research has begun to focus more on
R. J. Wilson (&)I. M. D. Maclean
Centre for Ecology and Conservation, University of Exeter,
Cornwall Campus, Penryn TR10 9EZ, UK
e-mail: R.J.Wilson@exeter.ac.uk
I. M. D. Maclean
e-mail: I.M.D.Maclean@exeter.ac.uk
123
J Insect Conserv (2011) 15:259–268
DOI 10.1007/s10841-010-9342-y
climate change as a threat to biodiversity. In the most well-
known study to recognise climate change as perhaps the
major twenty-first Century threat to biodiversity, biocli-
mate models of the distributions of 69 butterfly species in
Mexico, South Africa and Australia contributed to the
estimates of extinction risk from climate change for 1,103
species in total (Thomas et al. 2004). The ecological
importance and potential rapidity of insect responses imply
that much published research since Thomas et al.’s (2004)
paper would have focused on the effects of climate change
on this taxonomic group, and on how to adapt conservation
management in the light of the threat from climate change.
Here we review two sources of evidence for the threat of
climate change to insect biodiversity. First, we conduct a
comprehensive review of research published in ten high-
ranking journals in the past 5 years: (1) to determine whether
studies on insects make an important contribution to high
profile studies of climate change effects; (2) to quantify the
threat of climate change to insects, based on studies of
observed and modelled responses; (3) to show whether there
are taxonomic or regional biases in published research, and
what the implications of any such biases may be for estimates
of the threat of climate change. Second, given a lack of
research into the observed or potential impacts of climate
change on most insect groups, we review a wider range of
literature to determine what information is available con-
cerning the climate change risk to a group of threatened
insects comprising representatives from a range of different
taxa. We use insects listed by the Bern Convention on the
Conservation of European Wildlife and Natural Habitats as
our exemplar group. Where direct information for the effects
of climate change on threatened species is not available, we
consider how autecological data can be used to make pre-
liminary recommendations for conservation or research
needs for insects in a changing climate.
Methods
Evidence of climate change threat for insects
In this review we used the ISI Web of Science to find all
papers published between January 2005 and December 2009
in ten leading journals which included ‘‘climate change’’ in
the title, keywords or abstract. We reviewed journals of
general science (Nature,Proceedings of the National
Academy of Science of the USA,Science), general biology or
ecology (Ecology Letters,Proceedings of the Royal Society
of London B,), applied ecology (Ecological Applications,
Global Change Biology,Journal of Applied Ecology), and
conservation biology (Biological Conservation,Conserva-
tion Biology). We searched all papers with ‘‘climate change’’
and ‘‘biodiversity’’ in the title, abstract or keywords in
Global Change Biology and Proceedings of the National
Academy of Sciences, both of which have a very large
number of studies on climate change. We widened the search
to include any study with just ‘‘climate change’’ in the title,
abstract or keywords for the remainder of journals, leading to
a final total of 1,120 papers reviewed. In our quantitative
analysis, we only included those papers in which (a) IUCN
Red List categories were assigned to taxa; (b) a change in
population size or range were reported; or (c) extinction risk
was estimated; which greatly reduced the number of studies.
We also only included those studies in which the response
could be primarily attributed to climate change or where
climate change impacts were separated from other drivers of
change. We separated papers into those which provided
empirical evidence of recent change (‘‘observed’’) and those
which predicted future change (‘‘predicted’’). Papers could
include both observed and predicted estimates of change, and
a number of papers included measurements for more than
one taxonomic group.
The IUCN Red List criteria provided a means of
assigning each taxon studied to a particular threat category
on the basis of extinction risk or observed or projected
changes in population size over set time periods (see Mace
et al. 2008 for details). In the absence of better evidence we
assumed that a reduction in range size led to a corre-
sponding reduction in population size. We estimated the
proportion of taxa that would be in each IUCN Red List
category in 2100 (Table 3). To make this estimation, it is
necessary to convert the estimates of extinction risk and
population change for the time periods associated with
each study to a standard of 90-year representing the period
2010–2100. We did this by assuming that rates of popu-
lation changes and the frequency of extinction events
remained constant through time. Overall population
declines thus increase with time by an amount determined
by geometric decay, and overall extinction risks increase
with time by an amount determined by multiple-event
probability theory. Annual rates of population decline
(r) were estimated from each study as
r¼ln 1declinet
t
where decline
t
is the decline noted in the study and tis the
time period of the study. Total decline for the period
2010–2100 (90 years) is then given by:
decline90 ¼1expðr90Þ:
The extinction risk (E
90
) for the period 2010–2100 (90
years) can be calculated from the extinction risk (E
t
)
associated with each study over the period (t) of each study
as:
260 J Insect Conserv (2011) 15:259–268
123
E90 ¼11Et
ðÞ
90
t:
Climate change and protected insects
The 1979 Bern Convention on the Conservation of Euro-
pean Wildlife and Natural Habitats is an international legal
agreement which aims to promote European cooperation
for nature conservation. Appendix II of the Convention
includes 56 strictly protected insect species, of which 26
are Lepidoptera (Table 1). We carried out literature sear-
ches in May 2009 using the scientific name of each of these
species in ISI Web of Science, and in Google Scholar to
include a wider range of potential sources.
A recently published ‘‘Climatic Risk Atlas’’ for Euro-
pean butterflies (Settele et al. 2008) includes bioclimate
models of the future climatic ranges for 15 species of
butterfly listed by the Bern Convention. The atlas uses
three different climate change scenarios for the years 2050
and 2080, and models future potential ranges if species are
able to fully occupy future modelled climate space (‘‘Full
Dispersal’’), or only able to occupy those parts of the range
which overlap with their current suitable climate space
(‘‘No dispersal’’). The three scenarios were based on dif-
ferent trajectories of economic growth and environmental
policy, ranging from sustainability (SEDG, IPCC scenario
B1, ?2.4°C by 2080), through a business as usual approach
(BAMBU, IPCC scenario A2, ?3.1°C by 2080) to devel-
opment based on economic growth (GRAS, IPCC scenario
A1F1, ?4.1°C by 2080) (Nakicenovic et al. 2000; Span-
genberg 2007; Settele et al. 2008). We use the published
results of these models for our main quantitative data in
this part of the review, asking two questions for the mod-
elled effects of climate change on butterflies listed by the
Bern Convention: (1) what are the differences in modelled
future range size under different scenarios of climate
warming (?2.4°C, ?3.1°C, ?4.1°C)? and (2) what are the
differences in range size for full dispersal versus no dis-
persal? The two questions are relevant to determine the
importance for conserving threatened taxa by (1) mitigat-
ing levels of climate change, and (2) by promoting range
shifts by species into regions which become climatically
favourable.
We encountered very few direct estimates of climate
change effects on Bern Convention insects other than
Lepidoptera, so our Discussion relating to this part of the
review focuses on the evidence for sensitivity of the listed
species to climate change, based on their ecological char-
acteristics (e.g. dispersal propensity, range size, habitat
specialism), and uses this information to make recommen-
dations for research or conservation priorities for insects in
a changing climate.
Results
Evidence of climate change threat for insects
73 papers in the review included evidence of changes to
extinction risk linked to climate change: 39 papers includ-
ing observed changes, and 47 including predicted changes.
Only nine of these papers dealt with insects (4 observed, 5
predicted), of which eight included evidence for the Lepi-
doptera (Table 2). All papers linking climate change to
observed changes in extinction risk (usually range or pop-
ulation size) presented evidence for Lepidoptera (Table 2).
Projecting observed or predicted changes to extinction risk
over time did not lead to any estimates of decline over the
period 2010–2100 which would classify the study taxa as
Endangered or Critically Endangered, but 10% of taxa
Table 1 Insects listed as strictly protected by Appendix II of the
Bern Convention (Council of Europe 2010)
Orthoptera Ephemeroptera
Baetica ustulata Palingenia longicauda
Saga pedo Mantodea
Odonata Apteromantis aptera
Aeshna viridis Lepidoptera
Brachythemis fuscopalliata Apatura metis
Calopteryx syriaca Coenonympha hero
Coenagrion freyi Coenonympha oedippus
Coenagrion mercuriale Erebia calcaria
Cordulegaster trinacriae Erebia christi
Gomphus graslinii Erebia sudetica
Leucorrhinia albifrons Eriogaster catax
Leucorrhinia caudalis Euphydryas (Eurodryas) aurinia
Leucorrhinia pectoralis Fabriciana elisa
Lindenia tetraphylla Hyles hippophaes
Macromia splendens Hypodryas maturna
Ophiogomphus cecilia Lopinga achine
Oxygastra curtisii Lycaena dispar
Stylurus (=Gomphus) flavipes Maculinea arion
Sympecma braueri Maculinea nausithous
Coleoptera Maculinea teleius
Buprestis splendens Melanargia arge
Carabus bessarabicus Papilio alexanor
Carabus hungaricus Papilio hospiton
Carabus olympiae Parnassius apollo
Cerambyx cerdo Parnassius mnemosyne
Cucujus cinnaberinus Plebicula golgus
Dytiscus latissimus Polyommatus galloi
Graphoderus bilineatus Polyommatus humedasae
Osmoderma eremita Proserpinus prosperpina
Rosalia alpina Zerynthia polyxena
J Insect Conserv (2011) 15:259–268 261
123
would be classified as Vulnerable based on observed
changes (9% based on predicted changes) (Table 3).
Despite the small sample of studies of the climate
change threat to insects, there were examples from a rel-
atively wide range of geographical locations (Table 2). The
nine papers included three from northern Europe (Britain
and the Netherlands), one from north America (Utah), two
from mountain regions in the Mediterranean (Spain and
Egypt), and three from the Tropics (Borneo, Ecuador,
Costa Rica), again with a focus on elevational gradients.
Climate change and protected insects
Nine of the 15 butterfly species whose climate niches were
modelled by Settele et al. (2008) were categorised as
having High, Very High, or Extremely High climate
change risk, indicating that more than 70% of their current
European distributions are predicted to be climatically
unfavourable by 2080 in at least one climate change sce-
nario (Table 4; Fig. 1). Four of the remaining species were
categorised as experiencing some climate change risk,
because more than 50% of currently suitable cells were
predicted to be unsuitable by 2080 in one or more scenario.
Only one species (the scarce fritillary Hypodryas maturna)
did not show modelled losses of [50% in any 2080 sce-
nario (Lower Climate Change Risk), while the current
distribution of marsh fritillary (Euphydryas aurinia) was
only related to climate conditions to a limited extent
(AUC \0.75).
Increases in temperature by 2.4°C (SEDG scenario),
3.1°C (BAMBU scenario) or 4.1°C (GRAS scenario) pro-
duced clear differences in predicted distribution sizes of
Bern Convention butterfly species. In both 2050 and 2080
the highest warming scenario (GRAS) gave consistently
lower modelled distribution sizes than both other scenarios.
In 2080, the intermediate scenario (BAMBU) produced
consistently reduced distribution sizes relative to the most
conservative scenario (SEDG) (Fig. 1).
Future modelled climate space for all species included
areas which were not modelled as suitable for the species in
2000. Hence the Full Dispersal scenario of distribution
change always led to a less severe modelled decline in
Table 2 Papers in ten leading
journals between 2005 and 2009
which linked climate change to
observed or predicted changes
in distribution or population size
of insects
Reference Taxa Location
Observed
Chen et al. (2009) Lepidoptera (Geometridae ?Others) Borneo
Davies et al. (2005) Lepidoptera (Hesperia comma) England
WallisDeVries and Van Swaay
(2006)
Lepidoptera (Rhopalocera) Netherlands
Wilson et al. (2005) Lepidoptera (Rhopalocera) Spain
Predicted
Colwell et al. (2008) Lepidoptera (Geometridae) ?Hymenoptera (Formicidae) Costa Rica
Dangles et al. 2008 Lepidoptera (Gelechiidae) Ecuador
Goulson et al. (2005) Diptera (Calliphoridae, Muscidae) England
Hoyle and James (2005) Lepidoptera (Pseudophilotes sinaicus) Egypt
Logan et al. (2007) Lepidoptera (Lymantria dispar) Utah (USA)
Table 3 IUCN criteria for population decline and extinction risk
estimated for a period of 90 years (2010–2100) (see ‘‘Methods’’), and
the percentage of cases for observed and predicted effects of climate
change on insect taxa which classify as each of the three threatened
classes (CR, EN and VU)
Critically Endangered Endangered Vulnerable
Population decline
IUCN criteria/10 years 80% 50% 10%
Equivalent/90 years 99.99% 99.80% 95.96%
Extinction risk
IUCN criteria 50%/10 years 20%/20 years 10%/100 years
Equivalent/90 years 99.80% 63.36% 9.05%
% observed 0 0 10
% predicted 0 0 9
Data sources are shown in Table 2
262 J Insect Conserv (2011) 15:259–268
123
potential distribution size than the No Dispersal scenario;
and in seven out of 15 cases, the Full Dispersal scenario led
on average (across the three climate change scenarios) to
modelled increases in the size of the climate niche (Fig. 1;
Table 4).
Discussion
Our review suggests that relatively few papers published in
ten high-ranking journals in the past 5 years used insect
systems to show evidence of the extinction threat from
climate change (9 out of 73 papers showing a threat to
biodiversity, or 12%). A further 253 published papers in
the review related climate change to ecological change, but
did not provide data which satisfied the criteria for inclu-
sion in the review: 43 (17%) of these cases dealt with
insects. The relatively low proportion of insect-based
papers (relative to insect diversity) may reflect a wider
tendency of insect research to be somewhat marginalised
(Leather 2010). A wide range of journals now publish
research in ecology, conservation and global change, as is
evident from the reference list of the current article, and
inclusion of these journals would have increased the sam-
ple size. Nevertheless, journals focusing more specifically
on entomology or insect conservation are likely to play a
key role in presenting the evidence for climate change
effects on insects.
The few papers identified by the review illustrated some
important points about evidence for the threat of climate
change to insects. Evidence of observed change appeared to
support estimates of threat from predictions of the effects of
climate change: projecting the changes from the published
papers over the period 2010–2100 predicted that no taxa
would be categorised as Critically Endangered or Endan-
gered, but that 10% would be classified as Vulnerable based
on observed changes, or 9% based on predicted changes
(Table 3). Lepidoptera played a key role in providing evi-
dence of change, as would be expected, given the greater
availability of current and historical information on species
distributions and ecological requirements than for other taxa.
A number of studies were conducted at temperate latitudes
and on pest species (e.g. Goulson et al. 2005; Logan et al.
2007). The distributions of many ectothermic species
including insects are limited by low temperatures at high
latitudes, and so are expected (and observed) to expand their
distributions in northern Europe because of climate change
(e.g. Hickling et al. 2006), and widely dispersing generalists
like many pests may also be able to expand their distributions
relatively rapidly compared with more sedentary or habitat
Table 4 Modelled changes by 2080 to the climatically favourable range area in Europe for Bern Convention species of butterfly (after Settele
et al. 2008), averaged for three scenarios of climate change
Model accuracy (AUC) Climate risk category
?
Mean 2080 modelled change (%)*
Full dispersal No dispersal
Apatura metis 0.98 HHR ?47 -71
Coenonympha hero 0.88 HHR -44 -76
Coenonympha oedippus 0.95 R ?161 -50
Euphydryas aurinia 0.72 PR -24 -41
Hypodryas maturna 0.79 LR ?46 -37
Lopinga achine 0.81 R ?58 -43
Lycaena dispar 0.88 R ?54 -41
Maculinea arion 0.77 R -17 -52
Maculinea nausithous 0.91 HHHR -45 -84
Maculinea teleius 0.84 HHR -26 -74
Melanargia arge 0.98 HR ?119 -64
Papilio alexanor 0.94 HHR -53 -77
Parnassius apollo 0.80 HR -51 -66
Parnassius mnemosyne 0.77 HR -18 -65
Zerynthia polyxena 0.85 HR ?80 -59
Results for % change are based either on ‘‘Full dispersal’’ (entire area in 2080) or ‘‘No dispersal’’ (area of overlap between area in 2000 and in
2080)
* Mean modelled change is shown as the mean of the three climate change scenarios (SEDG, BAMBU, GRAS; see ‘‘Methods’’). Differences
among scenarios are shown in Fig 1
?
Climate Risk Category based on maximum decline: [95% Extremely High Risk (HHHR), [85% Very High Risk (HHR), [70% High Risk
(HR), [50% Risk (R), \50% Lower Risk (LR); AUC \0.75 Potential Risk (PR) (Settele et al. 2008)
J Insect Conserv (2011) 15:259–268 263
123
specialist species (Warren et al. 2001;Po
¨yry et al. 2009).
Hence a preponderance of studies of insect responses to
climate change at relatively high latitudes could lead to
underestimates of the threat of climate change (but see
Franco et al. 2006). Where studies have been carried out at
lower latitudes, such as in Mediterranean mountains, species
distributions have been observed to contract (Wilson et al.
2005) or predicted to do so (Hoyle and James 2005). Clearly
evidence from lower latitudes is vital for identifying the real
risk that climate change will erode species distributions, or
whether species are able to adapt to local or regional changes
in climate. Encouragingly, a few high profile studies have
begun to gather evidence of climate change effects on trop-
ical insects (e.g. Chen et al. 2009). In the Tropics, where
latitudinal gradients in climate are negligible, elevational
range shifts may play a major role in allowing species to track
the availability of suitable climatic conditions, and work
based to a large extent on insects has been important in
illustrating this point (Colwell et al. 2008).
Quantitative information on expected effects of climate
change on protected insects (those listed by the Bern
Convention) is also most forthcoming for the Lepidoptera
(Settele et al. 2008), and highlights two important points.
First, more marked increases in temperature lead to more
pronounced declines in the area of modelled climate space
for threatened species. Second, if species are able to colo-
nize areas which become climatically favourable, they are
likely to experience less drastic declines in distribution area
than if they are limited to future favourable areas which
overlap with their current distributions. Since most Lepi-
doptera listed by the Bern Convention have highly specialist
habitat requirements and therefore fragmented distributions
of habitat, it is unlikely that most will be able to colonize
regions that become climatically favourable (Warren et al.
2001; Mene
´ndez et al. 2006). Hence estimates of future
distribution sizes based on a No Dispersal scenario may be
more realistic for threatened species than those assuming
Full Dispersal, unless serious measures are put in place to
promote range shifts by species. In addition, the current and
future distributions of specific larval host plants will impose
a further constraint on the distribution of many herbivorous
insects (Arau
´jo and Luoto 2007; Schweiger et al. 2008).
Percentage change in area of modelled climate niche
(a) 2050
-100
-50
0
50
100
150
200
250
(b) 2080
-100
-50
0
50
100
150
200
250
Species:
Apatura metis
Coenonympha hero
Coenonympha oedippus
Euphydryas aurinia
Hypodryas maturna
Lopinga achine
Lycaena dispar
Maculinea arion
Maculinea nausithous
Maculinea teleius
Melanargia arge
Papilio alexanor
Parnassius apollo
Parnassius mnemosyne
Zerynthia polyxena
Fig. 1 Modelled changes to the
European climate niche space
for butterflies listed by the Bern
Convention (adapted from
Settele et al. 2008). Data show
percentage change in modelled
niche space between 2000 and
a2050, or b2080, assuming
either full dispersal (circles)or
no dispersal (triangles), and are
shown for three climate change
scenarios, for 2.4°C increase
(SEDG, white symbols), 3.1°C
increase (BAMBU, grey), and
4.1°C increase (GRAS, black)
264 J Insect Conserv (2011) 15:259–268
123
Protection and active management of current habitat and
populations represent vital parts of conservation pro-
grammes designed to facilitate species range shifts, being
necessary for the survival of species’ current populations
and the maintenance of future adaptive capacity. Habitat
fragmentation is likely to constrain range expansions by
species in many parts of the world, even when climate
conditions become more favourable (Wilson et al. 2009),
so the management of networks of habitat, or of landscapes
in which favourable habitats are maintained, is necessary
for species to be able to expand their distributions. Con-
servation interventions in the form of assisted colonization
may become an appropriate technique for those species
which are likely to suffer the most severe reductions in
distribution size, and which have the least chance of
reaching locations which become climatically suitable
(Hoegh-Guldberg et al. 2008). Insects have a combination
of characteristics (e.g. small size, relatively small area
requirements, relatively large population sizes) which may
make them appropriate for such techniques, and it is known
that introductions of insects to climatically favourable
regions may be successful (Mene
´ndez et al. 2006; Willis
et al. 2009). However, in situ conservation of extant pop-
ulations should remain the bedrock of species conserva-
tion; and habitat management based on sound ecological
science at introduction sites is crucial for the success of
translocations, such as in the case of the reintroduction of
Large Blue Maculinea arion to the UK (Thomas et al.
2009).
Research and conservation priorities for threatened
Lepidoptera
Many Lepidoptera listed by the Bern Convention possess
species traits which imply high climate change sensitivity,
or reduced capacity for adaptation. All eight butterfly
species listed by the Bern Convention but not modelled by
Settele et al. (2008) are European endemics, generally with
extremely restricted distributions to islands or mountain
ranges. Given their often small distributions or population
sizes, and their isolation to narrow suitable regions, these
species have very little opportunity for latitudinal range
shifts. For mountain-dwelling species, there may be some
limited scope for elevational range shifts, or for survival
under localised climate conditions provided by topographic
heterogeneity (Loarie et al. 2009). Research into the habitat
requirements and dispersal capacity of such species can be
extremely valuable in identifying the factors limiting spe-
cies distributions and their capacity to colonize locations
which become suitable. For example, movement patterns
have been quantified for Erebia sudetica in the Czech
Republic (Kuras et al. 2003), and habitat requirements for
Erebia calcaria have been modelled in Slovenia (de Groot
et al. 2009). For several protected butterfly species,
research into dispersal capacity suggests that distances of
10–50 km between habitat areas would almost certainly
prevent natural colonizations (for Parnassius apollo see
Brommer and Fred 1999; for Parnassius mnemosyne see
Meglecz et al. 1999; Valimaki and Itamies 2003; for
Coenonympha hero see Cassel-Lundhagen and Sjogren-
Gulve 2007,2008), again emphasising the importance of
managing whole landscapes or habitat networks.
Research showing how distribution patterns can be
predicted by a combination of the effects of climatic con-
ditions and landcover can be helpful in predicting the
locations and habitats which will be suitable for species in
the future, but also in targeting sites for management or
species searches in the present day. This approach has been
successfully applied for Parnassius mnemosyne (Heikkinen
et al. 2005,2007; Luoto et al. 2007) and Maculinea nau-
sithous (Jime
´nez-Valverde et al. 2008). At a finer scale,
understanding how climate conditions influence resource
availability for threatened species is vital to manage habitat
successfully in a changing climate. The host ants of Mac-
ulinea species have fine-scale distributions which depend
on local microclimate, and the habitat structure which
provides suitable microclimates varies depending on pre-
vailing regional climates (Thomas et al. 1998; Mouquet
et al. 2005). For the reintroduction of Maculinea arion to
the UK, management of sward structure in order to provide
suitable microclimates for the larval host ants (Myrmica
sabuleti) was an essential step (Thomas et al. 2009).
Continued appropriate management has allowed reintro-
duced populations to survive in the UK and colonize
additional suitable habitat nearby, in an exemplary case of
how detailed ecological knowledge and habitat manage-
ment can result in successful landscape-scale conservation.
Monitoring of habitat use and population responses to
habitat management are vital because changes to climate
conditions may mean that there are changes to the micro-
habitat types which satisfy the microclimate requirements
of species. Larvae in both Parnassius apollo and P. mne-
mosyne appear to thermoregulate by moving between areas
of litter, bare ground and more shaded vegetation (Vali-
maki and Itamies 2005; Ashton et al. 2009). In the case of
P. apollo, populations at lower elevations in central Spain
are associated with shrubbier, less-open conditions than
populations at higher elevations, implying that habitat
management can be adapted to local climates (Ashton et al.
2009). This mountain species has already suffered local
extinctions linked to climate warming in France (Descimon
et al. 2006) and Spain (Wilson et al. 2005; Ashton et al.
2009), with the low-elevation limits of the species shifting
uphill in both regions. ‘‘False-spring events’’ in France
appear to result in the early emergence of P. apollo larvae
from winter diapause, leading to starvation when
J Insect Conserv (2011) 15:259–268 265
123
subsequent conditions are too cold for feeding activity
(Descimon et al. 2006). The very small populations of
P. apollo which remain in some parts of its distribution
may have led to inbreeding depression which renders the
species less able to adapt to changing conditions, although
reintroductions for this species in Poland have had some
measure of success (Adamski and Witkowski 2007).
Priorities for other threatened Insecta
In common with the butterflies listed by the Bern Con-
vention, many narrow range European endemic species
from other taxa are likely to have very limited scope for
range expansion to future climatically favourable regions.
For example, Carabus olympiae (Coleoptera) is known to
have a restricted habitat distribution in the Italian alps, and
increased conversion of beech woodland or scrub to pasture
are likely to prevent the species from colonizing locations
outside its current narrow distribution (Negro et al. 2007,
2008). There are also a few studies which point to potential
climate effects on habitat associations. Saga pedo
(Orthoptera) is known to be restricted to the warm micro-
climates of steep south-west facing slopes at its northern
range limit (Kristin and Kanuch 2007), as is the case for
many butterfly species (Thomas 1993). Cerambyx cerdo
(Coleoptera) requires tree trunks receiving high levels of
solar insolation in central Europe, so semi-open pasture
landscapes favour the survival of this saproxylic beetle
(Buse et al. 2007,2008). If the habitat requirements of
many insect species vary across their latitudinal or eleva-
tional ranges, then future habitat management may need to
accommodate changes to habitat management to maintain
suitable microclimates (Davies et al. 2006). A safety-first
approach to management in the mean time would be to
maintain heterogeneous conditions, which have been
shown to stabilise insect population dynamics (Kindvall
1996; Sutcliffe et al. 1997; Oliver et al. 2010).
There is strong autecological evidence that the beetle
Osmoderma eremita will be affected by climate change.
This species is sensitive to increases in air temperature
(Renault et al. 2005). It also has extremely limited dispersal
(Hedin et al. 2008), so that persistence within woodlands is
dependent on aggregations of old growth oak trees, and in
fragmented habitats the species rarely moves between
woodlands (Ranius 2000,2007). Conservation of highly
sedentary, habitat specialist species which are sensitive to
climate represents a significant challenge under climate
change, but the information on habitat requirements such as
exists for O. eremita is a vital first step.
Species distribution models which incorporate the effects
of climate and landcover may be a greater challenge for
non-lepidopteran insect groups because less information on
species distributions and habitat associations are generally
available. However, where distribution data do exist, their
use in developing such models can play a key role in
identifying regions for targeted species searches, which can
help to determine the conservation status of species, as well
as strengthening evidence for the modelled effects of cli-
mate or habitat management. The Bern-listed Iberian
endemic spider Macrothele calpeiana provides an excellent
example of these points. The species’ distribution appears
to be determined mainly by climate variables, occurring in
regions with high precipitation and high precipitation
periodicity, but absent from regions where temperatures
reach extremes (Jime
´nez-Valverde and Lobo 2006). A
distribution model developed for M. calpeiana proved
accurate at identifying a region of southern Portugal in
which populations of the species occurred but had not
previously been found (Jime
´nez-Valverde et al. 2007). The
models also suggest that the species may require forest
cover in order for populations to persist, or shift their range
as the climate warms, hence identifying the key role that
habitat management may play in allowing the adaptation of
conservation management to climate change.
Conclusions
Strong evidence has been available for more than a decade
that the distributions and phenology of insects have chan-
ged in response to climate change (Mene
´ndez 2007; Wil-
son et al. 2007). Examples showing that the population or
distribution sizes of insects have declined are generally
more recent, and generally refer to Lepidoptera (Wilson
et al. 2005; Franco et al. 2006; Chen et al. 2009; but see
Parmesan 1996,2006). In a review of ten high-ranking
journals, insect examples represented only 12% of pub-
lished cases directly relating a changed extinction risk to
climate change. However, in these examples, predicted
estimates of threat corresponded closely to those shown by
empirical evidence for observed change. It is therefore
appropriate for conservation biologists to consider climate
change as an important threat to insects, and accordingly to
include climate change adaptation measures in conserva-
tion programmes.
In a review of the climate change threat to an exemplar
group of threatened insects (those listed by the Bern
Convention) we only found direct predictions of threat for
the butterflies (Settele et al. 2008), but autecological
information for a number of other taxa suggested a strong
sensitivity to climate change. Evidence for the influence of
climate on habitat associations and population—and
metapopulation-dynamics will be increasingly valuable for
conserving these and other threatened insects as the climate
changes.
266 J Insect Conserv (2011) 15:259–268
123
Acknowledgments We were funded by the European Social Fund
(Project ‘‘Climate Change and Landscape: Imagining the Future’’)
and by the Council of Europe. RJW would like to thank The Bern
Convention’s Group of Experts on Biodiversity and Climate Change
for assistance and discussion, particularly Deborah Procter, Vernon
Heywood, Carolina Lase
´nDı
´az and Ve
´ronique du Cussac.
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