CLIMATIC RISK ATLAS
OF EUROPEAN BUTTERFLIES
JOsEF SETTELE, OTAKAR KUDRNA, ALEXANDER HARPKE,
INGOLF KUHN, CHRIS VAN SWAAY, RUDI VEROVNIK, MARTIN WARREN,
MArTIN WIEMERS, JAN FH1ANSPACH, THOMAS HICKLER, ELISABETH KUHN,
INGE VAN HALDER, KARS VELING, ALBERT VLIEGENTHART,
IRMA WYNHOFF & OLIVER SCHWEIGER
Climatic Risk Atlas
of European Butterflies
by
JOSEF SETTELE, OTAKAR KuDRNA, ALEXANDER HARPKE,
INGOLF KUHN, CHRIS VAN SWAAY, RUDI VEROVNIK, MARTIN WARREN,
MARTIN WIEMERS, JAN HANSPACH, THOMAS HICKLER,
ELISABETH KUHN, INGE VAN HALDER, Kars VELING, ALBERT
VLIEGENTHART, IRMA WYNHOFF & OLIVER SCHWEIGER
Biorisk 1 (Special Issue)
> PENSOFT.
SOFIA—Moscow
2008
CLIMATIC Risk ATLAS OF EUROPEAN BUTTERFLIES
Josef Settele', Otakar Kudrna’, Alexander Harpke', Ingolf Kithn', Chris van Swaay’, Rudi Verovnik’,
Martin Warren’, Martin Wiemers®, Jan Hanspach', Thomas Hickler’, Elisabeth Kithn', Inge van
Halder’, Kars Veling’, Albert Vhegenthart’, Irma Wynhoff’? ¢ Oliver Schweiger'
1 UFZ, Helmholtz Centre for Environmental Research, Department of Community Ecology,
Theodor-Lieser-Str. 4, D-06120 Halle, Germany 2 Naturmuseum Sudtirol, Bindergasse 1,
1-39100 Bozen (Siidtirol), Italy 3 De Vlinderstichting — Dutch Butterfly Conservation, P.O. Box
506, 6700 AM Wageningen, The Netherlands 4 DruStvo za proucevanje in ohranjanje metuljev
Slovenije (DPOMS) , Stara Deckova cesta 14, 3000 Celje, Slovenia 5 Butterfly Conservation,
Manor Yard, East Lulworth, Wareham, Dorset BH20 5QP, United Kingdom 6 Department of
Population Ecology, Faculty of Life Sciences, University of Vienna, Rennweg 14, A-1030 Wien,
Austria 7 Department of Physical Geography & Ecosystems Analysis, University of Lund,
Sélvegatan 13, 223 62 Lund, Sweden
Biorisk 1 (Special Issue)
ISSN 1313-2652 (online), ISSN 1313-2644 (print)
dot: 10.3897 /biorisk.1
First published 2008
ISBN 978-954-642-454-9 (paperback)
ISBN 978-954-642-455-6 (HB)
ISBN 978-954-642-456-3 (e-book)
This is an open access book distributed under the terms of the Creative Commons Attribution
License, which permits unrestricted use, distribution, and reproduction in any medium, provided the
original author and source are credited.
Cover and book design: Zheko Akksiev
Pensoft Publishers
Geo Milev Str. 13a, Sofia 1111, Bulgaria
info@pensoft.net
www.pensoft.net
Printed in Bulgaria, November 2008
Climatic Risk Atlas of European Butterflies | 5
CONTENTS
Acknowledgements 7
Foreword by Butterfly Conservation Europe 8
Context and objectives of a climatic risk atlas of European butterflies 10
A. CLIMATE CHANGE, BIODIVERSITY,
BUTTERFLIES, AND RISK ASSESSMENT
A.1 Introduction: Butterflies as indicators 12
A.2 Scenarios and biodiversity 13
A.3 Climate change and Biodiversity Risk Assessment for butterflies 14
B. METHODOLOGY
B.1 The MEB project data as basis for the atlas 18
B.2 Scenarios used to assess climate change risks for European Butterflies 18
B.3 Climate niche modelling 20
Climatic factors of butterfly distribution 20
Modelling procedure 21
Assumptions for species dispersal 24
Visualisation of the multi-dimensional climatic niche 24
B.4 Climate change risk assessment for butterflies 24
Definitions of climate change risk categories for European butterflies 24
Integrated overall risk categories for European species —
integrating all scenarios and time steps 25
C. CLIMATE RISKS OF EUROPEAN BUTTERFLY SPECIES
C.1 Species inventory and taxonomy of European butterflies 28
C.2 Climatic fate of individual species 31
Hespertidae 32
Papilionidae 104
Pieridae 122
Lycaenidae 186
Riodinidae 340
Libytheidae 342
Nymphalidae 344
Danaidae 618
6 | Climatic Risk Atlas of European Butterflies
C.3 Non-modelled European butterfly species 620
C.4 Summary results 629
D. DISCUSSION OF METHODOLOGICAL LIMITATIONS
D.1 Limitations of species distribution models and future projections 634
D.2 Interaction of taxonomic status and modelling results 645
E. OUTLOOK: CLIMATE CHANGE AND BUTTERFLY
CONSERVATION
E.1 Direct and indirect climate change impacts on butterflies and biodiversity 652
Habitats and trophic interactions 652
Climate envelopes for European butterflies as a starting point for future
reseatch and conservation 653
Climate change and evolution 654
Biodiversity Risk Assessment 655
F..2 Butterflies as indicators of environmental change 656
Butterfly indicator developments 656
Butterflies as climate change indicators 657
E..3 Climate change and butterfly conservation 658
FE. APPENDICES, REFERENCES AND INDEX
Appendix 1: ‘Table on parameters for model suitability assessment 662
Appendix 2: ‘Table on scenario results: Changes in climatic niches 674
Appendix 3: Risk category statistics 696
References 698
Species index 706
Climatic Risk Atlas of European Butterflies | 7
Acknowledgements
For the provision of pictures of species (or the readiness to provide them), we are
indebted to Bernard Fransen, Eddie John, Heiner Ziegler, Helmut Hottinger, Henk
Bosma, Hermann Faas, Jean Delacre, Joe Belicek, Josef Pennerstorfer, Jostein Engdal,
Karl Heyde, Matt Rowlings, Michel Tarrier, Neil Thompson, Peter Ginzinger, Philippe
Mothiron, Thomas Kissling, and Zdravko Kolev.
For diverse discussions of modelling methodology we thank Risto Heikkinen and
for provision of basic species texts thanks go to Claire Hengeveld. For help in the
scenario chapters we acknowledge Joachim H. Spangenberg and Martin T. Sykes.
Zsolt Balint has contributed through the discussion of taxonomic questions.
We acknowledge the support of
¢ Helmholtz Centre for Environmental Research - UFZ
¢ MEB Mapping European Butterflies Project —in particular the many colleagues
who provided distribution data;
¢ European Commission (FP 6) Scientific Support to Policy project MACIS
(Minimization of and Adaptation to Climate change impacts on biodiverSity;
044399 (SSPI); Kuhn et al. 2008c);
¢ European Commission (FP 6) Integrated Project ALARM (Assessing LArge
scale environmental Risks with tested Methods; GOCE-CT-2003-506675;
www.alarmproject.net; Settele et al. 2005);
° Gesellschaft fiir Schmetterlingsschutz (GfS; www.curopean-butterflies.eu);
¢ Butterfly Conservation Europe (BCE; www.bc-europe.cu);
¢ Helmholtz Association of German Research Centres funding the Virtual
Institute for Macroecology (VH-VI-153; www.macroecology.org; Kuhn et al.
2008b);
¢ EEA - European Environment Agency (www.eea.cutopa.cu).
Special thanks are due to Pensoft Publishers (especially Teodor Georgiev and
Lyubomir Penev), who managed to design, layout, proofread and publish this book in
less than six weeks time.
8 | Climatic Risk Atlas of European Butterflies
Foreword by
Butterfly Conservation Europe
“
Butterfly \
CONSERVATION EUROPE
Climate change 1s a new and potent risk to biodiversity. The inevitable rise in global
temperatures predicted over the next decades and century 1s a serious threat to butterfly
and moth populations and is likely to exacerbate their decline.
Butterflies are a well-known and popular group of insects that can play a valuable
role as early warning indicators of environmental change. They have short life-cycles
and respond rapidly to change. Butterflies and moths have declined rapidly in recent
decades and are declining more rapidly than other well-known groups such as plants
and birds (which often depend on their caterpillars for food).
This Atlas is an early attempt to investigate the possible effects of climate change
on butterflies by modelling the impact of various future climate scenarios. The research
is based on a unique and comprehensive dataset on butterfly distributions in Europe,
derived from the Mapping European Butterflies (MEB) project. We believe that the
results are important because butterflies are one of the few groups of insects for
which such comprehensive data are available at a European level. As insects comprise
over two-thirds of all known species, the results are valuable to help understand the
possible impacts of climate change on biodiversity as a whole.
The results are alarming. The models suggest that the vast majority of European
butterflies will be badly affected by climate change. Most species will have to shift their
distributions considerably northwards and will lose a large amount of their suitable
climate space. Furthermore, many butterflies live in discrete colonies and have limited
powers of dispersal. Such species are restricted by available habitat and may not be
able to alter their distribution to keep in step with climate change.
The results have important implications for conservation and for EU and national
policies and for their funding and implementation. We have the chance to mitigate
some of the worst effects of climate change, including those on biodiversity, if we act
now. Specifically, we need:
¢ O02 000000000
soceecco0e
Cc0cceg
oo ee
1SOGS0000080 ©
@00000 op0c000
.@edecees
‘Geoo000e
©00000000
TITTY Tttty +
= S00000e eocccce
| C@8@0CCCCC00000
19 G6 © 0000000000066
eeooooqo0oo0o0
6006 S0000000
:
de
Fig. B.1.1. Geographical coverage and distribution of reference localities aggregated to 50 x
50 km? UTM grid (derived from the database which also was used for Kudrna, 2002).
The main source for future climate scenarios was a coupled Atmosphere-Ocean
General Circulation Model (HadCM3; New et al. 2000). The complete ALARM
explained in chapter A.2 cover a broad range of potential developments
scenarios as
in demography, socio-economics and technology during the 21* century. Specifically
for climate the following frame conditions apply in addition to the more general aspect
mentioned above:
20 Climatic Risk Atlas of European Butterflies
e SEDG (Sustainable Europe Development Goal) — a storyline for moderate
change:
The scenario of moderate change approximates the IPCC B1 climate change
scenario. Mean expected temperature increase in Europe until 2080 is 2.4°C.
e BAMBU (Business As Might Be Usual) —a storyline for intermediate change:
The scenario of intermediate change approximates the IPCC A2 climate change
scenario. Mean expected increase in temperature 1s 3.1°C.
e GRAS (GRowth Applied Strategy) — a storyline for maximum change:
The scenario of maximum change approximates the [PCC A1FI climate change
scenario. Mean expected increase in temperature is 4.1°C.
Based on the storylines, projections of future changes in climate were developed
on a 10 x 10 min grid of Europe. Monthly projected climate data (see chapter B.3)
were averaged for the two periods 2021-2050 and 2051-2080.
B.3 Climate niche modeling
Climatic factors of butterfly distribution
The climatic requirements of butterflies were modelled using monthly interpolated
climate data at the same 50 x 50 km? UTM grid (New et al. 2000, Mitchell et al. 2004)
that was used to present the distribution of the species (see chapter B.1). Mean values of
the following 22 climate variables (absolute values and annual variations) for the period
1971-2000 were considered for the analysis of climate requirements of the butterflies:
e annual temperature (°C);
range in annual temperature (°C);
quarterly temperature (e.g. March - May = spring; °C);
range in quarterly temperature (°C);
diurnal temperature range per year (°C);
diurnal temperature range per quarter (°C);
annual summed precipitation (mm);
range in annual precipitation (mm);
quarterly summed precipitation (mm);
range in quarterly precipitation (mm);
annual water deficiency (annual equilibrium evapotranspiration minus annual
precipitation; Sykes et al. 1996);
range in annual water deficiency;
soil water content for both upper and lower horizon retrieved from a dynamic
vegetation model (LPJ-GUESS; Smith et al. 2001, Rickebusch et al. 2008);
B. Methodology 21
e annual cloudiness (%);
e quarterly cloudiness (%);
e accumulated growing degree days with a base temperature of five degrees
until February, April, June, and August.
Many of these variables are partly redundant in their effects. ‘Thus, to avoid statistical
problems due to high levels of collinearity between climate variables we selected ecological
relevant and least correlated variables by means of cluster analysis. The threshold for
variable selection was a Pearson correlation coefficient lower than 0.3 (Graham 2003).
The remaining variables which have been used for the climate niche models of all
species within this atlas were
e accumulated growing degree days until August, which is highly representative
for general temperature gradients across Europe (Fig B.3.1);
e soil water content for the upper horizon, which 1s a realistic measure of water
availability and near surface microclimate (Fig B.3.2);
e ranges in annual precipitation (Fig B.3.3) and
e ranges in annual temperature (Fig B.3.4); with the two last ones reflecting
continentality and oceantty.
Modelling procedure
To assess species response to climate change, we first need to identify the ecological
niche that each species occupies with respect to key climatic variables. Climatic niche
models relating such variables to presence and absence data were developed using
generalized linear models (GLM) with a binomial error distribution and a logit link
function. We allowed for additive and curvilinear effects by incorporating second
order polynomials. Models were checked for spatial autocorrelation with Moran’s
I correlograms of model residuals, but none was detected. Initial models were
simplified by stepwise regression, while minimizing Akaike’s information criterion
(AIC; Sakamoto et al. 1986). Models were calibrated on an 80% random sample of the
initial data set and model accuracy was evaluated on the remaining 20%. Agreements
between observed presences and absences and projected distributions were evaluated
by the Area Under the Curve (AUC) of a Recetver Operating Characteristic (ROC) plot
which is independent of thresholds (Fielding & Bell 1997). Thresholds for calculating
presence-absence projections were obtained by a maximizing Kappa approach (Manel
et al. 2001). While the climatic niche models were developed at the 50 =< 50 km?
UTM erid, the future projections were downscaled to 10 < 10 min grid cells. Both
wete mapped within the geographical range of -10.417° to 31.917° (longitude) and
34.083° to 71.083° (latitude). All maps based on the WGS1984 coordinate system were
projected in the Miller cylindrical projection using ArcGIS software (ESRI 2006).
22
Climatic Risk Atlas of European Butterflies
Accumulated growing
degree days until August
ss 3.
Re
Ee
sy 1919-2593
|
2504 = d437
Soil water contents for
the upper horizon (%)
(LPJ-GUESS; Smith et al. 2001;
Rickebusch et al. 2008)
75,4 - 40,0
40, = 50,0
S10 - 65,5
65,65 100
Fig. B.3.1 Accumulated grow-
ing degree days until August. (a)
Current conditions (1971-2000);
(b) future conditions for 2050
under the intermediate scenario
(BAMBU); (c) future conditions
for 2080 under the intermediate
scenario (BAMBU).
Fig. B.3.2 Soil water content. (a)
Current conditions (1971-2000);
(b) future conditions for 2050
under the intermediate scenario
(BAMBU); (c) future conditions
for 2080 under the intermediate
scenario (BAMBU).
Fig. B.3.3 Range in annual pre-
cipitation. (a) Current conditions
(1971-2000); (b) future conditions
for 2050 under the intermediate
scenario (BAMBU); (c) future
conditions for 2080 under the in-
termediate scenario (BAMBU).
Fig. B.3.4 Range in annual tem-
perature. (a) Current conditions
(1971-2000); (b) future conditions
for 2050 under the intermediate
scenario (BAMBU); (c) future
conditions for 2080 under the in-
termediate scenario (BAMBU).
B. Methodology
Range in annual
precipitation (mm)
[3 » 44
45 - 67
68.99
10 - 153
| 54 « 285
Range in annual
temperature (C°)
Ta - 134
13,9 - [70
[7-2 te
2, -24,7
24,8. 29,9
24 Climatic Risk Atlas of European Butterflies
Assumptions for species dispersal
The next key factor we need to incorporate is the ability of a species to colonise new
potentially suitable areas in the course of climate change. For butterflies, this depends
closely on a species’ dispersal ability. However, detailed dispersal distances are not
available for most species and we thus examined two extreme assumptions:
1. Unlimited dispersal, such that the entire projected niche space denotes the
actual future distribution.
2. No dispersal, in which the future distribution results solely from the overlap
between current and future niche space.
Visualisation of the multi-dimensional climatic niche
To visualise the multi-dimensional climatic niche independent from the species’
geographic distribution, we provide a 4 x 4 panel of graphs. In each graph the
occurrence probability surface is presented, according to the climatic niche model
and the threshold beyond which occurrence is most likely, considering accumulated
erowing degree days until August (Gdd; x-axis) and soil water content (Swe; y-axis).
Additionally, for most species the relationship between occurrence probability, Gdd
and Swe varies with the other two considered variables annual temperature range and
annual precipitation range. Since a continuous visualisation of this four-dimensional
niche would be outside the scope of human perception, we provide 4 x 4 discrete
combinations of the latter two variables. Therefore, we depict the relationship
between occurrence probability, Gdd and Swe for combinations of minimum, lower
tercile, upper tercile and maximum values of annual temperature range and annual
precipitation range.
B.4 Climate change risk assessment for butterflies
Definitions of climate change risk categories for European butterflies
Each butterfly species assessed was placed in a risk category (see below) according
to the loss of grid cells in each scenario. Categories were only assigned for species
whose distributions were modelled reasonably accurately by the model (AUC > 0.75,
see chapter B.3). Species whose distributions were not modelled reasonable accurately
were assigned the category “PR — Potential climate change risk”.
B. Methodology 25
The categories of model quality are as follows:
AUC: > 0.95: Present distribution can be very well explained by climatic variables
AUC: > 0.85 — 0.95: | Present distribution can be well explained by climatic variables
AUC: < 0.75: Present distribution can be explained by climatic variables to only a
limited extent
AUC: > 0.75 — 0.85: Present distribution can be explained by climatic variables to a
moderate extent
The climate risk categories which have been derived from the analysis and which are
used throughout the atlas are as follows:
HHHR extremely high climate change risk 2.75
HHR very high climate change risk S65 95 Sal Os
al climate change risk Poi aay = O75
potential climate change risk 0 - 100 < 0.75
high climate change risk ica Sh
Integrated overall risk categories for European species —
integrating all scenarios and time steps
In the short description of the ecology of each species in chapter C.2, each species
was given an overall risk category. These are defined as follows:
HHHR (extremely high climate change risk): Climate change poses a very high risk
to the species because more than 95% of the grids with currently suitable climate
may no longer be suitable in 2080 under at least one scenario (under the “no
dispersal” assumption). Present distribution can be explained by climatic variables
at least to a moderate extent (AUC > 0.75).
HER (very high climate change risk): Climate change poses a very high risk to the
species because more than 85% of the grids with currently suitable climate may
no longer be suitable in 2080 under at least one scenario (under the “no dispersal”
assumption). Present distribution can be explained by climatic variables at least to
a moderate extent (AUC > 0.75).
26 Climatic Risk Atlas of European Butterflies
FIR (high climate change risk): Climate change poses a high risk to the species because
more than 70% of the grids with currently suitable climate may no longer be
suitable in 2080 under at least one scenario (under the “no dispersal” assumption).
Present distribution can be explained by climatic variables at least to a moderate
extent (AUC > 0.75).
R (climate change risk): Climate change poses a risk to the species because more than
50% of the grids with currently suitable climate may no longer be suitable in
2080 under at least one scenario (under the “no dispersal’ assumption). Present
distribution can be explained by climatic variables at least to a moderate extent
(AUG 0075):
LR (lower climate change risk): Climate change poses a lower risk to the species
because 50% or less of the grids with currently suitable climate may no longer be
suitable in 2080 under at least one scenario (under the “no dispersal” assumption).
Present distribution can be explained by climatic variables at least to a moderate
extent (AUC > 0.75).
PR (potential climate change risk): At the moment, climate change can only be regarded
as a potential risk for the species’ long-term survival in Europe. All species whose
present distribution can be explained by climatic variables to only a limited extent
(AUC: S$ 0.75) have been categorised as PR, independent of the rate of decline of
their climatic niche distribution.
C
ee
CLIMATE RISKS OF EUROPEAN
BUTTERFLY SPECIES
28 Climatic Risk Atlas of European Butterflies
C.1 Species inventory and taxonomy of European butterflies
Depending on different authors, the state-of-the-art in taxonomy and its interpretation,
Europe is thought to support around 450 butterfly species. For this atlas we have
modelled 294 species (in some cases only species complexes). The results on individual
species are shown in chapter C.2, with complete lists in Appendices 1 and 2. A further
149 species (a number which might change depending on the results of taxonomic
research) are listed in chapter C.3. Species covered in C.2 and C.3 only include those
from the geographical area of Europe, including the Azores, Madeira, the Canary
Islands, all Greek islands, Cyprus and the European part of Turkey, but excluding
territories of Belarus, Ukraine, Moldova, and Russia.
Generally, the classification of butterfly genera and species in the present work
is based on that utilized in “The distribution atlas of European butterflies’ (Kudrna
2002). Recent progress in systematic research (incorporating biological, genetic and
molecular aspects, but not ignoring traditional morphology) has led to changes in the
classification of many taxa. One always has to be aware, that taxonomic categories
are in principle scientific concepts and nominate taxa, here genera and species, are
scientific hypotheses. ‘Their authors may employ different views on the status of the
same taxon and their views may change as time goes by, subject to the application of
new methods and new research results.
Therefore the classification of European butterflies is bound to remain fluid — not
only if one considers that according to Desctmon & Mallet (in press) around 16% of
European butterfly species are known to hybridize in the wild. About half or more of
these hybrids are fertile, and show evidence of backcrossing. An example for the need
of reclassification on the genus level is the heterogeneity of the genus Pbejus KLUK,
1802, where new molecular data by Wiemers (2003) indicate that it is not monophyletic
and possibly consists of a few closely related genera).
The first draft of the original checklist used by Kudrna (2002) dates back to 1995;
it contained a number of questionable (micro)species in order to facilitate research
about their ranges and, perhaps, thetr overlapping zones. This helped, for instance, to
establish, that Ewphydryas glaciegenita 1s a high alpine ecological race of E. aurinia, not a
distinct species (Pelz, unpubl).
The nomenclature utilized here follows the International Code on Zoological
Nomenclature. Unfortunately the International Commission on Zoological Nomen-
clature does little at present to help stabilize zoological nomenclature by using its
plenary powers, the means created in the past for this purpose by the Commission.
Thus, for instance, the well established generic name Macuiinea van Ecker, 1915, must
be ‘sacrificed’, as it is a junior subjective synonym of Phengaris DOHERTY, 1891 (e.g.
Pech et al. 2004).
C. Climate Risks of European Butterfly Species
The main changes or adjustments for the present atlas are telegraphically reviewed
here, starting with general aspects and followed by a family-wise treatment:
General:
The authorship of all names proposed in the ‘Wiener Verzeichnis’ was
erroncously attributed to DENIS & SCHIFFERMULLER by most of the past authors.
It has been demonstrated by Kudrna & Belicek (2005) that the sole author of
the work was I. SCHIFFERMULLER. It has been shown that many names proposed
therein are nomina nuda, but except in one case they have been made available
by subsequent authors.
Hesperiidae:
Pyrgus malvoides 1s being treated as a distinct species by some and as a subspecies
by other authors (e.g. Higgins 1976, de Jong 1972). Following recent recording
(Kudrna 2002) and thus for technical reasons, it appears more appropriate to
treat both Pyrgus malvae and Pyrgus malvoides under the Pyrgus malvae complex.
Pieridae:
Following Braby (2005) and contrary to Klots (1933), Ponta Fasricius, 1807,
is provisionally recognized as a genus distinct from Pres SCHRANK, 1801.
Pontia daplidice was divided into two species by Geiger & Scholl (1982) due
to strong differences in allozyme pattern, but Porter et al. (1997) questioned
their specific distinctness after proving extensive hybridization in a contact
zone in Liguria, Italy. Because of these results and lack of information on the
distribution of both taxa in Central Europe, both taxa are treated as a complex
in the present atlas. Work on the identity and distribution of both P. edusa and
P. daphdice (ncluding molecular work with DNA markers) 1s in progress. It
seems that P. edusa 1s more widespread in Central Europe and currently the
more active migrant, while P. dapidice is restricted to the Western and Southern
Mediterranean region (Wiemers, unpubl. data).
Leptidea reah and L. sinapis have proven to be sibling species rather recently, but
they cannot be always reliably distinguished and most of old data often cannot
be referred to either species. This makes their treatment as L. senapis complex
unavoidable.
Lycaenidae:
Favonius quercus (LINNAEUS, 1758): Shirozu & Yamamoto (1956) have shown
in their taxonomic revision that Favonius SIBATANI & ITo, 1942, and Overcusia
Verity, 1943, are very closely related, whereas Neozephyrus is morphologically
and phylogenetically very distinct. Since Shirozu & Yamamoto (1956) following
the ‘fashion’ of their time afforded every species-group the status of a genus,
Quercusia 1s best treated as a junior subjective synonym of Favonius.
29
30 Climatic Risk Atlas of European Butterflies
Polyommatus eroides is a subspecies of P. eros Wiemers unpubl., Vodolazhsky &
Stradomsyk 2008a, 2008b).
Polyommatus caelestissimus 1s treated as a subspecies of P. cordon, because there ts
no evidence for genetic differentiation (Desctmon & Mallet in press, Wiemers
2003), therefore the UI'M data points of both species have been combined
into one map.
Plebejus glandon and P. aguilo are treated as separate allopatric species in this
atlas. However, many authors (e.g. Tolman & Lewington 2008) consider the
latter only as a subspecies of P. g/andon, often also including the local endemic
P. zuellichi from Sierra Nevada (which has not been modelled due to the few
data points).
Cyantris semiargus 1s provisionally removed from the genus Po/yommatus LXTREILLE,
1804. New molecular results indicate a closer relationship to Péebejus than to
Polyommatus (Wiemets 2003).
Some species of the genus Scoftantides HUBNER, 1819, used to be placed in
Pseudophilotes Bruret, 1958, which here is regarded as a junior subjective
synonym of the former. The use of these genera however keeps changing and
underlines the necessity of a revision.
Nymphalidae:
The genus Nymphals 1s divided into two genera: Aglas DALMAN, 1816, and
Nymphalis K.uK, 1802. (Wahlberg & Nylin 2003, Wahlberg pers. comm.).
Argyronome YIUBNER, [1819] is a junior subjective synonym of Argynnis
Faprictus, 1807 (Stmonsen 2006); thus: Argynnis laodice.
Coenonympha iphioides 1s a subspecies of C. ghcerton. (Wiemers 2007).
Coenonympha darwiniana 1s a subspecies of C. gardetta (Wiemers 1998, 2007,
Potteretals 1995).
Although Erebia arvernensis and E. carmenta are probably distinct species (Albre,
et al. 2008); at present their distribution data cannot be separated from E.
casstoides for technical reasons and thus they have to be treated as a “complex”
together with the “western” FE. cassioides.
Kudrna (2002) followed Miller (1968) and treated his ‘series’ as monophyletic
genera Hipparchia Fasricius, 1807, Maniola SCHRANK, 1801, Pararge TIUBNER
[1819] and Kzrinza Moore, 1893. Current research however shows, that Miller’s
(1968) ‘series’ ate not monophyletic (Pena et al. 2006). ‘Thus the provisional
classification employed here follows Pena et al. (2006); at present there is no
comprehensive revision or phylogenetic study of the higher classification
of the subfamily Satyrinae. The (provisionally?) resurrected genera ate:
Lastommata Westwoon, 1841, Lopinga Moore, 1893, Hyponephele MUSCHAMP,
1915, Pyronta HUBNER [1819], Brintesta FRUHSTORFER, [1911], Chazara Moore,
1893, Pseudochazara Lessr, 1951, Satyrus LATREILLE, 1810, Arethusana LESsE,
1951, Mznors HUBNER, [1819]. Whereas some of these genera (e.g. Hyponephele,
Lastommata, Maniola, Pararge) are ‘strong’, some other genera remain ‘weak’
C. Climate Risks of European Butterfly Species 31
and their taxonomic status may change in the course of new research results
becoming available. It is to be remembered that only an incomplete selection
of potentially congeneric species has been examined (by molecular and other
studies) so fat.
e Hipparchia alcyone 1s a junior subjective synonym of H. hermione (VERITY 1913)
(Kudrna 1977, 1984, Honey & Scoble 2001), based upon the lectotype of the
latter, designated by Kudrna (1977).
e Pseudochazara amalthea (FRIwALDszKy, 1845), placed by Kudrna (2002)
provisionally in the genus Hzpparchia, is a subspecies of Pseudochazara anthelea
(LEFEBVRE, 1831), as it is already treated by many authors.
In general we agree with Descimon & Mallet (in press), that “there 1s justification for
reviving the rather neglected (and misused) rank of subspecies, with the trend among
lepidopterists to consider only mote strongly distinct forms (in morphology, ecology,
ot genetics) as subspecies, and to lump dubious geographic forms as synonyms.” ‘This
provides “a useful compromise between descriptions of geographic variation, the
needs of modern butterfly taxonomy, and Darwin’s pragmatic use of the term species
in evolutionary studies.”
Although the definition of species will always be difficult, species will continue to
function as useful tools in biology. “Studies of gene exchange in the many hierarchical
layers of phenotype, genotype and genome in “bad” species of butterflies will illuminate
the nature of speciation and evolution at the species level more than discussions on the
“essence” of species.” (citations from Descimon & Mallet, in press).
C.2 Climatic fate of individual species
The present chapter is the core part of this atlas. It encompasses the largest part of
the book from page 32 to page 619. Here all species will be shown as pictures taken
in natural settings. Their ecology and biology is very briefly characterized, based on
general field guides and text books like Tolman & Lewington (2008), but also own
field experience. Information on ant-lycaenid relationships were largely drawn from
Fiedler (2006). For each species we give the statistics of changes in the climatic niche
distribution, the observed and modelled present distribution, the multidimensional
climatic niche, and the projected climatic niche space distribution under the different
scenarios SEDG, BAMBU, and GRAS for the years 2050 and 2080. On each of these
maps the distribution of climatic niche space is shown that a) remains stable (orange),
b) gets lost (grey), and c) 1s gained (dark brown). ‘Thus it is possible to optically deduct
the changes under the no and the full dispersal assumption (see chapter B.3 on page
20ff for further details on methodology).
32 Climatic Risk Atlas of European Butterflies
Erynnis tages (LINNAEUS, 1758) — Dingy Skipper
| =o
-2873 (-16.22%) | -4268 (-24.1%)
SEDG -4695 (-26.51%) | -6615 (-37.35%)
BAMBU_ | -5429 (-30.66%) | -8408 (-47.48%)
-7225 (-40.8%) | -10729 (-60.59%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 17709 cells)
The Dingy Skipper is a small, inconspicuous butterfly. It lays its eggs on the leaves of leguminous
plants such as Coronilla varia (Crown Vetch), Horseshoe Vetch (Hippocrepis comosa) and Common
Birdsfoot trefoil (Lous corniculatus), usually choosing plants growing near bare patches. The caterpillar
spins itself a small, tube-like shelter from leaves of the larval foodplant, living and feeding in it until
fully grown. It then builds itself a sturdier shelter 1n which to pass the winter. In the spring, without
further feeding, it pupates, either in the shelter, or in the moss layer. The adult butterfly is often
found on Bugle (4juga spp.) and, while visiting flowers, is easily observed. The Dingy Skipper has
one brood a year in central and northern Europe and two in the southern part.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.72).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
dd 4000 0 edd 4000 0 ad 4000 0 odd 4000
Minimum
Swe
06
1.0 02
0.6
Swe
1.0 0.2
Swe
0.6
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Erynnis tages (Hesperiidae) 33
(B1)
SEDG
GRAS
(A1Fl)
34 Climatic Risk Atlas of European Butterflies
Erynnis marloyi (BotspDuvaL, 1834) — Inky Skipper
Cc Full dispersal No dispersal
Be SEDG -123 (-13.61%) | -521 (-57.63%)
BAMBU_ | -240 (-26.55%) | -583 (-64.49%)
22 (2n5wr) | 0 (59
SEDG -72 (-7.96%) -578 (-63.94%)
BAMBU | -572 (-63.27%) | -807 (-89.27%)
a8 cram) | wa 835%
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 904 cells)
Inky Skippers are very dark little butterflies that fly rapidly, close to the ground. They are found on
dry grasslands, in dried-up riverbeds, on rocky slopes, and in woodland clearings. They can often be
seen basking in the sun, wings widespread, on light-coloured stones. Especially the females can also
often be seen drinking nectar on thyme. The larvae feed on bushy rosaceans particularly on Prunus
spinosa and P. cocomilla. Vhe Inky Skipper has one or two broods a year.
Present distribution can be very well explained by climatic variables (AUC = 0.98).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
toad 4000 0 odd 4000 0 ad 4000 0 add 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
35
Erynnis marloyi Hesperiidae)
(1a)
504s
(ZV)
nanva
(ld)
SVH9
2080
2050
36 Climatic Risk Atlas of European Butterflies
Carcharodus alceae (EsPEr, 1870) — Mallow Skipper
a ee
2554 (20.32%) | -1255 (-9.98%)
SEDG 1889 (15.03%) | -2106 (-16.75%)
BAMBU -222 (1.77%) -4815 (-38.3%)
-1188 (-9.45%) | -6139 (-48.83%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 12571 cells)
The Mallow Skipper is a butterfly of warm, grassy places, usually with rough vegetation. It is a
mobile butterfly, that strays outside its usual habitat, and thus can be met in unexpected places. In
warm summers, they migrate northwards and can be seen in warm, south-facing river valleys. The
resident populations are usually small. The butterflies are often seen visiting flowers for nectar. They
also spend a lot of their time basking in the sun, their wings widespread, showing to full advantage
their beautiful purple to oltve-green metallic sheen. Eggs are laid singly on the upperside of the
leaves of mallows (Madu ssp.). The caterpillars thrive on this food, growing very quickly. This
skipper has up to three or more broods per year. As winter approaches, the fully-grown caterpillars
make a cocoon in the litter layer. They pupate in the spring.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.71).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
1.6
ct
a a
=) a
= 7
Pm On
ef@eEs
o= mH
=e
£5 8
Oo. 4
ow 7
© Aad
cL 4 = a
ger
YH
e- :
qf. =
2 o
wv
x é Po]
= 4
o
0 2000 40000 2000 40000 2000 40000 2000 4000
Gd id Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
37
Carcharodus alceae (Hesperiidae)
2080
2050
38 Climatic Risk Atlas of European Butterflies
Carcharodus lavatherae (Esper, 1783) — Marbled Skipper
6c Full dispersal No dispersal
Be SEDG -21 (-0.64%) -1482 (-45.43%)
BAMBU -132 (-4.05%) | -1578 (-48.38%)
-379 (-11.62%) | -1893 (-58.03%)
SEDG 1101 (33.75%) | -1978 (-60.64%)
BAMBU 284 (8.71%) -2409 (-73.85%)
-454 (-13.92%) | -2792 (-85.59%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 3262 cells)
The green sheen on the upperside of the wings and body of the Marbled Skipper distinguish it from
other skippers in this genus. It lives 1n warm, flower-rich places. Populations are usually small, and it
is exceptional to see a large number of these butterflies gathered together. On very hot days, needing
to drink, they look for damp ground. The eggs are laid singly on the sepals of various woundworts
(Stachys spp.), especially Perennial Yellow Woundwort (S. rec/a). The young caterpillars spin a loose
shelter from leaves, under which they hide themselves while they feed, first eating the seeds and,
later, the leaves of the plant. They hibernate in the third or fourth instar and pupate at the foot of
the foodplant. The Marbled Skipper 1s single-brooded.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.73).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
ah
o
=
ze
o
™
o
oF 4000 0 ee 4000 0 oy 4000 0 ee 4000
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
39
Carcharodus lavatherae (Hesperiidae)
2080
2050
40 Climatic Risk Atlas of European Butterflies
Carcharodus flocciferus (ZELLER, 1847) — Tufted Marbled Skipper
a
104 (2.47%) -1661 (-39.43%)
SEDG 2641 (62.69%) | -1670 (-39.64%)
BAMBU 2427 (57.61%) | -2346 (-55.68%)
2972 (70.54%) | -2925 (-69.43%)
© Rudi Verovnik Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 4213 cells)
The Tufted Marbled Skipper can be found on flower-rich grasslands. It also occurs at the edges of
woodland and in bushy vegetation. The males usually perch on a tall plant and very actively defend
their territory. The female lays her eggs singly on the leaves of various woundworts (S/achys spp.),
Betony (5. officinahs), but also Hedge Woundwort (S. syfatica), Alpine Woundwort (5. a/pina), and
Marsh Woundwort (5. palustris). Species of horehounds (Marrubium spp.) ate also considered to
be larval foodplants. The young caterpillar lives in a curled-up leaf. When it is bigger, it spins two
neighbouring leaves together to make a shelter where 1t remains during the day, coming out at night
to feed on the leaves of the foodplant. It also pupates in such a shelter. This butterfly has one or two
broods a year. It overwinters either as a caterpillar or a pupa.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.79).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 odd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Carcharodus flociferus (Hesperiidae) 41
42 Climatic Risk Atlas of European Butterflies
Carcharodus orientalis (REVERDIN, 1913) — Oriental Marbled Skipper
.—hClcLw Full dispersal No dispersal
Be SEDG -26 (-2.11%) -611 (-49.59%)
BAMBU -119 (-9.66%) -594 (-48.21%)
-94 (-7.63%) -695 (-56.41%)
SEDG 255 (20.7%) -729 (-59.17%)
BAMBU -435 (-35.31%) -972 (-78.9%)
206 cso) | 1116 605806
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1232 cells)
The Ortental Marbled Skipper is mostly found on grasslands, on rocky slopes with grassy vegetation,
and occasionally on bushy or on low, shrubby vegetation. The butterflies fly quickly, close to the
ground. When they are at rest, they usually have their wings widely spread. Males can be commonly
found drinking on dump ground. The caterpillars feed on woundworts (Siachys spp.). It has two to
three generations a year, and hibernates as a caterpillar.
Present distribution can be well explained by climatic variables (AUC = 0.92).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 ood 4000 0 add 4000 0 add 4000
1.0
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Carcharodus orientalis (Hespertidae) 43
44 Climatic Risk Atlas of European Butterflies
Carcharodus baeticus (RamBuR, 1840) — Southern Marbled Skipper
6c Full dispersal No dispersal
Be SEDG 88 (21.26%) -239 (-57.73%)
168 (40.58%) -229 (-55.31%)
138 (33.33%) -288 (-69.57%)
248 (59.9%) -293 (-70.77%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 414 cells)
The Southern Marbled skipper occurs on dry, sparse vegetation, as found in the dunes, in dry,
erassy places, and on rocky slopes. Foodplants are various horehounds, including Horehound
(Marrubium vulgare), Black Horehound (Ballota migra ssp. foetida), and Ballota spp. The female lays her
eggs one by one on the leaves and shoots of the foodplant, seeming to prefer smaller plants. The
small caterpillars live hidden in a spun leaf. The older, larger caterpillars spin two neighbouring
leaves together to make a safe shelter. When they are fully-grown, the caterpillars go down to the
foot of the foodplant, and spin a few dried leaves together in which to pupate. The species was not
seen in the Alps in last 40 years, where it probably had just one generation per year, while in Spain
there are two or three generations a year. It hibernates as a caterpillar.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.8).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘dd 4000 0 Codd 4000 0 add 4000 0 odd 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
45
Carcharodus baeticus (Hesperiidae)
2080
2050
46 Climatic Risk Atlas of European Butterflies
Spialia phlomidis (HERRICH-SCHAFFER, 1845) — Persian Skipper
.—hCcLe Full dispersal No dispersal
He SEDG -59 (-15.4%) -195 (-50.91%)
-88 (-22.98%) -228 (-59.53%)
|GRAS | -110 (-28.72%) | -260 (67.89%)
SEDG 27 (7.05%) -229 (-59.79%)
BAMBU_ | -262 (-68.41%) | -352 (-91.91%)
-319 (-83.29%) | -381 (-99.48%)
© Rudi Verovnik Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 383 cells)
The Persian Skipper occurs on dry grasslands, on dry scrub, in rocky places and at woodland edges.
On hot days males and females congregate on dump ground where they are easy to spot. Although
the larval foodplant is not known for certain, probably various bindweeds (Conohulus spp.) are
used. Also, 1t 1s not clear how many broods it has a year, but 1t is probably two.
Present distribution can be well explained by climatic variables (AUC = 0.93).
Climate risk category: HHHR
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
-
edd 4000 0 odd 4000 0 Gad 4000 0 ada 4000
0.6
Minimum
Swe
02
Swe
1.0 O.2 0.6 1.0
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Spiala phlomidis (Hespertidae) 47
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
48 Climatic Risk Atlas of European Butterflies
Spialia sertorius (HOFFMANSEGG, 1804) — Red-underwing Skipper
ee
-426 (-6.29%) -1771 (-26.17%)
< -331 (-4.89%) -1755 (-25.93%)
! gy) ee SEDG 553 (8.17%) | -2424 (35.82%)
aomicch te yt BAMBU_ | -1006 (-14.86%) | -3188 (-47.1%)
ae
GRAS -2218 (-32.77%) | -4639 (68.54%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 6768 cells)
The Red-underwing Skipper is a small butterfly that likes warm habitats. It occurs on calcareous
and other dry grasslands, and also in dry, rough vegetation, as long as its larval foodplant, Salad
Burnet (Sanguisorba minor), 1s present. The eggs are laid between the buds on the flowerheads of this
plant and the caterpillars feed on the young leaves. Hibernation takes place as a caterpillar, in warm
areas when still small, and further north when fully grown. The caterpillars pupate in the litter
layer, in a sturdy cocoon made from plant remains. Because of its rapid flight and unremarkable
behaviour, this small butterfly 1s often not even noticed. However, they are often present in large
numbers over quite a small area. The butterflies like visiting flowers. It has one or two broods a
year, depending on the position of the breeding ground within the range.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.77)
Climate risk category: R
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
‘0.6
he
a a
=) a
es 7
Om Oe
cfes
o= mH
=f
£5 8
Oo. 4
os 7
oo
a ee ge
ger
te |
e- :
qf. =
2 o
wv
x é Po]
= 4
o
0 2000 4000 2000 40000 2000 40000 2000 4000
Gdd Gdd Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
49
Spiaha sertorius (Hesperiidae)
2080
2050
50 Climatic Risk Atlas of European Butterflies
Spialia orbifer (HUBNER, 1823) - Orbed Red-underwing Skipper
SEDG 2229 (81.71%) -341 (-12.5%)
BAMBU 528 (19.35%) -987 (-36.18%)
1211 (44.39%) | -976 (-35.78%)
SEDG 1025 (37.57%) | -1188 (-43.55%)
& | BAMBU -59 (-2.16%) -1848 (-67.74%)
1443 (52.9%) | -2000 (-73.31%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2728 cells)
The Orbed Red-underwing Skipper occurs on dry, flower-rich grasslands, on roadside verges, at the
edges of woods, on abandoned agricultural land, rocky slopes, and along dried-up river beds. The
butterflies have a rapid flight, close to the ground. The eggs are laid on the flowerheads of Salad
Burnet (Sanguisorba minor), and, in Eastern Europe, possibly also on Great Burnet (S. officenals).
The round flowerheads of the foodplant are the caterpillars’ first food, but as they grow larger,
they hide themselves between spun leaves. Hibernation takes place on the ground, in the litter
layer. They also pupate in rolled-up leaves of the foodplant. This skipper is double-brooded and
hibernates as a caterpillar.
Present distribution can be well explained by climatic variables (AUC = 0.89).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Ow
a]
wy
™
a
1.0
Minimum
Swe
06
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
o2
0 2000 40000 2000 40000 2000 40000 2000 4000
Gd dl Gadd Gdd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
51
Spiala orbifer (Hesperiidae)
2080
2050
52 Climatic Risk Atlas of European Butterflies
Syrichtus proto (OCHSENHEIMER, 1808) — Sage Skipper
.—CcLe Full dispersal No dispersal
| SEDG -1328 (-36.22%) | -1802 (-49.15%)
AMBU | -1862 (-50.79%) | -2012 (-54.88%)
a -1928 (-52.59%) -2248 (-61.32%)
SEDG -1929 (-52.62%) -2530 (-69.01%)
-3014 (-82.21%) -3242 (-88.43%)
-2945 (-80.33%) | -3553 (-96.92%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 3666 cells)
The Sage Skipper occurs on dry, flower-rich grassland, in open scrub, on rocky ground with sparse
vegetation. However, in its chosen habitat, the larval foodplants are usually abundant. Various species
of Phlomis are used, such as Jerusalem Sage (P. fructicosa), P. hchnitis, P. herba-vent, and perhaps also
Horehound (Marrubium spp.). Although there is probably only one generation a year, the butterflies
of the Sage Skipper can be seen from spring until autumn, due to prolonged emergence from the
pupa. This skipper passes the winter as a caterpillar inside the egg.
Present distribution can be well explained by climatic variables (AUC = 0.89).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
dd 4000 0 edd 4000 0 add 4000 0 Gd 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
1.0 02
0.6
Maximum Large (66%) Small (33%)
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
53
Syrichtus proto (Hesperiidae)
2080
2050
54 Climatic Risk Atlas of European Butterflies
Syrichtus tessellum (HUBNER, 1803) — Tessellated Skipper
6c Full dispersal No dispersal
SEDG 305 (28.8%) -806 (-76.11%)
3
S
-303 (-28.61%) | -938 (-88.57%)
eae -736 (-69.5%) -1059 (-100%)
-755 (-71.29%) | -1058 (-99.91%)
a -736 (-69.5%) -1059 (-100%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1059 cells)
The Tessellated Skipper occurs in open grassy and flower rich places where it feeds preferably on
thymes, vetches and yarrows. The butterflies have a rapid flight, often quite close to the ground. The
caterpillars feed on the labiates Phlomis tuberosa and P. samia, hiding in spun leaves. The Tessellated
Skipper has one or two generations a year, and passes the winter as a small caterpillar.
Present distribution can be well explained by climatic variables (AUC = 0.89).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
Ey
o
E zs
=
tl
= 6
o
x
Ow
=o
wo
tH
ro]
2 : . : E ; ;
o
=
w
ead 4000 0 odd 4000 0 oad 4000 0 odd 4000
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
55
Syrichtus tessellum (Hesperiidae)
2080
2050
56 Climatic Risk Atlas of European Butterflies
Pyrgus carthami (HUBNER, 1813) — Safflower Skipper
.—hCcLUwkE Full dispersal No dispersal
al SEDG 222 (3.38%) 1952 (-29.74%)
-770 (-11.73%) | -2415 (-36.8%)
-1787 (-27.23%) | -4285 (-65.29%)
-3285 (-50.05%) | -5563 (-84.76%)
© Rudi Verovnik Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 6563 cells)
The Safflower Skipper is the largest grizzled skipper in Central Europe. It prefers sheltered places,
occurring on dry, often calcareous grasslands, and on rough vegetation in places sheltered from
the wind. They often rest near bushes or at the edges of woods. They lay their eggs singly on the
upperside of the leaves of cinquefoils (Potentilla spp.), the caterpillar later using leaves as food.
However, the caterpillars are difficult to find, because they hide away in plant cushions during the
day, 1n order to survive the summer heat. The caterpillars hibernate in spun leaves in the litter layer,
probably in the last larval instar. However, in breeding experiments, each stage was found to be
capable of hibernating. Before they pupate, the caterpillars make a sturdy cocoon. The Safflower
Skipper is single-brooded.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.76).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘sad 4000 0 edd 4000 0 add 4000 0 odd 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
57
Pyrgus carthami (Hesperiidae)
2080
2050
58 Climatic Risk Atlas of European Butterflies
Pyrgus sidae (Esrrr, 1782) — Yellow-banded Skipper
AMBU -808 (-60.16%) | -1286 (-95.76%)
-384 (-28.59%) | -1328 (-98.88%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1343 cells)
The yellow and white bands on the underside of its hindwing make the Yellow-banded Skipper
unmistakable. These skippers can be found on flower-rich grasslands, flower-rich stony slopes
in gullies, and in open scrub. The butterflies fly slowly, close to the ground. Sulphur Cinquefoil
(Potentilla recta), Potentilla lirta, and perhaps also Abutilon theophrast ate used as larval foodplants. It
hibernates as a caterpillar and has one brood a year.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.85).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Ow
=o
ie]
is}
oO
= : : : : F ? :
ah
o
=
wv
o
dd 4000 0 edd 4000 0 add 4000 0 Gdd 4000
1.0
Minimum
Swe
06
1.0 02
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
59
Pyrgus sidae (Hesperiidae)
2080
2050
60 Climatic Risk Atlas of European Butterflies
Pyrgus andromedae (WALLENGREN, 1853) — Alpine Grizzled Skipper
a
511 (-26.46%) | -546 (-28.28%)
SEDG -425 (-22.01%) | -586 (-30.35%)
BAMBU -306 (-15.85%) | -542 (-28.07%)
-584 (-30.24%) | -777 (-40.24%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1931 cells)
The Alpine Grizzled Skipper occurs in damp, moist, grassy places, often near streams or bogs in
the Alps and Pyrenees. In Scandinavia, they are seen on dwarf scrub vegetation and also in rocky
places on steep slopes. The only reported foodplant 1s Dryas octopetala. The caterpillars live in a
communal shelter of spun leaves. They hibernate, pupating in a similar shelter the following spring.
The species’ development takes two years.
Present distribution can be well explained by climatic variables (AUC = 0.87).
Climate risk category: LR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 edd 4000 0 add 4000 0 add 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
61
Pyrgus andromedae (Hesperiidae)
2080
2050
62 Climatic Risk Atlas of European Butterflies
Pyrgus cacaliae (RAMBUR, 1840) — Dusky Grizzled Skipper
.—hlcL Full dispersal No dispersal
Be SEDG -272 (-23.59%) -368 (-31.92%)
BAMBU -207 (-17.95%) | -320 (27.75%)
-312 (-27.06%) | -400 (-34.69%)
SEDG 514 (-44.58%) | -630 (54.64%)
BAMBU -336 (-29.14%) | -538 (-46.66%)
523 (-45.36%) | -719 (-62.36%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1153 cells)
Damp grassland, at the edge of bogs or streams, is the habitat of the Dusky Grizzled Skipper,
although they sometimes occur on dry grasslands. The male show marked territorial behaviour. The
female lays her eggs on various species of cinquefoil (Polenii/la spp.), such as Tormentil (P. erecta),
and Alpine Cinquefoul (P. crantzi2), and also on Szbbaldia species. She prefers plants growing on drier
places, such as on hummocks in damp habitats, or at a little distance from its habitat. The caterpillars
live hidden between spun leaves, and it is 1n this stage that the Dusky Grizzled Skipper hibernates.
The development takes two years.
Present distribution can be well explained by climatic variables (AUC = 0.92).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
toad 4000 0 odd 4000 0 oad 4000 0 add 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Pyrgus cacaliae (Hesperiidae) 63
64 Climatic Risk Atlas of European Butterflies
Pyrgus centaureae (RAMBUR, 1840) — Northern Grizzled Skipper
a
-1565 (-27.17%) | -1588 (-27.57%)
SEDG -1836 (-31.87%) | -2073 (-35.99%)
BAMBU | -2932 (-50.9%) | -3022 (-52.47%)
GRAS -3722 (-64.62%) | -3811 (-66.16%)
© Bernard Fransen Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 5760 cells)
The Northern Grizzled Skipper prefers damp, moist and wet places. It is mostly found in open areas
of bogs or swamps, in scrub near swamps, and above the tree-line on stretches of wet ground with
dwarf shrubs. With its inconspicuous colours and rapid flight, the butterflies of this grizzled skipper
are difficult to follow. The female lays her eggs on Cloudberry (Rubus chamaemorus). Lattle is known
about the way the caterpillars live. This species is single-brooded.
Present distribution can be very well explained by climatic variables (AUC = 0.96).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘dd 4000 0 edd 4000 0 Cad 4000 0 ad 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
65
Pyrgus centaureae (Hesperiidae)
*l
1
oe
- |
ul “wf att
%
aby
Ls
2080
2050
66 Climatic Risk Atlas of European Butterflies
Pyrgus malvae (LINNAEUS, 1758) / malvoides
(ELWeEs & EpDwarps, 1897) (complex) — Grizzled Skipper
a
SEDG -5554 (-27.17%) | -10058 (-49.21%)
BAMBU_ | -4212 (-20.61%) | -7735 (-37.84%)
-2288 (-11.19%) | -5349 (-26.17%)
© Rudi Verovnik Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 20440 cells)
Due to data availability we here treat both Pyrgus malhae (LINNAEUS, 1758) and P. malhoides (ELWES
& Epwarps, 1897) as the Grizzled Skipper species complex, knowing that Pyreus mahoides is mostly
recognised as a distinct species.
The Grizzled Skipper occurs in many different habitats, mostly on dry, as well as moist, flower-rich
grasslands, and also on calcareous grasslands and heaths. The eggs are laid on the underside of the
leaves of cinquefoils (Pofentilla spp.) and strawberries (Fvagaria spp). The caterpillar builds a small
shelter from a leaf, in which it stays hidden, feeding on the leaves of the foodplant. The sturdy
cocoon in which it pupates is spun from plant remains. This species has one to two broods.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.71).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
‘dd 4000 0 odd 4000 0 add 4000 0 Gdd 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Swe
0.6
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
67
Pyrgus mahae | malvoides (Hesperiidae)
2080
2050
68 Climatic Risk Atlas of European Butterflies
Pyrgus serratulae (RAMBUR, 1840) — Olive Skipper
.6—ClcLw Full dispersal No dispersal
Be SEDG 52 (0.84%) -1871 (-30.1%)
-450 (-7.24%) | -2061 (-33.16%)
-1483 (-23.86%) | -3798 (-61.11%)
-3008 (-48.4%) | -4947 (-79.6%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 6215 cells)
The Olive Skipper occurs in areas that do not become too hot in the summer, on poor to rough,
flower-rich grasslands, which are either open or surrounded by woodland. The butterfly of the
Olive Skipper is quite mobile, and may be seen in unexpected places. The eggs are deposited on
the underside of the leaves of cinquefoils (Ponta spp.), and master-worts (Astrantia spp.). The
caterpillar lives in a small shelter, spun from a rolled-up leaf, feeding on the leaves of the foodplant.
It passes the winter in a cocoon, and pupates the following spring at the base of the foodplant. This
skipper is single-brooded.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.82).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 edd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
69
Pyrgus serratulae (Hesperiidae)
2080
2050
70 Climatic Risk Atlas of European Butterflies
Pyrgus onopordi (RamBur, 1840) — Rosy Grizzled Skipper
CL Full dispersal No dispersal
| SEDG 279 (-12.86%) | -1032 (-47.58%)
} | BAMBU -306 (-14.11%) -1069 (-49.29%)
GRAS -449 (-20.7%) | -1312 (-60.49%)
SEDG 88 (4.06%) -1395 (-64.32%)
BAMBU -98 (-4.52%) -1602 (-73.86%)
GRAS -373 (-17.2%) -2079 (-95.85%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2169 cells)
The Rosy Grizzled Skipper has a characteristic anvil-shaped spot on the underside of its
hindwing. These butterflies can be seen on different types of grassy vegetation and on rocky
slopes. Different foodplants are used in different parts of its range. In Switzerland the eggs are
laid on rockroses (Hehanthemum spp.) and cinquefoils (Potentilla spp.), in Spain, mallows (Maha
spp.) ate used. In most places, the Rosy Grizzled Skipper has two broods a year and in some
places three. It hibernates as a caterpillar.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.78).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
7 | |
‘add 4000 0 odd 4000 0 add 4000 0 Gdd 4000
Minimum
Swe
06
1.0 02
0.6
Swe
1.0 0.2
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
71
Pyrgus onopordi (Hesperiidae)
2080
2050
72 Climatic Risk Atlas of European Butterflies
Pyrgus carlinae (RaMBvrR, 1840) — Carline Skipper
SEDG -100 (-12.22%) -481 (58.8%)
BAMBU 22, (2.69%) -419 (51.22%)
-133 (-16.26%) | -506 (-61.86%)
SEDG -231 (-28.24%) | -714 (87.29%)
BAMBU_ | -197 (-24.08%) | -614 (-75.06%)
GRAS -362 (-44.25%) | -703 (85.94%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 818 cells)
The Carline Skipper prefers dry, south-facing slopes with quite short vegetation. However, they
can also be seen on damp grasslands, and in very open larch woods. Large numbers can sometimes
occut locally. The female lays her eggs singly on the underside of the leaves of various cinquefotls
(Potentilla spp.). The caterpillar remains in the egg during the winter, emerging in the spring. It then
spins a shelter by attaching a leaf of the foodplant to the ground, 1n which it lives, hidden. It pupates
close to the ground, and has one generation a year.
Present distribution can be well explained by climatic variables (AUC = 0.9).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 edd 4000 0 oad 4000 0 add 4000
1.0
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
73
Pyrgus carlinae (Hesperiidae)
2080
2050
74 Climatic Risk Atlas of European Butterflies
Pyrgus cirsit (RAMBUR, 1840)
[ie] _ 7 Full dispersal No dispersal
-472, (-10.85%) | -1306 (-30.01%)
524 (-12.04%) | -1326 (-30.47%)
3 | BAMBU
-1107 (-25.44%) | -1933 (-44.42%)
| | SEDG -94T (-21.76%) | -1936 (-44.49%)
BAMBU_ | -1773 (-40.74%) | -2764 (-63.51%)
-3080 (-70.77%) | -3713 (-85.32%)
——
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 4352 cells)
This grizzled skipper occurs on warm, flower-rich grasslands, preferring those in sheltered situations,
for example, on slopes, or near shrubs or woodland. In the northern part of its range, it is only
found in very warm places. The female lays her eges singly on the underside of the leaves of
vatious cinquefoils (Pofentilla spp.), choosing the smaller plants. This butterfly hibernates as a very
tiny caterpillar in the egg, in which it remains until the spring. It then emerges and spins itself a small
shelter from leaves. In the last larval instar, this is replaced by a parchment-like shelter. It pupates in
a cocoon. It is single-brooded.
Present distribution can be well explained by climatic variables (AUC = 0.9).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
06
tt
=) 2
os
ss
= = Pas
o= mH
:- 7 on
£5 8
Oo 4
os 7
a oo
a 4 ge
ger
™
ee
qf. =
3
nid ite
= 4
o
0 2000 40000 2000 40000 2000 40000 2000 4000
Gdd Gdd Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
75
Pyrgus cirsit (Hesperiidae)
2080
2050
76 Climatic Risk Atlas of European Butterflies
Pyrgus armoricanus (OBERTHUR, 1910) — Oberthiir’s Grizzled Skipper
.6—hlcLUwk Full dispersal No dispersal
Be SEDG 1076 (16.16%) -2050 (-30.79%)
362 (5.44%) -2404 (-36.11%)
-402 (-6.04%) | -5276 (-79.24%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 6658 cells)
Oberthtir’s Grizzled Skipper 1s generally limited to unimproved grasslands. In the north of its range,
the populations are small, but large numbers of butterflies make up the southern populations.
The butterflies can often be seen visiting flowers and seem to be especially fond of Globulana.
The female lays her eggs singly on the underside of the leaves of cinquefoils (Potenii/ia spp.) and
rockroses (Hehanthemum spp.). The caterpillars of the first instar only eat the surface layer, leaving
translucent “windows” in the leaf. The larger caterpillars eat the whole leaf, and live in a tent-like
shelter, spun from one or more leaves. It is the caterpillar that hibernates. It pupates in a cocoon
on the ground and there are two to three generations a year.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.78).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
-
ion
o
ch
o
dd 4000 0 odd 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
Swe
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
77
Pyrgus armoricanus (Hesperiidae)
2080
2050
78 Climatic Risk Atlas of European Butterflies
Pyrgus alveus (HUBNER, 1803) (complex) — Large Grizzled Skipper
.6—hCcLUwk Full dispersal No dispersal
Be SEDG -102 (-0.93%) -2549 (-23.36%)
BAMBU 580 (5.32%) -2013 (-18.45%)
-1796 (-16.46%) | -3665 (-33.59%)
SEDG 2539 (23.27%) | -3680 (-33.72%)
BAMBU 1087 (9.96%) } -4815 (-44.13%)
246 (2.25%) -6117 (-56.06%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 10912 cells)
Due to data availability and resolution we include P. frebevivensis, P. accretus and P. aleus 1n this
complex, while we are aware that these taxa are often treated separately. The Large Grizzled
Skipper is usually found in mountainous areas on dry, poor, flower-rich grassland, and rough
vegetation. Quick-flying and alert, it also likes basking on the ground and drinking from wet mud.
It is a very variable butterfly with many different subspecies and forms, making identification
difficult. Choosing the smaller leaves, the female lays her eggs singly on various rock-roses
(Hehanthemum spp.). At first, the caterpillar lives between spun leaves on plants, later on it spins
a tent-like shelter on the ground. It passes the winter in this stage and builds a spectal tube-like
structure in which to pupate. There are one or two generations a yeat.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.73).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
=
sh
a
a
=
Ow
Za
09
ie
Oo
eS : : : ; F 7 :
dd 4000 0 edd 4000 0 oad 4000 0 odd 4000
Minimum
Swe
06
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
79
Pyrgus alveus (Hesperiidae)
2080
2050
80 Climatic Risk Atlas of European Butterflies
Pyrgus belliert (OBERTHUR, 1910) — Foulquier’s Grizzled Skipper
CL Full dispersal No dispersal
Be SEDG -180 (-44.33%) -356 (-87.68%)
BAMBU_ | -172 (-42.36%) | -338 (-83.25%)
-109 (-26.85%) | -369 (-90.89%)
SEDG 242 (59.61%) -395 (-97.29%)
BAMBU -60 (-14.78%) -404 (-99.51%)
-113 (-27.83%) -406 (-100%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 406 cells)
This grizzled skipper likes flower-rich, grassy vegetation. It occurs on sub-alpine and alpine grasslands
as well as on damp, sometimes rough, grassy vegetation. Various cinquefoils (Potentilla spp.)
are used as foodplant, including Tormentil (P. erecta), Silverweed (P. anserina), Spring Cinquefoil
(P. tabernaemontam), and Creeping Cinquefoil (P. repfans). This butterfly hibernates in the larval stage
and has one generation a year.
Present distribution can be well explained by climatic variables (AUC = 0.94).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
add 4000 0 od 4000 0 Cad 4000 0 add 4000
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 O.2
Swe
0.6
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
2050
Pyrgus belieri (Hesperiidae)
SEDG
(B1)
BAMBU
GRAS
2080
(A2)
(A1Fl)
81
82 Climatic Risk Atlas of European Butterflies
Pyrgus warrenensis (VERITY, 1928) — Warren’s Skipper
Cc Full dispersal No dispersal
Be SEDG -447 (-31.81%) -568 (-40.43%)
BAMBU -437 (-31.1%) -541 (-38.51%)
537 (-38.22%) | -644 (-45.84%)
SEDG -725 (-51.6%) -954 (-67.9%)
BAMBU -655 (-46.62%) | -834 (-59.36%)
-796 (-56.65%) | -997 (-70.96%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1405 cells
Warren’s Skipper occurs mostly on flower-rich alpine grasslands. The males fly where flowers are
most abundant, along mountain streams, for example. From a perch on a bush or tall plant, they also
defend their territory. The females meanwhile are often found on warm, dry patches near bushes,
where they lay their eggs one by one on the rockrose Hehanthemum alpestre, preferring the smaller
plants. The small caterpillar lives hidden in a spun leaf, while caterpillars of later stages spin two
leaves together, and finally, make a larger shelter from several leaves. ‘The Warren’s Skipper is single-
brooded and passes the winter as a caterpillar.
Present distribution can be very well explained by climatic variables (AUC = 0.99).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
7 |].
dd 4000 0 edd 4000 0 ad 4000 0 odd 4000
Minimum
Swe
06
1.0 02
0.6
Swe
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Pyrgus warrenensis (Hesperiidae) 83
84 Climatic Risk Atlas of European Butterflies
Heteropterus morpheus (PAuas, 1771) — Large Chequered Skipper
aa ee
4028 (53.29%) | -541 (-38.51%)
SEDG 1760 (23.28%) -954 (-67.9%)
BAMBU 2305 (30.49%) | -834 (59.36%)
819 (10.83%) -997 (-70.96%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 7559 cells)
The conspicuous pattern on the underside of the wings of the Large Chequered Skipper is unlike
that of any other European butterfly. Furthermore, it has a characteristic, bouncing flight, low over
the vegetation. It occurs in damp, rough, grassy places, such as grasslands, road verges, edges of
streams, at the edges of raised bogs, and in woodland clearings. The eggs are laid singly or in small
batches on the blades of grasses, such as Purple Moor-grass (Modnea caeruka) or Purple Smallreed
(Calamagrostis canescens). The caterpillar makes a tubular shelter from grass, at first using a folded
blade, and later using spun grass-blades. It leaves its shelter in search of food. The caterpillars pass
the winter in the shelter. Pupation takes place in spring. The Large Chequered Skipper has one
generation a year.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.78).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
tea 4000 0 edd 4000 0 oad 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Heteropterus morpheus (Hesperiidae)
SEDG
GRAS
(B1)
(A1Fl)
85
86 Climatic Risk Atlas of European Butterflies
Carterocephalus palaemon (PALLAs, 1771) — Chequered Skipper
.6—hlcLUwk Full dispersal No dispersal
Be SEDG -1722 (-15.33%) | -3397 (-30.24%)
-1877 (-16.71%) | -3570 (-31.78%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 11232 cells)
The Chequered Skipper usually occurs on damp grassland at woodland margins, or where there is
woodland nearby, including road verges, also on the drier areas of raised bogs and at their edges.
Breeding grounds are also known 1n somewhat drier locations at higher altitudinal levels. Populations
are usually small. The female lays her eggs one by one on the blades of coarse-leaved grasses. The
caterpillars spin folded grass leaves together into a little tube, in which they spend most of ther life,
leaving it only to feed. Those who know what to look for, can find their traces, small half-moons
nibbled away from the edge of the grass-blade. The caterpillars hibernate in a more substantial
tube-like shelter. It pupates, its papery, pale-yellow pupa b suspended from withered grass by a silken
girdle. The Chequered Skipper is single-brooded.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.79).
Climate risk category: LR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
aad 4000 0 edd 4000 0 Gad 4000 0 ada 4000
1.0
Minimum
Swe
0.6
Swe
0.6 1.0 O2
1.0 6 O.2
Swe
0.6
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
87
Carterocephalus palaemon (Hesperiidae)
2080
2050
88 Climatic Risk Atlas of European Butterflies
Carterocephalus silvicolus (MEIGEN, 1829) —
Northern Chequered Skipper
6c Full dispersal No dispersal
f SEDG -1008 (-18.35%) | -2364 (-43.04%)
-1928 (-35.1%) | -3609 (-65.7%)
© Otakar Kudrna Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 5493 cells)
The Northern Chequered Skipper can be found flying on the damp, flower-rich grass of woodland
rides. They can often be seen drinking nectar from various speedwells (Veronica spp.). The eggs are
laid on various coarse-leaved grasses. The caterpillar lives in a shelter spun from a blade of grass.
In the last larval instar, the caterpillar makes a shelter from withered leaves in which it hibernates.
The pale-yellow pupa is suspended in a silken girdle in the vegetation. It pupates in the spring. The
Northern Chequered Skipper is single-brooded.
Present distribution can be well explained by climatic variables (AUC = 0.93).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 edd 4000 0 add 4000 0 odd 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
89
Carterocephalus silvicolus (Hesperiidae)
2080
2050
90 Climatic Risk Atlas of European Butterflies
Thymelicus lineola (OCHSENHEIMER, 1806) — Essex Skipper
a
-2839 (-14.61%) | -4045 (-20.82%)
SEDG -3425 (-17.63%) -5330 (-27.43%)
BAMBU_ | -5580 (-28.72%) | -7891 (-40.61%)
-7605 (-39.14%) | -10551 (-54.3%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 19431 cells)
The Essex Skipper occurs in many sorts of flower-rich places with tall grass, such as grasslands, road
verges and on banks of streams. Because these butterflies need quite a lot of nectar, they can often
be seen drinking on thistles, knapweeds and other purple or pink flowers rich in nectar. The female
deposits her eggs in groups in the leaf-sheaths of coarse grasses, and lays them when the foodplants
are already withered. It is the eggs that hibernate. The development of the caterpillar takes place
the following spring. At first, the caterpillar only feeds during the day, but later also at night. They
pupate in the vegetation. The pupa may be suspended from a grass blade by a silken girdle, but also
just by the tip. The Essex Skipper is single-brooded.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.75).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
a
dd 4000 0 edd 4000 0 fad 4000 0 odd 4000
Minimum
Swe
‘1.6
10 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 0.2
Swe
06
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
91
Thymelicus lineola Hesperiidae)
2080
2050
92 Climatic Risk Atlas of European Butterflies
Thymelicus sylvestris (Popa, 1761) — Small Skipper
a
-2283 (-18.52%) | -4097 (-33.23%)
SEDG -3060 (-24.82%) | -4995 (-40.51%)
BAMBU_ | _ -4395 (-35.65%) | -6533 (-52.99%)
-5406 (-43.85%) | -8045 (-65.25%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 12329 cells)
The Small Skipper occurs in all sorts of flower-rich places with tall grass, such as grasslands, road
verges and edges of streams. The butterflies are fond of visiting thistles, knapweeds and other
purple or pink flowers for their nectar. The female deposits her eggs in batches of three to twenty
in the leaf-sheaths of coarse-leaved grasses. Unlike the Essex Skipper, she only uses young, green
leaves. After about three weeks, the eggs hatch and the small caterpillars begin spinning a shelter
straightaway 1n which to hibernate. Only the next spring do they begin to feed and grow. At first,
they only feed during the day and later also at night. They pupate in a web of loosely spun white
threads. The Small Skipper is single-brooded, but the butterflies emerge over a long period.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.72).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
dd 4000 0 edd 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Swe
0.6
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
93
Thymelicus sylvestris (Hespertidae)
2080
2050
94 Climatic Risk Atlas of European Butterflies
Thymelicus acteon (ROTTEMBURGER, 1775) — Lulworth Skipper
.—ClcLUwA Full dispersal No dispersal
Be SEDG 222 (2.27%) -1744 (-17.79%)
BAMBU -456 (-4.65%) | -2089 (-21.31%)
-445 (-4.54%) | -2359 (-24.07%)
SEDG -827 (-8.44%) | -2875 (-29.33%)
BAMBU_ | -2148 (-21.92%) | -4352 (-44.4%)
-2860 (-29.18%) | -5473 (55.84%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 9801 cells)
The Lulworth Skipper can be found on warm, dry grassland, with bushes or scrub nearby, or at the
edge of woodland. In the north of its range, it is mostly found on calcareous grassland. Eggs are
laid on the withered leaves of many grasses. Directly after hatching, the small caterpillar spins itself
a cocoon in which to hibernate. It does not begin to eat and grow until the following spring. It then
builds itself a shelter by spinning blades of grass together, which it only leaves when looking for
food. When fully grown, it pupates, changing into a green pupa suspended in the vegetation by a
silken girdle. The Lulworth Skipper is mostly single-brooded.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.76).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
a
todd 4000 0 edd 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
1.0 02
0.6
Swe
1.0 0.2
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
95
Thymelicus acteon (Hesperiidae)
2080
2050
96 Climatic Risk Atlas of European Butterflies
Hesperia comma (LINNAEUS, 1758) — Silvet-spotted Skipper
eae
-2321 (-19.66%) | -3408 (-28.87%)
SEDG -4598 (-38.95%) | -6026 (-51.04%)
BAMBU 5466 (-46.3%) | -7294 (-61.78%)
-7069 (-59.88%) | -9066 (-76.79%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 11806 cells)
The Silver-spotted Skipper occurs on open, poor grasslands, heathlands, and on sparsely covered
blown sand. The habitat varies from dry to moist, but is never rich in nutrients. However, in their
search for nectar, the skippers do visit nutrient-rich areas where flowers are growing, usually not
so far from their breeding ground. Eggs are laid on various fine-leaved grasses, passing the winter
in this stage. In the spring, the caterpillars emerge and spin a shelter from blades of grass in which
they spend most of their time, usually coming out to feed at night. When fully-grown, the caterpillar
spins a cocoon of silk and grass blades near the ground in which it pupates. The Silver-spotted
Skipper has one generation a yeat.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.77).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
0.6
1.0 02
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
E 1.0 0.2 0.6 fi. 1.0
1} —}—
|
Swe
o2
t=]
2000 40000 2000 40000 2000 40000 2000 4000
Gdd
Gdd Gdd Gdd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
97
Hesperia comma (Hesperiidae)
2080
2050
98 Climatic Risk Atlas of European Butterflies
Ochlodes sylvanus (EsPER, 1777) — Large Skipper
hc Full dispersal No dispersal
Be SEDG -726 (-3.53%) -3276 (-15.94%)
BAMBU -953 (-4.64%) -3309 (-16.1%)
-1398 (-6.8%) | -4058 (-19.74%)
SEDG -750 (-3.65%) | -4881 (-23.75%)
BAMBU_ | -2082 (-10.13%) | -6893 (-33.54%)
-3892 (-18.94%) | -9349 (-45.49%)
© Peter Ginzinger Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 20554 cells)
The Large Skipper is found on sunny, grassy vegetation in or near woods and scrub. It often visits
flowers, and the development of some rough vegetation close to the breeding ground is thus
favourable. They are especially fond of bramble blossom for nectar. The female deposits her eggs
onto the blades of coarse-leaved grasses one by one. The caterpillars make a shelter by spinning
blades of grass together, and pass the winter in the third larval instar. When fully grown, they also
spin leaves and silk together, to form a tube-like shelter in which to pupate. The Large Skipper is
single-brooded.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.75).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
dd 4000 0 edd 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
10 02
0.6
Swe
1.0 0.2
Swe
06
10 0.2
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
06
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
99
Ochlodes sylvanus (Hesperiidae)
2080
2050
100 Climatic Risk Atlas of European Butterflies
Gegenes pumilio (HOFFMANSEGG, 1804) — Pigmy Skipper
[eaten [ neato
-15 (-2.86%) -201 (-38.29%)
SEDG 90 (17.14%) -220 (-41.9%)
BAMBU | -123 (-23.43%) | -342 (-65.14%)
-223 (-42.48%) -420 (-80%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 525 cells)
The Pigmy Skipper occurs in the same types of habitat as the Mediterranean Skipper (Gegenes
nostrodamus). Dry gullies, rocky slopes, sandy or stony riverbanks with sparse vegetation, dry grassland
and low scrub near the coast are all places where these butterflies can be seen, often basking 1n the
sun on stones and bare soil. In spring, there are fewer butterflies than in the summer and autumn.
The larvae feed on the grasses Hyparrhenia hirta and Sorghum halepensis. Vhe Pigmy Skipper has two
to three generations a year.
Present distribution can be very well explained by climatic variables (AUC = 0.96).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
-
dd 4000 0 odd 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 O.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 02
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
101
Gegenes pumilio (Hesperiidae)
(1a)
504s
(ZV)
nNanva
2080
2050
102 Climatic Risk Atlas of European Butterflies
Gegenes nostrodamus (Fasrictus, 1793) — Mediterranean Skipper
CL Full dispersal No dispersal
Be SEDG -628 (-51.27%) -805 (-65.71%)
BAMBU -626 (-51.1%) -828 (-67.59%)
-718 (58.61%) | -927 (-75.67%)
SEDG -731 (-59.67%) | -1005 (-82.04%)
BAMBU -912 (-74.45%) | -1142 (-93.22%)
-1035 (-84.49%) | -1216 (-99.27%)
© Thomas Kissling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1225 cells)
Like the Pigmy Skipper (G. pumilo), the Mediterranean Skipper can be found 1n rocky or stony
places, such as dried-up riverbeds, on dry, grassy vegetation or in thickets, and low scrub along
the coast. The males defend their territory perched on a stone or on the ground, returning to the
same spot if disturbed. These butterflies fly fast and close to the ground. The larvae feed on Reed
(Phragmites australis) and the grass Saccharum ravennae. It has one to three generations a year. There are
far fewer butterflies in the spring than in the summer or autumn.
Present distribution can be well explained by climatic variables (AUC = 0.94).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
-
‘dd 4000 0 edd 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Gegenes nostrodamus (Hesperiidae) 103
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
104 Climatic Risk Atlas of European Butterflies
Zerynthia rumina (LINNAEUS, 1767) — Spanish Festoon
a oe
-1378 (-39.37%) | -1932 (-55.2%)
-1187 (-33.91%) | -2196 (-62.74%)
SEDG -1574 (-44.97%) | -2617 (-74.77%)
BAMBU_ | -2119 (-60.54%) | -3078 (-87.94%)
-1967 (-56.2%) | -3405 (-97.29%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 3500 cells)
The Spanish Festoon can be found on scrub and dry, grassy vegetation. The butterflies are often
found in rocky areas, where they can be seen, wings widespread, basking on stones, warming
themselves in the sun. The female lays her eggs singly or in small groups on birthworts (Aristolochia
spp.), such as A. longa, A. rotunda, and A. pistolochia. The caterpillars feed on these poisonous
plants, pupating for the winter. It can sometimes take two to three years before the butterfly
emerges from the pupa. Apart form a few sites in southern Spain the species 1s single brooded.
Present distribution can be well explained by climatic variables (AUC = 0.91).
Climate risk category: HHHR
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
3 a p4 Pa _
is}
oc
S : . . , —_ : ? ;
_
ah
o
o
w
o
fad 4000 0 edd 4000 0 add 4000 0 Gd 4000
Minimum
Swe
06
Swe
0.6 1.0 Oo2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
105
Zerynthia rumina (Papilionidae)
2080
2050
106 Climatic Risk Atlas of European Butterflies
Zerynthia polyxena ({[SCHIFFERMULLER], 1775) — Southern Festoon
SEDG 4158 (84.87%) | -1612 (-32.9%)
BAMBU 3432 (70.06%) | -3156 (-64.42%)
4141 (84.53%) | -3917 (-79.96%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 4899 cells)
The caterpillars of the Southern Festoon live on various birthworts, such as Anstolochia clematilis, A.
rotunda, A. palhda, and A. pistolochia. Because their foodplants grow in different habitats, and because
the caterpillars also have different foodplants in different areas, this spring butterfly can be found in
quite different habitats. The eggs are laid singly or in small groups on the underside of the leaves,
where the caterpillars are usually also found. The caterpillars have a striking appearance. Otherwise
beige with black spots, it has some orange tubercles on each segment, each ending in a black, spiny
tuft. The Southern Festoon ts single-brooded and hibernates as pupa.
This species is listed in Annex IV of the Habitats’ Directive.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.85).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
‘1.6
Swe
E 1.0 0.2 0.6 10 O02 fh 1.0
q
| p< <
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 0.2
Swe
06
o2
L=]
2000 40000 2000 40000 2000 40000 odd 4000
Gdd Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
107
Zerynthia polyxena (Papilonidae)
2080
2050
108 Climatic Risk Atlas of European Butterflies
Zerynthia cerisyi (GopaRT, 1822) — Eastern Festoon
Cc Full dispersal No dispersal
Hi SEDG 1685 (291.02%) -11 (-1.9%)
BAMBU_ | 2079 (359.07%) -10 (-1.73%)
2080 (359.24%) -23 (-3.97%)
SEDG 2970 (512.95%) -35 (-6.04%)
BAMBU | 3022 (521.93%) | -73 (-12.61%)
4073 (703.45%) | -71 (-12.26%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 579 cells)
The Eastern Festoon occurs in warm, usually dry, places such as in dry grassland with scattered
bushes, in scrub, near hedges, on agricultural land, vineyards and olive groves. It is also found in river
valleys. Choosing plants growing near bushes or trees, the female lays her eggs mostly on Birthwort
(Anstolochia clematitis), but also on other birthworts. The Eastern Festoon has one generation a year
and passes the winter in the pupal stage.
Present distribution can be well explained by climatic variables (AUC = 0.91).
Climate risk category: LR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
-
faa 4000 0 odd 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 O.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
109
Zerynthia cerisyt (Papilonidae)
2080
2050
110 Climatic Risk Atlas of European Butterflies
Parnassius mnemosyne (LINNAEUS, 1758) — Clouded Apollo
.6—hlcLUw Full dispersal No dispersal
fe SEDG -465 (-5.1%) -2308 (-25.34%)
-1793 (-19.68%) | -2980 (-32.71%)
|GRAS | -1983 (-21.77%) | -3222 (-35.37%)
SEDG -175 (-1.92%) | -4413 (48.45%)
BAMBU_ | _ -2536 (-27.84%) | -6269 (-68.82%)
-2163 (-23.75%) | -7025 (-77.12%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 9109 cells)
The Clouded Apollo occurs in mountainous regions on damp to moderately dry grassland, usually
with woodland or scrub in the neighbourhood. The butterflies can often be seen visiting red or
purple flowers for the nectar they need. The foodplant is Corydads, that at the time of egg laying is
not yet above ground. The eggs are laid on its dried stems, on grass blades, or on other plants not
too far from the foodplants. The egg hibernates. In the spring, as soon as it has released itself from
the egg, the small caterpillar starts its search for a suitable foodplant. When fully-grown, it pupates
in a closely spun cocoon of fine threads, situated above the ground in the leaves of the foodplant.
The Clouded Apollo has one generation a year.
This species is listed in Annex IV of the Habitats’ Directive.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.77).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘dd 4000 0 odd 4000 0 add 4000 0 odd 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
111
Parnassius mnemosyne (Papilionidae)
2080
2050
112 Climatic Risk Atlas of European Butterflies
Parnassius phoebus (Fasrictus, 1793) — Small Apollo
.—hCcLw Full dispersal No dispersal
Be SEDG -275 (-24.71%) -455 (-40.88%)
BAMBU -198 (-17.79%) | -388 (-34.86%)
-314 (-28.21%) | -488 (-43.85%)
SEDG -526 (-47.26%) | -755 (-67.83%)
BAMBU -364 (-32.7%) -645 (57.95%)
574 (-51.57%) | -804 (-72.24%)
© Jostein Engdal Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1113 cells)
The Small Apollo occurs in the mountains, especially in damp places, such as beside streams and
where it gets flooded from time to time. Such spots are the habitat of the larval food plant Yellow
Mountain Saxifrage (Saxzfraga aixoides). However, 1n the Mercantour in the south-west of the Alps,
the foodplant is Roseroot (Rhodiola roseum). Although the eggs are sometimes laid on the foodplant,
they are also often laid not far from it. The very small caterpillar sometimes passes the winter in
the egg, sometimes outside it. The fully-grown caterpillars are black with a row of orange-red or
yellow spots along each side. At the beginning of the summer, they spin a fluamsy cocoon in which
to pupate, either low down on the larval plant, or on the ground. The Small Apollo is single-
brooded.
Present distribution can be very well explained by climatic variables (AUC = 0.99).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘dd 4000 0 edd 4000 0 oad 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
113
Parnassius phoebus (Papilionidae)
2080
2050
114 Climatic Risk Atlas of European Butterflies
Parnassius apollo (LiNNAEus, 1758) — Apollo
a
-2915 (-46.47%) | -3479 (-55.46%)
SEDG -2234 (-35.61%) | -3431 (54.69%)
BAMBU_ | -3319 (-52.91%) | -4177 (-66.59%)
-3972 (-63.32%) | -4758 (-75.85%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 6278 cells)
The Apollo occurs in areas on steep, sunny slopes with sparse vegetation. In Europe, there are
many different subspecies, forms and aberrations, because of the large isolation of populations.
The butterflies are fond of visiting thistles and other flowering plants for their nectar. The female
lays her eggs singly or in small groups on or near the foodplant stonecrop (Sedum spp.). The eggs
develop but the tiny caterpillar hibernates inside the eggshell or as newly hatched larvae. It emerges
in the spring, and starts feeding on the buds of the foodplant. The caterpillars of later instars also
eat the leaves. When it is time to pupate, the caterpillars look for a safe place between the stones,
where they then spin a flimsy cocoon for pupation. The Apollo is single-brooded.
This species is listed in Annex IV of the Habitats’ Directive.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.8).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
1.0
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
Swe
0.6
0 2000 40000 2000 40000 2000 40000 2000 4000
Gdd d Gdd
Gdd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Parnassius apollo (Papilionidae) 115
(B1)
SEDG
GRAS
(A1Fl)
116 Climatic Risk Atlas of European Butterflies
Iphiclides podalirius (LINNAEUus, 1758) — Scarce Swallowtail
a
:
GRAS
3058 (29.47%) | -1226 (-11.81%)
SEDG 4380 (42.2%) | -1186 (-11.43%)
BAMBU 3105 (29.92%) | -3182 (-30.66%)
3433 (33.08%) | -4169 (-40.17%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 10378 cells)
This large, conspicuous butterfly with its elegant gliding flight, is very impressive. It occurs in
warm, dry places with scrub and rough vegetation. The males of the Scarce Swallowtail congregate
on hilltops, dancing in the air and waiting for the females, a type of behaviour known as “hill-
topping”. They visit thistles and other flowers rich in nectar. The eggs are laid on the leaves of
small bushes or trees of Blackthorn (Prunus spinosa) and other species of Prunus. The caterpillars
feed on the leaves. When fully-grown, they pupate, the pupa suspended in a silken girdle in the
foodplant. Depending on its position in the range and on the altitude, the Scarce Swallowtail has
one to three generations a year.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.76).
Climate risk category: LR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
-
‘dd 4000 0 odd 4000 0 Gad 4000 0 odd 4000
Minimum
Swe
06
1.0 02
0.6
Swe
1.0 0.2
Swe
0.6
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Iphichides podahrius (Papilionidae) 117
118 Climatic Risk Atlas of European Butterflies
Papilio machaon LinNaAEus, 1758 — Swallowtail
oe
1206 (5.65%) -428 (-2.01%)
SEDG 2637 (12.36%) -809 (-3.79%)
BAMBU 1066 (5%) -2547 (-11.94%)
227 (1.06%) -4167 (-19.53%)
© Peter Ginzinger Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 21332 cells)
The Swallowtail is a species of flower-rich meadows and small-scale farmland. The males and
females meet at elevated places or above high trees, showing a type of behaviour known as “hill-
topping”. The butterfly of the Swallowtail needs a lot of nectar. Eggs are laid on various umbellifers,
including Wild Carrot (Daucus carota). The fully-grown larvae are very conspicuous, bright green
with black stripes and orange spots. Caterpillars from autumn broods leave the foodplant to pupate
on the ground and hibernation takes place as a pupa. The caterpillars of the summer broods pupate
low down on the larval foodplant. Depending on geographic setting, the Swallowtail produces one
to three generations a year.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.67).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
she
=|
toad 4000 0 edd 4000 0 ad 4000 0 odd 4000
1.0
Minimum
Swe
06
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe Swe
0.6 0.6
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Papilio machaon (Papilionidae) 119
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1FI)
120 Climatic Risk Atlas of European Butterflies
Papilio alexanor Esper, 1799 — Southern Swallowtail
.—hCcLUwk Full dispersal No dispersal
Be SEDG -224 (-32.61%) -404 (-58.81%)
BAMBU -277 (-40.32%) | -436 (-63.46%)
-278 (-40.47%) | -454 (-66.08%)
SEDG -257 (-37.41%) | -455 (-66.23%)
BAMBU -380 (-55.31%) | -534 (-77.73%)
62 cara) | 592 Coa
© Josef Pennerstorfer Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 687 cells)
The Southern Swallowtail is mostly found on warm, dry calcareous slopes with a flower-rich
vegetation and low-growing bushes. They prefer slopes that are steep and rocky. Different
foodplants are known, all of them umbellifers. Pahots saxifraga 1s the most important one in the
western part of its range, but eggs are also laid on Opopanan chironium, Seseh montanum, and Trinia
glauca. In the eastern part, the caterpillars feed mostly on various of fennels (Feria spp.), and
also on Opopanax hispidus, Burnet saxifrage (Pimpinella saxifraga), Scaligeria cretica, and Wild Parsnip
(Pastinaca sativa). Vhe caterpillars eat the flowers and ripening seeds. The Southern Swallowtail is
single-brooded and passes the winter in the pupa stage.
This species is listed in Annex IV of the Habitats’ Directive.
Present distribution can be well explained by climatic variables (AUC = 0.94).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
1.0
Swe
o.2 0.6
1.0
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
Swe
0.6
0.2
0 2000 40000 2000 40000 2000 40000 2000 4000
Gdd G Gdd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Papilio alexanor (Papilionidae) 121
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
2050 2080
122 Climatic Risk Atlas of European Butterflies
Leptidea sinapis (LINNAEUS, 1758) / reali REIssSINGER, 1990
(complex) — Wood White
SEDG 1829 (10.09%) | -2394 (-13.21%)
BAMBU 236 (1.3%) -2733 (-15.08%)
ose | ass co
SEDG 2744 (15.14%) | -3709 (-20.46%)
BAMBU 1117 (6.16%) | -5664 (-31.25%)
795 (4.39%) -7192 (-39.68%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 18126 cells)
At present, the Wood White is divided into two species, L. stnapis (LINNAEUS, 1758) and L. reah
REISSINGER, 1990, which are indistinguishable in the field. Their distribution is not yet entirely clear.
Their life cycles are similar. These fragile butterflies occur on damp, warm grassland near bushes and
scrub. They lay their thin, spindle-shaped eggs on different sorts of Lathyrus and birdsfoot-trefoil
(Lotus spp.). Cream, turning to bright yellow as they mature, they are easy to find. The pupa ts pale-
green, and the winter is spent in this stage. In the north of its range, it is single-brooded, but in the
middle it has two generations a year, and in the south sometimes three.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.63).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
=
sh
a
0
=
Ow
=a
ie]
sit}
oO
e Ec I
‘dd 4000 0 Codd 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Leptidea sinapis / reali (Pieridae) | 123
SEDG
(B1)
GRAS
(A1FI)
124 Climatic Risk Atlas of European Butterflies
Leptidea duponcheli (StauDINGER, 1871) — Eastern Wood White
6c Full dispersal No dispersal
Be SEDG 221 19.19%) -712 (-63.74%)
BAMBU_ | -186 (-16.65%) | -812 (-72.69%)
22 (1.97%) -856 (-76.63%)
SEDG 32 (2.86%) -782 (-70.01%)
BAMBU -628 (-56.22%) | -1009 (-90.33%)
-594 (-53.18%) | -1066 (-95.43%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1117 cells)
The Eastern Wood White occurs on dry grassland, often where scattered trees or bushes are growing,
and also in open woods. It is found on both calcareous and non-calcareous soils. The habitats of the
Eastern Wood White are drier and warmer than those of the Wood White (L. scmapis complex).
The eggs are laid on the leaves of Meadow Vetchling (Lathyrus pratensis), Yellow Vetchling (L.
aphaca) and birdsfoot-trefoils (Lois spp.), mostly on plants growing in the shade. The caterpillars
feed on the leaves of the foodplant. The Eastern Wood White is double-brooded and hibernates
in the pupal stage.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.69).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
pm
o
xh
o
aad 4000 0 odd 4000 0 add 4000 0 add 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
Swe
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
125
Leptidea duponcheh (Pieridae)
2080
2050
126 Climatic Risk Atlas of European Butterflies
Leptidea morsei (FENTON, 1881) — Fenton’s Wood White
SEDG 188 (12%) -1154 (-73.64%)
BAMBU 761 (48.56%) -988 (-63.05%)
370 (23.61%) -893 (-56.99%)
SEDG 293 (18.7%) -1416 (-90.36%)
BAMBU 196 (12.51%) | -1328 (-84.75%)
-133 (-8.49%) | -1198 (-76.45%)
© Helmut Héttinger Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1567 cells)
Apart from its greater size and slightly falcate forewings the species can be separated from the other
Wood Whites by conspicuous gliding flight sometimes displayed by males. The habitat of Fenton’s
Wood White is almost exclusively highly structured deciduous woodland. The only confirmed food
plant of this species in Europe is Black Pea (Lathyrus niger). This butterfly is double-brooded and
hibernates 1n the pupal stage.
This species is listed in Annexes II and IV of the Habitats’ Directive.
Present distribution can be well explained by climatic variables (AUC = 0.92).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
a
_
cH
=)
a
=
ow
|
io
m™
oO
a : : ; — : ;
cH
o
H
o
toad 4000 0 odd 4000 0 add 4000 0 Gdd 4000
Minimum
Swe
06
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Leptidea morsei (Pieridae) 127
128 Climatic Risk Atlas of European Butterflies
Anthocharis cardamines (LINNAEUS, 1758) — Orange-tip
a
-2928 (-15.58%) | -4787 (-25.47%)
SEDG -2159 (-11.49%) | -4447 (23.66%)
BAMBU_ | -4765 (-25.35%) | -7476 (-39.78%)
-6918 (-36.81%) | -10040 (-53.42%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 18795 cells)
In Western Europe, the first Orange-tip gives us the feeling that spring has arrived. However, high
in the mountains, these butterflies only appear in early summer. The Orange-tip occurs in damp to
quite wet grasslands at the edge of woods, or near thickets. The eggs are laid singly on flowerheads
of different crucifers. At first, the caterpillars feed on the flowerbuds of the foodplant and later on
the fruits. They live alone, which accords with their cannibalistic nature. When ready to pupate, they
move into rough vegetation, climbing up a little twig and turning into an attractive light-brown pupa,
suspended from the plant by a silken girdle. The Orange-tip hibernates in this stage and has one
generation a yeat.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.7).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
‘sad 4000 0 odd 4000 0 oad 4000 0 ad 4000
Minimum
Swe
0.6
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
129
Alnthocharis cardamines (Pieridae)
2080
2050
130 Climatic Risk Atlas of European Butterflies
Anthocharis euphenoides STauDINGER, 1869
6c Full dispersal No dispersal
Hee SEDG -989 (-33.3%) -1744 (-58.72%)
-1299 (-43.74%) -1763 (-59.36%)
-1948 (-65.59%) | -2614 (-88.01%)
-1956 (-65.86%) | -2887 (-97.21%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2970 cells)
This butterfly inhabits warm, dry places with flower-rich, grassy vegetation and scattered bushes.
Eggs are laid one by one on the flowerbuds of various crucifers, such as the buckler mustards
Biscutella laevigata and B. auriculata, London Rocket (Szsymwbrium irio), and Hedge Mustard (S. officinale).
The caterpillars feed mainly on ovaries, but at times are also cannibalistic. This species hibernates as
a pupa and has one generation a yeat.
Present distribution can be well explained by climatic variables (AUC = 0.91).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
fed 4000 0 odd 4000 0 add 4000 0 ada 4000
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 O.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
131
Alnthocharis euphenoides (Pieridae)
2080
2050
132 Climatic Risk Atlas of European Butterflies
Anthocharis gruneri HERRICH-SCHAFFER, 1851—Gruner’s Orange-tip
CL Full dispersal No dispersal
Be SEDG -266 (-26.68%) -613 (-61.48%)
-330 (-33.1%) -648 (-64.99%)
-356 (-35.71%) | -696 (-69.81%)
-666 (-66.8%) -888 (-89.07%)
-TT3 (-77.53%) | -971 (97.39%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 997 cells)
Gruner’s Orange-tip lives on dry, open grasslands, on rocky, often calcareous slopes, scrub and
clearings in dry woodland. The eggs are laid on Aethionema species, including Burnt Candytuft
(A. saxatile) and A. orbiculatum, the caterpillars eating both the leaves and ripening seeds. It is
single-brooded and the pupa hibernates.
Present distribution can be well explained by climatic variables (AUC = 0.92).
Climate risk category: HHHAR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
dd 4000 0 odd 4000 0 add 4000 0 add 4000
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 O.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
133
Alnthocharis gruneri (Pieridae)
(1a)
504s
(ZV)
nNanva
(ld bY)
SVH9
2080
2050
134 Climatic Risk Atlas of European Butterflies
Zegris eupheme (Esper, 1805) — Sooty Orange-tip
6c Full dispersal No dispersal
Be SEDG -411 (-47.9%) -520 (-60.61%)
BAMBU -536 (-62.47%) -562 (-65.5%)
-456 (-53.15%) -580 (-67.6%)
SEDG -765 (-89.16%) | -766 (-89.28%)
BAMBU -837 (-97.55%) | -837 (-97.55%)
-849 (-98.95%) | -849 (-98.95%)
© Neil Thompson Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 858 cells)
The Sooty Orange-tip is mostly seen in dry, flower-rich places, waste ground, and abandoned
agricultural land. Crucifers, such as London Rocket (Szsywbrium irio), the buckler mustard Bescutel/la
auriculata, Hoary Mustard (Harschfeldia incana), and radishes (Raphanus spp.), ate usually abundant in
its habitat. They are used by the butterflies for their nectar and as larval foodplants. The butterflies
have a quick, zigzageine flight. The Sooty Orange-tip hibernates as a pupa. This pupal stage may last
for one, two, or three years. It is single-brooded.
Present distribution can be very well explained by climatic variables (AUC = 0.97).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 odd 4000 0 Cad 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
135
Zegris eupheme (Pieridae)
(ZV)
Nganva
(ld LV)
SVH9D
2080
2050
136 Climatic Risk Atlas of European Butterflies
Euchloe belemia (EsPEr, 1798) — Green-striped White
hc Full dispersal No dispersal
Be SEDG -289 (-41.58%) -377 (-54.24%)
BAMBU -424 (-61.01%) | -432 (-62.16%)
-400 (-57.55%) | -446 (-64.17%)
SEDG -338 (-48.63%) | -495 (-71.22%)
BAMBU -607 (-87.34%) | -626 (-90.07%)
-649 (-93.38%) -670 (-96.4%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 695 cells)
The Green-striped White occurs in flower-rich places, between bushes, on waste ground, and on
abandoned agricultural land. It occurs locally, but can be numerous in its flight area. The butterfly
flies quickly and close to the ground. The caterpillars eat the ripening seeds of different crucifers,
such as the buckler mustard Bescutella auriculata, candytufts [bers spp.) including Annual Candytuft
(I. amara), and rockets (Sisymbrium spp.). The Green-striped White has two broods a year in the
spring, and passes the winter as a pupa.
Present distribution can be very well explained by climatic variables (AUC = 0.96).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
tod 4000 0 odd 4000 0 aad 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Euchloe belemia (Pieridae) 137
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
138 Climatic Risk Atlas of European Butterflies
Euchloe ausonia (HUBNER, 1806) (complex) — Dappled White
en ea
-91 (-2.03%) -1080 (-24.09%)
[axas —|er9
SEDG 338 (7.54%) -1439 (-32.1%)
BAMBU -327 (-7.29%) | -2058 (-45.91%)
GRAS -265 (-5.91%) | -2472 (-55.14%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 4483 cells)
Due to data availability we here treat Ewch/oe ausonia (HUBNER, 1806), Euchloe simplonia Preyer, 1829,
and Euchloe cramert BUTLER, 1869, as the Dappled White species complex, knowing that they are
normally being treated as distinct species.
The Dappled Whites are species of warm, dry places. They occur on flower-rich grasslands and
waste ground, along hedges and the edges of woods. Various crucifers are used as larval foodplant,
such as a Charlock (Szsywbrium arvensis), Woad (Isatis tinctoria), buckler mustards (Bzscutella spp.),
Evergreen Candytuft (Iberis sempervirens), and Crested Bunias (Bumias erucago). The caterpillars feed on
the flowers and ripening seeds. They are double-brooded and hibernate as pupa.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.85).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
did 4000 0 odd 4000 0 oad 4000 0 ada 4000
1.0
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
Swe
o6 10 02
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
139
E:uchloe ausonia (Pieridae)
2080
2050
140 Climatic Risk Atlas of European Butterflies
Euchloe tagis (HUBNER, 1804) — Portuguese Dappled White
6c Full dispersal No dispersal
Be SEDG -590 (-18.72%) -1537 (-48.76%)
BAMBU -999 (-31.69%) | -1507 (-47.81%)
-794 (-25.19%) | -1692 (-53.68%)
SEDG -920 (-29.19%) | -1943 (-61.64%)
BAMBU -1535 (-48.7%) | -2350 (-74.56%)
-1749 (55.49%) | -2743 (-87.02%)
© Jean Delacre Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 3152 cells)
This butterfly owes its specific name to the River Tagus in Portugal, along the banks of which
it was first found. The Portuguese Dappled White is found on warm, dry rocky places with
patches of flower-rich grassy vegetation, in dry scrub, and on abandoned agricultural land.
Various crucifers are used as foodplant, such as candytufts (/ders spp.), including [. chata,
I. saxatis and Annual Candytuft (I. amara), and buckler mustards (Bzscwiella spp.). The caterpillars
eat the ovaries and flowers. It has one brood a year and hibernates as a pupa.
Present distribution can be well explained by climatic variables (AUC = 0.86).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
‘dd 4000 0 edd 4000 0 add 4000 0 add 4000
Minimum
Swe
0.6
1.0 O02
Swe
0.6
1.0 0.2
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
141
E:uchloe tagis (Pieridae)
2080
2050
142 Climatic Risk Atlas of European Butterflies
Aporta crataegi (LINNAEUS, 1758) — Black-veined White
.—hlcLUw Full dispersal No dispersal
fi SEDG -661 (-3.52%) | -2426 (-12.91%)
-1530 (8.14%) | -3266 (-17.39%)
-2410 (-12.83%) | -6143 (-32.7%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 18785 cells)
The Black-veined White can be found on many different sorts of vegetation in a variety of landscapes,
but seems to prefer habitats in the neighbourhood of woods or scrub. Numbers of this migratory
species can fluctuate greatly and the reasons for this are not well understood. The species may build
up huge populations at unusual places such as hedges along motorways, but also disappear from
large areas for many years. The eggs are laid in large clusters of often sixty or more, on the leaves of
trees and bushes of the rose family (Rosaceae), such as hawthorn (Crafaegus spp.), Blackthorn (Prunus
spinosa), Cherry (Prunus spp.), Apple (Malus spp.), also in orchards. The caterpillars spend their time
ina communal silken nest until they pupate, the nest 1n which they hibernate being more substantial.
When they are ready to pupate, they disperse over the foodplant. The yellow pupa with black spots
suspends in a silken girdle. The Black-veined White has one brood a yeat.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.69).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
0.6
Swe
o2
0 2000 40000 2000 40000 2000 40000 2000 4000
Gd dl Gdd
Gdd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
143
Alporia crataegi (Pieridae)
2080
2050
144 Climatic Risk Atlas of European Butterflies
Pieris brassicae (LINNAEUS, 1758) — Large White
.—hlcLUw Full dispersal No dispersal
Be SEDG -1904 (-8.11%) -3228 (-13.75%)
BAMBU_ | -3133 (-13.34%) | -4027 (-17.15%)
-3241 (-13.8%) | -4419 (-18.82%)
SEDG -3429 (-14.6%) | -5640 (-24.02%)
BAMBU_ | -6368 (-27.12%) | -8473 (-36.08%)
-8906 (-37.93%) | -11751 (-50.04%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 23482 cells)
You can come across the Large White almost everywhere. It lays its eggs singly or in small clusters
on the underside of the foodplants, which are mainly broad-leaved Brassicaceae (like Brassica,
Crambe, Lunaria, Cakile) that offer enough food for the gregarious caterpillars, but also species of
the introduced genus Tropaeolum (Tropaeolaceae) which also contains glucosinolates On such a
nutritious diet, the caterpillar grows very quickly, reaching the pupal stage in three to six weeks. The
attractive white pupa is suspended in a silken girdle from the foodplant. The Large White has several
generations a year, the number depending on the geographical location and latitude of the flight
area, and the leneth of the summer.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.66).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
tad 4000 0 odd 4000 0 Cad 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 O.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 0.2
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Pieris brasswae (Pieridae)
SEDG
145
(B1)
GRAS
(A1Fl)
146 Climatic Risk Atlas of European Butterflies
Pieris krueperi (STAUDINGER, 1860) — Kriiper’s Small White
.—hCcLe Full dispersal No dispersal
Be SEDG 216 (16.44%) -619 (-47.11%)
BAMBU 11 (0.84%) -604 (-45.97%)
29%) | aR
SEDG 446 (33.94%) -748 (-56.93%)
BAMBU -141 (-10.73%) | -900 (-68.49%)
36 (2.74%) -1033 (-78.61%)
© Rudi Verovnik Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1314 cells)
Krueper’s Small White occurs 1n warm, dry places on calcareous, rocky slopes with scattered low,
herbaceous vegetation. Eggs are laid on the sepals of Golden Alyssum (44ssum saxatile), and
A. montanum. Vhe caterpillars feed on the ovaries. It hibernates as a pupa and has two or more
generations a yeat.
Present distribution can be well explained by climatic variables (AUC = 0.91).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
bs!
a
bie
2
aes
=
oH :
Oo
ba
ad
2
=]
bah}
o
2000, 4000 0 ord 4000 0 2000 4000 0 odd 4000
Minimum
Swe
06
Swe
0.6
Annual precipitation range
Swe
Maximum Large (66%) Small (33%)
Swe
16
Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
147
Pieris Rrueperi (Pieridae)
2080
2050
148 Climatic Risk Atlas of European Butterflies
Pieris mannit (MAYER, 1851) — Southern Small White
.—hCcLwe Full dispersal No dispersal
fi SEDG 214 (12.51%) -572 (-33.45%)
221 (12.92%) -716 (-41.87%)
639 (37.37%) | -1318 (-77.08%)
© Martin Wiemers Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1710 cells)
The Southern Small White, which looks very much like the Small White (P. rapae), occurs on both
calcareous and non-calcareous dry, grassy vegetation, in open scrub and open woodland. The
most important larval foodplants are the candytufts Evergreen Candytuft (Ibers sempervirens) and I.
saxatilsis,.although other crucifers, such as Adssoides utriculata, and mustards (Sznapis spp.) ate also
used. The eggs are laid singly or in small groups on the uppersides of the leaves. The caterpillars feed
on the leaves. It has several generations a year, pupating on plant stalks and stones. It overwinters as
a pupa, and the butterflies emerge in the spring.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.76).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
todd 4000 0 odd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
149
Pieris mannii (Pieridae)
2080
2050
150 Climatic Risk Atlas of European Butterflies
Pieris rapae (LINNAEUS, 1758) — Small White
.—hCcLUw Full dispersal No dispersal
Be SEDG 82 (0.34%) -1502 (-6.22%)
BAMBU -690 (-2.86%) -1742 (-7.22%)
-891 (-3.69%) -2258 (-9.35%)
SEDG -670 (-2.78%) | -3296 (-13.65%)
BAMBU_ | -3119 (-12.92%) | -6008 (-24.89%)
-4088 (-16.93%) | -8043 (-33.31%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 24148 cells)
The Small White is found in any sort of habitat where its larval foodplants grow, such as gardens,
parks, grasslands, heathland and woodland. The eggs are laid on the underside of the leaves of a
wide variety of crucifers, including cultivated brassicas, and also on Reseda species, such as Wild
Mignonette (R. 4wiea). The caterpillars grow very rapidly, sometimes pupating after two weeks. The
pupa hangs in a silken girdle, normally on vertical surfaces, and hibernates in this stage. The Small
White has several generations a year, depending on the geographical position and altitude of the
flight area, and the length of the summer.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.69).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
ead 4000 0 edd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
0.6
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 02
Swe
0.6
o.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
151
Pieris rapae (Pieridae)
2080
2050
152 Climatic Risk Atlas of European Butterflies
Pieris ergane (GEYER, 1828) — Mountain Small White
.—hClcLw Full dispersal No dispersal
He SEDG 94 (4.38%) -786 (-36.63%)
-101 (-4.71%) -932 (-43.43%)
-90 (-4.19%) -1043 (-48.6%)
SEDG 527 (24.56%) | -1157 (-53.91%)
BAMBU 223 (10.39%) | -1684 (-78.47%)
519 (24.18%) | -1894 (-88.26%)
© Rudi Verovnik Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2146 cells)
The Mountain Small White likes warm and dry places, and is found on dry, poor grasslands, in rocky
places, in open scrub and in large clearings in woods. The flight areas are usually on calcareous soil.
The males can sometimes be seen in large groups on damp ground. Compared to other whites, this
butterfly has a slow and measured flight. The main larval foodplant is Burnt Candytuft (Aeshionema
saxalile), butA. orbiculatum and Woad (Isatis tinctoria) are also used. The eggs are laid singly. The pupa
is normally fixed to stones and rocks. There are two or three generations a year, and the pupa of the
autumn generation hibernates.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.84).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
dd 4000 0 edd 4000 0 add 4000 0 add 4000
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Pieris ergane (Pieridae) 153
(B1)
SEDG
GRAS
(A1Fl)
154 Climatic Risk Atlas of European Butterflies
Pieris napi (LINNAEvs, 1758) — Green-veined White
[rear [Rea
-3275 (-12.38%) | -3819 (-14.43%)
SEDG -3817 (-14.43%) | -4424 (-16.72%)
BAMBU_ | -5386 (-20.36%) | -6063 (-22.91%)
-7583 (-28.66%) | -8253 (-31.19%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 26459 cells)
The Green-veined White occurs in many different habitats and landscapes with varying amounts of
shelter. It can be found on all types of grasslands and heathlands in open or more closed landscapes.
However, a too dry habitat is not favourable. This butterfly is very variable in appearance, having
several forms and subspecies. The eggs are laid singly or in small numbers on many, mostly wild,
crucifers It pupates on a stalk and overwinters in this stage. The Green-veined White has several
broods a year, the number depending on the geographical location and altitude of the flight area,
and the length of the summer.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.73).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
1.6
|
aq =
oy a
fee) i
ee
ef@eEs
o= mH
=e
£5 8
Oo. 4
of 7
D>
o mq = a
ger
a |
tage
a
@e-
2 o
wv
x é Pon]
= 4
o
0 2000 4000 2000 40000 2000 40000 2000 4000
Gd id Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
155
Pieris napi (Pieridae)
2080
2050
156 Climatic Risk Atlas of European Butterflies
Pieris bryoniae (HUBNER, 1791) — Mountain Green-veined White
a de
521 (-44.04%) | -594 (-50.21%)
SEDG -312 (-26.37%) | -637 (-53.85%)
BAMBU -197 (-16.65%) | -655 (-55.37%)
-303 (-25.61%) | -844 (-71.34%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1183 cells)
The butterfly of the Mountain Green-veined White strongly resembles that of the Green-veined
White (P. napi). The eggs, caterpillars and pupae are also almost indistinguishable. In some places,
such as in the Alps and the Jura Mountains, these species occur together. The Mountain Green-veined
White is found in flower-rich grasslands along river banks, and at woodland edges. Buckler Mustard
(Bescutella laevigata), pennycresses (Thlaspi spp.) and bittercresses (Cardamine spp.) ate used as foodplants.
The female, heavily dusted on her upperside with yellow or grey, lays her eggs singly on the flowers or
leaves of the larval foodplant. She shows a preference for smaller plants growing on poor ground. It
has one or two generations a year and hibernates 1n the pupal stage. In areas where this species occurs
together with P. napi (like in the SE Alps), one may often find hybrids of both species.
Present distribution can be well explained by climatic variables (AUC = 0.92).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
‘dd 4000 0 edd 4000 0 add 4000 0 ada 4000
Minimum
Swe
0.6
o2
Swe
0.6 1.0
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
Swe
0.6
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
157
Pieris bryoniae (Pieridae)
2080
2050
158 Climatic Risk Atlas of European Butterflies
Pontia callidice (HiBNER, 1800) — Peak White
CL Full dispersal No dispersal
Be SEDG -182 (-20.43%) -347 (-38.95%)
BAMBU -97 (-10.89%) -296 (-33.22%)
-211 (-23.68%) | -370 (-41.53%)
SEDG -452 (-50.73%) | -639 (-71.72%)
BAMBU -280 (-31.43%) | -509 (-57.13%)
-481 (-53.98%) | -682 (-76.54%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 891 cells)
The Peak White is found high in the mountains in flower-rich grasslands. The eggs are laid singly
on Alpine Bittercress (Cardamine belhdifola ssp. alpina), Hutchinsia alpina, Erysimum helveticum, and
Reseda glauca, especially on those growing on open stony patches. The caterpillars mostly eat the
lower leaves, and hide under stones when the weather is bad. In the Alps, the Peak White usually
has only one generation a year and hibernates as a pupa. However, in good summers or at lower
altitudes, a partial second generation is sometimes produced that passes the winter as a caterpillar.
In the Spanish Pyrenees, two generations a year are usual, and it hibernates either as a pupa or a
caterpillar.
Present distribution can be very well explained by climatic variables (AUC = 0.98).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
_
cH
=)
a
=
Ow
=a
io
m™
Oo
a : : : : i ;
4
o
=
os
o
‘aad 4000 0 edd 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
159
Pontia callidice (Pieridae)
2080
2050
160 Climatic Risk Atlas of European Butterflies
Pontia daplidice (LINNAEUs, 1758) / edusa (Fasrictus, 1777)
(complex) — Bath White
.—hCcLUwEe Full dispersal No dispersal
fi SEDG 4490 (29.33%) -364 (-2.38%)
S
Ys)
3091 (20.19%) | -2391 (-15.62%)
© Peter Ginzinger Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 15310 cells)
The Bath White complex has been split into two species in the 1980s, P. daphdice (LINNAEUS, 1758), which
mainly occurs 1n the Western Mediterranean, and P. edusa (Fasrictus, 1777), which 1s distributed over the rest
of Europe. However, these two species cannot be distinguished tn the field. Their life cycles are also very
similar. The adult butterflies are very mobile, fast flying and migrate northwards and westwards 1n warm
years. They occur in warm, stony places; often disturbed ground, such as road verges, abandoned agricul-
tural land or quarries. The females lay their eggs on such crucifers as mustards (Sznapis spp.), Adssum spp.,
and also on mignonettes (Reseda spp.). The caterpillars feed mainly on the flowers and seeds. Pupation takes
place on the stalk of the foodplant. These species have two or more broods a year and hibernate as pupae.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.72).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6.
Maximum Large (66%) Small (33%)
td
o
=I
ze
o
™
o
ae 4000 0 ad 4000 0 iy 4000 0 ee 4000
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Pontia daplidice | edusa (Pieridae) | 161
(B1)
SEDG
GRAS
(AFI)
162 Climatic Risk Atlas of European Butterflies
Colias phicomone (Esper, 1780) — Mountain Clouded Yellow
.6—hCcLUwe Full dispersal No dispersal
fi SEDG -306 (-21.5%) -513 (-36.05%)
BAMBU_ | -209 (-14.69%) | -450 (-31.62%)
-432 (-30.36%) | -611 (-42.94%)
SEDG -562 (-39.49%) | -890 (-62.54%)
BAMBU_ | -426 (-29.94%) | -781 (-54.88%)
-665 (-46.73%) | -1017 (-71.47%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1423 cells)
The Mountain Clouded Yellow has its breeding ground in flower-rich alpine meadows. Although
they are quick, strong flyers, they are not migrants and do not leave their habitat. Various leguminous
plants are used as larval foodplant, including Horseshoe Vetch (Hippocrepis comosa), White Clover
(Trifokum repens), Common Birdsfoot-trefoil (Lotus corniculatus), and vetches (Vicia spp.). This butterfly
species usually only has one generation a year, but in some years there is a partial second brood. The
caterpillars hibernate after their second moult.
Present distribution can be very well explained by climatic variables (AUC = 0.96).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
si
=|
° = : -
‘dd 4000 0 edd 4000 0 aad 4000 0 ada 4000
1.0
Minimum
Swe
0.6
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe Swe
0.6 0.6
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Colas phicomone (Pieridae) 163
164 Climatic Risk Atlas of European Butterflies
Colias palaeno (LINNAEUS, 1758) — Moorland Clouded Yellow
a
SEDG -3370 (-34.09%) | -3889 (-39.34%)
BAMBU -4202 (-42.5%) | -4562 (-46.15%)
-5087 (51.46%) | -5433 (54.96%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 9886 cells)
The Moorland Clouded Yellow occurs at the edges of raised bogs, and also on blanket bogs. At
high altitudes, it is found in drier habitats. Eggs are laid singly on Bog Whortleberry (Vaccinium
uliginosum). At first, the caterpillars only eat the upper layers of the leaf , producing “windows”, but
later, the whole leaf is eaten. The caterpillars hibernate among the dry leaves of the litter layer, and
the next spring feed and grow further, before finally pupating on a branch of the foodplant. It has
one generation a yeat.
Present distribution can be well explained by climatic variables (AUC = 0.9).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
‘1.6
sh
a i=)
=) a
ee
Pe Oe
ae =a
o= mH
=e
£5 3
Oo. 4
os
©
aes
Ga”
= = rm
ae
qf. =
22
wo
x é P=
= 4
o
0 2000 40000 2000 40000 2000 40000 2000 4000
Gd id Gadd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
165
Colas palaeno (Pieridae)
2080
2050
166 Climatic Risk Atlas of European Butterflies
Colias erate (EsPErR, 1805) — Eastern Pale Clouded Yellow
CL Full dispersal No dispersal
fi SEDG 2797 (116.83%) -331 (-13.83%)
BAMBU 815 (34.04%) | -1199 (-50.08%)
1982 (82.79%) | -989 (-41.31%)
SEDG 1379 (57.6%) | -1160 (-48.45%)
BAMBU | -317 (-13.24%) | -2059 (-86.01%)
44 (1.84%) -22.60 (-94.4%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2394 cells)
The Eastern Pale Clouded Yellow is a very mobile species and can therefore be seen in all sorts
of places. This species greatly extended its range during the 1970s and 80s. 50 years ago this
mainly Asian species was known in Europe only from the Black Sea coast. Lucerne (Medicago
saliva) 1s its most important larval foodplant and they are most numerous in areas where it 1s
grown. The caterpillars can also be found on other leguminous plants. This species has several
generations a year; three to five have been reported from Bulgaria and Romania. This species passes
the winter as a caterpillar (probably without diapause).
Present distribution can be well explained by climatic variables (AUC = 0.92).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
=o
wo
E35
=H
tl
= s
o
=,
ow
=o
wo
tH
o
z : = . : : ;
oa
=
w
dd 4000 0 edd 4000 0 add 4000 0 Gdd 4000
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Colas erate (Pieridae) 167
(B1)
SEDG
GRAS
(A1Fl)
168 Climatic Risk Atlas of European Butterflies
Colias croceus (GEOFFROY, 1785) — Clouded Yellow
© Chris van Swaay
.—hCcLUwk Full dispersal No dispersal
SEDG 1037 (6.41%) | -1750 (-10.81%)
BAMBU -123 (-0.76%) -2525 (-15.6%)
-420 (-2.59%) | -2931 (-18.11%)
SEDG -962 (-5.94%) | -3518 (-21.74%)
BAMBU -3091 (-19.1%) | -5896 (-36.43%)
-4329 (-26.75%) | -7460 (-46.09%)
Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 16185 cells)
In good summers, the Clouded Yellow is a very fast flyer and can be seen practically over the whole
of Europe, often on fields of clover or lucerne, or other flower-rich vegetations. It lays its eggs one by
one, on the leaves of such leguminous plants as Lucerne (Medicago sativa), clovers (Infolum spp.), and
vetches (Vaza spp.). It pupates, suspended in a silken girdle on the foodplant. In the south, it usually
passes the winter as a caterpillar. In the north, because it cannot tolerate the cold, 1t only appears as a
summer migrant, recolonizing from the south. This species has four to six broods a yeat.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.76).
Climate risk category: LR.
Observed species distribution (50 x 50 km? UTM grid; black
circles) and modelled actual distribution of climatic niche
(orange areas)
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
mM
o
‘dd 4000 0 odd 4000 0 oad 4000 0 ada 4000
1.0
Minimum
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe Swe
0.6 0.6
1.0 02
0.6
Swe
0.2
Multidimensional climatic niche. Occurrence probability defined by
accumulated growing degree days until August (Gdd) and soil water
content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
169
Colias croceus (Pieridae)
2080
2050
170 Climatic Risk Atlas of European Butterflies
Colias hecla LEFEBVRE, 1836 — Northern Clouded Yellow
.—ClcLw Full dispersal No dispersal
Be SEDG -736 (-75.1%) -785 (-80.1%)
-767 (-78.27%) | -791 (-80.71%)
© Jostein Engdal Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 980 cells)
The Northern Clouded Yellow occurs in the mountains of Lapland above the birch zone, on open,
grassy meadows, and stony slopes. The butterflies fly very quickly and are difficult to approach. Eggs
are laid singly or in small groups on the larval foodplants, or on neighbouring plants. Alpine Milk-
vetch (Astragalus alpinus) 1s the most important foodplant, but other milk-vetches (4stragalus spp.)
may also be used. The caterpillars feed on the flowers, leaves and stems, sometimes taking two years
to complete their life-cycle. Hibernation can take place as a caterpillar or a pupa.
Present distribution can be very well explained by climatic variables (AUC = 0.99).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘aad 4000 0 edd 4000 0 add 4000 0 add 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
2050
Colas hecla (Pieridae)
2080
171
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
172 Climatic Risk Atlas of European Butterflies
Colias myrmidone (EsPrER, 1780) — Danube Clouded Yellow
.—hlcLw Full dispersal No dispersal
He SEDG -329 (-7.74%) -1351 (-31.77%)
-2181 (-51.29%) | -2616 (-61.52%)
|GRAS | -1528 (-35.94%) | -2196 (-51.65%)
SEDG -1896 (-44.59%) | -3193 (-75.09%)
BAMBU_ | -2971 (-69.87%) | -3887 (-91.42%)
-2667 (-62.72%) | -4048 (-95.2%)
© Martin Wiemers Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 4252 cells)
The Danube Clouded Yellow occurs in dry, warm grassland where its foodplant, the broom
Chamaecylisus ratisbonensis, C. supnus, C. capitatus, C. austiracus ate abundant. However, the amount of
shelter from bushes can vary considerably. The female lays her eggs on the foodplant, the caterpillars
hibernate in the litter layer. It has two to three broods a year. In contrast to the Eastern Pale Clouded
Yellow, this species has disappeared from most of its former locations in Central Europe and 1s now
extinct in several countries.
This species is listed in Annexes II and IV of the Habitats’ Directive.
Present distribution can be well explained by climatic variables (AUC = 0.92).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
‘sad 4000 0 edd 4000 0 add 4000 0 ada 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
173
done (Pieridae)
LaS MMyVlit
Cob
2080
2050
174 Climatic Risk Atlas of European Butterflies
Colias chrysotheme (EsPrEr, 1780) — Lesser Clouded Yellow
.—hCcLw Full dispersal No dispersal
Be SEDG 416 (55.32%) -489 (-65.03%)
BAMBU -246 (-32.71%) | -565 (-75.13%)
463 (61.57%) -559 (-74.34%)
SEDG -519 (-69.02%) | -665 (-88.43%)
BAMBU -490 (-65.16%) | -707 (-94.02%)
-550 (-73.14%) -733 (-97.47%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 752 cells)
The Lesser Clouded Yellow is found on open, dry, steppe-like grassland and rocky slopes. Its main
foodplant is the Milk-vetch (Astragalus austriacus), but Milk-vetch (A. gheyphyllos) and vetches (Vicia
spp.) are also used. This species has three to four broods a year and hibernates as a caterpillar.
Present distribution can be well explained by climatic variables (AUC = 0.89).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
add 4000 0 odd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O2
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
175
Colas chrysotheme (Pieridae)
2080
2050
176 Climatic Risk Atlas of European Butterflies
Colias byale (LINNAEus, 1758) — Pale Clouded Yellow
.—hCcLUwk Full dispersal No dispersal
Be SEDG -437 (-3.49%) -2793 (-22.28%)
BAMBU -305 (-2.43%) | -2961 (-23.62%)
-786 (-6.27%) | -3393 (-27.07%)
SEDG -905 (-7.22%) | -5229 (-41.72%)
BAMBU_ | -1777 (-14.18%) | -7208 (-57.5%)
-3035 (-24.21%) -9284 (-74.06%)
© Rudi Verovnik Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 12535 cells)
The Pale Clouded Yellow is mostly found on fields of clover or lucerne, but also on meadows that
are lightly grazed where many leguminous plants are growing. It prefers open landscapes with few
trees or bushes. Eggs are laid singly on many species of Leguminosae. The caterpillar feeds on
young leaves and overwinters as half-grown caterpillar. It pupates, suspended in a girdle from a stalk
on the foodplant. The Pale Clouded Yellow has two or three generations a year. In the northern part
of its range, it is a migrant species, but in most of Central Europe it is a resident. The adult Pale
Clouded Yellow is very hard to distinguish from Berger’s Clouded Yellow (Codas adfacariensis), while
the larvae are very different.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.84).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
mM
o
2 : : : .
dd 4000 0 edd 4000 0 ad 4000 0 add 4000
1.0
Minimum
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe Swe
0.6 0.6
1.0 02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Colas hyale (Pieridae) 177
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
178 Climatic Risk Atlas of European Butterflies
Colias alfacariensis Risse, 1905 — Berger’s Clouded Yellow
a
SEDG -812 (-8.42%) | -4054 (-42.02%)
BAMBU_ | -1547 (-16.03%) | -5409 (-56.06%)
-2461 (-25.51%) | -6649 (-68.92%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 9648 cells)
The Berger’s Clouded Yellow is a butterfly of dry, open grasslands on calcareous soils. In the
northern part of its distribution range, it often occurs on the south-facing slopes of hills or
mountains. Eggs are mostly laid on Horseshoe Vetch (Hippocrepis comosa), also on Crown Vetch
(Coronilla varia). Hibernation takes place on the foodplant, or on the ground in the litter layer. For
pupation the caterpillar attaches itself to a foodplant, turning into a pupa, suspended by a silken
girdle. This species has two or three broods a year. The adult Berger’s Clouded Yellow and Pale
Clouded Yellow are so similar that it 1s not possible to identify them with certainty, while this 1s easy
for the caterpillars.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.84).
Climate risk category: R.
Annual temperature range
Minimum old (3350) Large (66%) Maximum
mM
o
° : : : ? :
fad 4000 0 Codd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe Swe
0.6 0.6
1.0 02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
179
Colas alfacariensis (Pieridae)
2080
2050
180 Climatic Risk Atlas of European Butterflies
Gonepteryx rhamni (LINNAEUS, 1758) — Brimstone
+X >. vy #& 6c Full dispersal No dispersal
ecm AY fi SEDG -1084 (-5.26%) | -3228 (-15.67%)
BAMBU -1593 (-7.73%) | -3440 (-16.7%)
-1973 (-9.58%) | -4092 (-19.87%)
SEDG -1165 (-5.66%) | -4835 (-23.47%)
BAMBU | -2555 (-12.4%) | -7022 (-34.09%)
GRAS -4146 (-20.13%) | -9486 (-46.05%)
© Peter Ginzinger Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 20599 cells)
For many, the Brimstone is the first sign of spring. The butterflies hibernate in heaps of twigs
or grass tussocks, and on any warm day, even in January, the males appear. The females, that are
paler in colour, are seen a little later. The eggs are laid apart on the young branches and leaves of
buckthorns (Rhamnus spp.). The caterpillars feed on the young leaves. They pupate, suspended from
the underside of a twig or nerve of a leaf. In the summer, when the butterflies emerge, they do not
mate but instead may become inactive for quite long periods before they hibernate. Courtship and
mating do not take place until the spring. The Brimstone always has just one generation a year.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.73).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
06
mM
a a
=) a
= =
Oe
-& =a
o= mH
=f |
£5 8
Oo. 4
of 7
28
a ng = a
ger
ae |
See
qf. =
Zo
wv
x a Po]
= 4
o
0 2000 4000 2000 40000 2000 40000 2000 4000
Gdd Gdd Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
181
Gonepteryx rhamni (Pieridae)
2080
2050
182 Climatic Risk Atlas of European Butterflies
Gonepteryx farinosa ZELLER, 1847 — Powdered Brimstone
=o
SEDG 334 (51.78%) -210 (-32.56%)
BAMBU -27 (-4.19%) -369 (-57.21%)
-144 (-22.33%) | -519 (-80.47%)
© Heiner Ziegler Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 645 cells)
The Powdered Brimstone larvae feed on the prickly Christ’s Thorne (Pahurus spina-christ) and on
various buckthorns, such as Rhamnus alpinus, RB. sibthorpianus and R. hvioides. They are therefore often
seen on bushy vegetation, namely in warm, dry places and on rocky slopes. However, in their search
for nectar, they also fly in other places. It has one generation a year and, like other brimstones,
hibernates as a butterfly.
Present distribution can be well explained by climatic variables (AUC = 0.94).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
1.0
Minimum
Swe
06
1.0 02
0.6
Swe
o2
MD
Se on
Ss
oy
ae
2a
2G 3
aH a4
O = =
Do
a 36
ze"
‘ td
.-
a 6 _
qe -
Zo
iw
= am
‘0
=
o rey 4000 0 od 4000 0 Cad 4000 0 odd 4000
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
183
Gonepteryx farinosa (Pieridae)
2080
2050
184 Climatic Risk Atlas of European Butterflies
Gonepteryx cleopatra (LINNAEUS, 1767) — Cleopatra
.—hCcLe Full dispersal No dispersal
Be SEDG -17 (-0.4%) -1417 (-33.35%)
-31 (-0.73%) -1653 (-38.9%)
-1013 (-23.84%) | -2621 (-61.69%)
-1293 (-30.43%) | -3250 (-76.49%)
© Peter Ginzinger Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 4249 cells)
The Cleopatra is found in light woodland, woodland edges and open bushy places. The males are
easily recognized in flight by the orange patches on their forewings; the female looks very much like
an ordinary brimstone. They are strong flyers that sometimes roam outside their breeding area. The
eges are laid on the young leaves of various buckthorns, such as Buckthorn (Rhawnus catharticus),
Mediterranean Buckthorn (R. a/aternus), and Alpine Buckthorn (R. a/pinus). The caterpillars feed on
the leaves, and pupate on the twigs of the foodplant. The Cleopatra hibernates as an adult butterfly,
and therefore can be seen flying for most of the year. Probably, it has only one generation a year.
Present distribution can be well explained by climatic variables (AUC = 0.92).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
7 {od
todd 4000 0 odd 4000 0 add 4000 0 Gdd 4000
Minimum
Swe
06
1.0 02
0.6
Swe
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
185
Gonepteryx cleopatra (Pieridae)
2080
2050
186 Climatic Risk Atlas of European Butterflies
Lycaena phlaeas (LINNAEUS, 1761) — Small Copper
eae
-4048 (-16.93%) | -4602 (-19.25%)
[ena cary [arene
SEDG -5057 (-21.15%) | -6765 (-28.29%)
BAMBU_ | -8526 (-35.66%) | -10086 (-42.18%)
-11453 (-47.9%) | -13177 (-55.11%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 23910 cells)
The Small Copper is a very common, widespread butterfly. It is found on all sorts of grasslands
and heathlands, roadsides and along the banks of canals. The butterflies often visit flowers, and the
males, perched on a tall blade of grass, defend their territory fiercely, flying out at other males of
its sort. Eggs are laid on different species of acidic sorrels, mainly Common Sorrel (Rumex acetosa).
This butterfly species overwinters as a caterpillar. In the north of its range, the Small Copper has
two generations a year, whereas in the southern part, 1t may have three or four.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.65).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
A
Minimum
Swe
1.6
|
a a
=) a
fe
ee
cfes
o= mH
=e
£5 3
Oo 4
of 7
© hd
o nq = i
ger
|
ae
qf. =
2 o
aw
x F Po]
= 4
o
0 2000 40000 2000 40000 2000 40000 2000 4000
Gd Gdd Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Lycaena phlaeas (Lycaenidae)
SEDG
(B1)
187
GRAS
(A1Fl)
188 Climatic Risk Atlas of European Butterflies
Lycaena helle ({ScCHIFFERMULLER], 1775) — Violet Copper
.—hClcLe Full dispersal No dispersal
He SEDG 159 (6.52%) -563 (-23.08%)
229 (9.39%) -487 (-19.97%)
Ta cra) | 686 (281399
SEDG -197 (-8.08%) | -1073 (-43.99%)
BAMBU -36 (-1.48%) -968 (-39.69%)
-466 (-19.11%) | -1201 (49.24%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2439 cells)
The Violet Copper is a rare butterfly and often confined to very small sites (where it may occur in
large numbers). It is found in swampy, wet grassland and rough vegetation bordering streams and
lakes. In Central Europe, eggs are laid on the underside of the leaves of Bistort (Po/gonum bistorta).
In the north of its range Viviparous Bistort (Po/gonum rivipara) 1s also used as larval foodplant. The
young caterpillars eat the lower epidermis, thus making the characteristic “windows”. It passes the
winter as a pupa. It has one, sometimes two, generations a year.
This species is listed in Annexes II and IV of the Habitats’ Directive.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.78).
Climate risk category: LR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
‘0.6
|
Qa a
=) a
>S 7
Pm Oe
cfes
o= mn
:- 7
£5 3
Oo. 4
om 6
D>
o 4 = a
ger
te |
ae
i=]
@e-
eo
wo
x é 5
= 4
o
0 2000 40000 2000 40000 2000 40000 2000 4000
Gd id Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
189
Lycaena helle (Lycaenidae)
2080
2050
190 Climatic Risk Atlas of European Butterflies
Lycaena dispar (HAworTH, 1803) — Large Copper
re ae
3583 (44.52%) | -1262 (-15.68%)
ee 2980 (37.03%) | -1764 (-21.92%)
SEDG 3656 (45.43%) | -2587 (-32.14%)
BAMBU 4992 (62.03%) | -3071 (-38.16%)
4375 (54.36%) | -4188 (-52.04%)
© Josef Settele Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 8048 cells)
The Large Copper occurs in marshy habitats, and on the peaty banks of lakes, rivers and streams, more
to the East also on waste lands. Nectar plants are important for the females, which lay more eggs when
there is more food available for them. Eggs are laid on large non-acidic sorrels (like Ramex crispus, R.
obtusifolus, but never R. acetosa or R. acetosella). The young caterpillars first eat from the underside of the
leaves, making the characteristic ‘windows’. Later caterpillars feed on the whole leaf. They hibernate
when half-grown between withered leaves at the foot of the foodplant and are sometimes associated
with ants (Myrmica rubra and Lasius niger). The Large Copper has several subspecies in Europe. The
subspecies L. d. bataia has one generation a year, and the others two and sometimes even three.
This species is listed in Annexes II and IV of the Habitats’ Directive.
Present distribution can be well explained by climatic variables (AUC = 0.88).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
‘1.6
sh
a a
=) a
i
ee
ef@eEs
o= mH
=f
£5 8
Oo 4
of 7
fo
cL a4 = a
ger
|
ae
a
@e-
2 o
wo
x é Po]
= 4
o
0 2000 40000 2000 40000 2000 40000 2000 4000
Gd id Gad Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
191
Lycaena dispar (Lycaenidae)
2080
2050
192 Climatic Risk Atlas of European Butterflies
Lycaena virgaureae (LINNAEUS, 1758) — Scarce Copper
.—CcLwe Full dispersal No dispersal
Be SEDG -780 (-5.26%) -2762 (-18.63%)
BAMBU | -2665 (17.98%) | -3876 (-26.15%)
-3078 (-20.77%) | -4622 (-31.18%)
SEDG -2694 (-18.18%) | -6097 (-41.13%)
BAMBU_ | -5416 (-36.54%) | -8571 (57.83%)
-7216 (-48.68%) | -10671 (-71.99%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 14822 cells)
It would be difficult to miss the beautiful and conspicuous Scarce Copper. The white patches on
the underside of the hindwing distinguish 1t from other coppers. It prefers sheltered, flower-rich
grassland in or near woods. The female visits a wide range of composites, especially Golden-rod
(Soddago virgaureae) from where it gets its species name. The female lays her eggs at the base of the
flowering stems of acidic sorrels like Ramex acetosa and R. acetosella. The eggs do not hatch until the
following spring when the small caterpillars make a good meal of the new, fresh vegetation. It has
one generation a year.
Present distribution can be well explained by climatic variables (AUC = 0.86).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 Codd 4000 0 add 4000 0 add 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Swe
0.6
Annual precipitation range
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
193
Lycaena virgaureae (Lycaenidae)
2080
2050
194 Climatic Risk Atlas of European Butterflies
Lycaena ottomana (LEFEBVRE, 1830) — Grecian Copper
6c Full dispersal No dispersal
Hi SEDG 103 (21.78%) -182 (-38.48%)
-54 (-11.42%) -346 (-73.15%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 473 cells)
The Grecian Copper can be found in low, scrubby vegetation, maquis and woodland clearings and
also occurs in dry and moist grassland at the bottom of valleys. It is the males that are usually seen;
the females seem to live a more sheltered existence and are difficult to find. However, the males also
take cover in trees if disturbed. They fly close to the ground. Sorrel, especially Sheep’s Sorrel (Ramex
acetosella), 1s used as larval foodplant. This butterfly species has two generations a yeat.
Present distribution can be very well explained by climatic variables (AUC = 0.98).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
‘1.6
tH
Qa a
=) a
ee
Pe Oe
a =a
o= mH
=e |
£5 8
oOo. 4
Oe 5
fo
a = = ba
ger
my
=
=!
Qe
2 y
w
é 5
= 4
o
0 2000 40000 2000 40000 2000 40000 2000 4000
Gd id Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
195
Lycaena ottomana (Lycaenidae)
2080
2050
196 Climatic Risk Atlas of European Butterflies
Lycaena tityrus (Popa, 1761) — Sooty Copper
.—ClcLe Full dispersal No dispersal
Be SEDG 119 (1.07%) -2721 (-24.36%)
-636 (5.69%) | -3400 (-30.43%)
-3278 (-29.34%) | -6816 (-61.01%)
-4194 (-37.54%) | -8250 (-73.85%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 11172 cells)
The Sooty Copper is found in a variety of habitats, damp and dry grassland, heathland, bogs, but
also scrub and clearings in woodland, mostly in small numbers. Eggs are laid on sorrels (Rumen
spp.), especially on Common Sorrel (R. acefosa). The caterpillar hibernates at the foot of the plant in
any of the instars. They pupate in the litter layer. It has up to three generations a yeat.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.82).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
‘dd 4000 0 odd 4000 0 oad 4000 0 odd 4000
Minimum
Swe
1.6
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
197
Lycaena tityrus (Lycaenidae)
2080
2050
198 Climatic Risk Atlas of European Butterflies
Lycaena alciphron (ROTTEMBURG, 1775) — Purple-shot Copper
CL Full dispersal No dispersal
Be SEDG -503 (-5.52%) -2545 (-27.93%)
BAMBU_ |_ -2988 (-32.79%) | -3855 (-42.3%)
-2959 (-32.47%) | -4292 (-47.1%)
SEDG -1861 (-20.42%) | -5247 (-57.58%)
BAMBU_ | _ -4297 (-47.15%) | -6740 (-73.96%)
-4303 (-47.22%) | -7612 (-83.53%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 9113 cells)
The Purple-shot Copper is found both in damp and dry places on many kinds of flower-rich
grassland. Only the male has the beautiful purple sheen on the upperside of its wings. Eggs are
laid on sorrels (Rumex spp.), mainly Common Sorrel (R. acefosa). This butterfly species hibernates
as a fully-developed caterpillar in the egg, or in the first instar. The caterpillars are nocturnal. They
pupate in the litter layer. It has one generation a year. It hibernates as a caterpillar outside the egg.
Across Europe there are different subspecies.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.8).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘sad 4000 0 edd 4000 0 ad 4000 0 odd 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
06
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
199
Lycaena akiphron (Lycaenidae)
2080
2050
200 Climatic Risk Atlas of European Butterflies
Lycaena hippothoe (LINNAEus, 1761) — Purple-edged Copper
AMBU | -6505 (-46.11%) | -8317 (-58.95%)
-8606 (-61%) -9862 (-69.9%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 14109 cells)
The Purple-edged Copper occurs on wet to damp grasslands, where the male butterflies attract the
attention, perched on a tall grass or other plant, watching over their territory. The populations are
mostly very local, but in a meadow, the butterflies can often be very numerous. The eggs are laid
on various sorrels (Rwmex spp.). At first the small caterpillar only shaves off a few cell layers on the
leaf surface, so making translucent “windows”, but later they feed on the whole leaf. The caterpillar
hibernates when still small, and completes its growth 1n the spring, pupating in the litter layer. It has
one brood a year. In southern part of its range the species has two generations and can be found on
dry grasslands as well. The Purple-edged Copper has a few subspecies.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.85).
Climate risk category: R.
Annual temperature range
Minimum Small ( (335%) Large (66%) Maximum
1.0
Minimum
Swe
06
|
a =]
=) a
es 7
ee
a =o
o= 0
=e
£5 3
Oo. 4
of 7
D io
a 4 ge
Ba”
= tH
ie
fe. -
Zo
w
= 4
o
0 2000 40000 2000 40000 2000 40000 2000 4000
Gdd d Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
201
Lycaena hippothoe (Lycaenidae)
2080
2050
202 Climatic Risk Atlas of European Butterflies
Lycaena candens (HERRICH-SCHAFFER, 1844) — Balkan Copper
6c Full dispersal No dispersal
Be SEDG 192 29.22%) -289 (-43.99%)
BAMBU -161 (-24.51%) | -376 (57.23%)
-75 (-11.42%) -414 (-63.01%)
SEDG 70 (10.65%) -343 (-52.21%)
BAMBU -391 (-59.51%) | -564 (-85.84%)
-394 (-59.97%) | -624 (-94.98%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 657 cells)
The Balkan Copper can be found in damp grassland, sub-alpine and alpine meadows, and in woodland
clearings with a well-developed herb layer. Different sorrels (Rumex spp.), including Common Sorrel
(R. acefosa) are used as larval foodplants. It has one generation a year. This butterfly looks very like
the Purple-edged Copper (Lycaena hippothoe), and used to be classified as a subspecies of it.
Present distribution can be well explained by climatic variables (AUC = 0.86).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
-
‘dd 4000 0 odd 4000 0 add 4000 0 ada 4000)
0.6
Minimum
Swe
Swe
a) 0.6 1.0 O02
1.0
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O2
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
203
Lycaena candens (Lycaenidae)
2080
2050
204 Climatic Risk Atlas of European Butterflies
Lycaena thersamon (Esper, 1784) — Lesser Fiery Copper
SEDG 1838 (40.44%) | -1584 (-34.85%)
1632 (35.91%) | -1421 (-31.27%)
a 2941 (64.71%) | -757 (-16.66%)
SEDG 2058 (45.28%) | -3651 (-80.33%)
BAMBU 574 (12.63%) | -3143 (-69.15%)
[oras | 2327 (51.2%) | -1739 (-38.26%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 4545 cells)
The Lesser Fiery Copper is mostly found on dry, flower-rich grasslands, both on calcareous and acid
soils. They are also seen on waste land, very open scrub and rocky slopes. The butterflies are often
seen drinking from the flowers of Danewort (Sambucus ebulus). Its most important larval foodplant
is Knotgrass (Polygonum aviculare), although other Polygonaceae species are perhaps also used. The
caterpillars feed on both the flowers and the leaves. This butterfly species hibernates as a caterpillar
and has several generations a year.
Present distribution can be well explained by climatic variables (AUC = 0.93).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
‘dd 4000 0 odd 4000 0 add 4000 0 odd 4000
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Lycaena thersamon (Lycaenidae) 205
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
206 Climatic Risk Atlas of European Butterflies
Thecla betulae (LINNAEUS, 1758) — Brown Hairstreak
SEDG -2243 (-17.38%) | -3619 (-28.04%)
BAMBU_ | -1298 (-10.06%) | -3214 (-24.9%)
-2925 (-22.66%) | -4462 (-34.57%)
SEDG -3697 (-28.64%) | -5889 (-45.62%)
BAMBU_ | -3110 (-24.09%) | -6912 (-53.55%)
-5029 (-38.96%) | -9532 (-73.85%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 12908 cells)
The Brown Hairstreak occurs in scrub, along woodbanks at the edge of deciduous woodland, and
in parks, but the butterfly is rarely seen. However, the eggs that are laid at the base of the buds of
various Prunus species ate easy to find. In the winter the pearly-white eggs show up on the bare
twigs of the shrubs or trees. The egg, thus, hibernates, hatching in the spring. The small caterpillar
bores its way into a bud, hollowing it out, later on also eating the leaves of Blackthorn (Prunus
spinosa), but also cultivated species of Prunus, such as plum and cherry trees. The caterpillars are
visited by ants of the genera Lasius and Formica. They pupate in the shrub layer under the tree
serving as foodplant. This butterfly species is single-brooded.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.79).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
m
o
2 ; : : ?
dd 4000 0 edd 4000 0 oad 4000 0 aad 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
Swe
0.6
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
207
Thecla betulae (Lycaenidae)
2080
2050
208 Climatic Risk Atlas of European Butterflies
Favonius quercus (LINNAEUS, 1758) — Purple Hairstreak
rea [Rea
-3105 (-22.36%) | -4170 (-30.03%)
SEDG -4962 (-35.73%) | -5822 (-41.93%)
BAMBU_ | -6242 (-44.95%) | -7466 (-53.77%)
-7703 (-55.47%) | -9184 (-66.14%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 13886 cells)
The Purple Hairstreak is only found around oaks, its only foodplant. Although the butterflies need
food, they do not visit flowers. Instead, they remain near the tree, looking for leaves with a layer
of honeydew left behind by aphids. This sugary secretion is the butterflies’ most important source
of energy. The eggs are laid at the base of the buds. The small caterpillar has developed by the
time winter comes, but it remains in the egg, emerging in the spring to feed on the buds and also
on the flowers of the oak. It does not eat the leaves and is sometimes visited by ants of the genus
Lasius. When mature, the caterpillars leave the foodplant, and pupate in the moss layer. The Purple
Hairstreak has one generation a yeat.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.72).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
ce
Minimum
Swe
1.6
10 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 0.2
0.6
Swe
o2
0 2000 40000 2000 40000 2000 40000 2000 4000
Gd dl Gdd
Gdd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
209
Fiavonius quercus (Lycaenidae)
2080
2050
210 Climatic Risk Atlas of European Butterflies
Laeosopis roboris (EsPER, 1793) — Spanish Purple Hairstreak
.—CcLe Full dispersal No dispersal
Be SEDG -912 (-22.66%) -2173 (-54%)
-1551 (-38.54%) | -2621 (-65.13%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 4024 cells)
The Spanish Purple Hairstreak occurs in damp, deciduous woods, on rough vegetation beside
streams and rivers, and in parkland. The butterflies spend most of their time in the top of ash trees,
the larval foodplant, and are difficult to see. The female lays her eggs on Ash (Fraxinus excelsior) and
other species of Fraxinus, and possibly also Common Privet (Ligustrum vulgare). The eggs remain on
the tree for the rest of the summer, autumn and winter. The small caterpillars that emerge in the
spring, feed on the flowers and new leaves of the foodplants. They are visited by ants of the genus
Lasius. Vhey pupate among the leaf litter. This species has one brood a yeat.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.81).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
fad 4000 0 edd 4000 0 add 4000 0 ada 4000)
1.0
Minimum
Swe
06
o2
Swe
0.6 1.0
1.0 0.2
Swe
0.6
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
211
Laeosopis roboris (Lycaenidae)
2080
2050
212 Climatic Risk Atlas of European Butterflies
Tomares ballus (Fasrictus, 1787) — Provence Hairstreak
.—CcLee Full dispersal No dispersal
Be SEDG -285 (-71.79%) -309 (-77.83%)
BAMBU -296 (-74.56%) | -315 (-79.35%)
-316 (-79.6%) -341 (-85.89%)
SEDG -309 (-77.83%) | -358 (-90.18%)
BAMBU -340 (-85.64%) -368 (-92.7%)
-386 (-97.23%) | -394 (-99.24%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 397 cells)
The Provence Hairstreak can be found on abandoned agricultural land, especially on calcareous
soils, in abandoned vineyards and bushy areas. Various leguminous plants are used as larval
foodplant, including Bladder Vetch (Authylis tetraphylla), Dorycnium hirsutum, Vberian Milk-vetch
(Astragalus lusitanicus), Hairy Medick (Medicago polymorpha), Black Medick (M. 4upukina) and birdsfoot-
trefoils (Lo/us spp.). The female deposits her eggs singly on the unopened leaves. The caterpillars
are sometimes attended by ants (e.g. Plagolepis pygmaea). This species has one generation a year, and
passes the winter as a pupa in the soil.
Present distribution can be well explained by climatic variables (AUC = 0.91).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
=
mm
a
a
=
Ow
=o
ee]
co
Oo
S : P - :
mH
o
2
Ow
$0
LP)
™
o
fad 4000 0 edd 4000 0 Gad 4000 0 Gdd 4000
Minimum
Swe
06
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.8
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
2050
Tomares ballus (Lycaenidae)
2080
SEDG
213
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
214 Climatic Risk Atlas of European Butterflies
Callophrys rubi (Linnakus, 1758) — Green Hairstreak
[rear | Neamt
-4049 (-18.94%) | -4842 (-22.65%)
SEDG -4212 (-19.7%) | -5250 (-24.56%)
BAMBU_ | -7570 (-35.41%) | -8595 (-40.2%)
-10212 (-47.76%) | -11497 (-53.77%)
© Peter Ginzinger Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 21380 cells)
The Green Hairstreak can be found on poor grassland, heathland and at the edges of bogs and
marshes. The butterflies like resting in trees and shrubs, such as Alder Buckthorn (frangula alnus),
where they are well hidden by their green colour. However, a sharp tap on a branch brings tens
of them out into the air. The Green Hairstreak is extremely polyphagous. The eggs are laid on a
variety of plants, including Heather (Eva spp.), rockroses (Hedanthemum spp.), and Alder Buckthorn
(Frangula alnus), \eguminous plants and species of Rosaceae. The caterpillars feed on the buds, flowers
and young leaves. They pupate in the litter layer and the pupa hibernates. The Green Hairstreak is
single-brooded.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.63).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
a
Minimum
Swe
06
10 02
Swe
0.6
1.0 O.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
td
o
S
ze
o
m
o
ot 4000 0 ns 4000 0 yh 4000 0 ae 4000
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Callophrys rubi (Lycaenidae)
vi
SEDG
(B1)
215
GRAS
(A1Fl)
216 Climatic Risk Atlas of European Butterflies
Callophrys avis CHAPMAN, 1909 — Chapman’s Green Hairstreak
CL Full dispersal No dispersal
Be SEDG -283 (-65.36%) -386 (-89.15%)
BAMBU -320 (-73.9%) -378 (-87.3%)
-275 (-63.51%) | -390 (-90.07%)
SEDG -294. (-67.9%) -408 (-94.23%)
BAMBU -303 (-69.98%) | -415 (-95.84%)
-243 (-56.12%) | -425 (-98.15%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 433 cells)
Chapman’s Green Hairstreak is mostly found in woods and scrubs where the Strawberry Tree
(Arbutus unedo) grows. The female chooses the young leaves of the older, larger bushes on which to
lay her eggs. Conaria myrtifola and the broom Cytsus malacitanus ssp. catalaunicus have also been named
as foodplants. It pupates on the ground at the foot of the foodplant. It is single-brooded.
Present distribution can be well explained by climatic variables (AUC = 0.9).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
aad 4000 0 odd 4000 0 ad 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 o.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
2050
Callophrys avis (Lycaenidae)
2080
SEDG
217
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
218 Climatic Risk Atlas of European Butterflies
Satyrium w-album (KNocu, 1782) — White-letter Hairstreak
SEDG -1917 (-17.27%) | -2641 (-23.79%)
-2141 (-19.28%) | -2955 (-26.61%)
ee -2827 (-25.46%) | -3629 (-32.68%)
SEDG -4836 (-43.56%) | -5477 (-49.33%)
BAMBU_ | _ -5644 (-50.83%) | -6698 (-60.33%)
-7380 (-66.47%) | -8657 (-77.97%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 11103 cells)
The White-letter Hairstreak usually occurs where there is a group of elm (U/mus spp.) trees, either
growing 1n a wood, or apart, sometimes even in the centre of a large town. However, there are
populations known that occupy a solitary tree. The eggs are laid on the wood at the base of the flower
buds, the female favouring terminal buds situated at the top of the tree. The small caterpillar stays
inside the egg during the winter months, emerging in the spring, boring its way into a flowerbud. It
feeds on flowerbuds and flowers, but not on leaves. A non-flowering tree is therefore not a suitable
foodplant. The caterpillars are sometimes visited by ants of the genus Lasius and Formica and pupate
in the litter layer under the tree. The White-Letter Hairstreak has one generation a year.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.78).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
o ck 4000 0 edd 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
219
Satyrium w-album (Lycaenidae)
2080
2050
220 Climatic Risk Atlas of European Butterflies
Satyrium pruni (LINNAEUS, 1758) — Black Hairstreak
.6—hCcLUwk Full dispersal No dispersal
Be SEDG -289 (-2.36%) -3101 (-25.27%)
BAMBU 1089 (8.88%) -22.45 (-18.3%)
-404 (-3.29%) | -3596 (-29.31%)
SEDG -255 (-2.08%) | -4671 (-38.07%)
BAMBU -702 (5.72%) | -6066 (-49.44%)
-2244 (-18.29%) | -7988 (-65.1%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 12270 cells)
The Black Hairstreak can be found where there are Blackthorn (Prunus spinosa) bushes growing.
This may be in moderately dry woodland, or in a hedgerow, or on a woodbank, but it also occurs
on solitary groups of Blackthorn bushes. Places with brambles are favorable, providing the nectar
the butterflies need. The eggs are laid on Blackthorn (Prunus spinosa), but sometimes other Prunus
species are used. The female chooses twigs that will flower the next spring. The small caterpillars
that develop, pass the winter in the eggshell. When they emerge, they feed on the buds and flowers.
When the caterpillars are fully-grown, they pupate. The black and white pupa, suspended in a
silken girdle from a twig, looks very like a bird dropping, an attempt to avoid predation. The Black
Hairstreak is single-brooded.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.83).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
fad 4000 0 edd 4000 0 add 4000 0 odd 4000
Minimum
Swe
O06
1.0 02
Swe
0.6
1.0 0.2
Swe
0.6
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
221
Satyrium pruni (Lycaenidae)
2080
2050
222 Climatic Risk Atlas of European Butterflies
Satyrium spini (FasrRicius, 1787) — Blue-spot Hairstreak
.—hCcLwe Full dispersal No dispersal
fe SEDG -1547 (-18.11%) | -3552 (-41.59%)
-2078 (-24.33%) | -3494 (-40.91%)
reras | sce | 2seacavame
SEDG -1885 (-22.07%) | -6706 (-78.52%)
BAMBU -2494 (-29.2%) | -5899 (-69.07%)
-695 (-8.14%) | -4290 (-50.23%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 8540 cells)
The Blue-spot Hairstreak can be found in sunny, warm places, on groups of bushes, or on scrub
at the edge of a wood, with some flowering vegetation as nectar source nearby. The female lays
her eggs on various buckthorns (Rhamnus spp.), choosing branches situated in the sun. The small
caterpillar only emerges in the spring, having passed the winter in the eggshell. It wastes no time
in beginning to eat buds and young leaves. Ants of the genera Lasius and Formica may visit the
caterpillar. When fully-grown, it pupates on the foodplant, the pupa suspended by a silken girdle.
The Blue-spot Hairstreak has one generation a yeat.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.71).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
06
sh
a a
=) a
eee
ee
a |
o= mH
= |
£5 3
Go. 4
os 6
28
a a4 = a
ger
|
ae
i=]
@e-
2 o
aw
* é 5
= 4
o
0 2000 40000 2000 40000 2000 40000 2000 4000
Gd id Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
223
Satyrium spini (Lycaenidae)
2080
2050
224 Climatic Risk Atlas of European Butterflies
Satyrium ilicis (EsPER, 1779) — Ilex Hairstreak
.—hCcLwee Full dispersal No dispersal
fi SEDG -164 (-1.51%) -2441 (-22.49%)
-613 (5.65%) | -2924 (-26.93%)
-3154 (-29.05%) | -5618 (51.75%)
-4333 (-39.91%) | -7136 (65.73%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 10856 cells)
The Ilex Hairstreak occurs locally at woodland edges, and in coppices with warm, dry oak scrub.
The female lays her eggs on the rather stunted oak trees, and certainly not on large oaks. The
tiny caterpillar hibernates in the egg, or 1n the first larval instar. It feeds on the young oak leaves,
eventually pupating in the litter layer under the foodplant. The caterpillars have been found in
association with ants of the genera Camponotus and Crematogaster. The adult butterflies need nectar,
feeding on flowering Common Privet (Ligustrum vulgare) and on bramble blossom (Rubus spp.). It
is single-brooded.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.71).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
1.6
10 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
10 0.2
0.6
Maximum Large (66%) Small (33%)
Swe
O 2000 40000 2000 40000 2000 40000 odd 4000
Gdd Gdd Gdd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
225
Satyrium iliis (Lycaenidae)
2080
2050
226 Climatic Risk Atlas of European Butterflies
Satyrium esculi (HUBNER, 1804) — False Hex Hairstreak
a
-1549 (51.46%) | -2151 (-71.46%)
SEDG -1726 (-57.34%) | -2448 (-81.33%)
BAMBU_ | -2208 (-73.36%) | -2811 (93.39%)
-2014 (-66.91%) -2998 (-99.6%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 3010 cells)
The False Ilex Hairstreak can be found in dry scrub, open, deciduous woodland, or in mixed
woodland. The butterflies can often be seen drinking nectar on thyme. The females lay their eggs
on various oak species, including the Holm Oak (Quercus ikx), Common or Pedunculate Oak (2.
robur), Kermes or Holly Oak (Q. coccsfera), and Pyrenean Oak (Q. pyrenaica). The small caterpillars
only emerge from the egg the following spring, when they start eating the young oak leaves. The ant
Camponotus cruentatus has been found in association with them. This species has one brood a yeat.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.82).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
fed 4000 0 odd 4000 0 add 4000 0 add 4000
Minimum
Swe
0.6
1.0 O02
Swe
0.6
1.0 0.2
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
227
Satyrium esiuli (Lycaenidae)
2080
2050
228 Climatic Risk Atlas of European Butterflies
Satyrium acactae (Fasricius, 1787) — Sloe Hairstreak
Cc Full dispersal No dispersal
fi SEDG 1205 (15.48%) -2303 (-29.59%)
-492 (-6.32%) | -2953 (-37.94%)
-1287 (-16.53%) | -5311 (-68.23%)
© Martin Wiemers Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 7784 cells)
The Sloe Hairstreak is found in dry scrub in woods, at wood margins, or in the open landscape. They
are also seen in abandoned vineyards on calcareous soils. When looking for nectar, the butterflies
seem to prefer white flowers. Blackthorn or Sloe (Prunus spinosa) is practically the only foodplant of
the Sloe Hairstreak, and eggs are laid on branches that are in the sun. The small caterpillars stay in
the ege until after the winter, emerging in the spring and feeding on the young Blackthorn leaves.
When fully-grown, they leave the foodplant to pupate on the ground in leaf litter.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.76).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
‘0.6
|
a a
=) 2
i
Pe Oe
ae =a
o= mH
=e
£5 8
Oo 4
ow 6
D>
a mG = a
ger
ee |
ae
=
Qe
2 y
wv
x é 5
= 4
o
0 2000 40000 2000 40000 2000 40000 2000 4000
Gd id Gadd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
229
Satyrium acaciae (Lycaenidae)
2080
2050
230 Climatic Risk Atlas of European Butterflies
Lampides boeticus (LINNAEUS, 1767) — Long-tailed Blue
aaa
815 (12.77%) | -1156 (-18.12%)
amas | mon [ci
SEDG -722 (-11.31%) | -3666 (-57.45%)
BAMBU 10 (0.16%) -2770 (-43.41%)
885 (13.87%) | -1663 (-26.06%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 638 1cells)
The underside of the wings of the Long-tailed Blue have an attractive pattern of fine, wavy lines
and two striking eye-spots near the rather long little tails on the hindwings. The habitats are typified
by the presence of many kinds of leguminous plants that serve as foodplants, and plenty of nectar-
rich flowers for the butterflies. The habitats are warm, dry places, sometimes in agricultural areas.
The main foodplant is Bladder Senna (Co/utea arborescens). The caterpillars feed on the ripening seeds
and on farms where peas and beans are grown, they can cause outbreaks. In natural situations, the
caterpillars are attended by various ant species. The life cycle of the Long-tailed Blue takes four to
six weeks. It does not go into hibernation, and can therefore only occur as a resident where it 1s
warm enough for all stages to survive.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.83).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
ted 4000 0 edd 4000 0 add 4000 0 odd 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
231
Lampides boeticus (Lycaenidae)
2080
2050
232 Climatic Risk Atlas of European Butterflies
Cacyreus marshalli (BUTLER, 1898) — Geranium Bronze
.—hCcLe Full dispersal No dispersal
Be SEDG -207 (-25.87%) -437 (-54.62%)
-277 (-34.62%) -464 (58%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 800)
Originally a species from southern Africa, the Geranium Bronze had probably been imported
on Pelargonium cultivars, the ordinary or garden geraniums. From there out, because of the large
popularity of these plants, it occurs e.g. in Spain and France, but is about to colonize larger parts
of Europe, using the Pe/argonium culttvars as foodplant. The caterpillars mostly eat the flowers and
buds of Pelargonium, but also the rest of the plants. In South Africa, the Geranium Bronze uses
wild Geranium species as foodplant, making it very probable that this butterfly will establish itself
in the wild in warm parts of Europe. However, in cooler areas, it would be impossible for it to
settle permanently, because it has no diapause, and could not survive the winter. It has numerous
generations a year, depending on the temperature.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.85).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Ow
|
w
is}
oO
Ps : : : : P : ;
ah
o
=
w
o
dd 4000 0 edd 4000 0 ad 4000 0 odd 4000
1.0
Minimum
Swe
06
1.0 02
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
233
Cacyreus marshal (Lycaenidae)
2080
2050
234 Climatic Risk Atlas of European Butterflies
Leptotes pirithous (LINNAEUS, 1767) — Lang’s Short-tailed Blue
a
654 (11.95%) | -1120 (-20.46%)
SEDG 800 (14.61%) | -1775 (-32.42%)
BAMBU -30 (-0.55%) -2922 (-53.37%)
838 (15.31%) -3855 (-70.41%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 5475)
The Lang’s Short-tailed Blue is a small, inconspicuous butterfly that occurs in flower-rich places and
on rough vegetation. It is often seen near scrub and on fields of Lucerne (Medicago sativa). The eggs
are laid on various leguminous plants, such as melilots (Medhius spp.), Purple Loosestrife (Lythrum
salicaria), and also species of Rosaceae and Plumbaginaeceae. The caterpillar feeds on the flowers
and seeds of the foodplant and has been found in association with ants of the genus Laszus. The
life cycle of the Lang’s Short-tatled Blue takes four to eight weeks, depending on the temperature.
It is not known whether this species enters diapause, but 1f 1t does, it should be either in the egg or
caterpillar stage.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.83).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘sad 4000 0 edd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
235
Leptotes pirithous (Lycaenidae)
2080
2050
236 Climatic Risk Atlas of European Butterflies
Zizeeria knysna (TRIMEN, 1862) — African Grass Blue
.—ClcLee Full dispersal No dispersal
Be SEDG -528 (-64.23%) -582 (-70.8%)
BAMBU -619 (-75.3%) -643 (-78.22%)
-619 (-75.3%) -666 (-81.02%)
SEDG -575 (-69.95%) | -708 (-86.13%)
BAMBU -759 (-92.34%) | -797 (-96.96%)
711 (265% | 215 e9915%9
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 822)
The African Grass Blue is a modest little blue, which flies close to the ground. It is a species of dry
erassland, road verges, waste ground, and gardens, but sometimes butterflies are found in damp,
shady places. Various medicks, are used as foodplant, such as Lucerne (Medicago sativa), Tree Medick
(M. arborea), Small Medick (M. minima), Black Medick (M. /upulna), and possibly various Oxals
species. The caterpillars are attended by ants of the genus Phezdok. The African Grass Blue has two
of more generations a year and passes the winter as a caterpillar. There are only a few butterflies in
the first generation.
Present distribution can be well explained by climatic variables (AUC = 0.94).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 odd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
237
Zizeeria knysna (Lycaenidae)
2080
2050
238 Climatic Risk Atlas of European Butterflies
Cupido minimus (FuEssty, 1775) — Small Blue
ae ae
-2256 (-16.79%) | -3275 (-24.37%)
SEDG -4157 (-30.94%) | -5451 (-40.57%)
BAMBU_ | -4491 (-33.42%) | -6672 (-49.65%)
-6299 (-46.88%) | -8702 (-64.76%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 13437)
This butterfly is well named. It is very small and its modest colours make it even seem smaller
than it really is. The Small Blue can only be found on calcareous soils. Open, mostly rather short
vegetation and a warm microclimate typify its habitat. The butterflies may occur in large numbers.
They lay their small, white eggs singly between the flowers and the sepals of Kidney-vetch (Authylis
vulneraria), where a practised eye can detect them. The creamy-white caterpillars feed on the flowers
and seeds, and are seldom seen. However, workers of various ant species attend them regularly.
When fully-grown, the caterpillars hibernate either between the withered petals of dead flowers, or
in the litter layer and pupate on the ground. The Small Blue has one or two broods a year.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.71).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘dd 4000 0 Codd 4000 0 add 4000 0 odd 4000
1.0
Minimum
Swe
1.6
10 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 0.2
Swe
06
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
239
Cupido minimus (Lycaenidae)
2080
2050
240 Climatic Risk Atlas of European Butterflies
Cupido osiris (MEIGEN, 1829) — Osiris Blue
.—hCcLwe Full dispersal No dispersal
Be SEDG -470 (-42.11%) | -810 (-72.58%)
BAMBU -506 (-45.34%) -847 (-75.9%)
-566 (-50.72%) | -961 (-86.11%)
SEDG -154 (-13.8%) | -1003 (-89.87%)
BAMBU_ | -427 (-38.26%) | -1039 (-93.1%)
-373 (-33.42%) | -1101 (-98.66%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1116)
The Osiris Blue lives on sunny, flower-rich grassland and road verges rich in sainfoin (Oxobrychis
spp.), which it uses for its nectar and as foodplant. The caterpillar feeds on various sainfoins
(Onobrychis spp.), such as Sainfoin (O. vzzfola), Mountain Sainfoin (O. ~ontana), and O. arenaria. In the
literature, other leguminous plants are mentioned, e.g. Bladder Senna (Co/tea arborescens), Kidney-
vetch (Authylhs vulneraria), and Bitter Vetch (Lathyrus montanus). The eggs are laid on the flowerheads.
The caterpillars feed on both the flowers and developing seeds and are attended by ants (e.g, Laszus
alienus). Vhe Osiris Blue has one or two broods a year, but the second may only be partial. The
caterpillars hibernate and pupate between leaves.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.76).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
taad 4000 0 odd 4000 0 add 4000 0 Gdd 4000
Minimum
Swe
‘0.6
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
241
Cupido osiris (Lycaenidae)
2080
2050
242 Climatic Risk Atlas of European Butterflies
Cupido argiades (PA.LAs, 1771) — Short-tailed Blue
.6—hCcLUwE Full dispersal No dispersal
Be SEDG 1445 (9.62%) -1822 (-12.13%)
BAMBU 1387 (9.23%) | -2139 (-14.24%)
746 (4.96%) -2692 (-17.92%)
SEDG 1257 (8.37%) | -3979 (-26.48%)
BAMBU 1482 9.86%) | -5685 (-37.83%)
626 (4.17%) -7883 (-52.46%)
© Josef Settele Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 15026)
Although the Short-tailed Blue looks rather like the Holly Blue (Ce/astrina argiolus), it has very fine, small
tails and prominent eye-spots on the hindwings. The Short-tatled Blue occurs in local populations
on damp grassland, heathland and flower-rich verges. It lays its eggs on Lucerne (Medicago satina)
and various clovers (Ijfokum spp.), vetches (Via spp.), birdsfoot-trefoils (Lo/us spp.) and melilots
(Meklotus spp.). When the caterpillars are fully-erown, they leave the foodplant, overwinter, and then
pupate in the litter layer. This butterfly species has two to three generations a yeat.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.81).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
sir;
o
= E ; : ~ :
_
ah
o
2
wv
o
sc
o
‘dd 4000 0 edd 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
1.0 02
Swe
0.6
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
Swe
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
243
Cupido argiades (Lycaenidae)
2080
2050
244 Climatic Risk Atlas of European Butterflies
Cupido decoloratus (STAUDINGER, 1886) — Eastern Short-tailed Blue
6c Full dispersal No dispersal
Hee SEDG ESOL IO YO} -940 (-46.08%)
-308 (-15.1%) | -1568 (-76.86%)
-933 (-45.74%) | -2007 (-98.38%)
-879 (-43.09%) | -2031 (-99.56%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2040)
The Eastern Short-tailed Blue occurs in clearings in deciduous woodland, and on flower-rich, grassy
vegetation with scattered bushes. The caterpillars feed on the flowerheads of Black Medick (Medicago
lupulina), and Lucerne (M. sativa). The Eastern Short-tatled Blue has three generations a year and
hibernates as a caterpillar.
Present distribution can be well explained by climatic variables (AUC = 0.93).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 Gadd 4000 0 add 4000 0 add 4000
1.0
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 O.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O2
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Cupido decoloratus (Lycaenidae) 245
246 Climatic Risk Atlas of European Butterflies
Cupido alcetas (HOFFMANSEGG, 1804) — Provencal Short-tailed Blue
.—hCcLwe Full dispersal No dispersal
fi SEDG 321 (14.54%) -714 (-32.34%)
BAMBU 796 (36.05%) -717 (-32.47%)
473 (21.42%) -933 (-42.26%)
SEDG 1589 (71.97%) | -1036 (-46.92%)
BAMBU 1782 (80.71%) | -1328 (-60.14%)
—foras | 1325 (60.01%) -1785 (-80.84%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2208)
The Provencal Short-tatled Blue occurs in woodland clearings, at the edges of woods, on grassy
vegetation near bushes, and along the banks of streams and edges of ditches. Goat’s-rue (Gakga
officinals) and Crown Vetch (Coronilla varia) are its most important foodplants, but other leguminous
plants, such as Common Birdsfoot-trefoil (Lots cornuulatus), Common Vetch (Via sativa), and
Lucerne (Mediago sativa) are also used. The caterpillars feed mostly on the flowers, but also on the
leaves, and are attended by ants (e.g. of the genus Formica). This butterfly species has two to three
generations a year and hibernates as a caterpillar.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.77).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
he
o
=I
ze
o
™
o
0 2000 40000 2000 40000 2000 40000 2000 4000
Gdd Gdd Gadd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
247
Cupido alcetas (Lycaenidae)
2080
2050
248 Climatic Risk Atlas of European Butterflies
Celastrina argiolus (LINNAEUus, 1758) — Holly Blue
a
-2795 (-13.69%) | -4341 (-21.26%)
SEDG -2427 (-11.88%) | -4943 (-24.2%)
BAMBU_ | -5101 (-24.98%) | -7524 (-36.84%)
-6682 (-32.72%) | -10036 (-49.14%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 20422)
The Holly Blue is widespread, although the populations are always small. This small butterfly is very
often seen in parks and gardens, as well as at woodland edges, and in bushy places. The eggs are laid
on the calyx or stem of the flowerbuds or on the ripe fruits of various sorts of plants, including
Holly dx aquifolium), \Wwy (Hedera hehx), Spindle-tree (Evonymus europaeus), Alder Buckthorn (Frangula
alnus), brambles (Rubus spp.) and heathers (Cal/una vularis and Erica spp.). At first, the caterpillars
feed on the buds and fruits of the foodplant, only later eating leaves. They are attended by ants of
different genera. The Holly Blue has two broods a year, and hibernates in the pupal stage.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.66).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
=
sh
a
0
=
Ow
=a
te]
sity
a
e = : = = 7 ;
‘dd 4000 0 edd 4000 0 oad 4000 0 odd 4000
Minimum
Swe
06
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Celastrina argiolus (Lycaenidae) 249
(B1)
SEDG
GRAS
(A1Fl)
250 Climatic Risk Atlas of European Butterflies
Scolitantides baton (BERGSTRASSER, 1779) — Baton Blue
ee
SEDG -81 (-3.06%) -1179 (-44.61%)
BAMBU -88 (-3.33%) -1433 (-54.22%)
-847 (-32.05%) | -2209 (-83.58%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2643)
The Baton Blue can be found on dry, grassy, and flower-rich vegetation, especially on warm patches.
The butterflies are often seen on flowers, drinking nectar. The females lay their eggs on the flowers
and leaf stalks of various species of thyme (I/ymus spp.), savory (Satureja spp.), lavender (Lavandula
spp.), and mint (Mentha spp.). The caterpillars feed on the flowers and developing seeds of the
foodplant. They are attended by ants (e.g. Myrmica scabrinodis and Lasius alenus). The Baton Blue
passes the winter as a caterpillar or a pupa, and has two generations a yeat.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.83).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘dd 4000 0 odd 4000 0 ad 4000 0 add 4000
1.0
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
251
Scolitantides baton (Lycaenidae)
2080
2050
252 Climatic Risk Atlas of European Butterflies
Scolitantides vicrama (Moore, 1865)
SEDG 2985 (55.35%) -353 (-6.55%)
BAMBU 383 (7.1%) -1599 (-29.65%)
1968 (36.49%) | -962 (-17.84%)
SEDG 2198 (40.76%) | -1457 (-27.02%)
309 (5.73%) -2974 (-55.15%)
a 2622 (48.62%) | -3135 (-58.13%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 5393)
This blue ts found on dry grassland on sandy or stony soils, south-facing slopes with grassy
vegetation, on steep slopes, in rocky gullies, on railway embankments, on road verges and in scrub.
Various sorts of thyme (I/ymus spp.) and savory (Sa/ureja spp.) are used as foodplant, the eggs being
mostly laid on the flowerheads. The caterpillars feed on the flowers and ripening seeds, and are often
found with ants (e.g. Myrmica sabulen). This butterfly species is double-brooded.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.85).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
dd 4000 0 odd 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
1.0 02
0.6
Swe
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
253
Scolitantides vicrama (Lycaenidae)
2080
2050
254 Climatic Risk Atlas of European Butterflies
Scolitantides abencerragus (PIERRET, 1837) — False Baton Blue
6c Full dispersal No dispersal
He SEDG -567 (-21.16%) -1068 (-39.87%)
-1287 (-48.04%) | -1366 (-50.99%)
GRAS | -1074 (-40.09%) | -1454 (54.27%)
-2301 (-85.89%) | -2392 (-89.29%)
-2531 (-94.48%) | -2614 (-97.57%)
© Bernard Fransen Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2679)
The False Baton Blue is restricted to the Iberian Peninsula and 1s found in stony areas with flower-
rich vegetation and scrub. Different plants are given in the literature, such as Thyme (Thymus vuleans),
Green Heather (Eva scoparia), and the labiate annual Chonia lusitanica. It is on the leaves of Choma
/usitanica that the female lays her eggs, although the caterpillars feed on the flowers. The False Baton
Blue hibernates as a caterpillar, and is single-brooded in Europe.
Present distribution can be well explained by climatic variables (AUC = 0.94).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
aad 4000 0 edd 4000 0 Cad 4000 0 aad 4000
1.0
Minimum
Swe
0.6
o2
Swe
0.6 1.0
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
255
Scolitantides abencerragus (Lycaenidae)
2080
2050
256 Climatic Risk Atlas of European Butterflies
Scolitantides bavius (EVERSMANN, 1832) — Bavius Blue
=
SEDG 142 (33.18%) -170 (-39.72%)
BAMBU 36 (8.41%) -182 (-42.52%)
rarras% | 210 asa
SEDG 307 (71.73%) -230 (53.74%)
BAMBU -124 (-28.97%) | -331 (-77.34%)
-70 (-16.36%) -406 (-94.86%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 428)
The Bavius Blue occurs often on flower-rich, dry grassland, on dry, stony slopes, and on open
patches in shrub and in vineyards on calcareous soil. Various species of Sa/za are used as foodplant,
including Sage (S. officinals), S. nutans, S. verbenaca, and Whorled Clary (S. vertici/lata). The caterpillars
feed mostly on the flowers, but sometimes also on the leaves. They are frequently found with ants.
The Bavius Blue 1s single-brooded. The pupa overwinters.
This species is listed in Annexes II and IV of the Habitats’ Directive.
Present distribution can be well explained by climatic variables (AUC = 0.86).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
‘0.6
1.0 02
Swe
0.6
1.0 0.2
Swe
0.6 H fh. 1.0
Annual precipitation range
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
o2
a
2000 40000 2000 40000 2000 40000 2000 4000
Gd Gdd
Gdd Gdd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
257
Scolitantides bavius (Lycaenidae)
2080
2050
258 Climatic Risk Atlas of European Butterflies
Scolitantides orion (PALLAS, 1771) — Chequered Blue
.—ClcLe Full dispersal No dispersal
fe SEDG -24 (-0.38%) -1835 (-29.27%)
291 canson | 2668 (a2.%
|GRAS | -1532 (-24.43%) | -2699 (-43.05%)
SEDG -1446 (-23.06%) | -3844 (-61.31%)
BAMBU_ | -2659 (-42.41%) | -4917 (-78.42%)
ews 42.25%) | S827 CRASH
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 6270)
The Chequered Blue, a small but conspicuous butterfly, can be seen on warm, and rocky slopes and
narrow ledges, where there 1s little vegetation, apart from its foodplants White Stonecrop (Sedum
album) and Orpine (Sedum telephium).’The females lay their eggs on the leaves of the foodplant near
the stem. The caterpillars are often found with different ants. The pupa overwinters, and is often
hidden under stones or in small hollows in the ground near the foodplant. The Chequered Blue
produces one or two generations a year, depending on its geographical position.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.78).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
-
‘dd 4000 0 edd 4000 0 oad 4000 0 odd 4000
Minimum
Swe
‘0.6
1.0 02
0.6
Swe
1.0 O.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 02
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
259
Scolitantides orion (Lycaenidae)
2080
2050
260 Climatic Risk Atlas of European Butterflies
Scolitantides panoptes (HUBNER, 1813) — Panoptes Blue
SEDG -867 (-58.07%) | -971 (-65.04%)
-1124 (-75.28%) | -1130 (-75.69%)
a -1146 (-76.76%) | -1207 (-80.84%)
SEDG -1324 (-88.68%) | -1344 (-90.02%)
BAMBU -1493 (-100%) -1493 (-100%)
-1493 (-100%) -1493 (-100%)
© Bernard Fransen Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1493)
At present the species is widespread in the Iberian Peninsula. It lives on dry grasslands between 200
and 1900m elevation. The caterpillars are visited by ants of the genus Camponotus. The species has
two generations and one can find adults on the wing from March until August. Larval foodplants
ate Thymus and Saturja species. The taxonomic status (whether subspecies of S. baton or a separate
species) is under discussion.
Present distribution can be well explained by climatic variables (AUC = 0.87).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
toad 4000 0 odd 4000 0 Aad 4000 0 add 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
2050
Scolitantides panoptes (Lycaenidae)
2080
SEDG
261
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
262 Climatic Risk Atlas of European Butterflies
Glaucopsyche alexis (Popa, 1761) — Green-underside Blue
-1448 (-9.24%)
-2065 (-13.17%)
AMBU 1588 (10.13%) | -5095 (-32.5%)
1591 (10.15%) | -6785 (-43.28%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 15676)
The Green-underside Blue is a pretty sight on both dry and damp flower-rich grassy vegetation, such
as meadows and woodland clearings. The females lay their eggs between the flowers of different
leguminous plants, including Sainfoin (Oxobrychis vatfola), brooms (Cylisus spp.), vetches (Via spp.),
Crown Vetch (Coronilla varia), Genista spp., and melilots (Med/oius spp.). The caterpillars feed on the
leaves and are frequently attended by the workers of various ants. Hibernation takes place as a pupa.
The Green-underside Blue is single-brooded.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.7).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
‘dd 4000 0 odd 4000 0 ad 4000 0 odd 4000
Minimum
Swe
‘1.6
10 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 0.2
06
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
263
Glaucopsyche alexis (Lycaenidae)
WE
4%
ahs “4
2080
2050
264 Climatic Risk Atlas of European Butterflies
Glaucopsyche melanops (BotsDUVAL, 1828) — Black-eyed Blue
.—ClcLw Full dispersal No dispersal
|
SEDG -720 (-20.13%) -2069 (-57.84%)
AMBU_ | -1708 (-47.75%) | -2238 (-62.57%)
a -1333 (-37.27%) -2464 (-68.88%)
SEDG -1767 (-49.4%) | -2884 (-80.63%)
=
B
AMBU | -2319 (-64.83%) | -3287 (-91.89%)
r -2032 (-56.81%) | -3524 (-98.52%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 3577)
The Black-eyed Blue occurs in open shrub, or open woodland, especially where Genisia or large
bushes of broom (Cyésus spp.) ate growing. Eggs are laid on the flowers of various leguminous
plants, such as Dorycnium spp., Birdsfoot-trefoils (Lotus spp.), Anthylis spp., brooms (Cylisus spp.) and
Genista spp.. The caterpillars feed on the flowers and are attended by ants of the genus Camponotus.
The Black-eyed Blue hibernates in the pupal stage. It usually has one generation a year.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.8).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
1.6
10 02
Swe
6 1002 O86 16 10
| |
oOo
Swe
06
Annual precipitation range
1d 02
06
Maximum Large (66%) Small (33%)
Swe
o2
L=]
2000 40000 2000 40000 2000 40000 2000 4000
Gdd
Gdd Gdd Gdd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
265
Glaucopsyche melanops (Lycaenidae)
2080
2050
266 Climatic Risk Atlas of European Butterflies
Tolana iolas (OCHSENHEIMER, 1816) — Iolas Blue
.6—hCcLwe Full dispersal No dispersal
Be SEDG -102 (-3.08%) -1371 (-41.46%)
BAMBU_ | -1610 (-48.68%) | -1909 (-57.73%)
-974 (-29.45%) | -1849 (-55.91%)
SEDG -1564 (-47.29%) | -2149 (-64.98%)
BAMBU_ | -2625 (-79.38%) | -2938 (-88.84%)
-2596 (-78.5%) | -3186 (-96.34%)
© Rudi Verovnik Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 3307)
The Iolas Blue is Europe’s largest blue. It occurs locally on calcareous soil, where shrubs of its
foodplant Bladder Senna (Colutea arborescens) are growing. In Greece, C. cicica is also used. This plant
is practically the most important source of nectar for the butterflies. At times, the males can be seen
some distance away from their habitat, but the females stay near the foodplants. They lay their eggs,
usually several at a time, on the inside of the calyx and inside the bladder-like fruits. The caterpillars
feed on the seeds and are visited frequently by ants (e.g. Tapinoma erraticum and Camponotus cruentatus).
Usually, they can easily be seen by holding a pod up to the light. When fully-grown, the caterpillars
pupate at the foot of the foodplant, passing the winter as a pupa. The Iolas Blue mostly has only
one generation a yeat.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.82).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘sad 4000 0 edd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
267
Tolana iolas (Lycaenidae)
2080
2050
268 Climatic Risk Atlas of European Butterflies
Phengaris arion (L1NNAEus, 1758) — Large Blue
-2206 (-16.66%) | -4172 (-31.51%)
SEDG -1865 (-14.08%) | -5421 (-40.94%)
BAMBU_ | -1732 (-13.08%) | -6599 (-49.83%)
-3006 (-22.7%) -8692 (-65.64%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 13242)
The Large Blue occurs locally on dry, open grasslands on limestone. It is one of the larger, more
conspicuous blues. The females lay their eggs on Marjoram (Onganum vuleare) and different species
of thyme (T/ymus spp.). The caterpillars feed on the buds and flowers of the foodplant until they
reach the last larval instar. They then leave their foodplant, and allow themselves to be taken by
workers of Myrmica sabukti, and sometimes of M. scabrinodis, to the ants’ nest. The caterpillars feed
on the ant grubs, hibernating and pupating in ants’ nests as well.
This species is listed in Annex IV of the Habitats’ Directive.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.77).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
=
sh
=
a
=
Ow
=a
i]
is}
Oo
: : ; : ; ‘ — :
dd 4000 0 ood 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
269
Phengaris arion (Lycaenidae)
2080
2050
270 Climatic Risk Atlas of European Butterflies
Phengaris teleius (BERGSTRASSER, 1779) — Scarce Large Blue
.—ClcLe Full dispersal No dispersal
Be SEDG -35 (-0.54%) -2184 (-33.69%)
BAMBU -551 (-8.5%) -2612 (-40.29%)
-1537 (-23.71%) | -3224 (-49.73%)
SEDG -1008 (-15.55%) | -3984 (61.45%)
BAMBU_ | -1587 (-24.48%) | -4723 (-72.85%)
-2476 (-38.19%) | -5731 (-88.4%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 6483)
The Scarce Large Blue can be found in moderately nutrient-rich meadows where its foodplant Great
Burnet (Sangusorba offiinals) 1s growing. In northern Europe, it occurs in open, short vegetation, but
in the warm, southern parts, it is also found in rough vegetation. The butterflies tend to keep near
the foodplants. The small caterpillars only feed on the flowerheads for two or three weeks. They
then go down to the ground where they wait to be picked up by worker ants of the genus Myrmica
and carried off to the ants’ nest. There they feed on ant grubs. The caterpillars also hibernate and
pupate in the ants’ nest. The species of host ant varies in different parts of its range. The Scarce
Large Blue is single-brooded.
This species is listed in Annexes II and IV of the Habitats’ Directive.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.84).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
dd 4000 0 edd 4000 0 add 4000 0 odd 4000
Minimum
Swe
1.6
1.0 02
0.6
Swe
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
271
Phengaris teleius (Lycaenidae)
2080
2050
272 Climatic Risk Atlas of European Butterflies
Phengaris nausithous (BERGSTRASSER, 1779) — Dusky Large Blue
.—hCcLwe Full dispersal No dispersal
Be SEDG -830 (-22.93%) -1999 (-55.24%)
BAMBU -699 (-19.31%) | -1960 (54.16%)
-1451 (-40.09%) | -2501 (-69.11%)
SEDG -789 (-21.8%) | -2643 (-73.03%)
BAMBU_ | -1716 (-47.42%) | -3061 (-84.58%)
-2334 (-64.49%) | -3460 (-95.61%)
© Josef Settele Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 3619)
The Dusky Large Blue occurs on damp, moderately nutrient-rich grassland and rough vegetation.
The butterflies are usually found on or near the foodplant Great Burnet (Sanguisorba officinals).
Having lived on the flowerheads of this plant for a few weeks, the small caterpillars go down to the
ground, in order to be carried away by workers of the ant Myrmica rubra to an ant nest. There, they
remain feeding on ant grubs, hibernating and pupating in the early summer. The newly-emerged
butterflies leave the nest. The Dusky Large Blue is one of the most specialized of the “ant blues”
being most adapted to one species of host ant. It is single-brooded.
This species is listed in Annexes II and IV of the Habitats’ Directive.
Present distribution can be well explained by climatic variables (AUC = 0.91).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
dd 4000 0 edd 4000 0 add 4000 0 Gdd 4000
Minimum
Swe
‘1.6
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
273
Phengaris nausithous (Lycaenidae)
2080
2050
274 Climatic Risk Atlas of European Butterflies
Phengaris alcon ({SCHIFFERMULLER], 1775) — Alcon Blue
SEDG -800 (-12.04%) | -2001 (-30.12%)
BAMBU 375 (5.65%) -1448 (-21.8%)
-1388 (-20.89%) | -2689 (-40.48%)
SEDG -741 (-11.15%) | -2794 (-42.06%)
BAMBU_ | -1062 (-15.99%) | -3525 (-53.06%)
-2097 (-31.57%) | -4883 (-73.51%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 6643)
The Alcon Blue occurs in local, scattered populations. Oftentimes it is referred to as two distinct
species: P. akon on low-lying wet heathland, on moist fen meadows and bogs, and P. rebeé on dry as
well as sub-alpine calcareous grasslands. Usually, only a few butterflies are seen. The bright, white
eges ate easy to find. Depending on the habitat, they are laid on the flowers and sepals of Marsh
Gentian (Gentiana pneumonanthe), Willow Gentian (G. asclepiadea), and Cross Gentian (G. cruciata). At
first, the small caterpillars feed on the ovaries of the foodplant, but they pass the last instar in the
nests of various Myrmica ants, which they parasitize by living like young cuckoos, being fed by the
worker ants. They hibernate and pupate in the ant nests. The Alcon Blue has one generation a year.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.83).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
‘1.6
sh
a a
=) a
ee
Pm On
cfes
o= mH
=
£5 8
Oo. 4
ow 6
fo
o ng = a
ger
|
ae
a
@e-
2 o
wv
x é Po]
= 4
o
0 2000 40000 2000 40000 2000 40000 2000 4000
Gd id Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
275
Phengaris alcon (Lycaenidae)
2080
2050
276 Climatic Risk Atlas of European Butterflies
Plebejus argus (LINNAEUS, 1758) — Silver-studded Blue
a
-2138 (-11.19%) | -4148 (-21.71%)
SEDG -865 (-4.53%) | -5024 (-26.29%)
BAMBU_ | -3038 (-15.9%) | -7385 (-38.65%)
12s conser) | 9598 (302579
© Martin Wiemers Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 19107)
The Silver-studded Blue can be found in warm places on short vegetation, ranging from dry to
quite damp, such as heath and poor grassland. Especially impressive are the roosting places with
enormous groups, sometimes of hundreds of butterflies, asleep in a very small area. The eggs are
laid on Cross-leaved Heath (Ema retrain), and on a wide range of leguminous plants. Usually, the
eggs are deposited low down on the foodplant or on its woody parts. The egg hibernates. The
caterpillars live on the leaves of the foodplants. They are often tended by ants of the genus Lasius.
Pupation often takes place in the outer passages of the ant nests. The ants also give the pupa a lot
of attention. Depending on the geographical position and altitude of occurrence, the Silver-studded
Blue has one or two generations a yeat.
Present distribution can be explained by climatic variables to only a moderate extent (AUC = 0.69).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
O06
1.0 02
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
E 1.0 0.2 0.6 fh. 1.0
Swe
o2
0 2000 40000 2000 40000 2000 40000 2000 4000
Gdd Gdd Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
277
Plebejus argus (Lycaenidae)
2080
2050
278 Climatic Risk Atlas of European Butterflies
Plebejus idas (LINNAEUS, 1761) — Idas Blue
a
-5296 (-28.72%) | -5309 (-28.79%)
SEDG -6083 (-32.99%) | -6192 (-33.58%)
BAMBU_ | _ -7986 (-43.31%) | -8006 (-43.42%)
-9992 (-54.19%) | -10023 (-54.36%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 18438)
The Idas Blue can be found on poor grassland and heath rich in herbaceous plants, both in open
places and woodland. Eggs are laid on many different species of leguminous plants, such as Broom
(Cytisus scoparius), Hairy Greenweed (Genzsta pilosa), Common Birdsfoot-trefoil (Coronilla varia), White
Melhilot (Med/otus alba), and the Ericaceae, Ling or Heather (Ca/luna vulgaris) and Bog Whortleberry
(Vaccinium uliginosum), and the Yellow Rocktrose (Hehanthemum oelandicum). The female deposits the
ege onto a woody part of the foodplant where the egg then hibernates. The caterpillars are attended
a lot by ants, especially those of the genera Formica and Lasius. When fully grown, the caterpillar
crawls into the passages of the ant nest to pupate. Depending on the geographical location the Idas
Blue produces one or two generations a yeat.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.74).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 Codd 4000 0 oad 4000 0 odd 4000
Minimum
Swe
‘1.6
1.0 02
0.6
Swe
1.0 0.2
Swe
0.6
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
O06
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
279
Plebejus idas (Lycaenidae)
2080
2050
280 Climatic Risk Atlas of European Butterflies
Plebejus argyrognomon (BERGSTRASSER, 1779) — Reverdin’s Blue
aa ee
1176 (21.05%) | -2065 (-36.95%)
ee 267 (4.78%) -2681 (-47.98%)
SEDG 760 (13.6%) -3178 (-56.87%)
BAMBU 297 (5.31%) -4064 (-72.73%)
-655 (-11.72%) | -4776 (-85.47%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 5588)
Reverdin’s Blue can be found on poor grassland in places ranging from dry to moist at the edges
of woodland or scrub. Eggs are laid on Crown Vetch (Coromilla varia) and Milk-vetch (Astragalus
ghcyphyllos). The female mostly chooses a woody part of the plant, where the egg may hibernate.
The small caterpillars feed on the young leaves. They are attended by Lasius and Myrmica as well as
Formica and Camponotus ants, and pupate deep down in the vegetation. Except 1n Scandinavia where
it is single-brooded, the Reverdin’s Blue has two generations a year.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.8).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘sad 4000 0 Codd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 0.2
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
281
Pkebejus argyrognomon (Lycaenidae)
2080
2050
282 Climatic Risk Atlas of European Butterflies
Plebejus optilete (KNocH, 1781) — Cranberry Blue
a se
-3065 (-30.37%) | -3153 (31.24%)
SEDG -4367 (-43.27%) | -4580 (-45.38%)
BAMBU_ | _ -4949 (-49.04%) | -5057 (-50.11%)
-6135 (-60.79%) | -6209 (-61.52%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 10092)
The Cranberry Blue occurs on raised bogs, heath and also in woodland clearings, where dwarf
shrubs are growing. Although populations are at times extremely small, they can also have
considerable numbers of butterflies. Eggs are laid on Marsh Andromeda (Andromeda palusins),
Erica tetrahx, Empetrum nigrum, and a small number of Vaccinium species, including Cranberry
(V. oxycoccus). The caterpillars eat both the flowers and leaves of the foodplants, moving from one
plant to another. When they are half-grown, they move into the litter layer and hibernate. However,
they pupate on the plant, spinning a silken girdle in which the pupa hangs. The Cranberry Blue
has one generation a year.
Present distribution can be well explained by climatic variables (AUC = 0.9).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Gadd 4000 0 edd 4000 0 oad 4000 0 ada 4000
1.0
Minimum
Swe
0.6
o2
Swe
0.6 1.0
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
Swe
0.6
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
283
Plkebejus optiletev (Lycaenidae)
2080
2050
284 Climatic Risk Atlas of European Butterflies
Plebejus glandon (PRUNNER, 1798) — Glandon Blue
CLL Full dispersal No dispersal
He SEDG 14 (3.29%) -229 (-53.88%)
94 (22.12%) -189 (-44.47%)
2eaamn | 237 655709
SEDG -221 (-52%) -348 (-81.88%)
BAMBU -93 (-21.88%) -283 (-66.59%)
-235 (-55.29%) | -363 (-85.41%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 425)
In Central Europe, the Glandon Blue is found up to the snow-line on grassy vegetation, where
the foodplants grow on open patches. On calcareous soils, Axdrosace chamacjasme is used, and on
non-calcareous soils, the caterpillars feed on A. oblusifoha and Vitakana primulflora. The caterpillars
feed on the leaves and sometimes the flowers of the foodplants. The Glandon Blue hibernates as
a caterpillar, and in all parts of its range has one generation a year. We treat Plebejus glandon as
separate from P. aguilo (BoIsDUVAL, 1832) (see next species) and P. zwellichi (HEMMING, 1933) @vhich
is endemic to the Sierra Nevada and was not modelled here due to its restricted distribution), while
other authors split the species into three subspecies.
Present distribution can be very well explained by climatic variables (AUC = 0.99).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
toad 4000 0 Codd 4000 0 oad 4000 0 aad 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
285
Plebejus glandon (Lycaenidae)
(1a)
5)q04s
(ZV)
nanva
2080
2050
286 Climatic Risk Atlas of European Butterflies
Plebejus aquilo (BoisDUVAL, 1832) — Arctic Blue
6c Full dispersal No dispersal
Hi SEDG -462 (-66.47%) -529 (-76.12%)
BAMBU -479 (-68.92%) | -537 (-77.27%)
-497 (-71.51%) | -553 (-79.57%)
SEDG -562 (-80.86%) | -637 (-91.65%)
BAMBU -646 (-92.95%) | -685 (-98.56%)
GRAS -682 (-98.13%) -695 (-100%)
© Bernard Fransen Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 695)
This species occurs exclusively in Northern Fennoscandia. It occurs on slate and shale rocks with
patches of grassy vegetation and with Crowberry (Empetrum nigrum), especially in areas sheltered
from the northwestern wind. It lives in elevations between 0 and 900 m asl. Larval foodplants are
Saxifraga aixoides and S. oppositifola. The small larvae feed on the flower buds and hibernate. Later the
larvae feed on the leaves. The species has one generation which is on the wing from late June until
early August. It is often considered as a subspecies of Plebejus glandon.
Present distribution can be very well explained by climatic variables (AUC = 1).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘sad 4000 0 odd 4000 0 add 4000 0 add 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Pkebejus aquilo (Lycaenidae) 287
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
2050 2080
288 Climatic Risk Atlas of European Butterflies
Plebejus orbitulus (PRUNNER, 1798) — Alpine Blue
.—hCcLUwe Full dispersal No dispersal
Be SEDG -234 (-18.84%) -356 (-28.66%)
BAMBU -226 (-18.2%) -333 (-26.81%)
-299 (-24.07%) | -397 (-31.96%)
SEDG -352 (-28.34%) | -505 (-40.66%)
BAMBU -365 (-29.39%) | -489 (-39.37%)
GRAS -493 (-39.69%) | -615 (-49.52%)
© Martin Wiemers Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1242)
In the Alps, the Alpine Blue is mostly found on sunny, rocky slopes, scree slopes with patches of
flower-rich vegetation, and often on the rough vegetation growing beside streams. These butterflies
usually fly to lower-lying areas in search of nectar. The foodplants are Alpine Milk-vetch (Astragalus
alpinus), Alpine Sainfoin (Hedysarum hedysaroides), and probably Mountain Beaked Milk-vetch
(Oxytropis jacquini). In Scandinavia, this blue occurs in the mountains on open, damp slate slopes
where its foodplant Alpine Milk-vetch grows. The Alpine Blue hibernates as a small caterpillar and
has one generation a year.
Present distribution can be very well explained by climatic variables (AUC = 0,98).
Climate risk category: LR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
Swe
0.6
0 2000 40000 2000 40000 2000 40000 2000 4000
Gdd d Gdd
Gdd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
289
Plebejus orbitulus (Lycaenidae)
2080
2050
290 Climatic Risk Atlas of European Butterflies
Plebejus sephirus (Frivacpszky, 1835)
.—CcLe Full dispersal No dispersal
Be SEDG 3 (0.25%) -833 (-70.06%)
BAMBU_ | -143 (-12.03%) | -760 (-63.92%)
743 (62.49%) -676 (-56.85%)
SEDG -100 (-8.41%) | -1115 (-93.78%)
BAMBU_ | -441 (-37.09%) | -1020 (-85.79%)
330 (27.75%) -853 (-71.74%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1189)
By some authors this species, described from Bulgaria, is treated as a subspecies of Pkbejus pylaon
(FISCHER, 1832) and as a distinct species by others. Other taxa besides P. pylaon, which are closely
related to P. sephirus, are P. hespericus (RAMBUR, 1839) from Spain and the Alpine P. frappi (VERITY,
1927). All of them are not treated here due to their limited distributions. It occurs eastwards of the
river Danube to the Dnepr region and southwards through Transylvania to the Balkans. Its habitats
are loess steppes and calcareous terrains in the forest steppe belt from sea level to 1600 m. Larval
host-plants are Astragalus species. The caterpillars are facultatively myrmecophilous and are garded
by many different ant species. It has one short generation in May, June or July, depending on altitude
and latitude. It hibernates as a small larva.
Present distribution can be well explained by climatic variables (AUC = 0.93).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
‘dd 4000 0 odd 4000 0 oad 4000 0 add 4000
Minimum
Swe
0.6
o2
Swe
0.6 1.0
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
Swe
0.6
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
291
Pkebejus sephirus (Lycaenidae)
2080
2050
292 Climatic Risk Atlas of European Butterflies
Aricia eumedon (Esrrr, 1780) — Geranium Argus
SEDG -3967 (-29.63%) | -4888 (-36.51%)
BAMBU -6493 (-48.5%) | -7131 (53.26%)
-7151 (53.41%) | -7854 (58.66%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 13389)
The Geranium Argus occurs locally, in meadows, stream valleys, and at woodland edges.
The female lays her eggs in the flowers of crane’s-bills (Geranium spp.), at the base of the ovary into
which the small caterpillar bores its way on hatching out. It stays there for a little more than a week.
It then starts feeding on leaves, first gnawing on the stems, which causes them to wilt. When half-
grown, the caterpillars hibernate in the litter layer, but pupate on the foodplant. They are sometimes
attended by ants of the genera Myrmica, Lasius and Tapinoma. The Geranium Argus has only one
generation a year.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.8).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
0.6
sh
a a
=) a
i e
Pe Oe
cfes
o= mH
=e |
£5 3
Oo. 4
oO r
28
a ng = a
ger
i |
ae
qf. =
eo
wo
x é 5
= 4
o
0 2000 4000 2000 40000 2000 40000 2000 4000
Gd id Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
293
Alricia eumedon (Lycaenidae)
2080
2050
294 Climatic Risk Atlas of European Butterflies
Aricia cramera (ESCHSCHOLTZ, 1821)
a
-1591 (-64.39%) | -1843 (-74.59%)
SEDG -1692 (-68.47%) | -2033 (-82.27%)
BAMBU_ | -2054 (-83.12%) | -2346 (-94.94%)
-1950 (-78.92%) | -2448 (-99.07%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2471)
This butterfly species can be found in flower-rich, places, in scrub or at woodland edges.
The most important foodplants are stork’s-bills (Erodium spp.) and crane’s-bills (Geranium spp.),
of which the caterpillars eat the leaves. Knapweeds (Cenfaurea spp.), tockroses (Hekanthemum spp.),
and Leguminous plants, such as clovers (Imfolum spp.) and brooms (Cytsus spp.) have also been
mentioned. This species has two to three generations a year and is often found together with ants
of the genera Myrmica and Lasius. Hibernation takes place as larva.
Present distribution can be well explained by climatic variables (AUC = 0.91).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
=
mm
a
a
=
Ow
=a
ee]
cH
oO
ne , 4 : il : ? :
mH
o
o
Ow
$0
LP)
™
o
faa 4000 0 ood 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
295
Alritia cramera (Lycaenidae)
(1a)
504s
(ZV)
nNanva
2080
2050
296 Climatic Risk Atlas of European Butterflies
Aricta agestis ({SCHIFFERMULLER], 1775) — Brown Argus
.6—hCcLUwk Full dispersal No dispersal
Be SEDG 299 (1.96%) -2499 (-16.38%)
BAMBU -354 (-2.32%) | -3124 (-20.48%)
-837 (5.49%) | -3631 (23.81%)
SEDG -2279 (-14.94%) | -4933 (-32.34%)
BAMBU_ | -3463 (-22.7%) | -6939 (-45.49%)
-4787 (-31.38%) | -8715 (-57.14%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 15253)
The Brown Argus occurs on warm grasslands, in heaths, and in dunes. It can also be found on
sandy, pioneer vegetation, as for example in sand-pits, or where houses are being built. It lays
its eggs on small crane’s-bill (Geranium spp.) and rockroses (Hedanthemum spp.). It may build up
numerous, small populations. At first, the small caterpillars only scrape off the undermost layer
of the leaves making little “windows” in the leaves. Later, the bigger caterpillars feed on the whole
leaf. They are often attended by Lasiws and Myrmica ants. The caterpillars can hibernate at any stage.
Pupation takes place in the litter layer. The Brown Argus has two to three generations depending
on the geographical location.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.7).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
m
a
- : : - ?
todd 4000 0 odd 4000 0 oad 4000 0 ad 4000
1.0
Minimum
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe Swe
0.6 0.6
1.0 02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Arricia agestis (Lycaenidae) 297
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
298 Climatic Risk Atlas of European Butterflies
Aricta artaxerxes (Fasricius, 1793) — Northern Brown Argus
a
-4476 (-46.3%) -4601 (-47.59%)
SEDG -4943 (51.13%) | -5186 (-53.64%)
BAMBU_ | -7110 (-73.54%) | -7262 (-75.11%)
-7886 (-81.57%) | -8037 (-83.13%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 9668)
The Northern Brown Argus likes open grasslands or dry vegetation at the edges of woods. It ts
more closely bound to calcareous soil than the Brown Argus. The eggs are laid on various rockroses
(Fekanthemum spp.), as well as on various crane’s-bills (Geranium spp.). The caterpillars feed on the
leaves, often attended by Laszus ants. When the caterpillars are half-grown, they hibernate. Pupation
takes place in the litter layer. There is always only one generation.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.72).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
0.6
Swe
0 2000 40000 2000 40000 2000 40000 2000 4000
Gd dl Gdd
Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
299
artaxerxes (Lycaenidae)
rita
A
2080
2050
300 Climatic Risk Atlas of European Butterflies
Aricta montensis (VERITY, 1928) — Southern Mountain Argus
.ClcLw Full dispersal No dispersal
SEDG -458 (-53.32%) -618 (-71.94%)
BU -606 (-70.55%) | -683 (-79.51%)
BAM
-593 (-69.03%) | -755 (-87.89%)
SEDG -591 (-68.8%) -804 (-93.6%)
BAMBU -605 (-70.43%) | -819 (-95.34%)
-464 (54.02%) | -840 (-97.79%)
© Matt Rowlings Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 859)
The taxonomic relationship of several Anca species is not yet cleat. Aria montensis is often regarded as
a subspecies of A. arfaxerxes. In the context of this atlas the species has been modeled independently,
but there also would have been good reasons to keep it within A. arfanxerxes. It occurs on the mountains
south of the range or A. arflaxerxes, 1t has no white spots on the upperside of the wings and distinct
black spots underneath them (like the subspecies A. a. a//ous, but 1t is bigger than those).
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.7).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
1.6
ct
Qa i=]
=) 2
= 7
Pm Oe
a =o
o= mH
=e
£5 8
Oo 4
Ome 6
oo
aes
= 0
SR 4
hee
qe. =
Pos
® é 5
= 4
o
2000 4000-0
Gdd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
301
Alritia montensis (Lycaenidae)
(1a)
04s
(ZV)
Nanva
(ld LV)
SVH9
2080
2050
302 Climatic Risk Atlas of European Butterflies
Aricta anteros (FREYER, 1838) — Blue Argus
=r Full dispersal No dispersal
z
S
-64 (-12.72%) -300 (-59.64%)
-300 (-59.64%) | -445 (-88.47%)
-355 (-70.58%) | -488 (-97.02%)
© Rudi Verovnik Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 503)
The Blue Argus is a species of flower-rich grasslands and rocky slopes with grassy vegetation.
Sometimes, they are also seen on scrub or in woodland clearings. This butterfly it is often found
where the soil is calcareous. Various sorts of crane’s-bills (Geranium spp.) are used as foodplant,
including G. asphodeloides, Bloody Crane’s-bill (G. sanguineum), and Rock Crane’s-bill (G. macrorrhizum).
The caterpillars feed on the leaves and are attended by ants. Depending on the altitude, the Blue
Argus has one to three generations a yeat.
Present distribution can be very well explained by climatic variables (AUC = 0.96).
Climate risk category: HHHAR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
|
pa
o
cil
o
aad 4000 0 odd 4000 0 add 4000 0 add 4000
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 o.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O2
Swe
od “4, o pe
‘=
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
303
Arricia anteros (Lycaenidae)
(1a)
504s
(ZV)
Nanva
2080
2050
304 Climatic Risk Atlas of European Butterflies
Aricta nictas (MEIGEN, 1829) — Silvery Argus
ae ee
-1380 (-32.09%) | -1528 (35.53%)
ams | aera [i
SEDG -207 (-4.81%) | -1031 (-23.97%)
BAMBU_ | -1611 (-37.46%) | -2242 (-52.13%)
GRAS -1952 (-45.38%) | -2625 (-61.03%)
© Bernard Fransen Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 4301)
The Silvery Argus occurs on flower-rich grasslands with some bushes or trees, in grass along the
hedges, at the edge of woodland, and in sunny, grassy woodland clearings. The grassland on which
the Silvery Argus occurs is usually damp with rather tall vegetation, but it is also seen on dry grassland.
The female lays her eges on the flowers of Wood Crane’s-bill (Geranium syhaticum) and Meadow
Crane’s-bill (G. pratense). The caterpillars eat the leaves as well as the flowers. They are attended by
ants. This butterfly hibernates as a larva, but in Scandinavia it is said to overwinter in the egg stage.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.82).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
_
cm
a
=
=
Ow
=o
uw
m™
Oo
a : ; : : .
4
o
=
1H
o
‘dd 4000 0 odd 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
305
(Lycaenidae)
ruta nicias
A
2080
2050
306 Climatic Risk Atlas of European Butterflies
Cyaniris semiargus (ROTTEMBURG, 1775) — Mazarine Blue
.—hlcLUw Full dispersal No dispersal
fi SEDG -131 (-0.75%) -2574 (-14.83%)
-349 (-2.01%) | -2498 (-14.39%)
-2792 (-16.08%) | -6166 (-35.52%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 17359)
The Mazarine Blue usually occurs on quite damp vegetation in flower-rich meadows and pastures
and at the edges of woodland. These butterflies are fond of basking together in groups and are
then easy to find and to be observed. The female deposits her eggs on the flowerheads of Red
Clover (Infolum pratense), eating the unopened buds. The first instar caterpillar only feeds on buds
and flowers, later stages also feed on leaves. The colours of the caterpillars are well adapted to
their surroundings, with pink in the first instar and then green in later stages. The caterpillars can
hardly be seen while feeding on the foodplants. Ants of the genera Lasius and Camponotus attend the
caterpillars. Depending on the altitude and position in the range, the Mazarine Blue has one to three
generations a year. The hibernation takes place as a larva.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.76).
Climate risk category: LR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
06
|
a =
=) a
= 7
Be
ee =a
o= mH
=
£5 8
Oo 4
of 7
vases a. es
eh ™ is on
te |
aoe
a
@e-
Zo
wo
x é Po]
= 4
o
0 2000 40000 2000 40000 2000 40000 2000 4000
Gd id Gadd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
307
Cyaniris semiargus (Lycaenidae)
ap] Th we
i
iT
a
as q
fa
OM,
2080
2050
308 Climatic Risk Atlas of European Butterflies
Polyommatus escheri (HUBNER, 1823) — Eschet’s Blue
cos
-1061 (-45.93%) -1633 (-70.69%)
ee -1243 (-53.81%) -1873 (-81.08%)
SEDG -788 (-34.11%) | -1988 (-86.06%)
BAMBU_ | -1188 (-51.43%) | -2116 (-91.6%)
-1178 (-51%) -2268 (-98.18%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2310)
Escher’s Blue occurs on dry, and flower-rich grassland, damp rough vegetation, in flower-rich rocky
places, on scrub and at woodland edges, and on agricultural land. The females lay their eggs singly
on milk-vetches (Astragalus spp.), in particular A. monspessulanus. Oxytropis hehetica may also be a
foodplant. When still small, the caterpillar goes into hibernation, and in the spring completes its
erowth, feeding on the leaves and flowerbuds of the foodplants. When fully grown, it leaves the
foodplant and pupates under stones. Both caterpillars and pupae are attended by ants of different
genera including Myrmica, Formica, Lasius and Plagiolepis. Eschet’s Blue is single-brooded. It hibernates
as a larva.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.8).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
-
dd 4000 0 edd 4000 0 add 4000 0 Gdd 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
309
Polommatus eschert (Lycaenidae)
2080
2050
310 Climatic Risk Atlas of European Butterflies
Polyommatus dorylas ({SCHIFFERMULLER], 1775) — Turquoise Blue
.—hlcLwE Full dispersal No dispersal
Be SEDG -949 (-21%) -2030 (-44.91%)
BAMBU_ | -1526 (-33.76%) | -2328 (-51.5%)
-1804 (-39.91%) | -2684 (-59.38%)
SEDG -372 (-8.23%) | -2706 (-59.87%)
BAMBU_ | -1383 (30.6%) | -3262 (-72.17%)
-1903 (-42.1%) | -3725 (-82.41%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 4520)
The Turquoise Blue likes the warmth, occurring on dry, flower-rich slopes and calcareous grassland,
often where there is shelter from a wood or from bushes. The butterflies are nearly always seen on
calcareous ground. The populations are usually small in mountainous areas. The female lays her eggs
on Kidney-vetch (Axnthylis vulnerana), depositing them onto the underside of the leaves and also
on the sepals. The newly-hatched caterpillars feed by scraping off the undermost layer of cells, so
making little “windows” in the leaves. Later, they feed on the whole leaf. They are attended by various
ant species, including those belonging to the genera Myrmica, Lasius, and Formica. The caterpillars
pupate in the litter layer. The Turquoise Blue mostly has two generations, but at higher altitudes and
in the north of its range, it is single-brooded. Hibernation takes place as a small larva.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.81).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
Bm
Gadd 4000 0 Codd 4000 0 Aad 4000 0 odd 4000
Minimum
Swe
‘1.6
1.0 02
0.6
Swe
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
311
Polyommatus dorylas (Lycaenidae)
2080
2050
312 Climatic Risk Atlas of European Butterflies
Polyommatus nivescens (KEFERSTEIN, 1851) — Mother-of-pearl Blue
.—ClcLe Full dispersal No dispersal
Be SEDG -378 (-80.94%) -392 (-83.94%)
BAMBU -434 (92.93%) | -434 (-92.93%)
-446 (-95.5%) -448 (-95.93%)
SEDG -466 (-99.79%) | -466 (-99.79%)
BAMBU -467 (-100%) -467 (-100%)
-467 (-100%) -467 (-100%)
© Hermann Haas Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 467)
The Mother-of-pearl Blue occurs in flower-rich grasslands and on warm, dry chalk rocks with
scattered patches of grassy vegetation and an occasional bush. The female lays her eggs on the
leaves of Kidney-vetch (Anthylis vulneraria), seeming to prefer smaller plants. Other leguminous
plants, such as clovers (Imfolum spp.) and birdsfoot-trefoils (Lo/ws spp.) are possibly also used as
foodplants. The small caterpillars go into hibernation, and after feeding and growing further, they
pupate at the end of the spring on the ground. The caterpillars are attended by ants of the species
Tapinoma erraticum. Vhe Mother-of-pearl Blue has one generation a yeat.
Present distribution can be very well explained by climatic variables (AUC = 0.98).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 Codd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
2050
Polhommatus nivescens (Lycaenidae)
2080
SEDG
313
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
314 Climatic Risk Atlas of European Butterflies
Polyommatus amandus (SCHNEIDER, 1792) — Amanda’s Blue
a
-4505 (-44.66%) | -5013 (-49.7%)
SEDG -6907 (-68.47%) | -7321 (-72.58%)
BAMBU_ | -8257 (-81.86%) | -8426 (-83.53%)
-8935 (-88.58%) | -9088 (-90.1%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 10087)
The German name for this butterfly, “Prachtiger Blaulinge” (Magnificent Blue), is well chosen, if
only for the colour of the males. They are a bright sky-blue, and they also attract attention by their
territorial behaviour. The females’ are modest brown in most of their distribution area and have a
ereenish-blue sheen. Amanda’s Blue occurs on flower-rich grassland with damp patches, that has
some shelter from bushes or a nearby woodland edge. The female lays her eggs on the leaves of
vetches (Vicia spp.) and possibly also vetchlings (Lathyrus spp.). The caterpillars hibernate in the
litter layer and are attended by ants of the genera Lasius, Myrmica, Formica, and Tapinoma. They also
pupate in the litter layer. Amanda’s Blue 1s single-brooded.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.83).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
-
‘dd 4000 0 odd 4000 0 add 4000 0 aad 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
Swe
0.6
0.
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
315
Polommatus amandus (Lycaenidae)
2080
2050
316 Climatic Risk Atlas of European Butterflies
Polyommatus thersites (CANTENER, 1834) — Chapman’s Blue
.—ClcLee Full dispersal No dispersal
Be SEDG 787 (11.31%) -2140 (-30.75%)
BAMBU -278 (-3.99%) | -2569 (-36.92%)
-140 (-2.01%) | -2824 (-40.58%)
SEDG 279 (4.01%) -3388 (-48.69%)
BAMBU_ | -746 (-10.72%) | -4622 (-66.42%)
-1437 (-20.65%) -5673 (-81.52%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 6959)
The Chapman’s Blue looks very much like the Common Blue (P. scarus). However, the two black
spots in the cell on the underside of the forewing are absent. It occurs in warm, dry places, such
as calcareous and poor grasslands, fields of sainfoin, and abandoned agricultural land. Eggs are
laid on Sainfoin (Oxobrychis vicitfola), and 1n Greece also on Cockscomb Sainfoin (O. caput-gall).
The caterpillars feed on soft parts of the leaflets, leaving the veins. The small caterpillars go into
hibernation, and in those parts of the range with hot climates, the egg or undeveloped larva possibly
goes into aestivation. The caterpillars are attended by ants of the genera Lasius, Formica, Myrmica, and
Tapinoma, and hide themselves in the litter layer and pupate. The Chapman’s Blue usually has two
generations a year, but in the south of the range also three.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.75).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
dd 4000 0 edd 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
317
Polbommatus thersites (Lycaenidae)
2080
2050
318 Climatic Risk Atlas of European Butterflies
Polyommatus icarus (ROTTEMBURG, 1775) - Common Blue
ear Ream
-3054 (-12.06%) | -4374 (-17.27%)
SEDG -2279 (-9%) -5242 (-20.7%)
BAMBU_ | -5208 (-20.57%) | -7950 (-31.4%)
-7231 (-28.56%) | -10608 (-41.9%)
© Martin Wiemers Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 25320)
The Common Blue is a very common butterfly. It looks rather like Chapman’s Blue (P. shersiles), but
the two black spots on the underside of its front wings distinguish it 1n most specimens. It can be
found on most types of grassy vegetation, ranging from quite dry, poor grassland to moderately
damp meadows. The female lays her eggs on a variety of leguminous plants, including Common
Birdsfoot—trefoil (Lo/us corniculatus). The caterpillars feed on the leaves. They are attended by ants of
the genera Lasius, Formica, Myrmica, Tapinoma, and Plagiolepis. When half-grown, the caterpillars may
hibernate in the litter layer. In hot climates, aesttvation possibly also takes place, in the egg or larval
stage. The caterpillars pupate in the litter layer. Depending on the geographical position and altitude
of the breeding ground, the Common Blue has one to three (or even more) generations a yeat.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.65).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
fod 4000 0 edd 4000 0 oad 4000 0 Gdd 4000
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 o.2
Swe
0.6
1.0 O2
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Polyommatus icarus (Lycaenidae)
SEDG
319
(B1)
GRAS
(A1Fl)
320 Climatic Risk Atlas of European Butterflies
Polyommatus eros (OCHSENHEIMER, 1808) — Eros Blue
a
RAS
-1162 (-43.83%) | -1322 (-49.87%)
G
SEDG -1071 (-40.4%) | -1530 (57.71%)
BAMBU_ | -1167 (-44.02%) | -1570 (-59.22%)
-1399 (-52.77%) -1855 (-69.97%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2651)
The Eros Blue is a mountain butterfly occurring on sub-alpine and alpine grasslands, on rocky
places with flower-rich vegetation and on scree slopes. Purple Beaked Milk-vetch (Oxytropis haller?)
and O. fetida are probably the major foodplants but other leguminous plants such as Astragalus
sempervirens, A, leontinus, and Common Birdsfoot-trefoil (Lots corniculatus) have also been named.
The caterpillars feed on the leaves of the foodplant and hibernate on it. In spring, they grow further
and when fully grown, pupate at the foot of the foodplants. They are attended by ants of the genera
Myrmica, Formica and Camponotus. The Eros Blue has one generation a year.
Polyommatus eroides, which 1s listed 1n the Annexes I] and IV of the Habitats’ Directive, 1s also is
included here.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.71).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
todd 4000 0 Codd 4000 0 oad 4000 0 odd 4000
Minimum
Swe
0.6
1.0 02
0.6
Swe
1.0 0.2
Swe
O46
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
2050
Polyommatus eros (Lycaenidae)
2080
321
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
322 Climatic Risk Atlas of European Butterflies
Polyommatus daphnis ({SCHIFFERMULLER], 1775) — Meleaget’s Blue
6c Full dispersal No dispersal
fi SEDG -599 (-7.06%) -2243 (-26.44%)
-2615 (-30.83%) | -3487 (-41.11%)
-4525 (53.34%) | -6338 (-74.71%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 8483)
The Meleager’s Blue is a local butterfly that occurs on poor, rough vegetation and poor grassland
often neat woods. The populations are usually small. The scalloped edge of the hindwing of the
female distinguishes this species from all other blues. The female lays her eggs on Horseshoe
Vetch (Hippocrepis comosa) and Crown Vetch (Coronilla varia), favouring plants that grow in the shade
provided by rocks or woodland. The caterpillars are attended by ants, including species of Lasts,
Formica, and Tapinoma. Either the egg or the small caterpillar hibernates, and pupation takes place in
the litter layer. The Meleager’s Blue has one generation a yeat.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.84).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
0.6
10 02
Swe
06
10 0.2
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
6 1002 O86 16 1.0
- _ i
1 1
Swe
o.2
L=]
2000 40000 2000 40000 2000 40000 2000 4000
Gadd Gdd
Gdd Gdd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
323
Polommatus daphnis (Lycaenidae)
2080
2050
324 Climatic Risk Atlas of European Butterflies
Polyommatus bellargus (ROTTEMBURG, 1775) — Adonis Blue
-405 (-3.31%) -5496 (-44.88%)
GRAS -1117 (-9.12%) | -6854 (-55.97%)
© Peter Ginzinger Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 12245)
The Adonis Blue is found on calcareous soil on moderately to very sparse grassland, that is often
sheltered by neighbouring woods or shrub. Its foodplants are Horseshoe Vetch (Hippocrepis comosa)
and Crown Vetch (Coronilla varia), and the eggs are laid on the leaves. It pupates in the litter layer.
The caterpillars are attended by ants of the genera Myrmica, Lasius, Plagiolepis, Tetramorium, Formica,
and Tapinoma. The Adonis Blue usually has two generations a year, and the caterpillars of the second
brood hibernate. There are also some single-brooded populations in the south of Greece.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.72).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
‘dd 4000 0 odd 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Polyommatus bellargus (Lycaenidae) 325
(B1)
SEDG
GRAS
(AFI)
326 Climatic Risk Atlas of European Butterflies
Polyommatus coridon (Popa, 1761) — Chalkhill Blue
AMBU -838 (-9.52%) | -5197 (-59.02%)
-1527 (-17.34%) | -6483 (-73.62%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 8806)
The Chalkhill Blue occurs on calcareous soil in dry, and flower-rich places with a short vegetation.
They seem to prefer sheltered places. Sometimes, populations can be extremely large which 1s
especially obvious in the late afternoon when the butterflies come together to roost. Hundreds of
butterflies can be seen, their heads pointing downwards into the vegetation, wings upright, the light-
coloured underwings gleaming in the evening sun. Horseshoe Vetch (Hippocrepis comosa) is its only
foodplant, the female laying her eggs on the leaves. The eggs hibernate. The caterpillars are attended
by ants of the genera Myrmica, Lasius, Formica, Plagiolepis, Tetramorium, Aphaenogaster, and Tapinoma.
The Chalk-hill Blue pupates in the litter layer. It usually only has one generation a year.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.82).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘aad 4000 0 odd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 60.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
327
Polyommatus coridon (Lycaenidae)
2080
2050
328 Climatic Risk Atlas of European Butterflies
Polyommatus hispanus (HERRICH-SCHAFFER, 1851) —
Provence Chalkhill Blue
SEDG 170 (21.46%) -557 (-70.33%)
BAMBU 68 (8.59%) -559 (-70.58%)
145 (18.31%) -608 (-76.77%)
SEDG 500 (63.13%) -656 (-82.83%)
BAMBU 214 (27.02%) -739 (-93.31%)
151 (19.07%) -787 (-99.37%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 792)
The Provence Chalkhill Blue occurs on dry, calcareous soils covered with a flower-rich grassy
vegetation, often with scattered bushes. Eggs are laid on Horseshoe-vetch (Hippocrepis comosa). The
caterpillars are attended by ants of the genera Plagiolepis and Crematogaster. The Provence Chalkhill
Blue has two generations a year and the butterflies can be seen as late as October.
Present distribution can be well explained by climatic variables (AUC = 0.95).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
|
edd 4000 0 edd 4000 0 add 4000 0 add 4000
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 O.2
Swe
0.6
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
329
Polbommatus hispanus (Lycaenidae)
2080
2050
330 Climatic Risk Atlas of European Butterflies
Polyommatus albicans (HERRICH-SCHAFFER, 1851) —
Spanish Chalkhill Blue
[etn [sean
-457 (-48.11%) | -598 (-62.95%)
SEDG -924 (-97.26%) | -926 (-97.47%)
BAMBU -946 (-99.58%) -950 (-100%)
-922 (-97.05%) -950 (-100%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 950)
The Spanish Chalkhill Blue can be seen in dry, calcareous rocky places with grassy vegetation, and in
dry, open scrub. Its most important foodplant is Horseshoe Vetch (Hippocrepis comosa), although H.
multisiliquosa is possibly also used. The female lays her eggs on the leaves. The caterpillars are often
found accompanied by Tapinoma ants. This butterfly species is single-brooded.
Present distribution can be well explained by climatic variables (AUC = 0.88).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
‘0.6
1.0 O02
Annual precipitation range
Swe
0.6
10 0.2
0.6
Swe
6 1002 O86 o6 1.0
oOo
S
Maximum Large (66%) Small (33%)
Swe
o2
0 2000 400 2 40000 2000 40000 2000 400)
Gdd Gdd Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
2050
Pobommatus albicans (Lycaenidae)
2080
SEDG
331
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
332 Climatic Risk Atlas of European Butterflies
Polyommatus admetus (EsPEr, 1785) — Anomalous Blue
Cc Full dispersal No dispersal
Hee SEDG 568 (36.96%) -842 (-54.78%)
-452 (-29.41%) -1020 (-66.36%)
-1004 (-65.32%) | -1357 (-88.29%)
aa -915 (-59.53%) | -1464 (-95.25%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1537)
The Anomalous Blue occurs in woodland clearings, open scrub, on sheltered slopes and in
agricultural areas. All the places are warm and have a flower-rich grassy vegetation. The females lay
their eggs on the flowers of Sainfoin (Oxobrychis vicufoha) and Cockscomb Sainfoin (O. caput-galh).
The caterpillars are attended by ants of the genera Crematogaster, Camponotus, and Tapinoma. The
Anomalous Blue has one generation a year and hibernation takes place on the ground, the small
caterpillars hiding under stones.
Present distribution can be well explained by climatic variables (AUC = 0.88).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
| ow
E =e
=H
= 6
o
=
=o
1
cH
oa
a : : , . , :
‘dd 4000 0 odd 4000 0 ad 4000 0 odd 4000
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O2
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Polyommatus admetus (Lycaenidae) 333
334 Climatic Risk Atlas of European Butterflies
Polyommatus ripartii (FREYER, 1830) — Ripart’s Anomalous Blue
6c Full dispersal No dispersal
fi SEDG -290 (-63.04%) -336 (-73.04%)
z BAMBU -342 (-74.35%) | -372 (-80.87%)
-349 (-75.87%) | -384 (-83.48%)
SEDG -359 (-78.04%) -408 (-88.7%)
BAMBU -406 (-88.26%) | -433 (-94.13%)
[oras | -447 (-97.17%) -456 (-99.13%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 460)
Ripart’s Anomalous Blue is mostly found on dry, grassy places with bushes. Different sainfoins are
used as foodplant, such as Sainfoin (Oxobrychis viciifoha), O. arenaria, Rock Sainfoin (O. sanxatiis), and
O. alba. The female lays her eggs on the flowers. The caterpillars hibernate when still small. They are
often attended by ants of the genera Crematogaster, Camponotus, and Lasius. The Ripart’s Anomalous
Blue has one generation a yeat.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.75).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
tea 4000 0 odd 4000 0 Cad 4000 0 odd 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 o.2
Swe
0.6
1.0 O02
Annual precipitation range
0.6
Maximum Large (66%) Small (33%)
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
2050
Polommatus riparti (Lycaenidae)
2080
335
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
336
Climatic Risk Atlas of European Butterflies
Polyommatus dolus (HUBNER, 1823) — Furry Blue
.—hCcLee Full dispersal No dispersal
SEDG -58 (-27.1%) -190 (-88.79%)
BU -90 (-42.06%) -192 (-89.72%)
2 BAM
-34 (-15.89%) -202 (-94.39%)
SEDG 105 (49.07%) -208 (-97.2%)
BAMBU -97 (-45.33%) -214 (-100%)
-79 (-36.92%) -214 (-100%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 214)
The Furry Blue occurs on dry, flower-rich, grassy vegetation with scrub, on waste ground, and in
clearings in deciduous or coniferous woods. The female lays her eggs on Sainfoin (Oxobrychis victifola).
The butterflies are attended by ants. Hibernation takes place when the caterpillars are still small.
Present distribution can be very well explained by climatic variablesnt (AUC = 0.97).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
1.0
Minimum
Swe
‘0.6
10 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
Swe
o6 10 02
o2
0 2000 40000 2000 40000 2000 40000 2000 4000
Gd dl Gdd
Gdd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
2050
Polommatus dolus (Lycaenidae)
2080
337
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
338 Climatic Risk Atlas of European Butterflies
Polyommatus damon ({ScHIFFERMULLER], 1775) - Damon Blue
SEDG -660 (-28.23%) -1169 (-50%)
BAMBU -850 (-36.36%) | -1277 (-54.62%)
-1112 (-47.56%) | -1526 (-65.27%)
SEDG -857 (-36.66%) | -1653 (-70.7%)
BAMBU_ | -1091 (-46.66%) | -1769 (-75.66%)
-1409 (-60.27%) | -2009 (-85.93%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2338)
The Damon Blue is found on calcareous soil, on grasslands, rough vegetation, such as small patches
at the edge of scrub or woodland, and also on abandoned Sainfoin (Oxobrychis vicufoha) fields. It lays
its eggs on different species of Oxobrychis, including Sainfoin (Oxobrychis vicufolia). The caterpillars seem
to like being between the flowers and only start to feed late in the afternoon. They are often attended
by ants of the genera Lasius or Formica. They pupate in the litter layer. The Damon Blue has one
generation a yeat.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.85).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
| ri 4 bel 7
sir}
oc
Fa : . F ? ;
_
ah
o
2
w
o
dd 4000 0 edd 4000 0 aad 4000 0 Gd 4000
Minimum
Swe
06
Swe
0.6 10 Oo2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
339
Polommatus damon (Lycaenidae)
2080
2050
340 Climatic Risk Atlas of European Butterflies
Hamearis lucina (LiNNAEus, 1758) — Duke of Burgundy Fritillary
.—hCcLUwk Full dispersal No dispersal
fe SEDG 5 (0.89%) -2192 (-35.31%)
529 (8.52%) -2161 (-34.82%)
Cer ee
SEDG -249 (-4.01%) | -3222 (-51.91%)
BAMBU -452 (-7.28%) | -4008 (-64.57%)
-1480 (-23.84%) | -5148 (-82.94%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 6207)
The Duke of Burgundy Fritillary occurs in woodland clearings, along the edges of paths in the
woods, and at woodland edges. Although the populations are often very local, the numbers of
butterflies can be considerable. Eggs are laid on the underside of the leaves of Primula species. The
caterpillars feed at night, hiding themselves during the day by lying along the main nerve of the
Primula \eaf. Vhey leave the larval foodplants to hibernate in the litter layer. Mostly, it has only one
generation a year, but 1n warm areas, there may be a second generation late in the year.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.77).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
tad 4000 0 edd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
341
Hamearis lucina (Riodinidae)
2080
2050
342 Climatic Risk Atlas of European Butterflies
Libythea celtis (LAICHARTING, 1782) — Nettle-tree Butterfly
.—hCcLee Full dispersal No dispersal
Be SEDG -582 (-15.8%) -1642 (-44.57%)
BAMBU_ | -1280 (-34.74%) | -1946 (52.82%)
-1165 (-31.62%) | -2165 (58.77%)
SEDG -813 (-22.07%) | -2341 (-63.55%)
BAMBU_ | -1635 (-44.38%) | -3068 (-83.28%)
-1312 (-35.61%) | -3409 (-92.54%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 3684)
The Nettle-tree Butterfly is one of the snout butterflies, so-called because of its long maxillary
palps, which point forward seemingly forming a snout. They are often found in scrub or woods,
where the major foodplant, the Nettle Tree (Ced#s australis) grows. The eggs are laid singly on the leaf
buds, early in the year because this butterfly hibernates in the adult stage, appearing again in March.
The green or brown caterpillars keep mostly to the underside of the leaves, and also pupate there.
The Nettle-tree Butterfly has one generation a year. The adult enters hibernation sometimes as soon
as August. The brown underside of its wings provides a good camouflage in the dense undergrowth
where it spends the winter looking very much like a dead leaf.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.8).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘dd 4000 0 ood 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
343
Libythea celts (Labytheidae)
2080
2050
344 Climatic Risk Atlas of European Butterflies
Argynnis paphia (LINNAEUS, 1758) — Silver-washed Fritillary
a
-2577 (-14.1%) | -4369 (-23.91%)
SEDG -2912 (-15.93%) | -5757 (-31.5%)
BAMBU_ | -4872 (-26.66%) | -8253 (-45.16%)
-6457 (-35.33%) | -11083 (-60.64%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 18276)
The Silver-washed Fritillary is a large, conspicuous butterfly that is often present in large numbers.
Needing a lot of nectar, they are often found on thistles at the edge of woodland. They also occur
on rough vegetation in woodland clearings. Unlike other butterflies, the eggs are not laid on the
foodplant. Instead, the female deposits them singly on the branches and trunks of trees growing at
woodland edges. As soon as they emerge from the egg in the late summer, the tiny caterpillar looks
for somewhere to hibernate. In the spring it starts looking for violets (20/4 spp.), on which it feeds
at night, hiding under the leaves of the foodplant during the day. It pupates on a stalk of a violet
plant, or in a crevice in the bark of a tree. The Silver-washed Fritillary has one generation a yeat.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.78).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
aad 4000 0 edd 4000 0 add 4000 0 add 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Arrgynnis paphia (Nymphalidae) 345
SEDG
(B1)
GRAS
(A1Fl)
346 Climatic Risk Atlas of European Butterflies
Argynnis pandora ({SCHIFFERMULLER], 1775) — Cardinal
a
-587 (-10.8%) | -2566 (-47.23%)
SEDG -1599 (-29.43%) | -3246 (-59.75%)
BAMBU_ | -3121 (-57.45%) | -4172 (-76.79%)
-2148 (-39.54%) | -4744 (87.32%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 5433)
The Cardinal 1s a woodland butterfly occurring at woodland edges and in glades with bushes and
grassy, flower-rich vegetations. Like the Silver-washed Fritillary (4. pap/ia), these butterflies are
often seen in places where plants rich in nectar, such as thistles, are plentiful. The Cardinal 1s a fast
and powerful flyer, vagrants being sometimes observed outside the distribution range. The eggs are
laid singly on the leaves of violets (V/zo/@ spp.). It hibernates as a small caterpillar and pupates in the
spring, suspended low down on the foodplant or on the nearby vegetation. It 1s single-brooded.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.83).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
‘0.6
10 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
100 0.2
0.6
Swe
o2
0 2000 40000 2000 40000 2000 40000 2000 4000
dl Gd Gdd
Gdd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
347
Arrgynnis pandora (Nymphalidae)
2080
2050
348 Climatic Risk Atlas of European Butterflies
Argynnis aglaja (LINNAEUS, 1758) — Dark Green Fritillary
.—hCcLUwE Full dispersal No dispersal
Be SEDG -1213 (-5.9%) -3328 (-16.19%)
BAMBU_ | _ -2865 (-13.94%) | -4599 (-22.38%)
-2885 (-14.04%) | -4947 (-24.07%)
SEDG -4162 (-20.25%) | -7016 (-34.14%)
BAMBU_ | -7493 (-36.46%) | -10512 (-51.15%)
-9906 (-48.2%) -13024 (-63.37%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 20551)
The Dark Green Fritillary can be found on many different types of flower-rich grasslands. The
erassland is often situated in or at the edge of woodland, and may be dry, calcareous or dune
grassland, or damp grasslands along the edges of bogs. The eggs are laid on the often already
withered leaf-stems of violets (zo spp.). Directly after hatching, the small caterpillar prepares for
hibernation, hiding itself in the litter layer until the spring. It then begins to feed on the fresh, new
growth of the violet plants, continuing into the summer, when it pupates either in the moss layer,
or under a tussock of grass. The caterpillars are quite mobile and visit several plants when feeding.
The Dark Green Fritillary is single-brooded.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.78).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
=
i
|
a
=
se | |
r=]
todd 4000 0 odd 4000 0 add 4000 0 ada 4000
Minimum
Swe
0.6
Swe
0.6
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
349
Arrgynnis aglaja (Nymphalidae)
2080
2050
350 Climatic Risk Atlas of European Butterflies
Argynnis adippe ({SCHIFFERMULLER], 1775) — High Brown Fritillary
a se
-3720 (-26.2%) | -4865 (-34.26%)
SEDG -3763 (-26.5%) | -5920 (-41.69%)
BAMBU -5935 (-41.8%) | -8143 (57.35%)
-8102 (-57.06%) | -10604 (-74.68%)
© Peter Ginzinger Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 14199)
The High Brown Fritillary occurs on woodland edges and in woodland clearings, where there is lush,
rough vegetation with plenty of nectar plants. The eggs are laid on the leaves of violets (Vola spp.)
and also on the bark of trees with violets growing near them. The tiny caterpillar remains within the
ege during the winter, and from about the beginning of March, leaves the egg and starts feeding on
violet leaves. It pupates on a twig or on a leaf close to the ground. The High Brown Fritillary has
one generation a yeat.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.78).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘dd 4000 0 odd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Swe
0.6
Annual precipitation range
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
351
Arrgynnis adippe (Nymphalidae)
2080
2050
352 Climatic Risk Atlas of European Butterflies
Argynnis niobe (LINNAEUS, 1758) — Niobe Fritillary
a
-2562 (-15.49%) | -3948 (-23.87%)
SEDG -3322 (-20.09%) | -6045 (-36.55%)
BAMBU_ | -5110 (-30.9%) | -7946 (-48.04%)
-7117 (-43.03%) | -10289 (-62.21%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 16539)
The Niobe Fritillary occurs on poor, dry grassland, often with woodland or scrub nearby. The eggs
are laid on the woody stock of violets (Vola spp.). The caterpillar develops quickly within the egg,
but does not hatch until after hibernation. Staying hidden during the day, it feeds on the violet plants
at night. It pupates low down in the vegetation. The Niobe Fritillary has one generation a year.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.74).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
ir}
o
_
ah
o
=
wv
o
sc
o
‘dd 4000 0 edd 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
1.0 02
Swe
0.6
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
Swe
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
353
Arrgynnis niobe (Nymphalidae)
2080
2050
354 Climatic Risk Atlas of European Butterflies
Argynnis laodice (PALLAS, 1771) — Pallas’ Fritillary
Cc Full dispersal No dispersal
fe SEDG -679 (-13.69%) -2209 (-44.53%)
201 (4.05%) -1794 (-36.16%)
|GRAS | -443 (-8.93%) | -2242 (-45.19%)
960 (19.35%) | -3083 (-62.14%)
167 (3.37%) -3849 (-77.59%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 4961)
The Pallas’s Fritillary can be easily recognized by the noticeable jagged, white line, running across
the underside of the hindwing. It can be found on damp, flower-rich grassland in open or deciduous
forests or mixed woods. Both males and females are fond of sucking nectar from bramble blossom.
The caterpillars live on Marsh Violet (Vola palustris) and Heath Dog Violet (V. canina). It has one
generation a year and passes the winter in the egg stage.
Present distribution can be well explained by climatic variables (AUC = 0.89).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
-
fad 4000 0 edd 4000 0 oad 4000 0 ad 4000
Minimum
Swe
0.6
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Arrgynnis laodice (Nymphalidae)
SEDG
GRAS
(B1)
(A1Fl)
355
356 Climatic Risk Atlas of European Butterflies
Issoria lathonia (LINNAEUus, 1758) — Queen of Spain Fritillary
.6—hcLUw Full dispersal No dispersal
fi SEDG -770 (-5.48%) 2836 (-20.17%)
-2031 (-14.45%) | -3944 (-28.05%)
-6471 (-46.02%) | -8591 (-61.1%)
Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 14060)
The large, silver-white mirrors on the underside of the hindwings, distinguish the Queen of Spain
Fritillary from other fritillaries. The butterflies of the first brood that emerge at the end of the
spring are quite small, but those of summer broods are often bigger. They can be found on a wide
range of dry, flower-rich grasslands, wasteland and fields. The female deposits her eggs singly on the
underside of the leaves of violets (V/zo/a spp.) on which the caterpillars later feed. On this nutritious
diet the caterpillars grow very quickly, pupating low down in the vegetation. The Queen of Spain
Fritillary has two to four generations a year, depending on the altitude and geographical position
of the habitat. Hibernation takes place in the larval stage in temperate areas, but probably it can
overwinter in other stages as well.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.78).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
06
10 02
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
6 1002 O86 6 10
oOo q
=
|
Swe
o2
O 2000 40m0 2 40000 2000 40000 2000 400)
Gdd Gdd Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
357
Issoria lathonia (Nymphalidae)
2080
2050
358 Climatic Risk Atlas of European Butterflies
Brenthis ino (ROTTEMBURG, 1775) — Lesser Marbled Fritillary
a
-2704 (-19.61%) | -5157 (-37.4%)
SEDG -4126 (-29.92%) | -7216 (-52.34%)
BAMBU_ | _ -4906 (-35.58%) | -8460 (-61.36%)
-6908 (-50.1%) -10472 (-75.95%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 13788)
The Lesser Marbled Fritillary occurs on damp to wet, flower-rich, rough vegetation, growing 1n the
shelter of woodland. This can be situated in a valley, or on the banks of a stream, in abandoned
meadows and swampy habitats. Because of changes in agricultural practices, this butterfly has been
able to expand into some abandoned wet meadows. After continuing succession, these meadows
will get overgrown by scrub and become unsuitable for the Lesser Marbled Fritillary. Eggs are laid
singly or in pairs on the leaves and flowerheads of Meadowsweet (F24pendula ulmania), Goat’s-beard
(Aruncus dioiwus), Great Burnet (Sanguisorba officinals) and brambles (Rubus spp.). It hibernates either
as an ege or a caterpillar. The caterpillars feed at night, and in bad weather, also during the day. They
pupate on the foodplant. The Lesser Marbled Fritillary is single-brooded.
Present distribution can be well explained by climatic variables (AUC = 0.87).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
—
fed 4000 0 Codd 4000 0 oad 4000 0 odd 4000
Minimum
Swe
‘1.6
1.0 02
0.6
Swe
1.0 0.2
Swe
0.6
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
359
Brenthis ino (Nymphalidae)
oa i
Fs ef
2080
2050
360 Climatic Risk Atlas of European Butterflies
Brenthis daphne (BrErcstTRAssER, 1780) — Marbled Fritillary
.—ClcLee Full dispersal No dispersal
Be SEDG 1518 (27.55%) -1190 (-21.6%)
BAMBU 1916 (34.78%) | -1365 (-24.78%)
73 (385%) | ATT aR
SEDG 2653 (48.16%) | -1927 (-34.98%)
BAMBU 2007 (36.43%) | -2868 (-52.06%)
1929 (35.02%) | -3676 (-66.73%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 5509)
The Marbled Fritillary occurs at the edges of woods or in scrub where brambles are growing. It
takes nectar from thistles growing in rough vegetation. Eggs are deposited singly on the leaves and
sepals of brambles (Rubus spp.), where they pass the winter, protected by withered leaves. In the
spring, the small caterpillar feeds on the new, young leaves in the leaf buds, the later larval stages
eating older leaves. The pupa is suspended from a leaf or branch of the foodplant. The Marbled
Fritillary 1s single-brooded.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.77).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Ow
=a
ee]
my
r=]
o
=
Ow
=o
w
my
o
todd 4000 0 ood 4000 0 ad 4000 0 odd 4000
1.0
Minimum
Swe
06
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
O68
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Brenthis daphne (Nymphalidae) 361
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
362 Climatic Risk Atlas of European Butterflies
Brenthis hecate ({SCHIFFERMULLER], 1775) — Twin-spot Fritillary
.—ClCcLee Full dispersal No dispersal
He SEDG 554 (14.53%) -1741 (-45.65%)
-589 (-15.44%) | -2135 (-55.98%)
|GRAS | -270 (-7.08%) | -2292 (-60.09%)
SEDG -217 (-5.69%) | -2542 (-66.65%)
BAMBU 554 (-14.53%) | -3243 (-85.03%)
-508 (-13.32%) | -3442 (-90.25%)
© Rudi Verovnik Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 3814)
The Twin-spot Fritillary can be found in flower-rich grassland, situated in the shelter of a woodland
edge, or in scattered bushes. The most important foodplant is Meadowsweet (F¢dpendula ulmaria),
although different species of Dorycnium, a leguminous plant, are named as a foodplant in Spain. It 1s
single-brooded, and passes the winter in the egg or caterpillar stage.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.79).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 edd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
363
Brenthis hecate (Nymphalidae)
2080
2050
364 Climatic Risk Atlas of European Butterflies
Boloria eunomia (Esrrr, 1799) — Bog Fritillary
eae ae
-1220 (-15.34%) | -1399 (-17.59%)
amas | ary | a
SEDG -1843 (-23.17%) | -2291 (-28.8%)
BAMBU -2410 (-30.3%) | -2747 (-34.54%)
-3171 (-39.87%) | -3443 (-43.29%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 7954)
The Bog Fritillary occurs on wet grasslands and marshy ground by streams, rivers, or lakes, and at
the edges of raised bogs. Sometimes, the area they occupy is very small. They can be seen beside
streams, flying slowly back and forth between small patches where its foodplant Common Bistort
(Polygonum bistoria) is growing. The small caterpillars live together in a loosely spun nest, and hibernate
when half-grown. However, the larger, later caterpillars are solitary and considerably more mobile,
frequently leaving their foodplants in order to bask in the sun on another plant. The Bog Fritillary
has one generation a year. There are two subspecies in Europe. The species’ range in central Europe
is highly fragmented.
Present distribution can be well explained by climatic variables (AUC = 0.89).
Climate risk category: LR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
‘1.6
sh
Qa a
=) a
ee
Pm Oe
ae =a
o= mH
=e
£5 8
Oo. 4
os 7
D>
a a4 = a
ger
|
ae
qf. =
eo
wv
x é 5
= 4
o
0 2000 40000 2000 40000 2000 40000 2000 4000
Gd Gdd Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
365
Boloria eunonia (Nymphalidae)
2080
2050
366 Climatic Risk Atlas of European Butterflies
Boloria euphrosyne (LINNAEUS, 1758) — Pearl-bordered Fritillary
a
-5707 (-26.51%) | -5782 (-26.86%)
SEDG -4372 (-20.31%) | -4649 (-21.59%)
BAMBU_ | -7044 (-32.72%) | -7118 (-33.06%)
-8912 (-41.4%) | -9059 (-42.08%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 21529)
The Pearl-bordered Fritillary occurs at the edges of woods and in clearings, in meadows and on
pastures near scrub. Its habitats are usually dry and moderately rich in nutrients. The butterflies
are quite mobile, leaving the habitat in search of nectar, visiting vegetation that ranges from very
dry to wet. Most species of violet (V/zo4a spp.) can be used as a foodplant. The female lays her eggs
singly, either on a foodplant or on a neighbouring plant. The caterpillars feed on the violet leaves,
hibernating in a rolled-up, withered leaf, when half-grown. The species pupates on the foodplant,
close to the ground. It has one to two broods per year.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.73).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
1.6
10 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 0.2
0.6
Swe
o2
0 2000 40000 2000 40000 2000 40000 2000 4000
Gd dl Gdd
Gdd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
367
Boloria euphrosyne (Nymphalidae)
2080
2050
368 Climatic Risk Atlas of European Butterflies
Boloria titania (Esrer, 1793) — Titania’s Fritillary
Cc Full dispersal No dispersal
fi SEDG -377 (-19.74%) -766 (-40.1%)
-151 (-7.91%) -525 (-27.49%)
-580 (-30.37%) | -874 (-45.76%)
570 (29.84%) -926 (-48.48%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1910)
Titania’s Fritillary occurs in grassy, open places at the edges of woodland, or in woodland clearings,
or on grassland with scattered trees, mostly on damp to swampy ground, where its foodplant Snake-
root or Common Bistort (Poheanum bistoria) 1s growing among tall flower-rich vegetation. Various
violets (toda spp.) are also used as a foodplant. The female deposits her rather large eggs singly on
either the foodplant, or on a plant nearby. The caterpillars go into hibernation in an early stage, and
pupate at the end of the spring on a stalk, near to the ground. The species is single-brooded.
Present distribution can be well explained by climatic variables (AUC = 0.9).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
Minimum
Swe
0.6
1.0 t.2
0.6
1.0 0.2
Annual precipitation range
Maximum Large (66%) Small (33%)
0 2000 40000 2000 49m0 2000 40000 2000 4000
Gdd Gdd Gdd Gdd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Boloria titania (Nymphalidae) 369
370 Climatic Risk Atlas of European Butterflies
Boloria selene ({SCHIFFERMULLER], 1775) —
Small Pearl-bordered Fritillary
.—hCcLUw Full dispersal No dispersal
SEDG -4708 (-23%) -5037 (-24.61%)
BAMBU -5114 (-24.99%) -5424 (-26.5%)
-6291 (-30.74%) | -6771 (-33.08%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 20468)
The Small Pearl-bordered Fritillary, is found in damp to wet meadows or on lightly-grazed pastures.
It is also found on raised bogs and in swampy habitats. Populations may occur in small, sheltered
locations, but also in an open landscape. The butterflies can often be seen taking nectar, seeming
to prefer thistles of various kinds. The eggs are laid on violets (Vzo/a spp.). When half-grown, the
caterpillars hibernate in a rolled-up leaf of the foodplant. They pupate low down on a stalk in the
litter layer. At high altitudes and in the north, the Small Pearl-bordered Fritillary is stngle-brooded,
but has two generations a year elsewhere.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.73).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
-
7 | il
|
|
|
dd 4000 0 odd 4000 0 ad 4000 0 odd 4000
Minimum
Swe
‘0.6
1.0 02
Swe
0.6
1.0 0.2
Swe
0.6
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
371
Boloria selene (Nymphalidae)
2080
2050
372 Climatic Risk Atlas of European Butterflies
Boloria chariclea (SCHNEIDER, 1794) — Arctic Fritillary
.—hlcLUwe Full dispersal No dispersal
He SEDG -519 (-98.3%) -523 (-99.05%)
518 (-98.11%) | -521 (-98.67%)
roras | steconiny | sa con20e
SEDG 512 (-96.97%) -528 (-100%)
BAMBU -526 (-99.62%) -528 (-100%)
re
© Jostein Engdal Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 528)
The Arctic Fritillary occurs in a harsh environment, the windy, dry, rocky tundra in the far north
of Europe, with a vegetation of grass and dwarf shrubs. The foodplant 1s not certain, but Casszope
tetragon (Ericaceae) and violets (Voda spp.) may be used. The butterflies, which only appear for about
two weeks a year, fly close to the ground, seeking the shelter of rocks or hollows.
Present distribution can be very well explained by climatic variables (AUC = 1).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 edd 4000 0 add 4000 0 add 4000
1.0
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
2050
Boloria chariclea (Nymphalidae)
2080
SEDG
373
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
374 Climatic Risk Atlas of European Butterflies
Boloria freija (BECKLIN, 1791) — Frejya’s Fritillary
a
-1581 (-28.71%) | -1593 (-28.93%)
SEDG -3654 (-66.35%) | -3670 (-66.64%)
BAMBU_ | -3324 (-60.36%) | -3340 (-60.65%)
-4189 (-76.07%) | -4197 (-76.21%)
© Jostein Engdal Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 5507)
Freyya’s Fritillary occurs in damp, peatland habitats. The female lays her eggs singly on Bog
Whortleberry (Vaccinium uliginosum), and possibly also on Cloudberry (Rubus chamaemorus), Bearberry
(Arctostaphylos uva-ursi), A\pine Bearberry (A. alpinus), and Crowberry (Empetrum nigrum). The
butterflies can also be seen on drier, grassy vegetation, searching for nectar, visiting Moss Campion
(Szlene acauhs) frequently. Frejya’s Fritillary 1s stingle-brooded.
Present distribution can be well explained by climatic variables (AUC = 0.95).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘dd 4000 0 odd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
375
Boloria freija (Nymphalidae)
2080
2050
376 Climatic Risk Atlas of European Butterflies
Boloria dia (LINNAEUS, 1767) — Weaver’s Fritillary
| EE
1436 (17.22%) -2450 (-29.38%)
anus [2105 | soca
SEDG 1325 (15.89%) -4030 (-48.33%)
BAMBU 892 (10.7%) -5229 (-62.71%)
605 (7.26%) -6394 (-76.69%)
© Karl Heyde Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 8338)
In the northern part of its range, the Weaver’s Fritillary can be found on warm slopes with open
woodland, scrub, and flower-rich grassland. More to the south, it 1s also found in damp, shady
places. It can be very common in a traditional South-European agricultural landscape. The eggs are
laid singly on various violets (Vzo/a spp.). The caterpillars hibernate when half-grown in the litter
layer. They pupate deep down in the vegetation. Weaver’s Fritillary has two to three broods a year.
Present distribution can be well explained by climatic variables (AUC = 0.86).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 odd 4000 0 ad 4000 0 ada 4000
1.0
Minimum
Swe
0.6
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
377
Boloria dia (Nymphalidae)
2080
2050
378 Climatic Risk Atlas of European Butterflies
Boloria thore (HtBNER, 1806) — Thor’s Fritillary
SEDG -200 (-12.32%) -396 (-24.38%)
-237 (-14.59%) -370 (-22.78%)
arg -365 (-22.48%) -471 (-29%)
SEDG 579 (35.65%) -405 (-24.94%)
BAMBU 229 (14.1%) -437 (-26.91%)
}GRAS 388 (23.89%) -549 (-33.81%)
© Rudi Verovnik Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1624)
In the Alps, Thor’s Fritillary can be found in flower-rich places, in clearings, or in sheltered “alcoves”
at the edges of woods. It prefers damp, north-facing slopes, and is often found beside streams or
in ravines. In the Scandinavian part of its range, the butterflies can be seen in clearings in birch
woods and coniferous forests, and near mountain lakes, swampy places, in gullies, and river beds.
The Yellow Wood Violet (Voda biflora) 1s 1ts major foodplant, but other violets are used as well. The
female lays her eggs either on the foodplant, or on a neighbouring plant, and the caterpillars take
nearly two years to develop.
Present distribution can be well explained by climatic variables (AUC = 0.95).
Climate risk category: LR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘dd 4000 0 odd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
379
Boloria thore (Nymphalidae)
2080
2050
380 Climatic Risk Atlas of European Butterflies
Boloria frigga (BECKLIN, 1791) — Frigga’s Fritillary
6c Full dispersal No dispersal
SEDG -1825 (-30.7%) -1919 (-32.28%)
BAMBU_ |} -1331 (-22.39%) | -1457 (-24.51%)
-1842 (-30.98%) | -1949 (-32.78%)
SEDG -3317 (-55.79%) | -3406 (-57.29%)
-2550 (-42.89%) | -2690 (-45.25%)
ae -3189 (-53.64%) | -3323 (-55.9%)
© Jostein Engdal Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 5945)
Frigea’s Fritillary is mostly found on open bogs and 1n swampy areas with shrubs of Birches (Be/wla
spp.) and willows (Sax spp.) scrub, in swampy areas in birch woods, and on bogs in coniferous
forests. It is a rare fritillary that occurs locally and always in low to very low numbers. Despite its
size, it is inconspicuous and can suddenly seem to vanish. The eggs are laid singly on Cloudberry
(Rubus chamaemorus).
Present distribution can be well explained by climatic variables (AUC = 0.96).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
Gadd 4000 0 edd 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 o.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
381
Boloria frigga (Nymphalidae)
an, | Wi wr
=s
bef
=
At
tal
2080
2050
382 Climatic Risk Atlas of European Butterflies
Boloria pales ({SCHIFFERMULLER], 1775) — Shepherd’s Fritillary
.hClcLw Full dispersal No dispersal
Be SEDG -359 (-23.68%) -493 (-32.52%)
BAMBU -286 (-18.87%) | -463 (-30.54%)
-520 (-34.3%) -631 (-41.62%)
SEDG -533 (-35.16%) -808 (-53.3%)
BAMBU | -417 (-27.51%) | -782 (-51.58%)
-631 (-41.62%) -1002 (-66.09%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1516)
The Shepherd’s Fritillary is a characteristic species of flower-rich sub-alpine and alpine grasslands
and can even be found on quite heavily grazed pastures. They can occur in high numbers. At the end
of the day, they often roost communally in damp vegetation with tall plants. Long-spurred Pansy
(Viola cakarata) 1s the major foodplant, but Alpine Plantain (P/aniago alpina) and valerians (Valriana)
are probably also used. Caterpillars are mostly found on plants growing in dry, rocky places. The
Shepherd’s Fritillary has one generation a year and passes the winter in the larval stage.
Present distribution can be well explained by climatic variables (AUC = 0.89).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘add 4000 0 edd 4000 0 ad 4000 0 aad 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O2
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Boloria pales (Nymphalidae) 383
384 Climatic Risk Atlas of European Butterflies
Boloria aquilonaris (STICHEL, 1908) — Cranberry Fritillary
.—CcLwE Full dispersal No dispersal
fi SEDG -2371 (-29%) -2668 (-32.63%)
-2362 (-28.89%) | -2625 (-32.1%)
-3717 (-45.46%) | -4025 (-49.22%)
© Peter Ginzinger Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 8177)
The Cranberry Fritillary inhabits raised bogs and wet heaths, mostly in sheltered places at the edges
of woods, or in clearings. The female lays her eggs singly on the leaves of Cranberry (Vaccinium
oxycoccos) and Marsh Andromeda (Andromeda polifoha). Vhe caterpillar goes into hibernation in
the moss layer just after hatching, only beginning to feed and grow the following year. However,
in adverse conditions, the caterpillar may hibernate a second time. It pupates low down in the
vegetation. It is usually single-brooded.
Present distribution can be well explained by climatic variables (AUC = 0.89).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
‘1.6
sh
Qa i=]
=) a
f= Ss
Oe Oe
ae =a
o= mH
=e
£5 3
Oo. 4
os 7
oo
aes
Ga”
= = rH
ae
qe. =
E gs
wv
x é Po]
= 4
o
0 2000 40000 2000 40000 2000 40000 2000 4000
Gdd Gdd Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
385
Boloria aquilonaris (Nymphalidae)
2080
2050
386 Climatic Risk Atlas of European Butterflies
Boloria graeca (STAUDINGER, 1870) — Balkan Fritillary
6c Full dispersal No dispersal
Be SEDG -239 (-49.69%) -353 (-73.39%)
BAMBU -281 (-58.42%) | -370 (-76.92%)
-271 (-56.34%) | -378 (-78.59%)
SEDG -197 (-40.96%) | -370 (-76.92%)
BAMBU -330 (-68.61%) -418 (-86.9%)
-259 (-53.85%) | -433 (-90.02%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 481)
The Balkan Fritillary can be found in the mountains on flower-rich grasslands, or on grassland with
scattered bushes, and in clearings in woods. These butterflies usually fly close to the ground. Various
violets (Vzo/a spp.) ate used as foodplant. The female lays her eggs either on the foodplant, or on a
plant nearby. The Balkan Fritillary has one generation a year and hibernates as a small caterpillar.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.75).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
tad 4000 0 odd 40000 2000 4000 0 odd 4000
1.0
Minimum
Swe
0.6
1.0 O02
Swe
0.6
1.0 o.2
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
0.
Gdd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Boloria graeca (Nymphalidae) 387
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
2050 2080
388 Climatic Risk Atlas of European Butterflies
Vanessa atalanta (LINNAEUS, 1758) — Red Admiral
6c Full dispersal No dispersal
Be SEDG -2683 (-11.89%) | -3874 (-17.16%)
BAMBU -4288 (-19%) | -4857 (-21.52%)
-4073 (-18.04%) | -5063 (-22.43%)
SEDG -5008 (-22.18%) | -6755 (-29.92%)
BAMBU -8963 (-39.7%) | -10381 (-45.99%)
-11874 (-52.6%) | -13684 (-60.62%)
© Peter Ginzinger Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 22574)
The Red Admiral is a wide-ranging, migratory butterfly that in temperate areas can only survive
mild winters. Each year, butterflies from southern Europe fly northwards, and in good years Red
Admirals can be seen practically everywhere. The butterflies need a lot of nectar, which they get
from flowers. They also feed on rotting fruit and at harvest time are often seen in orchards. They
are also attracted to the resin oozing from trees. The Red Admiral lays its eggs on Stinging Nettle
(Urtica diowa) and Pellitory (Panefaria spp.) in sunny, but not all too dry places. The caterpillars live
alone or in small groups in small tent-like shelters made by spinning leaves together. The caterpillars
pupate in a little tent of spun leaves.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.65).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Ow
Za
on |
4 |
a
e : ‘
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
O 2000 40000 2000 40000 2000 40000 odd 4000
Gdd Gdd Gdd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
389
Vanessa atalanta (Nymphalidae)
2080
2050
390 Climatic Risk Atlas of European Butterflies
Vanessa cardui (LINNAEUS, 1758) — Painted Lady
.6—hlcLw Full dispersal No dispersal
Be SEDG -1792 (-8.9%) -3883 (-19.28%)
BAMBU_ | -4224 (-20.98%) | -5073 (-25.19%)
-3350 (-16.64%) | -5180 (-25.73%)
SEDG -2799 (-13.9%) | -5567 (-27.65%)
BAMBU_ | _ -7054 (-35.03%) | -9170 (-45.54%)
-9272 (-46.05%) | -11805 (-58.63%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 20136)
The Painted Lady is a visitor from the south, which 1s difficult to confuse with other butterflies. The
black triangle on the tip of the forewing distinguishes it from the proper fritillaries. It is a migrant
and cannot survive the winter in temperate climates. Each year, Western Europe is recolonized by
butterflies from the south of Spain and Africa. In good years, the Painted Lady can be seen nearly
everywhere, but otherwise seems to be absent. It visits a variety of flowers for nectar. The Painted
Lady lays its eggs on very different foodplants. It prefers various thistles (C7rsium spp., Carduus spp.,
and Onopordum spp.), but also uses mallows (Masa spp.) especially in the south, Viper’s Bugloss
(Echium vuleare), and Stinging Nettle (Uria diowa). The eggs are laid singly on the upperside of the
leaves. The caterpillars feed in a shelter of loosely spun leaves. They pupate on the foodplant.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.63).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
aad 4000 0 edd 4000 0 oad 4000 0 add 400)
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Vanessa cardui (Nymphalidae) 391
(B1)
SEDG
GRAS
(A1Fl)
392 Climatic Risk Atlas of European Butterflies
Aglais io (LINNAEUS, 1758) — Peacock
nae
-1744 (-8.51%) -3602 (-17.59%)
aa -2035 (-9.94%) | -4094 (-19.99%)
SEDG -1718 (-8.39%) -5136 (-25.07%)
BAMBU_ | -3006 (-14.68%) | -7383 (-36.04%)
-5130 (-25.05%) | -10412 (-50.83%)
© Peter Ginzinger Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 20483)
In temperate Europe, the Peacock is one of the best-known butterflies, because of its striking
appearance and common occurrence. There is no other diurnal butterfly with such noticeable eye-
spots on the upperside of its wings. It is often seen 1n gardens and parks on herbaceous borders
and flowering shrubs, looking for nectar. Eggs are laid on Stinging Nettle (Urtica diowa) in damp,
shady places, sometimes at edges of woodland. Occasionally, Hop (Humulus lupulus) 1s also used. The
caterpillars live gregariously in flimsy webs. Sometimes, plants are completely covered with spun silk,
which serves as a home for tens of caterpillars. They leave the web to pupate on the foodplant. The
adult butterfly goes into hibernation, hiding itself away in cold lofts and sheds. The species usually
has one and sometimes a partial second generation per year.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.8).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
tod 4000 0 Coded 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
10 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
06
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
393
Auglais to (Nymphalidae)
2080
2050
394 Climatic Risk Atlas of European Butterflies
Aglais urticae (LINNAEUS, 1758) — Small Tortoiseshell
[rater | Neco
-3835 (-17.58%) | -4718 (-21.62%)
SEDG -4391 (-20.13%) | -5898 (-27.03%)
BAMBU_ | _ -7863 (-36.04%) | -9692 (-44.42%)
-10097 (-46.28%) | -12111 (-55.51%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 21818)
The Small Tortoiseshell is a common and welcome guest in parks and gardens, and is sometimes
very abundant. It occurs in low numbers in nearly all habitats. Its only foodplant is Stinging Nettle
(Urtia dioica). Vhe foodplants are often growing on nutrient-rich, disturbed ground in the sun, such
as in rough vegetation at the edges of meadows treated with manure or fertilizer. The eggs are laid
in large batches on the underside of the nettle leaves. The caterpillars are gregarious, living in flimsy
webs until they go their separate ways in the last larval instar. The species forms its pupa on the
foodplants. The Small Tortoiseshell hibernates as a butterfly, and can often be found in the cold
months hiding in houses or sheds. It is one of the first butterflies to be seen in the spring.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.8).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
toad 4000 0 odd 4000 0 ad 4000 0 fad 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Swe
0.6
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
395
Aglats urticae (Nymphalidae)
2080
2050
396 Climatic Risk Atlas of European Butterflies
Nymphalis c-album (LINNAEus, 1758) - Comma
6 hc Full dispersal No dispersal
fi SEDG -811 (-3.79%) -2936 (-13.72%)
-1943 (-9.08%) | -3916 (-18.3%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 21399)
The Comma ts a butterfly of wood edges and clearings in wood, except in very dry places. It 1s
easily recognized by the deep indentations 1n the margins of its wings, and the small white comma
on the otherwise dusky underside of the hindwing. Eggs are laid on many different plant species,
like Stinging Nettle (Urtica dioica), bramble (Rubus spp.), elm (U/mus spp.), willow (Sazx spp.), Hazel
(Corylus avellana), and Hop (Humulus lupulus). The caterpillar is solitary, brownish-black with long
spines and a broad white stripe on his back towards the rear, making it look like a bird dropping.
The pupa hangs from the foodplant on a small stalk. The butterflies hibernate in hollow trees,
hedgerows, and shrubs. In large parts of Europe it is double-brooded. However, in Scandinavia 1t
only has one generation, and in warm locations in Spain and Greece, it can have three.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.77).
Climate risk category: LR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 o.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0 2000 40000 2000 40000 2000 40000 2000 4000
Gd dl Gdd
Gdd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
397
Nymphals c-album (Nymphalidae)
2080
2050
398 Climatic Risk Atlas of European Butterflies
Nymphalis egea (CRAMER, 1775) — Southern Comma
CL Full dispersal No dispersal
Be SEDG 283 (15.9%) -368 (-20.67%)
309 (17.36%) -330 (-18.54%)
709 (39.83%) -784 (-44.04%)
1380 (77.53%) | -1161 (-65.22%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1780)
The Southern Comma is a species of dry grasslands and dry scrub. The major foodplant is Common
Pellitory (Panetana officinals), a plant that often grows on old walls, so that these butterflies are often
found near buildings. They can often be seen, wings widespread, basking in the sun on walls and
rocks, or on the ground. The caterpillars probably also feed on Stinging Nettle (Urtica dioica), willows
(Safx spp.), and elms (Udzus spp.). The Southern Comma has two to three generations a year, and
just as the ordinary Comma (P. c-a/bum), bibernates as an adult butterfly. It can therefore be seen
early in the spring.
Present distribution can be well explained by climatic variables (AUC = 0.94).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
=
ch
a
a
=
Ow
Za
uo
sir}
oO
PS : : = : : : :
ah
o
2
w
Oo
‘dd 4000 0 odd 4000 0 oad 4000 0 odd 4000
Minimum
Swe
06
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Nymphals egea (Nymphalidae) 399
(B1)
(A2)
(A1Fl)
400 Climatic Risk Atlas of European Butterflies
Nymphalis antiopa (LINNAEUus, 1758) — Camberwell Beauty
6c Full dispersal No dispersal
Be SEDG -886 (-5.5%) -2806 (-17.43%)
BAMBU -1047 (-6.5%) | -2840 (-17.64%)
-2083 (-12.94%) | -3979 (-24.71%)
SEDG -805 (-5%) -3615 (-22.45%)
BAMBU_ | -3635 (-22.57%) | -6079 (-37.75%)
-5251 (-32.61%) | -8017 (-49.79%)
© Peter Ginzinger Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 16102)
The Camberwell Beauty is an imposing butterfly, which can fly large distances, although this only
happens in some years. It occurs near patches of woodland in stream valleys, gullies, along woodland
edges and on scrub. Because they are fairly mobile, they can be seen in open countryside far away
from their foodplants. The female lays her eggs in large clusters around the twigs of birches (Be/u/a
spp.), willows (Sa4x spp.), and poplars (Populus spp.). At first, the caterpillars live together in a
communal web, becoming solitary when nearly fully-grown. The caterpillar is easily recognized, black
and spiny with a double row of red spots on its back. They pupate on the bark of the foodplants.
The adult butterflies hibernate in a hollow tree, wood pile or sust hidden in the vegetation, and wait
for spring before mating. The Camberwell Beauty has one generation a year.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.76).
Climate risk category: LR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
toad 4000 0 Codd 4000 0 add 4000 0 add 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
401
Nymphaks antiopas (Nymphalidae)
2080
2050
402 Climatic Risk Atlas of European Butterflies
Nymphalis polychloros (LINNAEUS, 1758) — Large Tortoiseshell
AMBU -6995 (-44.1%) | -8952 (-56.44%)
-8220 (-51.83%) | -10595 (-66.8%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 15861)
The Large Tortoiseshell is found in warm, sunny places in deciduous woods and near groups of trees.
Elms (U/mus spp.), willows (Sazx spp.) and sometimes fruit trees or Hawthorn (Crataegus monogyna)
are used as foodplants. The female, usually choosing a twig from the previous year, deposits a large
group of eggs in a band around it. The caterpillars live together in silken tents until the last larval
instar when they become solitary. They are fond of sitting on the sunny side of the foodplant or on
branches in the sun. The caterpillar is dull, dark-grey with rust-coloured stripes along its back and
sides, and long rust-brown spines. The pupa hangs from a twig in the foodplant, looking very like
a withered leaf. The newly-emerged butterflies often roam. The Large Tortoiseshell hibernates as a
butterfly in cool, dark places such as in a hollow tree. It has one generation a year.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.7).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
Minimum
Swe
06
Swe
0.6 10 02
1.0 0.2
Annual precipitation range
Swe
0.6
10 0.2
0.6
Maximum Large (66%) Small (33%)
Swe
o eck 4000 0 odd 4000 0 add 4000 0 Gdd 4000
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
403
Nymphaks polchloros (Nymphalidae)
2080
2050
404 Climatic Risk Atlas of European Butterflies
Nymphalis xanthomelas (EsPrr, 1781) — Yellow-legged Tortoiseshell
AMBU_ | -1738 (-42.28%) | -3683 (-89.59%)
-2297 (55.87%) | -4031 (-98.05%)
© Heiner Ziegler Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 4111)
The Yellow-legged Tortoiseshell occurs in lowland woodland in Eastern Europe, in damp, deciduous
woods growing on flood plains, or on the wooded banks of streams and rivers. Different trees are
used as foodplants, including willows (Saéx spp.), poplars (Populus spp.), elms (Udzus spp.), and
Nettle Tree (Cedtés austrais). Until nearly fully-grown, the caterpillars inhabit large communal nests,
which they spin in branches that hang over the water. The Yellow-legged Tortoiseshell hibernates as
a butterfly hidden away in such places as hollow trees or wood piles. It 1s single-brooded.
Present distribution can be well explained by climatic variables (AUC = 0.87).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
I
dd 4000 0 Coded 4000 0 ad 4000 0 Gdd 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
405
Nymphals xanthomelas (Nymphalidae)
2080
2050
406 Climatic Risk Atlas of European Butterflies
Nymphalis l-album (Esper, 1780) — False Comma
SEDG 101 (1.93%) -1761 (-33.63%)
1514 (28.91%) | -850 (-16.23%)
ae (1.72%) -1836 (-35.06%)
SEDG 3311 (63.22%) | -1611 (-30.76%)
BAMBU 3350 (63.97%) | -2324 (-44.38%)
3035 (57.95%) | -2884 (-55.07%)
© Zdravko Kolev Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 5237)
The False Comma occurs in the lowlands of Eastern Europe, in deciduous or mixed woods. It
prefers damp woods, and is found in clearings or at the wood edge. It is a mobile butterfly and a
strong migrant. The female lays her eggs in the spring, clustered around the twigs of the foodplants
which may be birches (Be/w/ia spp.), willows (Sax spp.), poplars (Populus spp.), or elms (U/vus spp.).
The False Comma has one generation a year, and because it hibernates as a butterfly, can be seen
for much of the year.
This species is listed in Annexes I and IV of the Habitats’ Directive, where it erroneously is named
N. vaualbum.
Present distribution can be well explained by climatic variables (AUC = 0.89).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘dd 4000 0 edd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
407
Nymphals l-album (Nymphalidae)
2080
2050
408 Climatic Risk Atlas of European Butterflies
Araschnia levana (LINNAEus, 1758) — Map
.—hCcLw Full dispersal No dispersal
Be SEDG -750 (-5.89%) -3045 (-23.93%)
BAMBU -231 (-1.82%) | -2980 (-23.42%)
1214 (-9.54%) | -3737 (-29.37%)
SEDG -2108 (-16.57%) | -5787 (-45.48%)
BAMBU_ | -1661 (-13.05%) | -7261 (-57.06%)
-2943 (-23.13%) | -9390 (-73.79%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 12725)
The Map its a common butterfly of woodland edges, coppices, hedgerows and wood banks. It
normally has two generations a year, and the butterflies of the spring brood are predominantly
orange, and those of the summer brood predominantly black on the upperside of the wings. The
Map gets its name from the intricate pattern on the underside of its wings. These butterflies enjoy
basking in the sun with their wings wide open, often on the ground. The Map has a dainty floating
flight, but it settles again quite quickly. The foodplants are Stinging Nettle (Uria dioica) and Small
Nettle (U. wrens). The eggs are deposited in short chains on the underside of the nettle leaves and
look very much like threaded pearls. The caterpillars are gregarious, but do not spin a nest. They
pupate on the foodplants, and pass the winter as a pupa.
Present distribution can be well explained by climatic variables (AUC = 0.87).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
‘did 4000 0 edd 4000 0 add 4000 0 Gd 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Swe
0.6
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
409
Alraschnia levana (Nymphalidae)
2080
2050
410 Climatic Risk Atlas of European Butterflies
Euphydryas iduna (DALMAN, 1816) — Lapland Fritillary
ae ee
-1047 (-79.32%) -1057 (-80.08%)
[ena a sry | as
SEDG -1211 (-91.74%) | -1223 (-92.65%)
BAMBU_ | -1276 (-96.67%) | -1280 (-96.97%)
-1315 (-99.62%) | -1315 (-99.62%)
© Otakar Kudrna Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1320)
At lower altitudes in Lapland, this lovely fritillary occurs on flower-rich swampy areas and wet
slopes with scattered birches. Above 500 m, it is found in drier rocky places. Compared to the other
fritillaries that occur in Lapland, the Lapland Fritillary 1s remarkably colourful. It is one of the
first arctic species to be seen, appearing practically as soon as the snow has melted. It flies quickly,
zigzageing low over the vegetation. The female lays her eggs in small groups on plantains (Plantago
spp.), Alpine Speedwell (Veronica alpina), Rock Speedwell (V._fruticans), and species of Vaccinium. The
caterpillars live communally in small silken webs and hibernate there. They pupate in the spring. It
has one brood a year.
Present distribution can be very well explained by climatic variables (AUC = 1).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 edd 4000 0 oad 4000 0 add 4000
1.0
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
2050
Euphydryas iduna (Nymphalidae)
2080
SEDG
411
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
412 Climatic Risk Atlas of European Butterflies
Euphydryas cynthia ({SCHIFFERMULLER], 1775) — Cynthia’s Fritillary
.6—hCcLUwk Full dispersal No dispersal
fe SEDG -77 (-19.49%) -147 (-37.22%)
-80 (-20.25%) -149 (-37.72%)
GRAS | -132 (-33.42%) | -180 (-45.57%)
SEDG -233 (58.99%) | -285 (-72.15%)
BAMBU | -165 (-41.77%) | -249 (-63.04%)
-210 (-53.16%) | -327 (-82.78%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 395)
Cynthia’s Fritillary distinguishes itself from other fritillaries by the large amount of white on the
basal parts of the upperside of the wings of the male. The butterflies occur on sub-alpine and
alpine grassland with short, grassy vegetation and low-growing shrubs and in rocky areas. They fly
quickly, close to the ground, and are fond of basking in the sun on rocks or bare patches of ground.
The female lays her eggs in clusters under leaves of Alpine Plantain (Planiago alpina) and Long-
spurred Pansy (Vola cakarata). The caterpillars are black and bristly with yellow bands between each
segment. They can be very numerous and take two years to develop. The first hibernation takes place
communally in a silken web. In the fourth larval instar, they hibernate a second time, solitarily under
stones, where they later pupate.
Present distribution can be very well explained by climatic variables (AUC = 0.98).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 Codd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
2050
Euphydryas cynthia (Nymphalidae)
2080
SEDG
413
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
414 Climatic Risk Atlas of European Butterflies
Euphydryas intermedia (MENETRIES, 1859) — Asian Fritillary
Cc Full dispersal No dispersal
Be SEDG -24 (-8.92%) -127 (-47.21%)
BAMBU -11 (-4.09%) -123 (-45.72%)
-69 (-25.65%) -155 (-57.62%)
SEDG -157 (-58.36%) | -231 (-85.87%)
BAMBU -45 (-16.73%) -193 (-71.75%)
[oras | -105 (-39.03%) -233 (-86.62%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 269)
The Asian Fritillary can be seen searching for nectar or foodplants on open scrub, rich in herbaceous
plants, or in clearings in open woodland, in flower-rich grassland, and on vegetation of dwarf
shrubs above the tree-line. These butterflies only occur locally, although at times in large numbers.
The female lays all her eggs at once, on the underside of a leaf of its foodplant, Blue Honeysuckle
(Lonicera caerulea). The caterpillars live in communal webs, and in the autumn make a more substantial
shelter of leaves, spun together with silk, in which to hibernate. They take two years to develop and
have to hibernate twice. The fully-grown caterpillar leaves the nest after the second hibernation, and
pupates on the foodplant or adjacent rocks.
Present distribution can be very well explained by climatic variables (AUC = 0.98).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
m™
o
° : : : : :
todd 4000 0 edd 4000 0 oad 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe Swe
0.6 0.6
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
2050
Euphydryas intermedia (Nymphalidae)
2080
SEDG
415
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
416 Climatic Risk Atlas of European Butterflies
Euphydryas maturna (LINNAEUS, 1758) — Scarce Fritillary
a
2526 (35.26%) | -1754 (-24.48%)
SEDG 3354 (46.82%) | -1971 (-27.51%)
BAMBU 3987 (55.65%) | -2446 (-34.14%)
2476 (34.56%) | -3429 (-47.86%)
© Rudi Verovnik Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 7164)
The Scarce Fritillary occurs in clearings, where young ash trees are growing in open, mixed wood-
land. The eggs are laid in one batch on a leaf of Ash (Fraxinus excelsior) or Aspen (Populus tremula),
preferably at a height of around 4 m. The caterpillars build a nest of silk and leaves, and feed to-
gether at first, while still quite small. They go into hibernation, remaining in the nest, which usually
falls onto the woodland floor. In spring, they leave the nest and separate, spreading out in search
of food. They use a variety of larval foodplants at this stage, including honeysuckle (Lonwera spp.),
plantains (P/antago spp.), or privets (Ligustrum spp.). They pupate in the litter layer. The species has
one generation a year although some of the caterpillars hibernate a second time before pupating.
This species is listed in Annexes II and IV of the Habitats’ Directive.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.79).
Climate risk category: LR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘dd 4000 0 ood 4000 0 oad 4000 0 ada 4000)
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
10 0.2
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
417
Euphydryas maturna (Nymphalidae)
2080
2050
418 Climatic Risk Atlas of European Butterflies
Euphydryas desfontainii (GovartT, 1819) — Spanish Fritillary
Cc Full dispersal No dispersal
eae -1003 (-61.12%) | -1132 (68.98%)
-1226 (-74.71%) | -1256 (-76.54%)
-1593 (-97.07%) | -1622 (-98.84%)
-1624 (-98.96%) | -1640 (-99.94%)
© Bernard Fransen Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1641)
The Spanish Fritillary occurs on open, grassy places with stony soil and lots of shrubs. It uses
various foodplants, such as Cephalania kucantha and C. syriaca, the teasels Dipsacus fullonum and D.
comasus, scabtous (Scabiosa and Knautia spp.) and possibly knapweeds (Cen/aurea spp.). The female lays
her eggs in small batches on the underside of leaves. The caterpillars live in a communal spun nest,
also hibernating there. There is one brood a year.
Present distribution can be well explained by climatic variables (AUC = 0.88).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
aad 4000 0 edd 4000 0 add 4000 0 aad 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
Swe
o6 10 02
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
2050
Euphydryas desfontainit (Nymphalidae)
2080
SEDG
419
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
420 Climatic Risk Atlas of European Butterflies
Euphydryas aurinia (ROTTEMBURG, 1775) — Marsh Fritillary
lc Full dispersal No dispersal
fe SEDG -276 (-4.24%) -976 (-14.99%)
335 (5.14%) -786 (-12.07%)
GRAS | -543 (-8.34%) | -1292 (-19.84%)
SEDG -1055 (-16.2%) | -1908 (-29.3%)
BAMBU_ | -1115 (-17.12%) | -2365 (-36.31%)
-2488 (-38.2%) | -3659 (-56.18%)
© Peter Ginzinger Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 6513)
The Marsh Fritillary occurs in very different types of habitat, like moist, sheltered grasslands, along
the edges of raised bogs and on dry, calcareous grasslands. The foodplants are Devil’s-bit Scabious
(Sacvisa pratense), Small Scabious (Scabiosa columbaria), Field Scabious (Knautia arvensis), and teasels
(Dipsacus spp.). The eggs are laid in large clumps under the leaves. The caterpillars spin a substantial
nest between the leaves of the foodplants, feeding in it, and also hibernating communally there.
However, later they are solitary, and look for places deep in the vegetation in which to pupate. The
Marsh Fritillary has one brood a year.
This species is listed in Annex I of the Habitats’ Directive.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.72).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Gadd 4000 0 Codd 4000 0 add 4000 0 aad 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
421
Euphydryas aurinia (Nymphalidae)
2080
2050
422 Climatic Risk Atlas of European Butterflies
Melitaea cinxia (LINNAEUS, 1758) — Glanville Fritillary
a
-2647 (-20.06%) | -3961 (-30.02%)
SEDG -4653 (-35.26%) | -5935 (-44.98%)
BAMBU_ | -5706 (-43.24%) | -7293 (-55.27%)
-6896 (-52.26%) | -8941 (-67.76%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 13195)
The Glanville Fritillary is found on many different types of flower-rich grasslands, both on calcareous
and acid soils. This butterfly can survive on meadows and pastures, as well as on road verges and
forgotten patches of vegetation, sometimes small habitats supporting large populations. Various
plantains (Plintago spp.), speedwells (Veronica spp,), and knapweeds (Cenfaurea spp.) ate used as
foodplants. The eggs are laid in large batches on the underside of the leaves. The caterpillars live
eregariously in a spun nest, also hibernating in a thicker one when half-grown. The Glanville Fritillary,
usually has one generation a year, partially a second one under favourable conditions.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.75).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
‘1.6
sh
a a
=) a
i
Pe Oe
ef@eis
o= mH
=e
£5 8
Oo. 4
of 7
D >
a = ge
ger
|
ae
qf. =
2 o
wv
x é Po]
= 4
o
0 2000 40000 2000 40000 2000 40000 2000 4000
Gdd Gdd Gadd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
423
Mehitaea cinxia (Nymphalidae)
2080
2050
424 Climatic Risk Atlas of European Butterflies
Melitaea phoebe (Gorze, 1779) — Knapweed Fritillary
SEDG 2383 (30.98%) | -1579 (-20.53%)
vais (837%) | 2181 (235%
ae 1267 (16.47%) | -2473 (-32.15%)
SEDG 2605 (33.86%) | -2630 (-34.19%)
BAMBU 1417 (18.42%) | -4209 (54.71%)
1843 (23.96%) -5324 (-69.21%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 7693)
The Knapweed Fritillary occurs in dry, open places with flower-rich vegetation. These grasslands
are often 1n a sheltered situation at the edge of woodland, or of groups of shrubs. In the north of
its range, the grasslands are mostly calcareous. The larval foodplants are knapweeds (Cen/aurea spp.).
The female lays her eggs in large batches on the undersides of the leaves. The small caterpillars live
eregariously in a silken nest, also hibernating together. Later, they become solitary, and when ready
to pupate, choose somewhere close to the ground. The Knapweed Fritillary usually has two broods,
but at higher altitudes and in the north of its range only one.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.77).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
‘said 4000 0 odd 4000 0 add 4000 0 oad 4000
Minimum
Swe
‘1.6
1.0 02
0.6
Swe
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Mektaea phoebe (Nymphalidae) 425
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
426 Climatic Risk Atlas of European Butterflies
Melitaea aetherie (HUBNER, 1826) — Aetherie Fritillary
.—hlcLw Full dispersal No dispersal
Be SEDG 51 (13.01%) -215 (-54.85%)
214 (54.59%) -248 (-63.27%)
2480 (632.65%) | -351 (-89.54%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 392)
In Spain, the Aetherie Fritillary is found 1n clearings in disturbed Holm Oak (Quercus tkx) woodlands,
where its foodplants, thistles (Czrsiwm spp.) and knapweeds (Cenlaurea spp.), such as Star Thistle
(Centaurea cakitrapa), Brown Knapweed (C. jacea), and C. carratracensis gtow. The female lays her eggs
in batches on the foodplants. The Aetherie Fritillary normally has one brood a year, and hibernates
as a caterpillar.
Present distribution can be well explained by climatic variables (AUC = 0.95).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
edd 4000 0 odd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
0.
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
427
Melitaea aetherte (Nymphalidae)
2080
2050
428 Climatic Risk Atlas of European Butterflies
Melitaea trivia ({SCHIFFERMULLER], 1775) — Lesser Spotted Fritillary
ri
AMBU | -1820 (-35.03%) | -3718 (-71.57%)
-1070 (-20.6%) -4285 (-82.48%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 5195)
The Lesser Spotted Fritillary occurs on flower-rich, grassy vegetation, in both dry and damp places,
and on waste ground. Various mulleins (Verbascum spp.) are used as foodplants, including Aaron’s Rod
(Verbascum thapsus), V. densiflorum and V. longifoium. The eggs are laid in batches on the underside of
the leaves. The young caterpillars feed, gregariously in a communal web, where they also hibernate.
After hibernation, they form smaller groups, and are often seen on the upper surface of the leaves.
This species has one or two broods a year, depending on the climatic zone of its flight area.
Present distribution can be well explained by climatic variables (AUC = 0.86).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘dd 4000 0 odd 4000 0 oad 4000 0 odd 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Swe
0.6
Annual precipitation range
Maximum Large (66%) Small (33%)
1.0 02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
429
Mehtaea trivia (Nymphalidae)
2080
2050
430 Climatic Risk Atlas of European Butterflies
Melitaea didyma (EsPkER, 1779) — Spotted Fritillary
a
1374 (11.08%) | -2686 (-21.66%)
SEDG 1522 (12.27%) | -3397 (-27.4%)
BAMBU -73 (0.59%) -5816 (-46.9%)
-347 (-2.8%) -7258 (-58.53%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 12400)
The Spotted Fritillary can be found on poor grasslands, steppe-like vegetation, and dry, rough vegetation
near rocks and on slopes. Further, it is also seen on fallow agricultural land, or at the edges of fields. It
uses a wide variety of plants as foodplant, plantains (P/aniago spp.), toadflaxes (Linara spp.), speedwells
(Verona spp.), foxgloves (Digilahs spp.), woundworts (S/achys spp.), valertans (Vakriana spp.), and
mulleins (Verbascum spp.). The female lays her eggs in clumps on the underside of the leaves, near to the
ground. At first, the caterpillars feed communally in a loosely spun shelter, but they separate quite soon,
either into smaller groups, or become solitary. The caterpillars hibernate alone, or in small groups, in a
spun web. They pupate low down on the foodplant or in the vegetation. The Spotted Fritillary has one
to three broods a year, depending on the geographical location and altitude of its breeding ground.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.77).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
fad 4000 0 Codd 4000 0 aad 4000 0 Gdd 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 O.2
Swe
0.6
1.0 O2
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
431
Mehtaea didyma (Nymphalidae)
2080
2050
432 Climatic Risk Atlas of European Butterflies
Melitaea diamina (LANG, 1789) — False Heath Fritillary
.—hCcLUw Full dispersal No dispersal
fi SEDG -1169 (-13.54%) | -2382 (-27.59%)
171 (1.98%) -1648 (-19.09%)
-1780 (-20.62%) | -4472 (-51.8%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 8633)
The False Heath Fritillary is found in sheltered, marshy habitats and damp woodland clearings. At
higher altitudes, it is also found on rough, calcareous grasslands. Different sorts of valerian (Valriana
spp.) are used as foodplants, the female depositing her eggs in large clusters on the underside of the
leaves. The small caterpillars only feed for a short time before hibernating communally in a silken
shelter. After hibernation, they separate, later pupating low down on the foodplant. The False Heath
Fritillary usually has one brood a year, but at low altitudes it sometimes has two.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.84).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
‘1.6
10 02
Swe
0.6 A i 1.0
|
« < < |
q |
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
10 02 6.6
10 0.2
Swe
06
o2
L=]
2000 40000 2000 40000 2000 40000 2000 4000
dl Gadd Gdd
Gdd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
433
Mehtaea diamina (Nymphalidae)
2080
2050
434 Climatic Risk Atlas of European Butterflies
Melitaea deione (GEYER, 1832) — Provencal Fritillary
.6—hCcLUw Full dispersal No dispersal
fe SEDG -832 (-34.3%) -1587 (-65.42%)
-1517 (-62.53%) -1792 (-73.87%)
fens | amar [sree cair9
-1236 (-50.95%) | -2071 (-85.37%)
-1062 (-43.78%) | -2213 (-91.22%)
© Martin Wiemers Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2426)
The Provengal Fritillary occurs in all sorts of flower-rich, grassy places; on flower-rich grasslands, both
in damp and dry places, and on calcareous as well as acid soil, on flower-rich, grassy vegetation on the
banks of rivers and streams, in scrub and at woodland edges. In Switzerland, at the extreme north of
its distribution range, it is only found in dry, warm, bushy places. Its foodplants are toadflax (Linana
spp.), snapdragons (Axtirrhinum spp.), and sometimes also foxgloves (Digitalis spp.). The female lays
her eggs in small batches on the underside of the leaves. The caterpillars hibernate in a spun shelter.
In most locations, this species has two broods a year, however in cooler places only one.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.76).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
‘sad 4000 0 ood 4000 0 add 4000 0 ada 4000
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 60.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
435
Mehitaea detone (Nymphalidae)
2080
2050
436 Climatic Risk Atlas of European Butterflies
Melitaea varia (MEYER-Dtr, 1851) — Grisons Fritillary
a
-66 (-13.04%) -298 (-58.89%)
SEDG -232 (-45.85%) | -438 (-86.56%)
BAMBU_ | -123 (-24.31%) | -355 (-70.16%)
-249 (-49.21%) | -434 (-85.77%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 506)
The Grisons Fritillary is a small fritillary that breeds on flower-rich sub-alpine and alpine grasslands
and on sunny, grassy slopes. The adult butterflies visit mostly low plants for nectar, and the males are
also seen on damp patches, excrement, and dead animals. The female lays her eggs in clusters on the
foodplant. The caterpillars have been found on Alpine Plantain (P/antago alpina) and Achillea species,
but Spring Gentian (Gentiana verna) and Stemless Trumpet Gentian (G. acaulis) are also named as
foodplants. The caterpillars hibernate, pupating in June or July of the following year, the pupa
usually hanging from a stone. However, at higher altitudes, development is slower, taking nearly two
years, and they hibernate twice.
Present distribution can be very well explained by climatic variables (AUC = 0.96).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘dd 4000 0 odd 4000 0 add 4000 0 add 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
437
Melitaea varia (Nymphalidae)
(1a)
504s
(ZV)
Nanva
2080
2050
438 Climatic Risk Atlas of European Butterflies
Melitaea parthenoides (KEFERSTEIN, 1851) — Meadow Fritillary
.6—hlcLw Full dispersal No dispersal
Be SEDG -263 (-8.26%) -1251 (-39.3%)
BAMBU -467 (-14.67%) | -1326 (-41.66%)
-805 (-25.29%) | -1756 (-55.17%)
SEDG -1137 (-35.72%) | -2098 (-65.91%)
BAMBU | -1391 (-43.7%) | -2457 (-77.19%)
-1842 (-57.87%) | -2969 (-93.28%)
© Henk Bosma Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 3183)
The Meadow Fritillary occurs in both dry and moderately damp habitats in many sorts of open,
erassy, flower-rich places near woodland. Plantains (P/aniago spp.) are the main foodplants, especially
Ribwort Plantain (P. anceolata). The eggs are laid in clusters on the underside of the leaves. The
caterpillars feed and hibernate communally in a silken shelter, only separating in the last larval instar.
They then look for a safe place to pupate, low down on the foodplant. The Meadow Fritillary usually
has two generations a year, but only one 1n cool breeding grounds.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.84).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
ce
a iy]
toad 4000 0 ead 4000 0 add 4000 0 odd 4000
Minimum
Swe
‘0.6
1.0 02
0.6
Swe
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
439
Mehitaea parthenoides (Nymphalidae)
2080
2050
440 Climatic Risk Atlas of European Butterflies
Melitaea aurelia (NickERL, 1850) — Nickerl’s Fritillary
.—hCcLUwk Full dispersal No dispersal
He SEDG 335 (7.57%) -1527 (-34.52%)
2241 (50.67%) | -1117 (-25.25%)
|GRAS | 801 (18.11%) -1641 (-37.1%)
SEDG 625 (14.13%) | -2076 (-46.94%)
BAMBU 1372 (31.02%) | -2361 (-53.38%)
490 (11.08%) -3288 (-74.34%)
© Martin Wiemers Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 4423)
Nickerl’s Fritillary occurs on open, dry grasslands and heaths on calcareous slopes and is very heat-
tolerant. The female deposits her eggs in clusters on the underside of the leaves of its foodplant
Ribwort Plantain (Plantago lanceolata). The caterpillars feed and also hibernate 1n communal silken
nests. They pupate low down in the vegetation. Nickerl’s Fritillary has one generation a year.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.82).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
‘1.6
sh
Qa a
=) a
=S 7
ean
cf@es
o= mH
=
£5 8
Oo. 4
ow 6
oo
a = ge
ger
te |
sme
qf. =
2 o
wv
x é Po]
= 4
o
0 2000 40000 2000 40000 2000 40000 2000 4000
Gdd Gdd Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
441
Mektaea aurelia (Nymphalidae)
2080
2050
442 Climatic Risk Atlas of European Butterflies
Melitaea britomartis (ASsMANN, 1847) — Assmann’s Fritillary
.6—hCcLwkE Full dispersal No dispersal
Be SEDG 1268 (29.63%) -1324 (-30.93%)
BAMBU 31 (0.72%) -2095 (-48.95%)
793 (18.53%) | -1987 (-46.43%)
SEDG -1646 (-38.46%) | -3144 (-73.46%)
BAMBU_ | -1961 (-45.82%) | -3747 (-87.55%)
-2251 (-52.59%) | -4035 (-94.28%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 4280)
Assmann’s Fritillary occurs on warm grasslands and heath that are often situated at the edges of
woodland, or neat groups of shrubs. Its foodplants are Ribwort Plantain (Plantago lanceolata), Yellow
Rattle (Rhinmanthus minor, and the Speedwell (Veronica austriaca). Vhe female lays her eggs in batches on
the underside of the leaves. The caterpillars feed communally in a silken nest, where they also hibernate.
They then separate, later pupating low-down in the vegetation. This fritillary is single-brooded.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.83).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
edd 4000 0 odd 4000 0 add 4000 0 odd 4000
1.0
Minimum
Swe
0.6
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Mektaea britomartis (Nymphalidae) 443
(B1)
SEDG
GRAS
(A1Fl)
444 Climatic Risk Atlas of European Butterflies
Melitaea athalia (ROTTEMBURG, 1775) — Heath Fritillary
.—hCcLUw Full dispersal No dispersal
Be SEDG 109 (0.7%) -2510 (-16.21%)
BAMBU -610 (-3.94%) | -2730 (-17.63%)
-1391 (8.98%) | -3658 (-23.63%)
SEDG 560 (3.62%) -3902 (-25.2%)
BAMBU -1542 (-9.96%) | -5975 (-38.59%)
-2980 (-19.25%) | -7785 (-50.28%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 15482)
The Heath Fritillary is found in many different sorts of biotope, ranging from dry to damp, grassy,
flower-rich places, often situated near bushes or in woodland, or in clearings and along the edges
of paths in woods. Its foodplants are plantains (Planiago spp.), cow-wheats (Melampyrum spp.),
speedwells (Veronica spp.), foxgloves (Digilahs spp.), and toadflaxes (Linara spp.). The female lays her
egos in clusters on the underside of the leaves. The caterpillars feed communally in silken nests, also
hibernating together when half-grown. They then disperse over the plant, either into small groups,
ot, mostly, become solitary. They pupate on the foodplant, and usually have one brood a year, except
in the south, where they have two.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.79).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
mM
o
: OLGiLa
tod 4000 0 Codd 4000 0 oad 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
445
Mehtaea athalia (Nymphalidae)
2080
2050
446 Climatic Risk Atlas of European Butterflies
Limenitis populi (LINNAEUS, 1758) — Poplar Admiral
SEDG -1234 (-10.89%) | -5291 (-46.69%)
BAMBU_ | -1580 (-13.94%) | -6352 (-56.05%)
-3162 (-27.9%) | -8315 (-73.38%)
© Rudi Verovnik Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 11332)
An encounter with the Poplar Admiral is one of those things that one never forgets. It is an impressive
butterfly, and the woods it inhabits are areas of natural beauty. They are found in mixed woodlands
with damp clearings, where its foodplants Aspen (Populus tremula) and Black Poplar (Populus nigra)
grow. The female deposits her eggs one by one on leaves that are preferably situated in the sun, with
more branches above them. The caterpillar feeds on the leaves and usually builds a hibernaculum,
which is fastened tightly onto a twig with spun thread. It hibernates in the second larval instar, and
in the spring eats large numbers of leaves before finally pupating, suspended from a leaf. The Poplar
Admiral has one generation a yeat.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.84).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
‘1.6
ct
a a
=) a
>= 7
Pe
cfes
o= mH
=e
£5 8
G. 4
of 7
D >
cL m4 = =
ger
ae |
re
a
Qe
2 o
aw
x é 5
= 4
o
0 2000 40000 2000 40000 2000 40000 2000 4000
Gd id Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
447
Limenitis popul (Nymphalidae)
2080
2050
448 Climatic Risk Atlas of European Butterflies
Limenitis camilla (Linnaeus, 1764) — White Admiral
=o
890 (10.08%) -1716 (-19.43%)
SEDG -979 (-11.08%) | -3580 (-40.53%)
BAMBU -161 (-1.82%) | -4232 (-47.91%)
-1447 (-16.38%) | -6253 (-70.79%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 8833)
The White Admiral is a true woodland butterfly and is found in clearings and woodland rides. The
female lays her eggs on young honeysuckle (Lonwera spp.) leaves, preferring those which are half in
the shade. The caterpillars feed on the leaves in a characteristic way. Starting at the tip and working
towards the stalk, they leave the main nerve intact, and use it to rest upon. The hibernation is in the
second larval instar. To make the hibernaculum the remaining part of the leaf is spun into a little
tube, and the leaf stalk is spun tightly onto a twig. After hibernating, the caterpillar feeds on the
newly-emerged leaves. It pupates upside down on a twig, and has one brood a year.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.81).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘dd 4000 0 edd 4000 0 oad 4000 0 odd 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
1.0 02
0.6
Maximum Large (66%) Small (33%)
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Limenitis camilla (Nymphalidae)
SEDG
GRAS
(B1)
(A2)
(A1Fl)
449
450 Climatic Risk Atlas of European Butterflies
Limenitis reducta (STAUDINGER, 1901) — Southern White Admiral
[raat [ ne ateena
735 (12.86%) -1332 (-23.3%)
SEDG 1180 (20.64%) | -1764 (-30.86%)
BAMBU 897 (15.69%) | -2390 (-41.81%)
ra ananny _| -9062 (ST 0TH
© Peter Ginzinger Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 5716)
The Southern White Admiral occurs in warm to very warm places in woodland and scrub, often
situated near streams, springs or other damp places. The eggs are laid on nearly all species of
honeysuckle (Lonwera spp.). The caterpillars feed on the leaves in the manner characteristic of this
genus, nibbling at the soft tissues while leaving the main nerve free. For hibernation, a small cradle
is made from the remains of the leaf, which is secured to a twig with silk. When they come out of
hibernation, they begin feeding on the young honeysuckle leaves. The caterpillar pupates suspended
from a twig of the foodplant. This species has one brood a year North of the Alps but two in the
Mediterranean region.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.82).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
si
=|
° : : ? : ?
todd 4000 0 odd 4000 0 Sad 4000 0 ada 4000
1.0
Minimum
Swe
0.6
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe Swe
0.6 0.6
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
451
Limenitis reducta (Nymphalidae)
2080
2050
452 Climatic Risk Atlas of European Butterflies
Neptis sappho (Pattas, 1771) - Common Glider
=o
ae 1157 (44.38%) -1389 (-53.28%)
SEDG 876 (33.6%) -1769 (-67.86%)
BAMBU 1506 (57.77%) -2254 (-86.46%)
}GRAS 1649 (63.25%) | -2435 (-93.4%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2607)
The Common Glider inhabits damp, deciduous woodland, especially in river valleys. The butterflies
often settle in the top of the trees to rest or bask 1n the sun, with their wings wide open, just like the
Hungarian Glider (N. mulans). This butterfly has a characteristic flight, elegantly gliding from perch
to perch. The caterpillars feed on Spring Pea (Lathyrus vernus) and Black Pea (L. niger). The Common
Glider has two broods a year and hibernates as a small caterpillar.
Present distribution can be well explained by climatic variables (AUC = 0.92).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 odd 4000 0 add 4000 0 ada 4000
0.6
Minimum
Swe
Swe
Zt 0.6 1.0 02
Swe
0.6 1.0
1.0 0.2
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Neptis sappho (Nymphalidae) 453
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
454 Climatic Risk Atlas of European Butterflies
Neptis rivularis (Scoro.t, 1763) — Hungarian Glider
a
-1082 (-50.68%) | -1391 (65.15%)
SEDG 74 (3.47%) -1285 (-60.19%)
BAMBU | -458 (-21.45%) | -1520 (-71.19%)
-121 (5.67%) | -1763 (-82.58%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2135)
The Hungarian Glider is a butterfly of light, damp woodland, occurring in deciduous, as well as 1n
mixed woods. It 1s similar to the White Admiral (Limenitis camilla), which is also a woodland butterfly.
The butterflies are rarely seen drinking nectar from flowers. It glides from perch to perch with only
an occasional flap of the wings. Goat’s-beard (Aruncus dioicus), Meadowsweet (Fikipendula ulmaria),
Bridewort (Spiraea sahiifolia), and S. chamaedryfoha are used as foodplants. The female deposits the
eges singly on the upperside of the leaves. After hatching, the tiny caterpillar first eats up the
egoshell and then makes itself a shelter in the tip of the leaf. It only leaves the shelter to feed. In the
autumn, it makes another shelter 1n which to hibernate. It has one brood a year.
Present distribution can be well explained by climatic variables (AUC = 0.87).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 Codd 4000 0 ad 4000 0 add 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
455
Neptis rivularis (Nymphalidae)
2080
2050
456 Climatic Risk Atlas of European Butterflies
Charaxes jasius (LINNAEUS, 1767) —'Two-tailed Pasha
.—ClcLe Full dispersal No dispersal
Be SEDG -760 (-38.4%) -1174 (-59.32%)
BAMBU -879 (-44.42%) | -1198 (-60.54%)
-840 (-42.45%) | -1367 (-69.08%)
SEDG -795 (-40.17%) -1504 (-76%)
BAMBU | -1121 (56.64%) | -1737 (87.77%)
-1012 (51.14%) | -1918 (-96.92%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1979)
With the whimsical patterning on the undersides of its wings and elegant little tails, the Two-tailed
Pasha is one of the most beautiful European butterflies. It breeds 1n warm, dry places with many
shrubs and trees where its foodplant, the Strawberry Tree (Arbutus unedo), 1s mostly abundant. The
males defend their territory, attacking other insects. The butterflies can often be seen feeding on
the juices of rotting fruit, such as figs, and are also attracted to alcoholic drinks. They are strong
flyers, often roaming into areas where their foodplant is absent. The Two-tatled Pasha has mostly
two broods a year. It hibernates as a caterpillar and pupates suspended from the foodplants. On
Gibraltar, Samos and Cyprus, it also uses other foodplants.
Present distribution can be well explained by climatic variables (AUC = 0.94).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o dd 4000 0 odd 4000 0 add 4000 0 odd 4000
1.0
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Charaxes jasius (Nymphalidae) 457
458 Climatic Risk Atlas of European Butterflies
ame metis (FREYER, 1829) — Freyer’s Purple Emperor
|| | Fattaispersat | No aispersa
He SEDG 3014 (144.14%) | -141 (6.74%)
1346 (64.37%) | -589 (-28.17%)
fas aio | em a0
212 (10.14%) | -1703 (-81.44%)
787 (37.64%) -1885 (-90.15%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2091)
Freyer’s Purple Emperor is a butterfly of very warm, damp places. The butterflies can be found
along wooded riverbanks, where its foodplant, White Willow (Sax alba) grows. The males and
females meet each other at the tops of tall trees, and the females lay their eggs in small batches at
the top of the tree, on leaves in the crown. The caterpillars grow quickly, and pupate suspended
under a leaf or on a twig. The caterpillars from the brood that follows, hibernate. The butterflies
have a varied diet. The females visit flowers for nectar, aphids for honeydew, and ripe fruit for the
sugars. [he males are often found on damp ground, on dung and on carrion. The butterflies are also
attracted to sap oozing from wounded trees. This species has two generations a yeat.
This species is listed in Annex IV of the Habitats’ Directive.
Present distribution can be very well explained by climatic variables (AUC = 0.98).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘edd 4000 0 odd 4000 0 add 4000 0 fad 4000
1.0
Minimum
Swe
0.6
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
459
Apatura metis (Nymphalidae)
2080
2050
460 Climatic Risk Atlas of European Butterflies
Apatura ilia ({SCHIFFERMULLER], 1775) — Lesser Purple Emperor
a
-633 (-6.36%) | -3412 (-34.26%)
SEDG 1042 (10.46%) | -4137 (-41.54%)
BAMBU 526 (5.28%) -5548 (-55.71%)
-37 (-0.37%) -7270 (-73.01%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 9958)
The Lesser Purple Emperor is found in clearings and along paths in damp, deciduous woodland that
is often situated in river valleys or on the banks of rivers. The butterflies are often seen near puddles
on the road, and on dung from other animals. Espectally the males are attracted to strongly smelling
cheese. Its foodplants are poplars (Populus spp.), and willows (Sadx spp.). The female, choosing
rather small, not so robust trees, deposits her eggs on the upperside of leaves that are half in the
shade, or in the sun. The half-grown caterpillar spins itself a little pad in the fork of a twig on which
it htbernates. In the spring, it returns to the fresh buds and leaves to feed. It pupates suspended from
the underside of a leaf or branch. The Lesser Purple Emperor has one to two broods a yeat.
Present distribution can be well explained by climatic variables (AUC = 0.86).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 ood 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Swe
0.6
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
461
Apatura ila (Nymphalidae)
2080
2050
462 Climatic Risk Atlas of European Butterflies
Apatura iris (LINNAEUS, 1758) — Purple Emperor
a sc
-2504 (-28.51%) | -4708 (-53.6%)
SEDG -2825 (-32.16%) | -5511 (-62.74%)
BAMBU -2969 (-33.8%) | -6641 (-75.6%)
-3732 (-42.49%) | -8007 (-91.15%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 8784)
The Purple Emperor inhabits damp, mature, deciduous woods with clearings in them. These woods
often have different sorts of trees, stream valleys, and woodland rides. Male butterflies are often
seen near puddles on the road and on the dung of other animals. They are also attracted to strongly
smelling cheese. Various willows (Sax spp.) are used as foodplants, the female depositing her eggs
preferably at the top of the tree, on the upperside of leaves that do not get the sun. The half-grown
caterpillar spins itself a small cushion in the fork of a twig on which it hibernates. In the spring, it
resumes feeding on the buds and fresh young leaves. It pupates, suspended from the underside of a
leaf, and has one generation a yeat.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.85).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
sad 4000 0 edd 4000 0 add 4000 0 odd 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Apatura iris (Nymphalidae) 463
(B1)
SEDG
BAMBU
(A2)
oe
aS
Pa if
GRAS
(A1Fl)
464 Climatic Risk Atlas of European Butterflies
Kirinia roxelana (CRAMER, 1777) — Lattice Brown
.6—hCcLUwE Full dispersal No dispersal
fi SEDG 257 (21.45%) -459 (-38.31%)
177 (14.77%) -438 (-36.56%)
© Peter Ginzinger Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1198)
The Lattice Brown inhabits open, dry woodland and dry scrub. In the hottest part of the day, the
butterflies settle on tree trunks, hiding themselves away in the deep shade of bushes and thickets.
They are also sometimes seen on dry riverbeds. The female deposits her eggs in crevices in the bark
of trees and bushes. The caterpillars feed on grasses, such as meadow-grass (Poa spp.), quaking grass
(Briza spp.), brome (Bromus spp.), and foxtail (Adopecurus spp.). The Lattice Brown has one brood a
year. It hibernates as a caterpillar.
Present distribution can be explained by climatic variables to quite some extent (AUC = 0.91).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
‘1.6
sh
Qa =
=) 2
i
Pe Oe
ces
o= mn
=o
£5 8
Go. 4
of 6
D >
a a4 = a
ger
a |
ale
qe. =
2 o
w
é 5
= 4
o
0 2000 4000 2000 40000 2000 40000 2000 4000
Gdd Gdd Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
465
Kirinia roxelana (Nymphalidae)
2080
2050
466 Climatic Risk Atlas of European Butterflies
Pararge aegeria (LINNAEUS, 1758) — Speckled Wood
hc Full dispersal No dispersal
Be SEDG 111 (0.49%) -1693 (-7.4%)
BAMBU -359 (-1.57%) -2025 (-8.85%)
mas caer) | 2002 11.69
SEDG -686 (-3%) -3718 (-16.25%)
BAMBU | -2115 (-9.24%) | -5993 (-26.19%)
-3117 (-13.62%) -8080 (-35.31%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 22882)
The Speckled Wood 1s a typical woodland butterfly. The woods may be deciduous, coniferous or mixed,
and the butterfly occurs along woodland rides, in clearings and at wood edges. In the north, it occurs
mostly in quite extensive areas of woodland, but in the south a line of trees can be sufficient to support
a population. Each male claims his territory on a branch that gets the sun, projecting out of the crown
of the tree, and waits there for a female to fly along. The foodplants are grasses that grow in woodland
and damp grassland, such as fescues (Festuca spp.), false-bromes (Brachypodium spp.), meadow-grasses
(Poa spp.), cock’s-foot (Dactylus spp.), Purple Moor-grass (Modinea caeruka) and Hokus spp. The female
deposits her eggs singly on the blades of grass. Hibernation takes place as a caterpillar or pupa, deep
down in a tussock of grass. The Speckled Wood has two to three generations a year.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.65).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
=
sh
J
0
sal
Ow
Za
uo
“
a
tod 4000 0 edd 4000 0 oad 4000 0 Gdd 4000
Minimum
Swe
0.6
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Pararge aegeria (Nymphalidae)
SEDG
(B1)
467
GRAS
(A1Fl)
468 Climatic Risk Atlas of European Butterflies
Lastommata megera (LINNAEUS, 1767) — Wall Brown
a se
-2133 (-12.9%) | -4415 (-26.71%)
SEDG -3806 (-23.03%) | -5677 (-34.35%)
BAMBU_ | -6087 (-36.83%) | -8167 (-49.41%)
-7558 (-45.73%) | -9777 (-59.15%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 16529)
The Wall Brown inhabits many different sorts of grassland, natural grasslands, as well as not all too
intensively farmed land. The butterflies are mostly very active, the males very alert. They fly up when
disturbed from rocks or walls on which they often bask in the sun, and are therefore conspicuous.
The males have a broad scent-brand on their forewings. The female deposits her eggs on the blades
of many different grasses, including fescues (Festuca spp.), false-bromes (Brachypodium spp.), cock’s-
foot (Dactylus spp.), bromes (Bromus spp.), and Crested Dog’s-tail (Cynosurus aristatus). When half-
grown, the caterpillar hibernates in the litter layer, where it later, deep down, pupates. The Wall
Brown has two to three broods a year.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.75).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Swe
O46
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
0 2000 40000 2000 40000 2000 40000 2000 4000
Gd dl Gdd
Gdd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
469
Lasiommata megera (Nymphalidae)
2080
2050
470 Climatic Risk Atlas of European Butterflies
Lasiommata petropolitana (Fasrictvs, 1787) — Northern Wall Brown
hc Full dispersal No dispersal
SEDG -2647 (-28.91%) | -3215 (-35.11%)
-3026 (-33.05%) | -3477 (-37.98%)
ee -3135 (-34.24%) | -3656 (-39.93%)
SEDG -2440 (-26.65%) | -3621 (-39.55%)
BAMBU_ | -3879 (-42.37%) | -4711 (-51.45%)
-4594 (50.17%) | -5721 (-62.48%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 9156)
The Northern Wall Brown is found in woodland clearings and meadows in woodland, usually in
relatively warm places with large rockmasses. The butterflies are fond of basking in the sun on
warm rocks or on the ground. The eggs are laid on a number of grasses, including fescues (Festuca
spp.), small-reeds (Ca/amagrostis spp.), and cock’s-foot (Dactylus spp.). The Northern Wall Brown
hibernates either in the last larval instar, or as a pupa, and has one brood a year.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.84).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
tad 4000 0 edd 4000 0 ad 4000 0 odd 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
471
Lasiommata petropolitana (Nymphalidae)
2080
2050
472 Climatic Risk Atlas of European Butterflies
Lastommata maera (LINNAEUS, 1758) — Large Wall Brown
.—hlcLw Full dispersal No dispersal
Be SEDG -126 (-0.86%) -2562 (-17.45%)
-1019 (-6.94%) | -6194 (-42.18%)
-1106 (-7.53%) | -7698 (-52.42%)
© Rudi Verovnik Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 14684)
The Large Wall Brown occurs in quite different habitats. It is mainly found in warm, dry places near
rocks, and poor grassland, or on rough vegetation near woodland edges. However, it can also occur
on damp grassland and rough vegetation near wood margins. The butterflies need a lot of nectar,
often visiting purple or pink flowers of thistles and other nectar-rich plants. The female lays her eggs
on the blades of grasses that include Hokus spp., bents (Agrostis spp.), and small-reeds (Calamagrostis
spp.). The caterpillars hibernate when half-grown in a grass tussock, and pupate later deep down in
the vegetation. The Large Wall Brown has one brood a year in the north of its distribution range,
and two a year in the south.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.75).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
‘sad 4000 0 edd 4000 0 oad 4000 0 Gdd 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 0.2
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
473
Lasiommata maera (Nymphalidae)
2080
2050
474 Climatic Risk Atlas of European Butterflies
Lopinga achine (LINNAEUS, 1763) — Woodland Brown
6c Full dispersal No dispersal
Hee SEDG -645 (-9.66%) -2182 (-32.66%)
2968 (44.43%) | -893 (-13.37%)
4790 (71.71%) | -2501 (-37.44%)
3726 (55.78%) | -3777 (-56.54%)
© Rudi Verovnik Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 6680)
The Woodland Brown is fond of warm, open places in damp, deciduous or mixed woods with well-
developed shrub and herbaceous layers. The butterflies rarely visit flowers, preferring to feed on
honeydew, moisture on buds, and sap oozing from wounded trees. The males often settle on puddles
on the ground, while the females tend to stay in the very top of the trees. Eggs are laid on all sorts of
erasses, including fescues (Fesiwca spp.), meadow-grasses (Poa spp.), small-reeds (Calamagrostis spp.),
and false-bromes (Brachypodium spp.), and also on sedges (Carex spp.). The half-grown caterpillar
hibernates in a grass tussock, where later in the year it also pupates. The Woodland Brown has one
brood a yeat.
This species is listed in Annex IV of the Habitats’ Directive.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.81).
Climate risk category: R
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
i
=|
o : : - :
‘dd 4000 0 edd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe Swe
0.6 0.6
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Lopinga achine (Nymphalidae)
SEDG
GRAS
(B1)
(A1FI)
475
476 Climatic Risk Atlas of European Butterflies
Coenonympha tullia (MULLER, 1764) — Large Heath
SEDG -4213 (-32.25%) | -6226 (-47.66%)
BAMBU_ | -4915 (-37.63%) | -7441 (56.97%)
-6014 (-46.04%) | -8747 (-66.97%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 13062)
The Large Heath inhabits raised bogs, wet heaths, swampy habitat and wet grasslands, often
occurring on quite rough vegetation. As a consequence of land drainage remaining habitat patches
are often small, and populations they contain very small indeed, making them difficult to detect.
Main foodplants are Cotton-grass (Enophorum vaginatum), as well as other Eriophorum species. The
female deposits her eggs singly on, or in the neighbourhood of, the foodplant. The caterpillars
hibernate in the third or fourth larval instar, hidden in tussocks of grass. They also pupate in the
tussocks of the foodplant. The Large Heath has one generation a year. The species already seems to
have been affected by climate change and has declined severely at the southern edge of its range.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.83).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
mM
o
tod 4000 0 Codd 4000 0 add 4000 0 Gdd 4000
1.0
Minimum
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe Swe
0.6 0.6
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Coenonympha tulia (Nymphalidae) 477
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
478 Climatic Risk Atlas of European Butterflies
Coenonympha oedippus (Fasricius, 1787) — False Ringlet
iE Full dispersal No dispersal
fe SEDG (-0.47%) -269 (-25.5%)
1170 (110.9%) | -109 (-10.33%)
|GRAS | 168 (15.92%) -331 (-31.37%)
SEDG 732 (69.38%) -387 (-36.68%)
BAMBU_ | 2128 (201.71%) | -481 (-45.59%)
wae @riswry, | 713 a7
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1055)
The False Ringlet is one of the rarest butterflies of Europe and ts declining at an alarming rate. Seeing
one is therefore a very special event. The False Ringlet inhabits low-lying, grassy marshes and reed-
beds that are usually situated in the shelter of woodland, creating a warm and humid environment.
The butterflies fly very slowly and hardly ever migrate to nearby habitats. The eggs are deposited
one by one on the blades of grasses, like meadow-grasses (Poa spp.), rye-grasses (Lofum spp.), hair-
grasses (Deschampsia spp.) and Purple Moor-grass (Modinea caeruka). The caterpillars hibernate half-
grown in the tussock, where they pupate as well. The False Ringlet has one generation a year.
This species is listed in Annexes II and IV of the Habitats’ Directive.
Present distribution can be well explained by climatic variables (AUC = 0.95).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
a
~
oH
r=
a
=
Ow
=o
uw
m4
Oo
a : : : , : ; = '
4
o
=
cs
o
m4
o
‘dd 4000 0 edd 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
Swe
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Coenonympha oedippus (Nymphalidae) 479
480 Climatic Risk Atlas of European Butterflies
Coenonympha rhodopensis Ewes, 1900 — Eastern Large Heath
Cc Full dispersal No dispersal
Be SEDG 332 (43.74%) -514 (-67.72%)
BAMBU 125 (16.47%) -592 (-78%)
197 (25.96%) -603 (-79.45%)
SEDG 351 (46.25%) -606 (-79.84%)
BAMBU 432 (56.92%) -705 (-92.89%)
322 (42.42%) -737 (-97.1%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 759)
The Eastern Large Heath occurs mainly on grasslands above the tree-line. It is also sometimes
found in clearings in damp woodland. The caterpillars feed on fescues (Festuca spp.), Blue
Moor-grass (Sesleria albicans), cotton-grasses (Eriophorum spp.), White Beak-sedge (Rhynchospora
alba), and Beaked Sedge (Carex rostrata). This species has one generation a year and passes the
winter as a caterpillar.
Present distribution can be well explained by climatic variables (AUC = 0.95).
Climate risk category: HHHAR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
he
o
2
be
o
sad 4000 0 Codd 4000 0 add 4000 0 odd 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 O.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Coenonympha rhodopensis (Nymphalidae) 481
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
482 Climatic Risk Atlas of European Butterflies
Coenonympha arcania (LINNAEUS, 1761) — Pearly Heath
.6—hCcLUwk Full dispersal No dispersal
fe SEDG -389 (-2.87%) -2771 (-20.46%)
-1104 (-8.15%) | -3319 (-24.51%)
fens [as can [ area
-3556 (-26.26%) | -7474 (-55.18%)
-4819 (-35.58%) | -9350 (-69.03%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 13544)
The Pearly Heath is found on dry, to moderately damp, grasslands and grassy places at the edges
of woodland or scrub. The Pearly Heath is common in some areas. The males can often be found,
perched in scrub, basking in the sun, from where they chase females that pass by. In the evening, the
butterflies gather together to roost communally in scrub or at wood margins. Meadow-grasses (Poa
spp.), bents (4grosts spp.), melicks (Medca spp.), fescues (Festuca spp.), and many other grasses are
used as foodplants, the preferred species differing between regions. The female lays her eggs one by
one or in short rows, on blades of grass. When half-grown, the caterpillar hibernates 1n a tussock of
grass, also pupating there later, deep down in the tussock. This species has one brood a year.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.83).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘sad 4000 0 odd 4000 0 ad 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
0.6
Swe
o.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
483
Coenonympha arcania (Nymphalidae)
2080
2050
484 Climatic Risk Atlas of European Butterflies
Coenonympha glycerion (BORKHAUSEN, 1788) — Chestnut Heath
Cc Full dispersal No dispersal
SEDG 735 (5.73%) -2415 (-18.82%)
BAMBU -174 (-1.36%) -2688 (-20.94%)
90 (0.7%) -2849 (-22.2%)
SEDG 2334 (18.18%) | -3666 (-28.56%)
BAMBU 609 (4.74%) -5395 (-42.03%)
-1143 (-8.91%) | -7723 (-60.17%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 12835)
The Chestnut Heath inhabits dry to damp grasslands in woods, meadows, poor grassland, calcareous
grasslands, and open marshy habitats. These grasslands are sometimes quite intensively grazed, as
can happen on calcareous grassland. However, if grazing is absent, for a few years, change in the
grassland does not seem to affect the butterflies. The butterflies do not fly very much, and only
cover limited distances. The eggs are laid one by one in short rows on the blades of grasses, such as
fescues (Festuca spp.), Lor-grass (Brachypodium pinnatum), Purple Moor-grass (Moknea caerulea), Upright
Brome (Bromus erectus), and Crested Dog’s-tail (Cynosurus cristatus). Pupation takes place deep down in
the vegetation. The Chestnut Heath mostly has one or two generations a year, depending on altitude
and latitude.
Present distribution can be well explained by climatic variables (AUC = 0.86).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘sad 4000 0 Codd 4000 0 add 4000 0 aad 4000
1.0
Minimum
Swe
0.6
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
485
Coenonympha ghcerton (Nymphalidae)
2080
2050
486 Climatic Risk Atlas of European Butterflies
Coenonympha gardetta (PRUNNER, 1798) — Alpine Heath
.—hCcLe Full dispersal No dispersal
Be SEDG -167 (-17.75%) -312 (-33.16%)
-97 (-10.31%) -279 (-29.65%)
354 (37.6280
-213 (-22.64%) | -475 (-50.48%)
-466 (-49.52%) | -662 (-70.35%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 941)
The Alpine Heath is mostly found on open, sub-alpine and alpine grasslands, and on grasslands with
scattered bushes and trees. It can occur in high numbers in some places. The female lays her eggs,
mostly one by one, on grass stalks. The caterpillars hibernate, and pupate at the beginning of the
summer, the pupa hanging from a plant, usually close to the ground. It is single-brooded.
Present distribution can be very well explained by climatic variables (AUC = 0.97).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
1.0
Minimum
Swe
06
tH
a a
f= = -
6 2
i
ces
5=n
=o
£5 3 : ,
Ba. a : : ay u
os 7
©
aes
= wih
SR 4
seh
a
= =
2 o
iw
Sa
‘0
=
o2
edd 4000 0 odd 4000 0 Cad 4000 0 odd 4000
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
487
Coenonympha gardetta (Nymphalidae)
2080
2050
488 Climatic Risk Atlas of European Butterflies
Coenonympha dorus (EsPER, 1782) —- Dusky Heath
6c Full dispersal No dispersal
Be SEDG -508 (-18.27%) -1864 (-67.03%)
-856 (-30.78%) | -2128 (-76.52%)
-927 (-33.33%) | -2297 (-82.6%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2781)
The Dusky Heath is a butterfly of dry, grassy vegetation, such as found on rocky slopes in low scrub,
thickets and in woodland clearings. Different grasses are used as foodplant, including bent (Agrostis
spp.), and Sheep’s-fescue (Festwa ovina). It has one generation a year and hibernates as a caterpillar.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.83).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
=
ms
a
a
=
ow
=a
ie]
c
oa
3 i : f 7 :
tT!
o
2
Ow
$0
LP)
™
o
dd 4000 0 edd 4000 0 Cad 4000 0 Gd 4000
Minimum
Swe
06
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Coenonympha dorus (Nymphalidae) 489
490 Climatic Risk Atlas of European Butterflies
Coenonympha hero (LINNAEuvs, 1761) — Scarce Heath
.—hCcLwe Full dispersal No dispersal
SEDG -1416 (-39.97%) | -1929 (-54.45%)
BAMBU 941 (26.56%) -845 (-23.85%)
-882 (-24.89%) | -2331 (-65.79%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 3543)
The Scarce Heath occurs in damp to wet grassy meadows in or at the edges of woods. Sometimes,
they occur away from woods in drier places or in flower-rich grassland. The butterflies are fond
of settling in grass, and do not fly far, nor very often. Among the grasses they use as food are
Tufted Hair-grass (Deschampsia cespitosa), and Bearded Couch (EAmus caninus). When half-grown,
the caterpillar hibernates in a grass tussock, where it later also pupates. This species has one
generation a year. It should be noted that this species 1s severely declining. (Many dots on the map
actually represent already extinct populations.)
This species 1s listed in Annex IV of the Habitats’ Directive.
Present distribution can be well explained by climatic variables (AUC = 0.88).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
‘1.6
10 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
10 0.2
0.6
Maximum Large (66%) Small (33%)
Swe
0 2000 40000 2000 40000 2000 40000 2000 4000
Gdd Gdd Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Coenonympha hero (Nymphalidae) 491
492 Climatic Risk Atlas of European Butterflies
Coenonympha leander (Esper, 1784) — Russian Heath
CL Full dispersal No dispersal
| SEDG 225 (27.21%) | -425 (51.39%)
| BAMBU | -399 (-48.25%) | -597 (-72.19%)
g 574 (-69.41%) | -746 (-90.21%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 827)
The Russian Heath is found both on dry, as well as somewhat damp, grassy vegetation, at wood
margins and in woodland clearings and on grasslands. The butterflies have a rather low flight. At
rest, just like other heaths, the Russian Heath keeps its wings closed. Sheep’s-fescue (Festuca ovina)
and Slender False-brome (Brachypodium syhaticum) are among the grasses used as foodplants. This
species has one brood a year and hibernates as a caterpillar.
Present distribution can be well explained by climatic variables (AUC = 0.89).
Climate risk category: HHHAR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
‘0.6
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 02
0.6
Swe
0 2000 40000 2000 40000 2000 40000 2000 4000
Gdd Gdd Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
493
Coenonympha leander (Nymphalidae)
2080
2050
494 Climatic Risk Atlas of European Butterflies
Coenonympha pamphilus (Linnatus, 1758) — Small Heath
.—hCcLUwkE Full dispersal No dispersal
SEDG -1709 (-7.66%) -3476 (-15.58%)
BAMBU -2727 (-12.22%) | -3878 (-17.38%)
-2920 (-13.09%) | -4439 (-19.9%)
SEDG -1994 (-8.94%) | -5095 (-22.84%)
BAMBU_ | -4197 (-18.81%) | -7614 (-34.13%)
-6105 (-27.36%) | -10438 (-46.79%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 22310)
The Small Heath is a common species of nearly all types of grassland. It is mostly found on fairly
open, poor meadows and pastures. The female deposits her eggs one by one, or in rows, on the
blades of most grasses, including meadow-grasses (Poa spp.), Sweet Vernal-grass (Anthoxanthum
odoratum), fescues (Festuca spp.), and bents (Agrostis spp.). The caterpillar grows very quickly on
nutritious grasses, but can also use poor grasses, growing then more slowly. In the third or fourth
larval instar, the caterpillar hibernates deep down in a tussock of grass, where it also later pupates.
The number of broods a year is between one and three and depends on the geographical position
of the habitat.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.66).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
a
dd 4000 0 Codd 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Coenonympha pamphilus (Nymphalidae) 495
(B1)
SEDG
GRAS
(A1Fl)
496 Climatic Risk Atlas of European Butterflies
Pyronia tithonus (LINNAEUS, 1771) — Gatekeeper
.—hCcLUw Full dispersal No dispersal
fi SEDG -385 (-6.01%) -1451 (-22.63%)
-48 (-0.75%) -1437 (-22.41%)
-1165 (-18.17%) | -2649 (-41.32%)
-2622 (-40.9%) | -3899 (-60.82%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 6411)
The Gatekeeper occurs on many different types of grassland. Mostly, it chooses rather rough, dry
to damp vegetation, situated beside woodland or scrub, or not far from them. The butterflies are
fond of basking in the sun on scrub, and are often seen visiting flowers. The female lays her eggs
singly on the leaf-blades of nearly all soft grasses, such as Cock’s-foot (Dactylus spp.), fescues (Festuca
spp.), bents (Agrostis spp.), and Rye-Grass (Lokum spp.). The caterpillar avoids bright sunlight, and
feeds mostly on grasses growing 1n the shade. When half-grown, it hibernates in a tussock of grass.
It completes its growth the following year, and pupates low down in the vegetation. The Gatekeeper
is single-brooded.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.82).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
1.6
Swe
E 10 0.2 0.6 10 o2 fi. 1.0
oOo
= |
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 0.2
0.6
Swe
o2
O 2000 400 2 40000 2000 40000 2000 400)
Gdd Gdd Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
497
Pyronia tithonus (Nymphalidae)
2080
2050
498 Climatic Risk Atlas of European Butterflies
Pyronia cecilia (VALLANTIN, 1894) — Southern Gatekeeper
.—hCcLee Full dispersal No dispersal
fe SEDG -794 (-33.79%) -1382 (-58.81%)
-925 (-39.36%) | -1406 (-59.83%)
|GRAS | -939 (-39.96%) | -1613 (-68.64%)
SEDG -891 (-37.91%) | -1781 (-75.79%)
BAMBU_ | -1206 (-51.32%) | -2042 (-86.89%)
-1037 (-44.13%) | -2280 (-97.02%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2350)
The Southern Gatekeeper can be found on dry grasslands, rocky slopes with grassy vegetation, in
open scrub, and now and then in woodland clearings. The foodplants are grasses, probably Tufted
Hair-grass (Deschampsia cespitosa), although this is still uncertain. This butterfly species has one or two
broods a year, and passes the winter as a caterpillar.
Present distribution can be well explained by climatic variables (AUC = 0.91).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘dd 4000 0 odd 4000 0 Cad 4000 0 add 4000
1.0
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
499
Pyronia cecilia (Nymphalidae)
2080
2050
500 Climatic Risk Atlas of European Butterflies
Pyronia bathseba (Fasrictus, 1793) — Spanish Gatekeeper
6c Full dispersal No dispersal
He SEDG -696 (-20.11%) -1954 (-56.46%)
-1440 (-41.61%) -2070 (-59.81%)
-2178 (-62.93%) | -3150 (-91.01%)
-2079 (-60.07%) | -3404 (-98.35%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 3461)
The markings on the Spanish Gatekeeper are brighter than those of the other gatekeepers. The
butterflies are mostly found on dry, rather rough, grassy vegetation with trees or bushes in the
neighbourhood. The caterpillars feed on grasses, especially false-bromes (Brachypodium spp.), but
also probably bromes (Bromus spp), and meadow-grasses (Poa spp.). The Spanish Gatekeeper is
single-brooded, and passes the winter in the caterpillar stage.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.83).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
‘1.6
1.0 02
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 o2
0.6
Swe
6 1002 O86 o6 1.0
oOo
=
|
7 |
S
Swe
o2
i=]
2000 40000 2000 40000 2000 40000 2000 4000
Gdd
Gdd Gdd Gdd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
501
Pyronia bathseba (Nymphalidae)
2080
2050
502 Climatic Risk Atlas of European Butterflies
Aphantopus hyperantus (LINNAEUS, 1758) — Ringlet
CO Full dispersal No dispersal
SEDG -1583 (-9.49%) -3790 (-22.73%)
-1875 (-11.25%) | -4274 (-25.64%)
-2340 (-14.04%) | -4840 (-29.03%)
-7118 (-42.69%) | -12005 (-72.01%)
© Peter Ginzinger Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 17662)
The Ringlet can be found in grassy places with bushes, woodland clearings, and on grasslands
bordering woods. The habitats vary from dry to rather wet, such as at the edge of a raised bog.
The butterflies are avid visitors of such flowers as thistles, knapweed, and other plants rich in
nectar. The caterpillars feed on nutrient-rich grasses, such as cock’s-foot (Dactylus spp.), false-bromes
(Brachypodium spp.), fescues (Festuca spp.), and cat’s-tails (Phkum spp.). The female usually just releases
her eggs, letting them fall into the vegetation, either while perched or in flight. The caterpillars
hibernate, hidden in a grass tussock, and when they resume feeding, only do so at night. They also
pupate deep down in the vegetation. The Ringlet has one brood a year.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.84).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
sad 4000 0 ood 4000 0 oad 4000 0 Gad 4000
Minimum
Swe
06
1.0 02
0.6
Swe
1.0 0.2
Swe
6
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Abphantopus lyperantus (Nymphalidae) 503
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
504 Climatic Risk Atlas of European Butterflies
Maniola jurtina (LINNAEUus, 1758) — Meadow Brown
.6—hCcLUwkE Full dispersal No dispersal
Be SEDG -1330 (-6.1%) -3022 (-13.85%)
BAMBU_ | -2884 (-13.22%) | -4188 (-19.19%)
-2986 (-13.68%) | -4489 (-20.57%)
SEDG -4349 (-19.93%) | -6262 (-28.7%)
BAMBU_ | -6627 (-30.37%) | -9090 (-41.66%)
-9317 (-42.7%) | -11852 (-54.32%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 21820)
The Meadow Brown is a common butterfly of many different sorts of grassland, both natural
grasslands and not too intensively used farmland, and semi-natural grassland. The butterflies are
noticeable by their lively behaviour. The female deposits her eggs on a large number of grasses,
including species of fescue (Festuca spp.), false-brome (Brachypodium spp.), cock’s-foot (Dactylus spp.),
dog’s-tail (Cynosurus spp.), and brome (Bromus spp.). Sometimes, she also just drops the eggs in flight.
The caterpillar goes into hibernation 1n the litter layer when half-grown. It also pupates deep in the
litter layer. The Meadow Brown has one generation a year.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.7).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
a
oa
=]
ze
o
™
o
te 4000 0 ns 4000 0 oo 4000 0 ee 4000
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Maniola jurtina (Nymphalidae) 505
"a ae
r
| cr
(B1)
SEDG
GRAS
(A1Fl)
506 Climatic Risk Atlas of European Butterflies
Hyponephele lycaon (Kin, 1774) - Dusky Meadow Brown
.6—hlcLw Full dispersal No dispersal
SEDG -805 (-11.69%) -2069 (-30.06%)
BAMBU -3974 (-57.73%) | -4245 (-61.66%)
-2698 (-39.19%) | -3454 (50.17%)
SEDG -3582 (-52.03%) | -4792 (-69.61%)
BAMBU_ | _-5739 (-83.37%) | -6138 (-89.16%)
-5461 (-79.33%) | -6406 (-93.06%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 6884)
The Dusky Meadow Brown occurs in many different sorts of grassland. It is important that its habitat
is dry and warm. In the north of its range, the grassland is steppe-like, and more to the south the
butterfly 1s found on closer vegetation. It uses various grasses as foodplant, including fescues (Festuca
spp.), false-bromes (Brachypodium spp.), and Stipa species. When the female is about to lay an egg, she
first alights on the ground and walks to a foodplant, then she chooses a low-hanging grass blade on
which to deposit an egg. The caterpillar hibernates in the first larval instar and only starts feeding in
the spring. At first, it feeds mainly during day, but when the weather gets too warm, it only feeds at
night. Pupation takes place deep in the vegetation or litter layer. The Dusky Meadow Brown 1s single-
brooded.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.78).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
=
ms
r=
a
=
Ow
Za
i]
sir}
oO
: : : es : . :
Ow
$0
oo
~
o
dd 4000 0 edd 4000 0 ad 4000 0 odd 4000
Minimum
Swe
06
Annual precipitation range
Maximum Large (66%) Small (33%)
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
507
HAlyponephele hraon (Nymphalidae)
2080
2050
508 Climatic Risk Atlas of European Butterflies
Hyponephele lupina (Costa, 1836) — Oriental Meadow Brown
6c Full dispersal No dispersal
fe SEDG -389 (-13.75%) -1308 (-46.24%)
-1228 (-43.41%) | -1547 (-54.68%)
|GRAS | -981 (-34.68%) | -1690 (-59.74%)
SEDG -1383 (-48.89%) | -1845 (-65.22%)
BAMBU | -2198(-77.7%) | -2482 (-87.73%)
-2354 (-83.21%) | -2720 (-96.15%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2829)
The Oriental Meadow Brown is a butterfly of warm, dry places. The butterflies can be found on
dry grasslands with scattered bushes, or dry scrub vegetation, and in light woodland. At rest, their
wings are almost always closed. They have a rapid flight, often keeping to the shadow cast by trees or
bushes. Grasses, such as meadow-grasses (Poa spp.) and fescues (Fes/uca spp.) are used as foodplants.
The species has one generation a year and hibernates in the larval stage.
Present distribution can be well explained by climatic variables (AUC = 0.88).
Climate risk category: HHHAR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
=
sh
=]
o
=
Ow
Za
uo
pity
oa
= : : : 2 : :
he
o
2
w
o
aad 4000 0 Coded 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
509
Hyponephele lupina (Nymphalidae)
2080
2050
510 Climatic Risk Atlas of European Butterflies
Erebia ligea (Linnaeus, 1758) — Arran Brown
6c Full dispersal No dispersal
SEDG -2958 (-23.74%) | -3875 (-31.09%)
BAMBU -2415 (-19.38%) | -3187 (-25.57%)
-4075 (-32.7%) | -4737 (-38.01%)
SEDG -2798 (-22.45%) | -3895 (-31.26%)
BAMBU_ | _ -4435 (-35.59%) | -5388 (-43.24%)
-5485 (-44.01%) | -6418 (-51.5%)
© Martin Wiemers Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 12462)
The Arran Brown occurs in sheltered, grassy, flower-rich places in woodland. These places are
usually rather damp and in deep shade. The female deposits her eggs on the withered blades of
various grasses, including cock’s-foots (Dactylus spp.), hatr-grasses (Deschampsia spp.), fescues (Festuca
spp.), and Heath-grass (Danthonia decumbens). Sedges (Carex spp.) ate also used. The egg hibernates,
and the following year, the caterpillar feeds and grows, and in the last larval instar hibernates again.
In the summer, the caterpillar pupates. The Arran Brown 1s single-brooded, but the life cycle takes
two years to complete.
Present distribution can be well explained by climatic variables (AUC = 0.86).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
ie
=|
- . = : :
as
|
oad 4000 0 edd 4000 0 oad 4000 0 add 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
Swe
0.6
Annual precipitation range
Maximum Large (66%) Small (33%)
1.0 02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
511
Erebia lgea (Nymphalidae)
2080
2050
512 Climatic Risk Atlas of European Butterflies
Erebia euryale (Esrrr, 1805) — Large Ringlet
6c Full dispersal No dispersal
Be SEDG -524 (-25.05%) -825 (-39.44%)
BAMBU -435 (-20.79%) | -769 (36.76%)
-791 (-37.81%) | -1100 (-52.58%)
SEDG -326 (-15.58%) | -1147 (54.83%)
BAMBU | -275 (-13.15%) | -1218 (-58.22%)
-500 (-23.9%) | -1541 (-73.66%)
© Rudi Verovnik Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2092)
The Large Ringlet occurs in light woodland, in grassy clearings 1n woods, and above the tree-line
on grassland with rather tall vegetation. In the Jura Mountains, these butterflies can be found at the
edge of raised bogs, and at an altitude of 1000 m, also in light, damp woods. The Greek populations
also seem to prefer damp places. Both the males and females visit flowers for their nectar and the
males can often be seen drinking on damp ground or on dung. Various grasses are used as foodplant,
including Blue Moor-grass (Seskria albicans), \Wood Meadow-grass (Poa nemoralis), Red Fescue (Festuca
rubra), Sheep’s-fescue (F. ovina), Calamagrostis varia, and also sedges (Carex spp.). It takes two years for
the development from egg to butterfly.
Present distribution can be well explained by climatic variables (AUC = 0.89).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
‘sad 4000 0 edd 4000 0 add 4000 0 add 4000
Minimum
Swe
0.6
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
513
Erebia euryale (Nymphalidae)
2080
2050
514 Climatic Risk Atlas of European Butterflies
Erebia eriphyle (FRreYER, 1836) — Eriphyle Ringlet
.—ClcLw Full dispersal No dispersal
Be SEDG -57 (-14.81%) -144 (-37.4%)
BAMBU -33 (-8.57%) -141 (-36.62%)
-99 (-25.71%) -172 (-44.68%)
SEDG -168 (-43.64%) | -250 (-64.94%)
BAMBU -76 (-19.74%) -251 (-65.19%)
-121 (-31.43%) | -304 (-78.96%)
© Josef Pennerstorfer Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 385)
The Eriphyle Ringlet occurs locally in the Alps and 1s one of the smaller ringlets. It is a characteristic
species of the north-facing slopes, occurring on rough, herbaceous vegetation in the alpine and sub-
alpine zones, where there are plenty of alder (A/mus spp.) saplings. They are often found in places
where the conspicuous plants Adenostyks aliariae and Masterwort (Peucedanum ostruthium) are growing.
It uses Sweet-Vernal-grass (Anthoxanthum odoratum) and Tufted Hair-grass (Deschampsia cespitosa) as
foodplants. Some of the caterpillars leave the egg before the winter, but the rest hibernate in the
egg and emerge in the spring, when they all start feeding on grasses. In the autumn, they hibernate
again and moult twice before pupating at the end of May.
Present distribution can be very well explained by climatic variables (AUC = 0.99).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
‘dd 4000 0 edd 4000 0 oad 4000 0 odd 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
515
Erebia eriphyle (Nymphalidae)
(1a)
04s
(ZV)
Nanva
(ld)
SVH9
2080
2050
516 Climatic Risk Atlas of European Butterflies
Erebia manto ({SCHIFFERMULLER], 1775) — Yellow-spotted Ringlet
CL Full dispersal No dispersal
fe SEDG -258 (-19.07%) -466 (-34.44%)
ns cam | a9 can
|GRAS | -357 (-26.39%) -548 (-40.5%)
SEDG -368 (-27.2%) -668 (-49.37%)
BAMBU -228 (-16.85%) | -661 (-48.85%)
-402 (-29.71%) | -850 (-62.82%)
© Josef Pennerstorfer Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1353)
The Yellow-spotted Ringlet is a species of cool and cold conditions. Above the tree-line it occurs on
open mountain meadows, and below it inhabits damp, flower-rich grasslands and woodland glades.
The female deposits her eggs one at a time on the blades of various grasses, such as fescues ( Festuca
spp.) and cat’s-tails (Phkum spp.). It hibernates twice, the first time as an egg or a tiny caterpillar,
without having fed at all. During the next summer, the caterpillar develops as far as the penultimate
stage and then hibernates again. It pupates in the following summer.
Present distribution can be well explained by climatic variables (AUC = 0.9).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
fad 4000 0 odd 4000 0 add 4000 0 add 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
517
Erebia manto (Nymphalidae)
2080
2050
518 Climatic Risk Atlas of European Butterflies
Erebia epiphron (KNocu, 1783) — Mountain Ringlet
.—ClcLw Full dispersal No dispersal
fe SEDG -747 (-23.03%) -1068 (-32.93%)
-761 (-23.47%) | -1057 (-32.59%)
|GRAS | -980 (-30.22%) | -1321 (-40.73%)
SEDG 524 (-16.16%) | -1317 (-40.61%)
BAMBU_ | -717 (-22.11%) | -1526 (-47.06%)
-679 (-20.94%) | -1837 (-56.65%)
© Rudi Verovnik Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 3243)
The Mountain Ringlet inhabits mountainous areas that are snow-covered in winter. It is found on
damp grasslands and rough vegetation, often in the shelter of a slope or woodland edge. The female
lays her eggs on various grasses and sedges, including fescues (Festuca spp.), Mat-grass (Nardus stricta),
and Heath-grass (Danthonia decumbens). The caterpillar hibernates twice. It completes the first larval
instar before hibernating the first tume. By the end of the summer it reaches the last but one stage
before hibernating again. The following year, it pupates in the summer, and the butterfly emerges. The
Mountain Rinelet is a very variable butterfly with many local and regional subspecies and forms.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.84).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘sad 4000 0 edd 4000 0 add 4000 0 add 4000)
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
519
Erebia epiphron (Nymphalidae)
2080
2050
520 Climatic Risk Atlas of European Butterflies
Erebia pharte (HUBNER, 1804) — Blind Ringlet
CL Full dispersal No dispersal
fe SEDG -284 (-20.94%) -487 (-35.91%)
-212 (-15.63%) | -429 (-31.64%)
|GRAS | -337 (-24.85%) | -542 (-39.97%)
SEDG -428 (-31.56%) | -679 (-50.07%)
BAMBU_ | -269 (-19.84%) | -666 (-49.12%)
-426 (-31.42%) | -832 (-61.36%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1356)
This Ringlet owes its rather odd name to the absence of eye-spots on both surfaces of the wings. The
Blind Ringlet occurs both on dry and damp grasslands, especially where the vegetation is tall, and
in grassy places at the edge of woodland. Above the tree-line, they occur mainly on dry vegetation.
They have various foodplants, depending on the habitat. In dry places, the caterpillars can be found
on Mat-grass (Nardus stricta), Festuca quadnflora, and Sheep’s Fescue (F ovina), and in damper places
in the shade on Red Fescue (F rubra), Glaucous Sedge (Carex flacca), and C. ferruginea. The caterpillar
hibernates twice, pupating at the end of May between spun grass stalks.
Present distribution can be well explained by climatic variables (AUC = 0.89).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
fea 4000 0 Cored 4000 0 oad 4000 0 add 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 60.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Erebia pharte (Nymphalidae)
SEDG
BAMBU
GRAS
(B1)
(A2)
(A1Fl)
521
522 Climatic Risk Atlas of European Butterflies
Erebia melampus (FuEss.i, 1775) — Lesser Mountain Ringlet
.—ClcLee Full dispersal No dispersal
Be SEDG -333 (-23.43%) -444 (-31.25%)
BAMBU -244 (17.17%) | -389 (-27.38%)
-413 (-29.06%) | -520 (-36.59%)
SEDG -499 (-35.12%) | -740 (52.08%)
BAMBU -379 (-26.67%) | -686 (-48.28%)
588 (-41.38%) | -917 (-64.53%)
© Martin Wiemers Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1421)
The Lesser Mountain Ringlet occurs in many different biotopes. It can be found on wet grasslands,
pastures and meadows, on dry, nutrient-poor grasslands in light woodland, on dry grasslands with
bracken, and above the tree-line on alpine grasslands. Because of the wide choice in biotope, this
butterfly is common throughout the Alps. In the time up to hibernation, the caterpillars feed during
the day. After hibernating, they are only active at night. They can be found on Wood Meadow-grass
(Poa nemorahs), Sweet Vernal-grass (Anthoxanthum odoratum), and Sheep’s Fescue (Festuca ovina). The
development from egg to butterfly takes one year.
Present distribution can be very well explained by climatic variables (AUC = 0.98).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘edd 4000 0 edd 4000 0 add 4000 0 add 4000
1.0
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
523
Erebia melampus (Nymphalidae)
2080
2050
524 Climatic Risk Atlas of European Butterflies
Erebia aethiops (EsPer, 1777) — Scotch Argus
6—hlcLUw Full dispersal No dispersal
Be SEDG -968 (-21.4%) -1883 (-41.63%)
BAMBU | -519 (-11.47%) | -1542 (-34.09%)
7395 cane) | 2206 922%
SEDG 491 (10.86%) | -1899 (-41.99%)
BAMBU 316 (6.99%) -2318 (-51.25%)
548 (12.12%) | -2876 (-63.59%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 4523)
The Scotch Argus can be found in a wide variety of habitats. It inhabits flower-rich grasslands and
grassy woodland clearings, and can also be found on heathland, bogs and marshy areas with very
open woodland. Many different grasses can be used as a foodplant, such as Purple Moor-grass
(Molinea caerulea), Blue Moor-grass (Seslria albicans), Tor-grass (Brachypodium pinnatum), Cock’s-foot
(Dactylus glomerata), fescues (Festuca spp.), cat’s-tails (Phkum spp.), Sweet Vernal-grass (Anthoxanthum
odoratum), and Quaking Grass (Briza media), and also sedges (Carex spp.). The caterpillar hibernates
in the second or third larval instar. It pupates in the litter layer, and is single-brooded.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.82).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
‘dd 4000 0 edd 4000 0 oad 4000 0 odd 4000
Minimum
Swe
06
1.0 02
0.6
Swe
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
525
Erebia aethiops (Nymphalidae)
2080
2050
526 Climatic Risk Atlas of European Butterflies
Erebia triaria (PRUNNER, 1798) — de Prunnet’s Ringlet
CL Full dispersal No dispersal
SEDG 70 (5.99%) -590 (-50.47%)
BAMBU 7 (0.6%) -608 (-52.01%)
-66 (-5.65%) -690 (-59.02%)
-255 (-21.81%) | -877 (-75.02%)
© Bernard Fransen Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1169)
In the Alps, de Prunner’s Ringlet is found on very dry grasslands with scattered rocks, whereas
more to the south it occurs in clearings in woods growing on rocky ground, in Spain in clearings in
pine forests on limestone. Its foodplants are Sheep’s-fescue (Festuca orina), Smooth Meadow-grass
(Poa pratensis), Alpine Meadow-grass (P. a/pina), and Feather-grass (Supa pinnata). \t hibernates as a
caterpillar and has one brood a year.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.81).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
=
sh
ra
o
=
Ow
a]
uw
sir
ao
a
‘edd 4000 0 Codd 4000 0 add 4000 0 add 4000
Minimum
Swe
06
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
SwG
08
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
527
Erebia triarna (Nymphalidae)
ia
| |
ara
alysis
(1a)
504s
Wo
» am
(ZV)
Nanva
(4b)
SVH9
2080
2050
528 Climatic Risk Atlas of European Butterflies
Erebia embla (BEcKLIN, 1791) — Lapland Ringlet
.—hClcL Full dispersal No dispersal
Be SEDG -835 (-21.95%) -1141 (-29.99%)
BAMBU -782 (-20.56%) | -1145 (-30.1%)
-806 (-21.19%) | -1207 (-31.73%)
SEDG -1583 (-41.61%) | -1864 (-49%)
BAMBU_ | -1605 (-42.19%) | -2010 (52.84%)
-2238 (-58.83%) -2592 (-68.14%)
© Jostein Engdal Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 3804)
The Lapland Rineglet inhabits bogs and marshes with sedges and grasses, and scattered groups of
willows and myrtle. The marshes are often situated in coniferous or birch woods. Its foodplants
are probably sedges (Carex spp.), Tufted Hatr-grass (Deschampsia cespitosa), and Bog Hait-grass (D.
selacea). These butterflies are difficult to find because they are shy and always occur in low numbers.
Also, their habitat is very inaccessible. The development of the Lapland Ringlet takes two years.
Present distribution can be very well explained by climatic variables (AUC = 0.97).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
aad 4000 0 ood 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 O.2
Annual precipitation range
Swe
0.6
1.0 O02
0.6
Maximum Large (66%) Small (33%)
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
529
Erebia embla (Nymphalidae)
2080
2050
530 Climatic Risk Atlas of European Butterflies
Erebia disa (BECKLIN, 1791) — Arctic Ringlet
.—CcLEe Full dispersal No dispersal
Be SEDG -1068 (-72.41%) -1242 (-84.2%)
BAMBU -956 (-64.81%) | -1166 (-79.05%)
-1025 (-69.49%) | -1249 (-84.68%)
SEDG -1285 (-87.12%) | -1446 (-98.03%)
BAMBU_ | -1347 (-91.32%) | -1470 (-99.66%)
rats 952% | 472 (990%
© Jostein Engdal Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1475)
Just like the Lapland Rineglet (E. emb/a), the Arctic Ringlet is a marsh butterfly, but the two species do
not overlap, the Arctic Ringlet avoiding lowland marshes. It occurs above 350 m altitude, on open
marshes in woods, namely marshes with a low cover of grasses and sedges without trees or bushes,
although surrounded by birch trees. It also occurs at lakesides and near small streams and is also
seen in woodland clearings in its search for nectar. Its foodplants are grasses. It takes two years for
the egg to develop into a butterfly.
Present distribution can be very well explained by climatic variables (AUC = 0.99).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 edd 4000 0 add 4000 0 add 4000
1.0
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 o.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
2050
Erebia disa (Nymphalidae)
2080
SEDG
531
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
532 Climatic Risk Atlas of European Butterflies
Erebia medusa (Fasrictus, 1787) — Woodland Ringlet
.6—hCcLUwkE Full dispersal No dispersal
fe SEDG -943 (-22.87%) -1907 (-46.24%)
-151 (-3.66%) | -1635 (-39.65%)
GRAS | -1416 (-34.34%) | -2279 (-55.26%)
SEDG 304 (7.37%) -2199 (-53.32%)
BAMBU 106 (2.57%) -2660 (-64.5%)
-111 (-2.69%) | -3288 (-79.73%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 4124)
The Woodland Ringlet occurs in many different biotopes. It can be found on damp, flower-rich
grasslands and rough vegetation near or in woodland, in marshes, but also on calcareous grasslands.
Its foodplants are various grasses, such as Purple Moor-grass (Modnea caerulea), fescues (Festuca spp.),
Tor-grass (Brachypodium pinnatum), Upright Brome (Bromus erectus), and Wood Millet (Mikum effusum),
and also sedges (Carex spp.). The caterpillar is active at night and mostly hibernates when half-grown,
but at very high altitudes it may hibernate twice. It pupates in the litter layer. The development takes
between one and two years.
Present distribution can be well explained by climatic variables (AUC = 0.9).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 Cried 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
0.6
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
533
Erebia medusa (Nymphalidae)
2080
2050
534 Climatic Risk Atlas of European Butterflies
Erebia alberganus (PRUNNER, 1798) — Almond-eyed Ringlet
6c Full dispersal No dispersal
fe SEDG -208 (-20.29%) -328 (-32%)
35.02% | 258 (257%
|GRAS | -244 (-23.8%) -352 (-34.34%)
SEDG -427 (-41.66%) | -585 (57.07%)
BAMBU_ | -276 (-26.93%) | -483 (-47.12%)
513 (-50.05%) | -685 (-66.83%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1025)
The Almond-eyed Ringlet is found on sub-alpine grasslands, and on, sometimes damp, grasslands,
in woodland clearings or in the shelter of woods. The butterflies can often be seen on flowers,
drinking nectar. On warm days, the males congregate to drink on damp patches. In the Alps, the
caterpillars feed on Sheep’s Fescue (Festuca ovina) and Sweet Vernal-grass (Anthoxanthum odoratum).
They hibernate when half-grown, and pupate at the end of the following spring.
Present distribution can be well explained by climatic variables (AUC = 0.87).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
fad 4000 0 odd 4000 0 add 4000 0 add 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 O.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
535
Erebia alberganus (Nymphalidae)
2080
2050
536 Climatic Risk Atlas of European Butterflies
Erebia pluto (PRUNNER, 1798) — Sooty Ringlet
SEDG -189 (-21.4%) -306 (-34.65%)
-145 (-16.42%) | -274 (-31.03%)
-239 (-27.07%) | -353 (-39.98%)
SEDG -437 (-49.49%) | -597 (-67.61%)
BAMBU_ | -285 (-32.28%) | -467 (-52.89%)
-468 (53%) -648 (-73.39%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 883)
The Sooty Ringlet is a butterfly of mountainous areas with everlasting snows and glaciers. It occurs
high up, on steep screes and moraines, breeding on patches with a mosaic of sparse vegetation and
small stones. The female deposits the pale-coloured eggs on mostly light-coloured stones. Once the
caterpillar has emerged from the egg, it sometimes has to travel some distance before finding one
of the grasses it uses as a foodplant. Caterpillars have been found on the fescues Festuca halleri and
F. guadriflora, as well as on Annual Meadow-grass (Poa annua). The caterpillars take two or possibly
three years to develop.
Present distribution can be very well explained by climatic variables (AUC = 0.98).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘sad 4000 0 edd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
2050
Erebia pluto (Nymphalidae)
2080
SEDG
537
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
538 Climatic Risk Atlas of European Butterflies
Erebia gorge (EsPrr, 1805) — Silky Ringlet
ae
a A .—hlcLw Full dispersal No dispersal
He SEDG -452 (-26.49%) -578 (-33.88%)
-375 (-21.98%) | -538 (-31.54%)
-602 (-35.29%) | -717 (-42.03%)
SEDG -587 (-34.41%) | -884 (51.82%)
BAMBU -534 (-31.3%) -900 (-52.75%)
GRAS -721 (-42.26%) | -1128 (-66.12%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1706)
The Silky Ringlet can be found on screes, sunny, rocky slopes, and dry grasslands with scattered
rocks. The butterflies often bask in the sun on stones, but, their wings closed, they are hardly
noticeable. They are rarely seen visiting flowers. The caterpillars are found on fescues (Fes/uca spp.),
Alpine Meadow-grass (Poa alpina), P. minor, and Blue Moor-grass (Sesleria albicans). The caterpillars
take two years to develop.
Present distribution can be well explained by climatic variables (AUC = 0.88).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Gadd 4000 0 odd 4000 0 oad 4000 0 aad 4000
1.0
Minimum
Swe
0.6
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
539
Erebia gorge (Nymphalidae)
2080
2050
540 Climatic Risk Atlas of European Butterflies
Erebia mnestra (Esper, 1805) — Mnestra’s Ringlet
.—hClcLw Full dispersal No dispersal
Be SEDG 72 (25.71%) -119 (-42.5%)
-169 (-60.36%) | -216 (-77.14%)
© Martin Wiemers Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 280)
Mnestra’s Ringlet occurs on sub-alpine and alpine grasslands, especially dry grasslands on steep,
sunny slopes. The butterflies are seldom seen on flowers, drinking nectar. The female lays her eggs
close to the ground on dry stalks of grass. The plain green caterpillars have been found on fescues
(Festuca spp.) and also on Blue Moor-grass (Ses/eria albicans). They hibernate twice.
Present distribution can be very well explained by climatic variables (AUC = 0.96).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
BEE
Ow
=o
on
a]
a
fad 4000 0 odd 4000 0 oad 4000 0 ada 4000
Minimum
Swe
0.6
1.0 O02
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
2050
Erebia mnestra (Nymphalidae)
2080
SEDG
541
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
542 Climatic Risk Atlas of European Butterflies
Erebia epistygne (HUBNER, 1819) — Spring Ringlet
.—hlcLUwe Full dispersal No dispersal
Be SEDG -137 (-79.65%) -162 (-94.19%)
-165 (-95.93%) | -170 (-98.84%)
-166 (-96.51%) -172 (-100%)
© Bernard Fransen Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 172)
The Spring Ringlet appears in the early spring in grassy, rocky clearings in light woodland. The
Spanish populations in the Montes Untversales occur in clearings or on level ground in light
pinewoods on calcareous soil, on short, grassy vegetation with low shrubs and scattered rocks. ‘The
main foodplant is Sheep’s-fescue (Festuca orina), but other fescues and meadow-grasses (Poa spp.)
have also been named as foodplants. FE. epzstygne has one generation per yeat.
Present distribution can be well explained by climatic variables (AUC = 0.91).
Climate risk category: HHHAR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
=
ms
=]
a
=
Ow
=o
uo
cu
oa
a kt f
dd 4000 0 odd 4000 0 oad 4000 0 add 4000
Minimum
Swe
06
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.8
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
2050
Erebia epistygne (Nymphalidae)
2080
SEDG
543
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
544 Climatic Risk Atlas of European Butterflies
Erebia ottomana (HERRICH-SCHAFFER, 1847) —
Ottoman Brassy Ringlet
CCL Full dispersal No dispersal
is SEDG -69 (-10.94%) -240 (-38.03%)
|GRAS | -95 (-15.06%) -304 (-48.18%)
193 (30.59%) -212 (-33.6%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 631)
In the mountains, the Ottoman Brassy Ringlet occurs on slopes and level ground with grassy
vegetation. At lower altitudes, it is found in woodland clearings. The caterpillars feed on Sheep’s-
fescue (Festuca ovina) and other fine-leaved grasses. It is single-brooded.
Present distribution can be explained by climatic variables to quite some extent (AUC = 0.81).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 odd 4000 0 add 4000 0 odd 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
o2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Erebia ottomana (Nymphalidae) 545
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
546 Climatic Risk Atlas of European Butterflies
Erebia tyndarus (Esper, 1781) — Swiss Brassy Ringlet
CL Full dispersal No dispersal
fi SEDG -24 (-5.52%) -205 (-47.13%)
BAMBU 51 (11.72%) -160 (-36.78%)
125i) | 20a C4099
SEDG -251 (57.7%) -357 (-82.07%)
BAMBU -80 (-18.39%) -272 (-62.53%)
-200 (-45.98%) | -339 (-77.93%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 435)
The Swiss Brassy Ringlet occurs in woodland clearings on grassy, rocky slopes, and on screes. The
caterpillars feed on different fescues (Pes/uca spp.) and on Mat-grass (Nardus stricta). The caterpillars
hibernate when they are still small. They develop further the following spring, pupating sometime
between June and August.
Present distribution can be very well explained by climatic variables (AUC = 0.99).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
aad 4000 0 Gadd 4000 0 Cad 4000 0 add 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 O.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
2050
Erebia tyndarus (Nymphalidae)
2080
SEDG
547
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
548 Climatic Risk Atlas of European Butterflies
Erebia cassioides (REINER & HOHENWARTH, 1792) (complex) —
Common Brassy Ringlet
© Kars Veling
SEDG -337 (-28.66%) -404 (-34.35%)
-320 (-27.21%) -378 (-32.14%)
-457 (-38.86%)
SEDG -656 (-55.78%) | -746 (-63.44%)
BAMBU -525 (-44.64%) | -630 (-53.57%)
-647 (-55.02%) | -747 (-63.52%)
Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1176)
The Common Brassy Ringlet encompasses 4 taxa which are included in this species complex: arvernensis,
casstoides, macedonica, carmenta. Vhey represent isolates in different glacial refugia. Their separate species
status however is still uncertain and requires verification. The butterflies live on dry, grassy slopes,
rocky slopes with patches of grassy vegetation, and scree. The females lay their eggs on dry stalks of
grass, close to the ground. The species’ foodplants are various fescues (Fes/uca spp.). The caterpillars
hibernate in the first or second larval instar and pupate the following year between June and August.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.8).
Climate risk category: R.
Observed species distribution (50 x 50 km? UTM grid; black
circles) and modelled actual distribution of climatic niche
(orange areas)
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
faaid 4000 0 odd 4000 0 oad 4000 0 add 4000)
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 60.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O2
Swe
0.6
0.2
Multidimensional climatic niche. Occurrence probability defined by
accumulated growing degree days until August (Gdd) and soil water
content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
549
Erebia cassioides (Nymphalidae)
2080
2050
550 Climatic Risk Atlas of European Butterflies
Erebia pronoe aa 1780) — Water Ringlet
a
fe SEDG -407 (-24.23%) -570 (-33.93%)
25 cay | aT2 cana
-586 (-34.88%) | -692 (-41.19%)
SEDG -315 (-18.75%) | -721 (-42.92%)
BAMBU -319 (-18.99%) | -774 (-46.07%)
-466 (-27.74%) | -1011 (-60.18%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1680)
The Water Ringlet is found especially on flower-rich grasslands, rocky, grassy slopes, and near
woodland. The female usually deposits her eggs close to the ground, on dry stalks of grass. The
caterpillars feed on Sheep’s Fescue (Festuca ovina), EF: quadriflora and vatious meadow-grasses (Pod spp.).
The caterpillars hibernate in the first larval instar and pupate in June or July of the following year.
Present distribution can be well explained by climatic variables (AUC = 0.89).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
todd 4000 0 odd 4000 0 oad 4000 0 add 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
551
Erebia pronoe (Nymphalidae)
2080
2050
552 Climatic Risk Atlas of European Butterflies
Erebia styx (FREYER, 1834) — Stygian Ringlet
.—ClcLe Full dispersal No dispersal
Be SEDG -200 (-46.62%) -227 (-52.91%)
BAMBU -186 (-43.36%) | -206 (-48.02%)
-255 (-59.44%) | -264 (-61.54%)
SEDG -247 (-57.58%) | -282 (-65.73%)
BAMBU_ | -242 (-56.41%) | -280 (-65.27%)
-287 (-66.9%) -361 (-84.15%)
© Martin Wiemers Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 429)
The Stygian Ringlet occurs on warm, steep, rocky slopes with grassy vegetation and sometimes
scattered bushes, often on limestone. Its foodplant is Blue Moor-grass (Seslra albwans). In
Switzerland, it takes two years for the egg to develop into a butterfly. The tiny caterpillar goes into
hibernation immediately after leaving the egg, only starting to feed the next spring. In the autumn,
they again hibernate, pupating in June or July of the following year. In other areas at lower altitudes,
they can complete their life cycle 1n one year.
Present distribution can be well explained by climatic variables (AUC = 0.95).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘said 4000 0 Codd 4000 0 Gad 4000 0 ada 4000
1.0
Minimum
Swe
06
o2
Swe
0.6 1.0
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
553
Erebia styx (Nymphalidae)
(1a)
504s
(ZV)
nNanva
(ld bY)
SVH9
2080
2050
554 Climatic Risk Atlas of European Butterflies
Erebia montana (PRUNNER, 1798) — Marbled Ringlet
CL Full dispersal No dispersal
Hi SEDG -77 (-12.13%) -282 (-44.41%)
BAMBU 25 (3.94%) -224 (-35.28%)
-112 (-17.64%) | -303 (-47.72%)
SEDG -349 (54.96%) | -530 (-83.46%)
BAMBU_ | -186 (-29.29%) | -409 (-64.41%)
-332 (-52.28%) -512 (-80.63%)
© Neil Thompson Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 635)
The Marbled Ringlet is mainly found on warm, rocky slopes with patches of grassy vegetation
in flower-rich grasslands, and occasionally in woods, for the most part on limestone. When the
butterflies settle on stones or on the ground with their wings closed, they are hardly noticeable.
However, they can be seen visiting flowers, which they do regularly. The caterpillars hibernate in the
first larval instar and in the spring begin feeding on fescues (Festwca spp.), or on Mat-grass (Nardus
stricta). Vhe caterpillars have been found on the larger tussocks growing in the shelter of rocks. They
pupate in June or July.
Present distribution can be very well explained by climatic variables (AUC = 0.98).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
‘dd 4000 0 edd 4000 0 add 4000 0 ada 4000
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
555
Erebia montana (Nymphalidae)
(ZV)
Nanva
(ld)
SVH9
2080
2050
556 Climatic Risk Atlas of European Butterflies
Erebia neoridas (BotsDUVAL, 1828) — Autumn Ringlet
CL Full dispersal No dispersal
SEDG -10 (-1.48%) -473 (-69.87%)
BAMBU -1 (-0.15%) -488 (-72.08%)
219 (32.35%) -560 (-82.72%)
© Matt Rowlings Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 677)
The Autumn Ringlet occurs in woodland clearings and on grassy vegetation with scattered bushes.
The caterpillars feed on different grasses, such as Crab-grass (Digilaria sanguinalis), Alpine Meadow-
Grass (Poa alpina), Smooth Meadow-grass (Poa pratensis), Annual Meadow-grass (P. annua), Sheep’s-
fescue (Festuca ovina), and Meadow Fescue (FP. pratensis). This species is single-brooded and hibernates
as a caterpillar.
Present distribution can be well explained by climatic variables (AUC = 0.93).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
Ey
wo
=
i
tl
= a
o
= aa
m
re)
Ges
a?
& 3
= -
fim
wo
© ow
o
fo
iomnill oa
=
c 7
5
Es
fa
=
oO
‘dd 4000 0 edd 4000 0 oad 4000 0 odd 4000
Annual precipitation range
Swe
0.6
Swe
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
557
Erebia neoridas (Nymphalidae)
(1a)
504s
(ZV)
Nanva
(4b)
SVH9
2080
2050
558 Climatic Risk Atlas of European Butterflies
Erebia melas (HERBST, 1796) — Black Ringlet
CL Full dispersal No dispersal
Be SEDG 179 (27.54%) -158 (-24.31%)
121 (18.62%) -231 (-35.54%)
© Rudi Verovnik Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 650)
The Black Ringlet can be found on rocky slopes with patches of grassy vegetation, in dry and sub-
alpine and alpine grasslands, and sometimes in clearings in coniferous woods. The butterflies fly
close to the ground, often resting with wings widespread. The caterpillars feed on Sheeps’-fescue
(Festuca ovina), and possibly other grasses. The Black Ringlet has one generation a year and hibernates
in the larval stage.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.84).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
a
-
ms
a
a
=
Ow
Za
wo
sir
Oo
3 : ; ; ;
tH
o
2
Ow
so
LP)
™
o
‘dd 4000 0 odd 4000 0 add 4000 0 add 4000
Minimum
Swe
06
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
559
Erebia melas (Nymphalidae)
2080
2050
560 Climatic Risk Atlas of European Butterflies
Erebia oeme (Esrer, 1805) — Bright-eyed Ringlet
CL Full dispersal No dispersal
Be SEDG -195 (-17.81%) -357 (-32.6%)
BAMBU -32, (-2.92%) -276 (-25.21%)
-298 (-27.21%) -427 (-39%)
SEDG -260 (-23.74%) | -538 (-49.13%)
BAMBU | -155(-14.16%) | -523 (-47.76%)
-413 (-37.72%) | -732 (-66.85%)
© Rudi Verovnik Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1095)
The Bright-eyed Ringlet is often found on damp to wet grasslands, sometimes with lots of sedges
in the vegetation. However, it also occurs on dry grasslands, rocky slopes and in woodland clearings.
The grass in some of these habitats may be quite tall. Several different foodplants are known,
including sedges (Carex spp.), rushes (Juncus spp.), Purple Moor-grass (Moknea caerulea), Quaking
Grass (Briza media), Red Fescue (Festuca rubra), and meadow-grasses (Poa spp.). The caterpillars take
one or two years to develop, depending on the altitude of the breeding ground.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.89).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘aad 4000 0 edd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
561
Erebia oeme (Nymphalidae)
(ZV)
Nanva
(ld LV)
SVH9
2080
2050
562 Climatic Risk Atlas of European Butterflies
Erebia meolans (PRUNNER, 1798) — Piedmont Ringlet
.—hcLw Full dispersal No dispersal
Be SEDG -89 (-3.23%) -1130 (-40.97%)
;
BAMBU -153 (5.55%) | -1103 (-39.99%)
-342 (12.4%) | -1389 (-50.36%)
SEDG -360 (-13.05%) | -1687 (-61.17%)
BAMBU -644 (-23.35%) | -1813 (-65.74%)
-1145 (-41.52%) | -2198 (-79.7%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2758)
a
Z
=
a
+
»
“3
The Piedmont Ringlet occurs on a variety of grassy places in and near woods. The female lays
her eges on many different grasses, including fescues (Festuca spp.), Wavy Hair-grass (Deschampsia
flexuosa), Mat-grass (Nardus stricta), and several sorts of bent (4grosts spp.). At very high altitudes,
when the caterpillars are half-grown, they hibernate in the litter layer. They may hibernate twice
before pupating deep down in the vegetation.
Present distribution can be well explained by climatic variables (AUC = 0.92).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
tad 4000 0 odd 4000 0 Cad 4000 0 Gdd 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
563
Erebia meolans (Nymphalidae)
2080
2050
564 Climatic Risk Atlas of European Butterflies
Erebia pandrose (BORKHAUSEN, 1788) — Dewy Ringlet
.6—hCcLwkE Full dispersal No dispersal
fi SEDG -950 (-21.14%) -1006 (-22.39%)
-779 (-17.34%) | -830 (-18.47%)
-1128 (-25.11%) | -1265 (-28.15%)
© Rudi Verovnik Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 4493)
The Dewy Ringlet is a common species. In Scandinavia, it is mostly found in damp places where
grass and bushes are growing, often near small streams, and in more mountainous areas on grassy
slopes. In Central Europe, the butterflies occur on stony, alpine meadows with a vegetation of
short grasses and dwarf shrubs. These butterflies have a characteristic, undulating flight. This gives
the impression that they are not using their hindwings, and are just about to fall to the ground.
The female lays her eggs on different grasses, such as fescues (Festuca spp.), meadow-grasses (Poa
spp.), Blue Moor-grass (Sesleria albicans), ot Mat-grass (Nardus stricta). In most parts of its range, the
caterpillars hibernate twice.
Present distribution can be well explained by climatic variables (AUC = 0.86).
Climate risk category: LR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
06
sh
a a
=) a
>S 7
Pm On
ae |
o= mH
=e |
£5 3
Go. 4
of 7
D>
o ng = a
Ba”
i=]
Qe
eo
)
* é Po]
= 4
o
O 2000 40000 2000 40000 2000 40000 2000 4000
Gd id Gadd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
565
Erebia pandrose (Nymphalidae)
2080
2050
566 Climatic Risk Atlas of European Butterflies
Melanargia russiae (Esrrr, 1784) — Espetr’s Marbled White
6c Full dispersal No dispersal
Be SEDG -781 (-51.55%) -1158 (-76.44%)
BAMBU_ | -988 (-65.21%) | -1169 (-77.16%)
-982 (-64.82%) | -1275 (-84.16%)
SEDG -752 (-49.64%) | -1352 (-89.24%)
BAMBU_ | -1026 (-67.72%) | -1441 (-95.12%)
-752 (-49.64%) | -1504 (-99.27%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1515)
Esper’s Marbled White is a butterfly of grassy vegetation, occurring both on acid and calcareous
soils. The butterflies are often found on dry, rocky slopes but also in grassy glades in woods. They are
fond of visiting thistles and like plants for their nectar. Various meadow grasses (Poa spp.) and false
bromes (Brachypodium spp.), Feather Grass (Stipa pinnata), and Aegilops geniculata are used as foodplants.
The caterpillars hibernate and pupate on the ground in the spring. This species is single-brooded.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.85).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 odd 4000 0 add 4000 0 add 4000
1.0
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 0.2
Swe
0.6
1.0 O2
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
567
Melanargia russiae (Nymphalidae)
2080
2050
568 Climatic Risk Atlas of European Butterflies
Melanargia galathea (LINNAEUs, 1758) — Marbled White
hc Full dispersal No dispersal
fi SEDG 884 (6.33%) -2681 (-19.21%)
912 (6.53%) -3015 (-21.6%)
-22 (0.16%) -6633 (-47.52%)
© Peter Ginzinger Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 13958)
The Marbled White inhabits many different types of poor grassland that has not been fertilized.
It is common in the southern part of its range, also on road verges and in agricultural areas. The
butterflies need a lot of nectar, and are often seen on plants in bloom. Various grass species, mostly
fine-leaved, are used as foodplants. The female usually just drops her eggs into the grass, sometimes
while flying. The first instar caterpillar hibernates without eating first. In periods of hot weather, the
other instars can also go without food. They pupate deep in the vegetation. The Marbled White has
one generation a yeat.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.81).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
1.6
|
a a
=) a
eee
ee
cf@es
o= mH
=e
£5 8
Oo. 4
of 7
oo
cL nq = =
ger
|
e- :
a
@e-
2 o
wv
x é 5
= 4
o
0 2000 4000 2000 40000 2000 40000 2000 4000
Gd id Gadd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
569
Melanargia galathea (Nymphalidae)
2080
2050
570 Climatic Risk Atlas of European Butterflies
Melanargia lachesis (HUBNER 1790) — Iberian Marbled White
.—hClcL Full dispersal No dispersal
fe SEDG -863 (-61.91%) -1047 (-75.11%)
-1054 (-75.61%) | -1131 (-81.13%)
|GRAS | -1045 (-74.96%) | -1232 (-88.38%)
SEDG -1055 (-75.68%) | -1324 (-94.98%)
BAMBU_ | -1129 (-80.99%) | -1374 (-98.57%)
-910 (-65.28%) | -1394 (-100%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1394)
The Ibertan Marbled White occurs in flower-rich, dry to damp grasslands, near dry scrub, and
along woodland edges and hedgerows. Grasses such as Annual Meadow-grass (Poa annua), Meadow
Fescue (Festuca pratensis), Compact Brome (Bromus madritensis), and Brachypodium retusum are used as
foodplants. This species is single-brooded and hibernates as a caterpillar.
Present distribution can be well explained by climatic variables (AUC = 0.89).
Climate risk category: HHHAR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
‘sad 4000 0 odd 4000 0 aad 4000 0 add 4000
Minimum
Swe
0.6
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
571
Melanargia lachesis (Nymphalidae)
(1a)
504s
(ZV)
Nanva
(ld)
SVH9
2080
2050
572 Climatic Risk Atlas of European Butterflies
Melanargia larissa (EsPErR, 1784) — Balkan Marbled White
.—hCcLe Full dispersal No dispersal
fe SEDG IZA 1.33%} -497 (-44.34%)
-25 (-2.23%) -534 (-47.64%)
roras | 2e@iwy | Som cans
SEDG 246 (21.94%) -586 (-52.27%)
BAMBU -410 (-36.57%) | -873 (-77.88%)
-249 (-22.21%) | -1008 (-89.92%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1121)
The Balkan Marbled White occurs on dry grasslands, rocky slopes, in open scrub, and in grassy
clearings in woods. Sometimes, these butterflies are even seen in villages. The caterpillars feed on
different grasses. This butterfly is single-brooded and hibernates as a caterpillar.
Present distribution can be well explained by climatic variables (AUC = 0.89).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
“4
o
‘dd 4000 0 ed 4000 0 add 4000 0 ada 4000
Minimum
Swe
0.6
o2
1.0
Swe
0.6
1.0 0.2
Swe
0.6
a
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
573
Melanarsia larissa (Nymphalidae)
2080
2050
574 Climatic Risk Atlas of European Butterflies
Melanargia arge (SULZER, 1776) — Italian Marbled White
a
719 (116.53%) | -305 (-49.43%)
SEDG 776 (125.77%) | -331 (-53.65%)
BAMBU 767 (124.31%) | -409 (-66.29%)
655 (106.16%) | -447 (-72.45%)
© Otakar Kudrna Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 617)
The Italian Marbled White occurs locally in small populations. It occurs 1n rocky, calcareous places
on patches of dry, grassy vegetation among loose thickets of Prickly Juniper (Juniperus oxycedrus),
Rosa sempervirens, the bramble Rubus ulmifolus, and Thymus capitatus, with a few scattered trees. The
caterpillars feed on grasses such as Feather Grass (Spa pinnata), and the false-brome Brachypodium
retusum. he Italian Marbled White has one generation a year, and only flies for three weeks per year
in any one location.
This species is listed in Annexes II and IV of the Habitats’ Directive.
Present distribution can be very well explained by climatic variables (AUC = 0.98).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
Gadd 4000 0 edd 4000 0 add 4000 0 add 4000
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 0.2
Swe
0.6
Annual precipitation range
Maximum Large (66%) Small (33%)
1.0 O02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
575
Melanargia arge (Nymphalidae)
2080
2050
576 Climatic Risk Atlas of European Butterflies
Melanargia occitanica (EsPER, 1793) — Western Marbled White
CLL Full dispersal No dispersal
Hi SEDG -958 (-68.19%) -1076 (-76.58%)
-1141 (-81.21%) | -1246 (-88.68%)
-1196 (85.12%) | -1402 (-99.79%)
© Martin Wiemers Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1405)
The Western Marbled White is mostly found in dry, rocky places with grassy vegetation and scattered
rocks, although it is also seen in damper locations. Its foodplants are various grasses, such as Tor-
erass (Brachypodium pinnatum), Bermuda-grass (Cynodon dactylon), and Cock’s-foot (Dactylus glomerata).
This species is single-brooded and hibernates as a caterpillar.
Present distribution can be well explained by climatic variables (AUC = 0.86).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
O 2000 40000 2000 40000 2000 4000 2000 40%)
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Swe
0.6
1.0 O02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
0.2
Gdd Gdd Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Melanargia occitanica (Nymphalidae) 577
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
2050 2080
578 Climatic Risk Atlas of European Butterflies
Melanargia ines (HOFFMANSEGG, 1804) — Spanish Marbled White
.—hClcLwe Full dispersal No dispersal
fe SEDG -460 (-41.52%) -547 (-49.37%)
-648 (-58.48%) | -659 (59.48%)
|GRAS | -657 (-59.3%) -690 (-62.27%)
-1019 (-91.97%) | -1019 (-91.97%)
-1073 (-96.84%) | -1073 (-96.84%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1108)
The Spanish Marbled White is found in warm, dry places with low vegetation on rocky slopes, on
flower-rich grassy vegetation, and in scrub. Its foodplants are grasses, such as Tor-grass (Brachypodium
pinnatum) and Compact Brome (Bromus madritensis). The caterpillars hibernate, pupating on the
ground in the early spring.
Present distribution can be well explained by climatic variables (AUC = 0.95).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 odd 4000 0 add 4000 0 add 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Melanargia ines (Nymphalidae) 579
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
580 Climatic Risk Atlas of European Butterflies
Satyrus ferula (Fasrictus, 1793) — Great Sooty Satyr
.—hCcLe Full dispersal No dispersal
Be SEDG -703 (-31.91%) -1206 (-54.74%)
BAMBU -741 (-33.64%) | -1333 (-60.51%)
-941 (-42.71%) | -1572 (-71.36%)
SEDG -378 (-17.16%) | -1706 (-77.44%)
BAMBU_ | -601 (-27.28%) | -1846 (-83.79%)
GRAS -631 (-28.64%) | -2036 (-92.42%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2203)
In the mountains, the Great Sooty Satyr is mostly found in open, dry, rocky places with grassy
vegetation. At lower altitudes, the butterflies occur in open clearings in woods, or at wood edges. They
can often be seen drinking nectar on such flowers as thistles and scabious (Kuautia spp.). The female
lays her eggs low down on dry grass stems. The caterpillars hibernate when still quite small, and pupate
in May or June. Sheep’s-fescue (Festwca ovina) 1s the most important foodplant, but other fescues (Festuca
spp.), false-bromes (Brachypodium spp.), and bromes (Bromus spp.) ate probably also used.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.76).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
m
o
a : : : :
dd 4000 0 odd 4000 0 add 4000 0 add 4000
1.0
Minimum
Swe
06
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe Swe
0.6 0.6
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
581
Satyrus ferula (Nymphalidae)
2080
2050
582 Climatic Risk Atlas of European Butterflies
Satyrus actaea (EsPER, 1780) — Black Satyr
BAMBU_ | -1016 (-62.79%) | -1579 (-97.59%)
GRAS -1015 (-62.73%) | -1611 (-99.57%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1618)
The Black Satyr can be found on grassy vegetation, rocky slopes, and in scrub. The caterpillars
feed on various grasses, such as fescues (Fes/uca spp.), meadow-grasses (Poa spp.), false-bromes
(Brachypodium spp.), and bromes (Bromus spp.). They hibernate and then pupate on the ground at the
end of the spring in May to June. The Black Satyr is single-brooded.
Present distribution can be well explained by climatic variables (AUC = 0.92).
Climate risk category: HHHAR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
1.0
Minimum
Swe
06
Swe
; 6 10 02
Swe
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
2 oO.
0 2000 40000 2000 40000 2000 40000 2000 4000
Gdd Gdd Gadd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
583
Satyrus actaea (Nymphalidae)
2080
2050
584 Climatic Risk Atlas of European Butterflies
Minois dryas (ScopPo.t, 1763) — Dryad
6 hc Full dispersal No dispersal
Hi SEDG 2262 (53.35%) -1957 (-46.16%)
BAMBU 3822 (90.14%) | -1934 (-45.61%)
2375 (56.01%) | -2311 (-54.5%)
SEDG 3230 (76.18%) | -2611 (-61.58%)
BAMBU 3810 (89.86%) | -3066 (-72.31%)
4305 (101.53%) | -3463 (-81.67%)
© Martin Wiemers Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 4240)
The Dryad inhabits grassy, rather rough vegetation, often located at the edge of woodland or scrub,
and mostly quite damp. The foodplants are relatively broad-leaved, nutritious grasses, such as Purple
Mootr-grass (Modnea caerulea), small-reeds (Calamagrostis spp.), and bromes (Bromus spp.). The female
drops her eggs in flight into the grass. The tiny caterpillars emerge and, without feeding, enter a
period of inactivity during the summer months. In the autumn, they begin to feed, hibernating
in the first or second larval instar. The caterpillars make a little hollow in the ground in which to
pupate, but do not spin a cocoon. The Dryad is single-brooded.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.83).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘edd 4000 0 odd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
Swe
o6 10 02
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
585
Minois dryas (Nymphalidae)
2080
2050
586 Climatic Risk Atlas of European Butterflies
Hipparchia fagi (Scoro.t, 1763) — Woodland Grayling
een a
2258 (34.54%) | -1710 (-26.16%)
SEDG 2059 (31.5%) -2487 (-38.04%)
BAMBU 1747 (26.72%) | -3478 (-53.2%)
1334 (20.41%) | -4425 (67.69%)
© Rudi Verovnik Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 6537)
The Woodland Grayling is a butterfly of open woods that occurs on grassy vegetation, along
woodland rides, and in woodland glades. The butterflies are fond of settling on tree trunks, head end
up, where, with their wings tightly closed and the eye-spots hidden, they are very well camouflaged.
The eggs are laid on the blades of such grasses as fescues (Festuca spp.), false-bromes (Brachypodium
spp.), and bromes (Bromus spp.). The caterpillars grow very slowly and, when they are half-grown,
they hibernate in the litter layer. The caterpillar pupates in a little hollow in the ground, spinning
itself a sort of cocoon. The Woodland Grayling has one generation a year.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.83).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘dd 4000 0 Coded 4000 0 add 4000 0 ada 4000)
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
587
Fipparchia fag (Nymphalidae)
2080
2050
588 Climatic Risk Atlas of European Butterflies
Hipparchia hermione (LINNAEus, 1764) — Rock Grayling
ee
-1810 (54.7%) | -2266 (-68.48%)
amas [wien [ane
SEDG -906 (-27.38%) | -2535 (-76.61%)
BAMBU_ | -2091 (-63.19%) | -2968 (-89.69%)
-2086 (-63.04%) | -3191 (-96.43%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 3309)
The Rock Grayling can be found at the edge of very open woodland on poor steppe-like grasslands,
as found, for example, near open coniferous woods growing on sandy ground. In these habitats,
there are usually rock masses or other sorts of open ground. The butterfly often rests on the ground
or against tree trunks, where the colours and pattern of the underside of its wings provide excellent
camouflage. The eggs are laid on different grasses, including fescues (Fes/uca spp.) and false-bromes
(Brachypodium spp.). The caterpillars grow very slowly and hibernate when half-grown in the litter
layer. They pupate in a little hollow in the ground, after spinning a sort of cocoon. The Rock
Grayling has one brood a year.
Present distribution can be explained by climatic variables to only a limited extent (AUC = 0.72).
Climate risk category: PR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
dd 4000 0 odd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
0.6
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
10 0.2
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
589
Hipparchia hermione (Nymphalidae)
2080
2050
590 Climatic Risk Atlas of European Butterflies
Hipparchia syriaca (STauDINGER, 1871) — Eastern Rock Grayling
CLL Full dispersal No dispersal
He SEDG 774 (40.33%) -595 (-31.01%)
481 (25.07%) -646 (-33.66%)
|GRAS | 534 (27.83%) -773 (-40.28%)
SEDG 652 (33.98%) -893 (-46.53%)
BAMBU -201 (-10.47%) | -1231 (-64.15%)
15 Ga | -1as2 CTH
© Josef Pennerstorfer Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1919)
The Eastern Rock Grayling looks very much like the Woodland Grayling (F7. fag), and they occur
together in some places.. This butterfly is a species of scrub and light woodland of all sorts,
deciduous, coniferous and mixed. At the hottest part of the day, the butterflies hardly fly at all,
resting with closed wings on the shadow side of the tree trunk, or on the ground where they blend
into their surroundings. They hardly ever visit flowers. Their flight is quick and powerful. The
caterpillars feed on grasses. Although single-brooded, they can be seen from May until September.
The butterflies may be inactive during the hot, dry summer.
Present distribution can be well explained by climatic variables (AUC = 0.95).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
ai
o
=]
faa 4000 0 odd 4000 0 add 4000 0 ada 4000
0.6
Minimum
Swe
0.2
Swe
at 0.6 1.0
1.0
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Flipparchia syriaca (Nymphalidae) 591
(B1)
SEDG
GRAS
(A1Fl)
592 Climatic Risk Atlas of European Butterflies
Hipparchia semele (Linnaeus, 1758) — Grayling
Se
-3269 (-23.63%) -4031 (-29.14%)
ams | ate [om a
SEDG -5233 (-37.83%) | -6024 (-43.55%)
BAMBU_ | _ -6634 (-47.96%) | -7748 (-56.01%)
-8424 (-60.9%) | -9748 (-70.47%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 13833)
The Grayling 1s found in dry, infertile surroundings, occurring on poor, dry grasslands, dry heaths,
and also often at the coast. The males and females meet each other above a solitary tree in a wide
open landscape, such as a pine on heathland. The female lays her eggs on various fine-leaved grasses,
including fescues (Festuca spp.), bents (Agrostis spp.), and bromes (Bromus spp.). Walking over the
bare ground, she approaches a grass tussock, and deposits an egg on a withered grass blade a
few centimetres above the ground. The caterpillar grows very slowly, feeding mostly at night. It
hibernates deep down in a grass tussock. When it is ready to pupate, it spins itself a sort of cocoon
in a little hollow in the ground. The Grayling has one brood a year.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.79).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
m
o
e : : : -
tod 4000 0 odd 4000 0 add 4000 0 add 4000
1.0
Minimum
Swe
06
1.0 02
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe Swe
0.6 0.6
1.0 02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
593
Hipparchia semele (Nymphalidae)
2080
2050
594 Climatic Risk Atlas of European Butterflies
Hipparchia volgensis (MAZOCHIN-PORSHNYAKOV, 1952) —
Delattin’s Grayling
.—COcLe Full dispersal No dispersal
| SEDG 426 (43.43%) -382 (-38.94%)
2 = -96 (-9.79%) 549 (-55.96%)
155 (15.8%) -575 (-58.61%)
SEDG 152 (15.49%) -465 (-47.4%)
& | BAMBU -432 (-44.04%) | -735 (-74.92%)
-467 (-47.6%) -856 (-87.26%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 981)
Delattin’s Grayling is a butterfly of warm and dry places. It is found on rocky slopes, on dry
grasslands, and in light woodland. It is stngle-brooded. Adults fly from June to July. Details of the
ecology of the species are unknown.
Present distribution can be well explained by climatic variables (AUC = 0.94).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
0.6
ct
Qa a
=) a
ae
Pe Oe
cfes
o= mn
=e |
£5 8
Go. 4
Oe
D ©
ger
mH
-s
qe. =
2 o
w
= 4
o
0 2000 40000 2000 40000 2000 40000 2000 4000
Gdd Gdd Gad Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
595
Hipparchia volgensis (Nymphalidae)
2080
2050
596 Climatic Risk Atlas of European Butterflies
Hipparchia statilinus (HUFNAGEL, 1766) — Tree Grayling
.—hCcLUw Full dispersal No dispersal
fi SEDG 1004 (16.41%) -1947 (-31.83%)
-56 (-0.92%) -2416 (-39.5%)
-506 (-8.27%) | -4017 (-65.67%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 6117)
The Tree Grayling occurs in very warm, dry and nutrient-poor areas with much open ground and
sparse vegetation. The size and markings of this butterfly are very variable. While remaining on
the ground, the female deposits her eggs one by one on withered blades of grass. Grey Hatr-grass
(Corynephorus canescens), Sheep’s-fescue (Festuca ovina), Brown Bent-grass (Agrostis rineals), bromes
(Bromus spp.), Feather grass (Supa pinnata), and other grasses are used as foodplants. The small
caterpillar passes the winter in a grass tussock and, if it does not freeze, remains active during the
winter. However, growth only begins after hibernation. As development ts very slow, the caterpillar
only pupates in the summer. The flight period of the Tree Grayling, that only has one brood a year,
is therefore very late.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.76).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
‘1.6
sh
a a
=) a
ee
Pe Oe
-& =o
o= mH
=e
£5 3
Oo. 4
of 7
oo
a = ge
ger
me
e-
qf. =
eo
wv
x é 5
= 4
o
0 2000 4000 2000 40000 2000 40000 2000 4000
Gdd Gdd Gdd Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
597
Fipparchia statiiinus (Nymphalidae)
2080
2050
598 Climatic Risk Atlas of European Butterflies
Hipparchia fatua (FRYER, 1845) — Freyer’s Grayling
6c Full dispersal No dispersal
Be SEDG -89 (-8.69%) -482 (-47.07%)
BAMBU -46 (-4.49%) -439 (-42.87%)
aE avery | 8 (S019
SEDG 235 (22.95%) -527 (-51.46%)
BAMBU -245 (-23.93%) | -693 (-67.68%)
GRAS -426 (-41.6%) -827 (-80.76%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 1024)
Freyer’s Grayling can be found in dense thickets, on rocky slopes, on dry, grassy vegetation, in light
woodland and in olive groves and orchards. There are trees in most habitats. In appearance and
choice of biotope, Freyer’s Grayling is very similar to the Tree Grayling (H. séauiinus). The butterflies
are fond of resting on the ground or on tree trunks, and hardly ever visit flowers. It uses different
grasses as foodplants, and has one generation a yeat.
Present distribution can be very well explained by climatic variables (AUC = 0.98).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
-
todd 4000 0 edd 4000 0 add 4000 0 add 4000
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
599
Hipparchia fatua (Nymphalidae)
2080
2050
600 Climatic Risk Atlas of European Butterflies
Hipparchia fidia (LINNAEUs, 1767) — Striped Grayling
.—hClcLwAee Full dispersal No dispersal
Hi SEDG -731 (-33.35%) -1623 (-74.04%)
-1294 (59.03%) | -1644 (-75%)
=5 : oe : BAMBU_ | -1518 (-69.25%) | -2110 (-96.26%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2192)
The Striped Grayling occurs mostly on dry, rocky slopes with bushes and grassy vegetation, and in
light woodland. The butterflies often rest on the ground or on tree trunks. Foodplants are various
grasses, such as Bermuda-grass (Cynodon dactylon), Cock’s-foot (Dactylus slomerata), mweadow-grasses
(Poa spp.), and false-bromes (Brachypodium spp.). This butterfly 1s single-brooded and hibernates as a
caterpillar, on or just in the ground.
Present distribution can be well explained by climatic variables (AUC = 0.88).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
tad 4000 0 edd 4000 0 add 4000 0 add 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
601
Hipparchia fidia (Nymphalidae)
2080
2050
602 Climatic Risk Atlas of European Butterflies
Hipparchia senthes (FRuHstToRFER, 1908) — Balkan Grayling
.ClcLw Full dispersal No dispersal
Hi SEDG -62 (-7.38%) -350 (-41.67%)
BAMBU -2 (-0.24%) -293 (-34.88%)
a5 cn | 386 C4595)
SEDG 207 (24.64%) -421 (-50.12%)
BAMBU -280 (-33.33%) | -631 (-75.12%)
-444 (52.86%) | -767 (-91.31%)
© Neil Thompson Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 840)
The Balkan Grayling replaces the Grayling (1. semed) in the Southern Balkan Peninsula, the Greek
island of Levkas and the Aegean islands. It lives in dry grasslands, scrubland, and open woodlands
and occurs from 0-1600m elevation. The species has one generation per year with adults on the
wing from May until October.
Present distribution can be well explained by climatic variables (AUC = 0.94).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
todd 4000 0 edd 4000 0 add 4000 0 ad 4000
1.0
Minimum
Swe
06
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
603
Fipparchia senthes (Nymphalidae)
2080
2050
604 Climatic Risk Atlas of European Butterflies
Arethusana arethusa ({SCHIFFERMULLER], 1775) — False Grayling
a oe
-64 (-1.14%) -2666 (-47.43%)
SEDG -616 (-10.96%) -3665 (-65.2%)
AMBU -1332 (-23.7%) | -4595 (-81.75%)
-2335 (-41.54%) | -5381 (-95.73%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 5621)
The False Grayling is an inconspicuous butterfly. It occurs on warm, poor grasslands, often situated
near the edges of woodland or scrub, on both calcareous and acid soils. ‘The female drops her eggs
at random into the vegetation, leaving the young caterpillars to choose what to eat. They feed on
different grasses, including Upright Brome (Bromus erectus), Gray Hatr-grass (Corynephorus canescens),
Tor-grass (Brachypodium pinnatum), Crested Dog’s-tail (Cynosurus cristatus), and fescues (Festuca spp.).
The caterpillars hibernate in the first larval instar. They pupate in a grass tussock and have one
brood a yeat.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.77).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
=
sh
=
a
=
Ow
Za
ee]
is}
oO
3 == ' : : ? :
‘dd 4000 0 ood 4000 0 add 4000 0 odd 4000
Minimum
Swe
06
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Alrethusana arethusa (Nymphalidae)
SEDG
GRAS
(B1)
(A1Fl)
605
606 Climatic Risk Atlas of European Butterflies
Brintesta circe (Fasrictus, 1775) — Great Banded Grayling
.6—hlcLUwkE Full dispersal No dispersal
Be SEDG 1057 (14.02%) -2308 (-30.61%)
BAMBU -443 (5.88%) | -2864 (-37.99%)
-42 (0.56%) -3143 (-41.69%)
SEDG 185 (2.45%) -3680 (-48.81%)
BAMBU -961 (-12.75%) | -5018 (-66.56%)
-1518 (-20.14%) | -6183 (-82.01%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 7539)
The Great Banded Grayling is immediately noticeable by its size, and is one of the largest butterflies
of Europe. It glides more than it flies, and can be seen on dry grasslands at the edges of woodland,
and on poor and moderately nutrient-rich agricultural land. The butterflies need quite a lot of
nectar, and are easily observed on the purple flowers of thistles and other plants. The females release
their eggs into the vegetation, sometimes while perched, often while in flight. The caterpillars can
use most grasses as a foodplant. The tiny caterpillar first hibernates in a grass tussock, only in the
spring beginning to feed and grow. The caterpillars pupates in a sort of cocoon in a little hollow in
the ground. This butterfly species is single-brooded.
Present distribution can be explained by climatic variables to a moderate extent (AUC = 0.81).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘dd 4000 0 odd 4000 0 oad 4000 0 ada 4000)
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
607
Brintesia circe (Nymphalidae)
Ly
ee
2080
2050
608 Climatic Risk Atlas of European Butterflies
Chazara briseis (LINNAEUS, 1764) — The Hermit
.—hCcLUwE Full dispersal No dispersal
SEDG 645 (8.85%) -1902 (-26.08%)
BAMBU -27 (-0.37%) -2256 (-30.94%)
-349 (-4.79%) -2727 (-37.4%)
SEDG -402 (5.51%) | -3226 (-44.24%)
BAMBU_ | -1170 (-16.04%) | -4231 (-58.02%)
-22.63 (-31.03%) | -5429 (-74.45%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 7292)
The Hermit inhabits steppe-like grasslands in dry, warm places. In the northern part of its distribution
range, it is restricted to dry chalk hills. Although it 1s a very large butterfly, when its wings are closed,
it blends perfectly into its surroundings. Especially when resting so on a rock, they are almost
invisible. The female deposits her eggs one at a time on the withered blades of many different
erasses, including false-bromes (Brachypodium spp.), bromes (Bromus spp.), and fescues (Festuca spp.).
The caterpillars hibernate in the first larval instar. They pupate in a sort of cocoon, in a little hollow
in the ground, or sometimes low down on the foodplant. The Hermit has one brood a year and is
one of the most seriously threatened species in central Europe.
Present distribution can be explained by climatic variables to quite some extent (AUC = 0.8).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
2
=
sh
=
a
=
Ow
=o
i]
sity
oO
= E : : . ? :
dd 4000 0 odd 4000 0 oad 4000 0 Gdd 4000
Minimum
Swe
06
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
609
Chazara briseis (Nymphalidae)
2080
2050
610 Climatic Risk Atlas of European Butterflies
Pseudochazara anthelea (LEFEBVRE, 1831) — White-banded Grayling
.—ClcL Full dispersal No dispersal
Hee SEDG -175 (-21.39%) -360 (-44.01%)
-134 (-16.38%) | -332 (-40.59%)
| -231 (-28.24%) -481 (-58.8%)
-322 (-39.36%) | -623 (-76.16%)
© Albert Vliegenthart Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 818)
With the white bands on each wing and the dark narrow scent-brand, the males of the White-banded
Grayling are unmistakable. The butterflies occur on dry, grassy vegetation in open scrub, and on rocky
slopes. Sometimes, they are seen in open woods. They often drink nectar from thistle-like plants. The
males defend their territory, perching on a rock or other prominent features in the surroundings.
This butterfly 1s single-brooded.
Pseudochazara amalthea (FRIWALDSZKY, 1845), placed by KupDRNA (2002) provisionally in the genus
Hipparclia, 1s a subspecies of Pseudochazara anthela (LEFEBVRE, 1831), as it is already treated by
many authors.
Present distribution can be well explained by climatic variables (AUC = 0.9).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
‘edd 4000 0 edd 4000 0 oad 4000 0 add 4000
1.0
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O2
0.6
Swe
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Pseudochazara anthelea (Nymphalidae)
SEDG
(B1)
611
BAMBU
(A2)
GRAS
(A1Fl)
612 Climatic Risk Atlas of European Butterflies
Oenets norna (BECKLIN, 1791) — Norse Grayling
.—hCcLwE Full dispersal No dispersal
fi SEDG -1683 (-81.03%) | -1691 (-81.42%)
-1680 (-80.89%) | -1687 (-81.22%)
-2026 (-97.54%) | -2030 (-97.74%)
© Bernard Fransen Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 2077)
The Norse Grayling occurs in both dry and wet biotopes, on marshes with a vegetation of mosses
and grasses, at the edges of swampy habitats and streams, in low birch scrub, on sparse vegetation in
clearings in birch woods, and on sunny, rocky slopes with low shrubs. The butterflies spend much of
the day resting on tree trunks or on the ground with their wings closed and are so hardly noticeable.
They are quick flyers and difficult to approach. The female lays her eggs on various sedges and
grasses. The egg takes two years to develop into a butterfly, and the caterpillars hibernate twice.
Present distribution can be very well explained by climatic variables (AUC = 0.98).
Climate risk category: HHHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
Swe
0.6
0 2000 40000 2000 40000 2000 40000 2000 4000
Gdd d Gdd
Gadd
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
Oenezs norna (Nymphalidae) 613
SEDG
(B1)
BAMBU
(A2)
GRAS
(A1Fl)
2050 2080
614 Climatic Risk Atlas of European Butterflies
Oeneis glacialis (MOLL, 1785) — Alpine Grayling
.—hlcLw Full dispersal No dispersal
fe SEDG -192 (-20.65%) -368 (-39.57%)
-114 (12.26%) | -323 (34.73%)
|GRAS | -247 (-26.56%) | -400 (-43.01%)
SEDG -430 (-46.24%) | -637 (-68.49%)
BAMBU_ | -272 (-29.25%) | -529 (-56.88%)
-480 (-51.61%) -710 (-76.34%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 930)
At altitudes of about 1500 m, the Alpine Grayling occurs on dry, scrubby vegetation. Above the
tree-line, they can be seen in dry, stony alpine grasslands, and on dry, open sunny slopes. Most
habitats have a stream in the vicinity. Perched on a stone, the males defend their territory, chasing
away other butterflies, as well as other insects. The female lays her eggs one at a time on dry grass
stalks close to the ground. The caterpillar hibernates in the first larval instar, and having fed during
the growing season, hibernates again in the last instar. Eventually, some time between April and
June, it pupates. Its main foodplant is Sheep’s-fescue (Fes/aca ovina) but other fescues are also used.
This butterfly 1s single-brooded.
Present distribution can be very well explained by climatic variables (AUC = 0.99).
Climate risk category: HR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
o
‘dd 4000 0 edd 4000 0 add 4000 0 add 4000
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 60.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 O02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
615
Oenets glacialis (Nymphalidae)
2080
2050
616 Climatic Risk Atlas of European Butterflies
Oenets jutta (HUBNER, 1806) — Baltic Grayling
y .—CcLee Full dispersal No dispersal
Hee SEDG -1165 (-19.45%) | -1791 (-29.89%)
-1434 (-23.94%) | -2057 (-34.33%)
.
“ -2096 (-34.99%) | -3090 (-51.58%)
-2839 (-47.39%) | -3887 (-64.88%)
© Chris van Swaay Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 5991)
Both in lowland and mountains, the Baltic Grayling occurs in peaty and swampy habitats. Most
butterflies are found in damp habitats with a vegetation of grasses and sedges, often with open
water in the middle and surrounded by coniferous woodland. The butterflies frequently visit the
wood edge looking for flowers, because the peat vegetation is poor in nectar plants. They also
rest on the branches or trunks of trees, or on dead wood. The Baltic Grayling shares its habitat
with hardly any other butterfly. The female lays her eggs on various grasses, but which ones the
caterpillars feed on is not known. The caterpillar’s development takes nearly two years.
Present distribution can be very well explained by climatic variables (AUC = 0.96).
Climate risk category: R.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
a
‘edd 4000 0 odd 4000 0 ad 4000 0 add 4000
Minimum
Swe
06
1.0 O02
Swe
0.6
1.0 0.2
Annual precipitation range
Swe
0.6
Maximum Large (66%) Small (33%)
1.0 02
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
617
Oeneis jutta (Nymphalidae)
2080
2050
618 Climatic Risk Atlas of European Butterflies
Danaus chrysippus (LINNAEUS, 1758) — Plain Tiger
a
-209 (-50.73%) | -304 (-73.79%)
SEDG -74 (-17.96%) -273 (-66.26%)
BAMBU -195 (-47.33%) | -334 (-81.07%)
-203 (-49.27%) | -369 (-89.56%)
© Kars Veling Changes in climatic niche distribution
(in 10’x10’ grid cells; present niche space: 412)
The Plain Tiger occurs in coastal areas on warm, rocky places with scrub, on agricultural land and
in gardens. The Plain Tiger can fly great distances, and in this way can found new populations. The
caterpillars feed on the milkweeds Asclepias curassavica and Cynachum procera. \t has several broods a
year and does not hibernate. In areas with cold winters, populations can therefore only be temporary.
It is difficult to establish whether the species 1s present as a resident the whole year, or as a migrant,
only breeding in the summer.
Present distribution can be well explained by climatic variables (AUC = 0.89).
Climate risk category: HHR.
Annual temperature range
Minimum Small (33%) Large (66%) Maximum
Ow
Za
on
mH
a
dd 4000 0 edd 4000 0 add 4000 0 ada 4000
1.0
Minimum
Swe
0.6
1.0 02
Swe
0.6
1.0 O2
Annual precipitation range
Maximum Large (66%) Small (33%)
Swe
0.6
0.2
Observed species distribution (50 x 50 km? UTM grid; black Multidimensional climatic niche. Occurrence probability defined by
circles) and modelled actual distribution of climatic niche accumulated growing degree days until August (Gdd) and soil water
(orange areas) content (Swc) for combinations of minimum, lower tercile, upper
tercile and maximum values of annual temperature range and annual
precipitation range. Climatic conditions: orange — unsuitable; green —
hostile; black line — modelled threshold
619
Danaus chrysippus (Danaidae)
(1a)
504s
(ZV)
Nanva
(4b)
SVH9
2080
2050
620 Climatic Risk Atlas of European Butterflies
C.3 Non-modelled European butterfly species
Due to our methodological restrictions, it was not possible to model the climate
change risk for a large number of species with a very restricted European distribution
(149 species, see Table C.3.1). Most of these species would expect to be categorized
as being at extremely high risk (THHR) from climate change because they have
such a limited distribution and because any change in climate space is likely to have
a strong impact.
Better models should be developed for these species in the future, but it 1s
important not to ignore them in the current discussion on climate change impacts
on butterflies. We have therefore listed them below to highlight European endemics
and give an indication of the European distribution. We also have listed the closest
relative of the taxon concerned in those cases where taxonomic discussion is ongoing.
In some cases, the species mentioned in the first column may eventually be included
as subspecies of this closest relative.
621
C. Climate Risks of European Butterfly Species
WD (Od SMW “TV ‘su ‘val Wa DIUSDINDD SIO") MOTPA PEPNO[D uro—eg
MOTPA Pepnoyy) ysorH
dry, ssurs¢ usa\svy]
UOO}SI+] ULIOI)
[PRIMOT[LAS ULITSIO’)
COG] “dy BuuvyHg svy0yD
(OG8 WHAVHOS-HORRIAE]) Vuasounv svy07y
(GLLT ‘saioraav yy) yyasoyf vyesdojvD
(9¢8] “IVANASIOg) auounp supqooqjupy
ovpHoId
<
(POGL “Iaaay) noyas viquudsey
6E8I “ANH wojedsoq oyzdog
(68L1 “ISauap]) sauggodr uoqopy
ovpruorideg
(TOgT ‘uausaw’]) xnug suoyouly |
(PORT “INaMy) wsege sus aulg [.
(TPQT ‘NNVINSHAAY) Wryadqui supqous
(zegy ‘unanvy) eadruegs vynids
(OPRT UNaNVY) exuvus subi
(L781 “WANaQEY) xeuqy sopidopag
(CIOL ‘NIGUAAMY) Mapunys suposwguny
(CEQ “IVANASIOg) Mzuogsoq oquog
¢
(apoo g-eudye 1-991€ OS]) SeujuUNOD UesdoOJN® Jo sapod AuJUNOD 9y} UM Z'E'D JIGe] 99s :uONNquISIp Uesdouny
(dlwapuUs Ule]JUNOW = || ‘OlWapPuUsd pues! = |) DIWaPUS UBadOINy = J
‘BuloBuo si UOISSNOSIP DIWIOUOXE} BJOUM SO9SED BSOY} UI PA9UJBDUOD UOXE}] OU} JO DAI}E/9J ISASOJD :dAITE[9J ISOSO}D
(elssny pue
‘BAOPIOW ‘OUIeIYA ‘SNuejag JO S9019} By] Bulpnjoxe) seyre Siu} UIYUM pd||@apOW U99q 1OU BABY YOIYM saideds AyolNg ueedon®z :1"E"D sqeL
Climatic Risk Atlas of European Butterflies
622
PS EE |
i anja soydea
onjg Areur7)
ong sumbrorT
ssouardg YN SSA Wa
SS SES el Sa TTR
wONNQINISIp ERRATA [ae |
(Pel UAMINAAQ) Zaswayq vuavat [
OZ6L ‘NVWaVH’) Sogdod agalsdomyyy
(OPT “arinug) suuprqgan suudaty
(LpQ] WHdAVHOS-HoRIa py) “uenbag opiduy
L7G WHANALS yaassv9 opiduy
(PrQy WAAAY) Supqoou sepyyiyD
(ZL UAINVUD) Supjogn suuvx}y
(OLG] SAMA] SASWAUOLLOUL DID
(PESl “ONTS)) seman syungd ye
sepruavoAT
¢
(TST ANAT) apuorqo vyuog
(SOT “WANAQTT) syuundsaq secary
<
B96] ‘OIAOMUO’T VunyVg Stat]
¢
TOQT UNATAY Sasuascapou Xia Pauory
(GZ8T ANANEY) yuqoys xiajqouor
(ZSQT “AHAMUY) Veuad aojqouTy
LOST UHONICOAVLS Saujusus aojgouy
CI61 ‘TUHOSHLOY Wupuadsag aojqonry
COG] SININV.LS asada a07qoury
8007 “MOVE 2UgULISUOI a0;GINF
(THB “IHAZNOC) vuopvYo a0;GONTT
(6781 “ONTS) acocnza szojoy
(ZIST “adog aa) agaly seyoD
623
C. Climate Risks of European Butterfly Species
vIUIpsIeS :TI| [7A UOPLLOI SUJDUUUMOM Of
Ajeay yINoS :T I] A wyAo Gea SuvULUMOL Og
eTUOCTeIe’) :S eae SHOP SHIDUUUMON OT
SIN “WD SO) W: :
vUOpsryy WO
ureds WIION :Si
BPVADN VIF :Sq] WA i
onyg, (Gvyeg) orureavey
(CLR, ‘MANaAT]) M708 supounuoyog
(LQ6] “AHHOIN]) Yuacuvuuas suouuoljog
(6L6L ‘OSOT, 29 OLLATIVG) 20708 sujouuol]ogy
(SZ6L ‘VUNVOVS AC) Suan suppumMolog
(O16 AHIAAAQ) Zassaagnl supouluMoly og
(9Z6] ‘ALINE A) SUnLaquxa suDUUMULO] Og
SOOT ‘SNR AC 2 SISLAO’) aveuaja sujouuMoljog
(QR 1 ‘NNVINSUMAR) Suuspsala09 supvULUULOGOT
(9L61 ‘NMOUG) Seswaeuvowv sHjoULUOL OI
(S661 ‘SVIVAVHS) 29 SISLNOT)) SMMOMPUD SUIDULUUOY OT
(L961 SUAISUOY) apsuew supouumod og
(CGT ‘ONIN Goan suloqary
SOOT “IV LA UATAZLAL 29700 sulogag
(LZ6T ‘Alaa A) 2/fnuy sulogag
(OPSt “IvAnasiog) smmuaid suloqaty
(zEgy ‘uaHOosq) wood sulaqar gy
(SpST waaay) suoplsd sulogag
(Lpgy Maria) suagy suloqary
(6EQ] ‘Una Y) smnadsaq suloqay
(ZGQ] SudAaEyq) sayedina suloqar gy
(ZEQ] AAA) svuvpsvp suloqagy
(OLG6L ‘UAHLINAQ) 2eayzaq sulogag
(PEST “ONTS]) syaqs Duval T
([SQ] ‘uaAdas) vuedyqus vuavakT
Climatic Risk Atlas of European Butterflies
624
Id “AS SON ‘etavurtpuros onary aS Axelay Fepog (6781 “IvAnasiog) samo vuojog
< ¢
LV ‘LI ‘HO “WA
: Aseyniayy ureyuno ‘SOASNVINAAOP]) vavdpu v12070
Sy :saouardg 29 sdry ‘T4 “AS ‘ON (eels an (FOST 1H) PALO
Arey] posurm-Aysnq (LL UNLLNg) vqosdu vuojog
Arey[AU | ULoTsIO7) (CZQI SINVAOD) vsya stuuloupy
sepreydurAyy
(ZOG6T “TAaeD) Logadly) vuvuvanT,
(QGQ] SAAAMM YY) womulpua Duvuviny.
(ZSGQ] AHATVHOS-HORIAAE]) 4//ad0U SasvUuco |,
(C6LT ‘snloniavy) smsruqGooqy snonavy,
(GhQT UHAMU) SMUG SHIMAVT,
(Z86] ‘NEIMOO 2 SNRIQ) awispgavg sapyunjyods
(QR “IvVANasIog) Masapay winzulzv 5
(OL6L AHIIAIQ) swaecuer supouuoljog
ureds yynog :Sy Ea ; (6L61 “IV Lt OTILISNG-ZAINON)) avpajont suspumoljog
a
soupy TW AD
625
C. Climate Risks of European Butterfly Species
uredg yenuay :Sy Wa |
ZO ‘OW SAV Mel HOY We |
MHS SLLSLV Sdqv] WW |
Sy Wy sseouerdg Eiecd
LE wt S4qV] IN|
SU UO SIN Od] Wa
Id SAS (ON ‘elavurpuros onary fat
VIFqUIUL’) :Ci]
LV SLL HO 'sdty] Ww |
SH SY ssoouarkg iiecd
BITAS 29 soouaTA :S7 fsoouarkg YAY ee
ir
Sy YW :seouerhg
LV SLLSHO dry] 22 |
IS “LV SLL sdy} Wea
LE Wt S4pV] Wa |
Ld 4svoo YyANOs “‘spuvys]| Ayeut’) SSH feat
Sa UD ‘Va “IV] WA
a
wonnqmsip uvadomy| | a |
nosy dada vigaay
jo[SuTY stoiedey
Jo[SuNT Urjopns
opsuny ues
opsuny Amoq osyey
JO[SUNT syore'T
JO[SUNT SJOYSIN
JO[SUNT pur[poo,\ ono7y
jopsuny s,uewdey’)
jo[suNy uvisesng
jopsuny Assvsg s.ossoy oq
JO[SUNT S,dFAqoajoT
jopsuny Assvig, ystueds
JO[SUNT ouTVAvL
JO[SUNT popurg A\oT]O_
jopsuTY yoods sy)
JO[SUNY $.50zIVY
jopsuny Assvrg Ss oTAOyIO'T
JO[SUNT VISOUPY OSTey
PIA “YOIvuopy
yIwoy] [ews urjos)
wrap] uryyeg
7?
pvap] ULqTH
YIP] ULITSIO’)
C/8T UOHINNIQC Mads viqay
LOS] SWHONICOVLS M7japus viqady
(PZT “IVaOD) ves Diqasry
OS8T ‘NIISVUS Oluuaqys viqaay
TERT “IVANASIOg Osis vigasry
O0G6L “TIOHOIN Sawadopogs viqaay
LOST WHONICAV.LS surpod vigasy
SOG6L ‘NVNdVH) Mund vig
OOGT “SHANTY Symjuano Diqaary
$G6I “ASSAY 299 OIAOMUO’T SYnIU DIgaly
B78 “IVANAsIOg zaaiqafay vIgaaTy
SORT UILLAg veundsay viqasy
CCRT “IVANASIOG aos108 viqasry
CO8T “INANE ypnsvfoanyf viqaay
(68L 1 ‘NUSAVHNUOG) Puspuyys viqaly]
OG68T UAZLVY Ysugo viqary
€G6I ‘OIAOMIO’T Lunn vIqas7]
(9081 SDOUSNVINAHOP]) 2fadouqyjan vigaary
(QGL ‘SQavNNIq) saddixad suvung,
(OPQ WHAM) seeugy vg duduouso’y
OLGL “ay syvjuauo vyduuous0’y
LOGL SWAONIAAV.LS vuvgzsa vyduuoua0’y
(9081 ‘wANaAyz]) vuuatos vy duuous0D
Climatic Risk Atlas of European Butterflies
626
PUYIST SOFISSIN AO a UAMOIG MOPVIJ[ S,UOSUTOY J,
vue PT Sa} La
eIeUe’) ULILD) :Sy] gee]
La
SPULIST VZUO ‘TJ
“velfay WIUEPILS 4 LIISIOD A]
surAvrgy snsddv)
VAIPLY + Ld baa
SpULIST ULTOY : [J
SuTpAVIL) ULIOI)
SuTAorE) sarozy
soyjedse) ATH
OxIOTE] TA rior
uonngqmsip uvadoma Rese
SsuyAvrs) uJOYyINOS
(6061 SMOHILIAAQ) wvoeu vjowuDyy
0661 ‘NOSNOH[, SasspusnIyvg DIOLUDIAT
O76] ‘SHAVUD Yrorud njowupyyy
(L861 ‘NOSIOH]) 72g vrouDyy
(6881 SISIHHD) zssha viqounddapy
(F861 “UNVIN) soe riqourddiy
Z661 “ELLINS 29 NING aqupoun vigaodgayy
$361 ‘VNYaNy wzuopsogs viqowwdgayy
(CZ6] “Aaanv.LS) vpoupod riqawwddayy
(6ZQ] “INVaO) suaaoau viqasvddyy
(IZ6] ‘uaanv.Ls) vunzyodveu vigauwdgayy
(LL8] SMaDNIanv.Ls) vuasuau viqowwddayy
(L681 MANVG-ANAHLLAg) sauasapou riqowwddayy
961 ‘VNNGNY sgaqsry viqoswdgayy
L9G] ‘SNIDOIPFY Biawos viqoswddayy
(2061 AIIO.LSHAN) Bevuad viqowddepy
(6FG6L “MIOFA) Sasuaudia viqawwddapy
(OGL “taaay) vyew riqounddapy
LLOL SVNuansy Zuasuasega vigauwd gee]
(206[ “WaTMOLSHAA]) eaqonyg vigaswd apy
L9G ‘SNIDOIPT Suqoong viqosod ayy
(6681 ‘WaNOUULS) vuuorv wiqowwddeyy
(9Z8] ‘ITIANOG) suanpsuw viqouvd dip y
627
AQspuysrurwop-w} = | SsuRY UoLyy (ZEgl “ONS adosagsy ven dy
VITOPLW i Ld ‘spurysy Aseursy oq | 1a | DIupUt DSSOUD A, [esupy poy Azeur) (6181 “IaVaOyD) veuvyna pssauv A
(CLL SANNUC) SasuarusGaed VssaUD A
(LZ6L ‘ISWOY) suavyyia DivSDYI0pNas g
Surpsoryg yeq (O861 ‘NMOUG) aoqdesy vavsvqr0puas gy
SuyArsryy) spd (L861 ‘NEMO 2 SNR) $afsa4o DADSDYIOPHAS |
SuTAoFE) ULTIDOIL (OL8T UHONIAGNVLG) BMT DADSDYIOPNAS J
SulAory uvisy Aor (GPQ] “WadYHOS-HORRIAP]) Léalae nepspqsopnas J
C. Climate Risks of European Butterfly Species
SuTAvrey ULTUOPoIVIY (CLO ‘SSOUD) wyS108ULI DADSDYIOPNas J
JO[SuNT uenvUeC (L8L1 ‘snlontavy) ade piqasapory
poo, popyoods Azeur’) (LL8L SYHONIANAV.LS) saprorq fax adupang
eITOPC] iL.d par poor, pappeds uvmopryy (GLLI ‘sntontavy) weg gex atanuny
VIUIPILS : TT SVOISIOT) YY DAITOUM DIDULUOISD [. UMOI [[VA\ ULITSIO7) (PZQ] ANAAP]) weavsounund VIDIUMOISY
WL WY “IV “MIN SM Og * UMOIG 9d}Iw'T Jossy] (P8LI “WHdsy]) aways vuury
IH SAS SON izravurpuros onary 2 Suykerg onory (Z6LT SUHAIANHODS) adog szaaQ)
VIVIPILS 3] {OISIOT) YT avayjln my (9ZQ] SITIHNOG) vsnugon swap
ERE AS INE 2S och NL DIEDYID DAVIYIW (F061 UOHRAAIQC) sSeswaporau vavyqyay\y
Aseyas yy Op wy] (Q7R] UHAMAY) Valapsy VaDIY aT
Arey] ssokory] (PSL Adds) Luuimpav vavIy A
ONY PITFeJA] ULTPOTS (CEgy “IvAnasalog) vsmsagd vidvUD] ayy
UAOTY MOpva] ULIsoV (LQ UNTIN'Z,) vessaudyay VIOLUDINT
UMOIG AOPLI|\] ULTUTPIVS (JST SINVITIH) seumu rjowupyy
Climatic Risk Atlas of European Butterflies
Table C.3.2: Country codes of European countries (ISO 3166-1 alpha-2 code)
i a
Eo
[a [aban
[at [amie
is [pane
Per [page
Pi [Bae
Fd
[cat [inetd
[oe [ome
[2 [emchnepiie
aaa
[bx [Deore
[ee [anon
[as [aie
Pa [int
[ro [eee
[em [Rows
[ae [Orie
[ar [oimier
[en [ome
CS
fiw [roms
[ie [inte
Pc oN
a
2
Dries
a
7
re
2
=e
Nolios
eee
os i
Mie
Ni feNatcae
im
my [toed
Pr [Posh
50 mes
Sc
RU [Ros Racaion
2
a ee
seas
mds
m [tg
ts [Uinins
BEB BEeE
<
Sree eee
O 4
C. Climate Risks of European Butterfly Species 629
C.4 Summary results
The results of our analyses are summarized in Figures C.4.1 and C.4.2 (and in tables
App. 3.1 and App. 3.2). The study shows clearly that climate change poses a considerable
additional risk to European butterflies. However, the risk varies considerably under the
three storylines considered in this atlas.
Under the extreme, no dispersal GRAS scenario for 2080, 24% of the modelled
species lose more than 95% of their present climatic niche and 78% lose more than
50%. A further 20% are under potential risk and only 6% of species can be rated as
being at lower risk.
Under the intermediate, no dispersal BAMBU scenario for 2080, 9% lose more
than 95% of their climatic niche and 66% lose more than 50%. While under the best
case SEDG scenario, only 3% lose more than 95% of their climatic niche and 48%
lose mote than 50%.
The results also show that there is a considerable time lag in the effects of climate
change on European butterflies. Until 2050, the effects across different scenarios are
still moderate. Under the no dispersal GRAS scenario (excluding the PR category),
around 60% of species are still rated as experiencing a lower risk until 2050, while in
2080 these are a mete 6 %. The relative differences are similar across all the scenarios.
Dispersal ability is one of the major factors that will affect a species ability to survive
under future climatic conditions. This ability is a proxy not only for mobility itself, but
also for the availability of suitable habitats in the new areas of the respective climatic
niches. Under the very moderate SEDG scenario, 33% of species could experience a
net increase in climate niche space until 2050, and even 30% of the species have this
option until 2080. Under the other two scenarios, around a quarter of species could
experience an increase in climate space until 2050, while by 2080 this is only the case
for 18%. Thus, until 2050, there may be better conditions for some warmth-loving
species, provided they can respond, but these subsequently get worse.
The most striking differences in the results are the ones between the different
scenarios. Considering the no dispersal scenarios again for 2080, the number of
species of the lower risk category ranges between 43% for SEDG and 6% for GRAS
(with 20% for BAMBU).
On page 630 we present the results of climate risk analysis for both time steps
(left and right column) under the three scenarios SEDG (first row), BAMBU (second
row), and GRAS (third row) for full dispersal, while on page 631 the same is shown
under the assumption of no dispersal (for definition of risk categories see chapter B.4,
pages 24ff.).
630 Climatic Risk Atlas of European Butterflies
© tail ispersal
00 SF IL |
i —_—-- - = + : ~~ -
| UU | Ti | | Re | LA deer | LR imer MEIER. | 1 He | Lib Beer | LE incr
Bhi dispersal = a u a. ah? bes Fe] aS Gull disjecreal, Lh SI
Sd SEG | | | MAB SETS
ell dlinperrsal J
_ 2050 HAR THA 2 (AAD
[amen | Hi in | & | UR deer] LE iner J vannmere | ten HK Lit deer | Lt imer
Biel digersel | UH | wd Siidapersd | a: | as | 45
Dee0 AME a6 AMIE
Bell Sepersall bell dispersal
SHO Ti RAS SOD ta HAS
fl — ~
Haat | HMe = i [LR eee | LR ince | enn FLEE Hie. | Lim deer (Eine
Stull fayaraatl] 1t il 7 Stull | aT 225 44
TSE GA Be CALAn
Figure C.4.1: Percentages of climate risk categories of European butterflies under the
full dispersal assumption for 2050 (left column) and 2080 (right column) under the three
scenarios SEDG (first row), BAMBU (second row), and GRAS (third row).
(without “PR”; for category definitions see pp. 24ff.)
C. Climate Risks of European Butterfly Species 631
mi dia piers | nai dia pemrcal
2 oF [MG 20a SF [ay
i -— = f
HEE = HE. ! Litdeer | 10 baer | anne | HER 1a
Sno ilispepeal e4 Lift. Gna | 43 2H
NS6 SEIN IRS SEIN.
| penne | Wan disperal
20 BAL PL ed
UA deer | LR deer
1-5
a+
HHI | HHR Hk
= Bz 4)
Wiididla | ik B Lider LE ier
Bins dikpersal | oa | if | | len ae | f Sinn degerent 14.7 tei
ben hasin lea FAATAA
Sinn ibis spereal Bon dispersal
F “3 eral
E ye
| 482
=
a
T #
HH | HAR le R | LR dees | Utiner | anna | i ao LL incr
Bieteere| it | 4) | 7 | coe | ai | os ee ee
MES GLAS DORE GRAS
Figure C.4.2: Percentages of climate risk categories of European butterflies under the
no dispersal assumption for 2050 (left column) and 2080 (right column) under the three
scenarios SEDG (first row), BAMBU (second row), and GRAS (third row).
(without “PR”; for category definitions see pp. 24ff.)
D
ee
DISCUSSION OF METHODOLOGICAL
LIMITATIONS
634 Climatic Risk Atlas of European Butterflies
D.1 Limitations of species distribution
models and future projections
General suitability of species distribution projections
Species distribution models (Guisan & Zimmermann 2000) are widely used in
assessing possible future distribution patterns. They correlate current environmental
conditions with observed presences/absences (or only presences) of single species.
Species distribution models are based on the assumption that a species occupies all
areas with a suitable environment, i.e. the species fills its so called fundamental niche.
In reality, species realize only parts of their fundamental niches and thus are rarely
in equilibrtum with their environment. This may be caused by several reasons, e.g.
biotic interactions or dispersal ability may restrict the niche filling or slow it down in a
changing environment.
Though the approach of species distribution models is purely correlative and
does not imply causality, species distributions can usually be described well enough
to draw some useful conclusions. Based on the assumption that the relation between
environment and species distribution holds in a changing environment, distribution
models are used to project the potential impacts on species distribution and biodiversity
corresponding to climate and land use change scenarios.
Species distribution models are validated by comparing the fit between the
modelled distribution, based on current environmental conditions, and the observed
distribution. Such validation helps estimating the model reliability and uncover
soutces and patterns of discrepancy between modelled and real species distribution.
Having assessed the modelling errors, it is possible to make a careful and sound
interpretation of the results.
The most important causes of modelling errors for species distribution models
ate as follows:
¢ Data quality: A low sampling effort, duc tolow accessibility orinconspicuousness
of species, underestimates the true species distribution and truncates
environmental niches. Data should cover the whole range of a species and thus
the full niche of species, otherwise tolerance to environmental conditions may
be misjudged (Kadmon et al. 2003).
¢ Data resolution: Data resolution may not be appropriate to capture
environmental conditions that are experienced by organisms (Pearson &
Dawson 2003). Mean climatic conditions taken over a larger area may not
reflect the local climatic conditions experienced in a species’ microhabitat.
For example conditions in mountains and peat bogs may change substantially
D. Discussion of methodological imitations
within short distances due to variations in local topography and land-form
(Trtvedi et al. 2008).
Modelling algorithm: Choice of modelling algorithm affects prediction
error (Pearson et al. 2006). Some methods may give a better fit to nonlinear
data, but are subject to over-fitting (Randin et al. 2006) (1.e. they may describe
calibration data well but are useless for temporal or spatial projections). To
reduce the influence of the chosen technique on the modelling results Araujo
& New (2007) suggested to combine the results of many different (good)
models by ensemble forecasting. Ensemble methods are surely appropriate
in the absence of detailed ecological knowledge of the species modelled.
However, for species with known ecological requirements, model results
can be judged according to their sensibility. Using ensemble methods and
average across all results Gncluding insensible models) would lead to worse
instead of better model results.
Species’ ecology: Model performance may be influenced in a variety of ways
by species characteristics (McPherson & Jetz 2007). Range characteristics
resemble the ecological niche of a species in space. Species with clumped
distribution and narrow niches have a good predictability; results for range
size itself were ambiguous and may even be an statistical artefact (for
discussion see: Hernandez et al. 2006). Species of higher trophic levels are
expected to be modelled least well, since they strongly depend on other species
(Huntley et al. 2004). A high temporal or spatial mobility (e.g. good dispersal,
migratory behaviour) may lead to reduced detectability or misjudgement of
habitat preferences (Pulliam 2000). Low performance of inconspicuousness
ot scarce species is opposed by a good performance of large and easily
detectable species (Seoane et al. 2005).
Intraspecific variation: In species distribution models (and usually in
dynamic models as well) it is assumed that species are not differentiated and
do not have phenotypic or behavioural plasticity. lgnoring the adaptation
to local conditions leads to overestimation of the environmental niche
(Harte et al. 2004).
Evolutionary adaptation: Usually, both types of modelling approaches ignore
the ability of species for (micro-)evolutionary changes, i.e. species can adapt
to novel conditions only within the limits of their genetically predetermined
potential. However, it was shown that species have the evolutionary potential
to adapt to novel environmental conditions (Thomas et al. 2001). This will
underestimate the potential environmental niche in the future.
Lacking parameters: Limited knowledge on environmental conditions
influencing species distribution seriously reduces modelling performance
(Barry & Elith 2006). Inclusion of land use increased model performance on
small scale (Luoto et al. 2007) as well as large scale (Pompe et al. 2008). The
influence of biotic interactions (e.g. competition, facilitation, diseases, prey)
are generally ignored in species distribution models though they are assumed
635
636 Climatic Risk Atlas of European Butterflies
to play a large role in shaping species distribution. Recent applications are
promising at least to describe simple interactions (Heikkinen et al. 2007,
Schweiger et al. in press).
¢ Changing biotic and environmental interactions. Species distribution
models based on correlations between current distribution environment may
be invalidated by uncoupling and rearrangement of biotic interactions (see
above) or the appearance of unprecedented environmental conditions (e.g.
CO, fertilization of plants). For example, increased environmental CO, may
lead to increased stomatal conductance and hence to less evapotranspiration
and subsequently higher soil water availability than can be assumed from
purely climatic conditions. Ignoring this can largely affect species distribution
models (Rickebusch et al. 2008).
This list makes clear that biodiversity models are not able to replicate all the complex
patterns of reality and are valid only under its simplifying assumptions. The extent of
this limitation might question the usefulness of such models. However, the aim of
such modelling procedures is not to capture the full complexity of the systems but
to gain a better understanding of the basic patterns and underlying natural processes.
The simplifications allow us to reduce the complexity of the real world to a level that
can be handled with the available techniques and knowledge. It is better to achieve
some limited knowledge, even with imperfect methods and the related uncertainties,
than to remain in complete ignorance.
Hence predictive species distribution models cannot forecast the future but are
able to give potential reactions of species under environment change scenarios, which
may help in dertving appropriate policy recommendations.
Approaches used in the present atlas
We used generalised linear models (GLMs) to relate climatic variables to species
distributional data using a binomial error structure, which means that we need both
reliable presence and absence data and to assess the full climatic niche of a species. It
would hence be best if both northern and southern range margins are at least to some
extent defined by climate and not solely by other factors such as e.g. geographical
borders. ‘To ensure high quality models we excluded countries with low levels of
sampling and highly uneven coverage.
The southern range margins of many Mediterranean species (e.g. Danaus chrysippus,
Gegenes nostrodamus, Leptotes pirithous, Libythea celtis, Tomares ballus see Figs. D.1.1-D.1.5)
ate, however, often limited by the geographical borders of Europe and not necessarily
by climate. Here, future projections may be treated with caution; since extrapolations
to future warmer and dryer climates might be overly pessimistic in some cases as such
species’ potential performance in such climates (e.g. species that also occur in North
Africa) was not included in model parameterisation.
D. Discussion of methodological limitations 637
Figure D.1.1: Actual distribution Figure D.1.2: Actual distribution
and modelled present climatic and modelled present climatic
niche of Danaus chrysippus niche of Gegenes nostrodamus
Figure D.1.3: Actual distribution Figure D.1.4: Actual distribution
and modelled present climatic and modelled present climatic
niche of Leptotes pirithous niche of Libythea celtis
Figure D.1.5: Actual distribution Figure D.1.6: Actual distribution
and modelled present climatic and modelled present climatic
niche of Tomares ballus niche of Vanessa cardui
|
638 Climatic Risk Atlas of European Butterflies
Vanessa cardui (Fig, D.1.6) for example, is even more difficult, as this species seems to
sutvive the winter only in South Mediterranean regions while populations might vanish
there in summer, depending on re-migrants from the North. Here it would be most
important to model the future overwintering area. Unfortunately this area seems to be
situated mostly outside Europe, and little is known on this (apart from the Canaries
where this species can be found all year round). It therefore 1s of limited use to model
the total area where this species can be found, because this is the whole of Europe.
Within this atlas we present ecological niche models based on climatic variables. It
is crucial to note that the depicted current and future climatic niches do not represent
predictions of future species distributions but are projected distributions of the potentially
suitable climate space for the species. Of course there are many more factors that determine
whether a species is actually able to occupy the potentially suitable climate spaces, including
landscape structure, habitat quality, resource availability or predation and parasitic pressures
and the dispersal ability of a species. However, as applied at the coarse scale across the
whole of Europe, climate is usually best suited to explain such large scale distributional
patterns (Uhuiller et al. 2004). ‘This is supported by the generally high accuracy of our
models (1c. high AUC values for most species). However, there are exceptions such as
Lycaena helle (Fig. D.1.7), Coenonympha oedippus (Fig. D.1.8), and Euphydryas maturna (Fig, D.1.9),
whose distributions are tied closely to particular habitat conditions. Such local habitats with
specific structural and microclimatic conditions are averaged out at the resolution used
in this analysis and thus do not contribute to model development. In these cases where
habitat and microclimatic conditions are much more important determinants of species’
distributions, large scale bioclimatic niche models often fail to adequately describe and
consequently to project future species distributions. We address these concerns by taking
AUC values into account when assessing species risk (see section B4).
A high temporal or spatial mobility (e.g. good dispersal, migratory behaviour,
invasive species) may lead to misjudgement of habitat preferences. This means that
results for species like Vanessa cardui (Fig. D.1.14), Vanessa atalanta (Fig. D.1.10) or
Cacyreus marshal (Fig. D.1.11) have to be treated with caution. In many such cases
it would be much better to only include the range of permanent populations (e.g. in
Colkas croceus, Fig. D.1.12; and Lampides boeticus, Fig. D.1.13). Unfortunately, this is a
rather difficult task which might be tackled in the future, because it is often not known
whether records represent permanent populations.
Another example is Boloria titania (Fig. D.1.15), where the model was not able to
predict the occurrence of this species in the Baltic States. The inclusion of constraints
of the larval host plant Pohgonum bistorta in the process of model development
results in much better models (Schweiger et al. in press). This shows that including
other essential abiotic and biotic environmental factors can help to improve model
accuracy, but in most cases we lack crucial information about the multitude of species
interactions that may add to the effects of climate. Consequently, to be consistent
across the species, we relied solely on climatic variables in the current atlas.
There ate many different methodological approaches to develop ecological
niche models and ways to deal with issues such as host-plant constraints (Guisan &
D. Discussion of methodological limitations
Figure D.1.7: Actual distribution Figure D.1.8: Actual distribution
and modelled present climatic and modelled present climatic
niche of Lycaena helle niche of Coenonympha oedippus
Figure D.1.9: Actual distribution FigureD.1.10:Actualdistribution
and modelled present climatic and modelled present climatic
niche of Euphydryas maturna niche of Vanessa atalanta
FigureD.1.11:Actualdistribution FigureD.1.12:Actualdistribution
and modelled present climatic and modelled present climatic
niche of Cacyreus marshalli niche of Colias croceus
639
640 Climatic Risk Atlas of European Butterflies
FigureD.1.13:Actualdistribution
and modelled present climatic
niche of Lampides boeticus
FigureD.1.14: Actualdistribution
and modelled present climatic
niche of Euphyadryas cynthia
eee ore
oo my
FigureD.1.15: Actualdistribution
and modelled present climatic
niche of Boloria titania
Zimmermann 2000, Elith et al. 2006, Heikkinen et al.
2006, ‘Thuiller et al. 2008). We used GLMs despite the
fact that they did not always provide the best model
fit for the current distribution. However, their clear
and simple mathematical formulation allows highly
accurate extrapolations into new environmental space
(Elith et al. 2006, Kuhn et al. in press).
There are also many ways to evaluate model
accuracy. [hemostcommonly accepted measutes ate
Cohen’s Kappa (Cohen 1960) and AUC (Hanley &
McNeil 1982). Both measure the agreement between
observed and modelled occurrence (threshold
dependent occurrence for Kappa and threshold
independent occurrence probability for AUC) and
depend on four cases of agreement: cases where
actual occurrences were modelled correctly (right
presence) or incorrectly (false absence) and cases
where actual absences were modelled correctly
(right absences) or incorrectly (false presences).
Unfortunately, all four cases are weighted equally,
which may result in extraordinarily high accuracy
values when accuracy is dominated by just one of
the four cases (Lobo et al. 2008). For instance, a
number of species like Lycaena helle (Fig. D.1.7)
and Coenonympha oedippus (Fig. D.1.8), or Euphydryas
cynthia (Fig. D.1.14) and Bolona titania (Fig. D.1.15)
and many others (as listed in table D.1.3) have a
very restricted distribution with a high number of
absences and a low number of presences.
The model was able to predict most of the
absences correctly (right absence rate) but not
many of the presences (false absence rate).
As a consequence, the models give high AUC
values because of the dominating effect of the
right absences. However, for ecological niche
modelling, the false absence rate is of most
interest since it shows how well the model fits the
actual occurrence, while false presences can easily
be a consequence of other interacting limiting
factors. To take this into account, we provide
false absence rates as well as other values for all
modelled species in a table in Appendix 1 and the
most extreme cases in Tables D.1.1-4.
D. Discussion of methodological limitations 641
Table D.1.1: 38 European butterflies with the highest AUC values (AUC > 0.95):
(incl. overall risk categories with full and without dispersal)
i Risk
page Species categ. no
disp disp
372
410
6
0.99
2 0.99
| ER
i:
1
1
1
N
ee)
—
N
pas
ee)
170 HHH 0.99
84 | Pkebejus glandon (PRUNNER, 1798) 0.99
514 | Erebia erplyle (Peever, 1836) | ERR | 0.99
530 0.99
14
i)
0.99
ea
1S 0.98
8 ne]
194 | Lycaena ottomana (LEFEBVRE, 1830) 0.98
288 a
8
HH
HH
0.98
HH
LE
Soa
ae)
[ie
[Pontia calldice (Hownex, 1800) LR
| LR
Phebus orbitulus Pwunnen, 1798) | ER
| ER
312
Euphydryas cynthia ([SCHIFFERMULLER], 1775)
412
414
Euphydryas intermedia (MENETRIES, 1859)
R 0.98
458 R 0.98
Apatura metis (FREYER, 1829)
reed
Erebia melampus (FUESSLI, 1775)
536 0.98
554
Erebia montana (PRUNNER, 1798)
0.98
574 0.98
59 0.98
LR
LR
| AR |
| HR |
LR | HER |
ir | R | 098
|
Sree
rR R |
| AR |
612 HHHR 0.98
LR
LR
LR
LR
LR
LR
Hipparchia fatua (FREYER, 1845)
0.97
486 | Coenonympha gardetta (PRUNNER, 1798)
28 | Erebia embla (BECKLIN, 1791)
Oi
R
R
R 0.96
| oR | 097
100 | Gegenes pumilio (HOFFMANSEGG, 1804) fon
0.96
HITE 0.98
Sie]
=
Ses
| LR
[Erebia pluto Pruner, 1798) LR
Ss
ea
| LR
| ER
R
R
ted
IB ake
HR
HR
HR
R
HR 0.98
HR
HR
HR
HR
HR
HR
fed
HR
Melanargia arge (SULZER, 1776)
134. | Zeoris eupheme (EspER, 1805) HHHR 0.97
f
4 | Erynnis marloyi (BotspuvaL, 1834) jal
(L
336 | Polyommatus dolus (HUBNER, 1823)
162 | Cokas phicomone (EsPER, 1780)
YE >
R
R
R
R
R
R
R
ine
R
R
R
4 | Pyreus centaureae (RAMBUR, 1840)
R
S R
es)
| ER
[aang
ee)
| LR
136 0.96
a
Ea)
ue
esi
Sas
642 Climatic Risk Atlas of European Butterflies
Table D.1.2: 34 European butterflies with the highest Kappa values (> 0.54):
(incl. overall risk categories with full and without dispersal)
i Risk
page Species categ. full] categ.no | Kappa
dis disp
286 | Pkbejus aquilo (BOISDUVAL, 1832) HHHR | HHHR 0.78
170 | Cokas hecla LEFEBVRE, 1836 FELELRY | EEE: 0.77
372 | Boloria charicla (SCHNEIDER, 1794) fel teiebe) | MERE: OS
4 | Pyreus centaureae (RAMBUR, 1840) Ore
28 | Erebia embla (BECKLIN, 1791)
0.70
530. | Erebia disa (BECKLIN, 1791) EVER Ss) ete: 0.72
N
0.68
0.64
eae ee)
Ses)
16 | Oeneisjutta Hopnen, 1806) ERR
PR | OR | 066
a
| ER
0.64
164 | Cohas palaeno (LINNAEUS, 1758)
514 0.64
R
288
Plebejus orbitulus (PRUNNER, 1798)
0.64
380 0.64
i es 0.63
0.63
0.63
0.62
0.62
} R
R
} R
R
R
Enis ne
ea ares
Es ors
a
en eo
[|
[ener rd
ar [|
as pe
Tae |
es
: Te [om
: Tae | me _
: en eres
ee ee
ars eel
Cae | io
R
R
R
R
R
R
282
Plebejus optilete (IKNOCH, 1781)
284 H
564 Re
374
0
158
384
4
Erebia pandrose (BORKHAUSEN, 1788)
0.60
0.60
0.60
0.59
059
410 | Euphydryas iduna (DALMAN, 1816) HHHR
Erebia eriphyle (FREYER, 1836)
184
Gonepteryx cleopatra (LINNAEUS, 1767)
192
Bolona freija (BECKLIN, 1791)
Lycaena virgaureae (LINNAEUS, 1758)
N
1h
Oecnets glacials (MOLL, 1785)
Parnassius phoebus (Fprictus, 1793)
88 | Carterocephalus silicolus (MEIGEN, 1829)
0.59
0.58
358 | Brenthis ino (ROTTEMBURG, 1775)
554 | Erebia montana (PRUNNER, 1798)
502 | Aphantopus hyperantus (LINNAEUS, 1758)
R
R
R
R
Ts
R
R
R
R
R
R
R
0.58
0.58
612 | Oeners norna (BECKLIN, 1791) HHHAR | HHHAR
408 | Araschnia levana (LINNAEUS, 1758) 0.58
364 | Boloria eunomia (EsPER, 1799) R
0.57
Boloria aquilonaris (STICHEL, 1908)
484 | Coenonympha ghverion (BORKHAUSEN, 1788)
R
R
Bolonia frigga (BECKLIN, 1791)
R
R
R
R
R
R
H
ike
Ke
Ke
510 | Erebia iigea (LINNAEUS, 1758) 0.55
Plebejus glandon (PRUNNER, 1798)
ih
ike in
il H
ile ile
dy H
ae ib
H H
eh
Al A!
Ee H
H
iE ial
H
isl
Aly H
ile H
All
ile H
ale
ile H
re ae
Ss
486_| Coenonympha gardetta (Prunnen, 1798) | ER TOR | 057
p IR | OR
| IR | WR
162 | Cokas phicomone (EsPER, 1780) R 0.55
498 | Pyronia cecilia (VALLANTIN, 1894) | oR | HHHR | 055
D. Discussion of methodological limitations 643
Table D.1.3: 44 European butterflies with the highest percentage of false absences (> 0.55):
(incl. overall risk categories with full and without dispersal)
Risk
Species categ. full | categ. no
i di
diel
false
Lote abs
R
-f
_
D
N
OQ
56
386
542
4
H 0.90
0.84
| HER |
ER
0.83
eal
0.82
H
H
Scolitantides bavius (EVERSMANN, 1832)
Bolonia graeca (STAUDINGER, 1870)
Erebia epistygne (HUBNER, 1819)
Carcharodus baeticus (RAMBUR, 1840)
2 | Ania anteros (FREYER, 1838)
6 | Speaka phlomidis (HERRICH-SCHAFFER, 1845)
34 | Polommatus ripartii (FREYER, 1830)
80 | Coenonympha rhodopensis Exwes, 1900
544
202 | Lycaena candens (HERRICH-SCHAFFER, 1844)
174 | Cohas chrysotheme (EsPER, 1780)
240 | Cupido osiris (MEIGEN, 1829)
188
558
PA
8
94
108
124
332
290
148
8
.
9
36
z
iE
H
dist
Pp
z
Erebia ottomana (HERRICH-SCHAFFER, 1847)
R
‘el
| 544 |
| 202. |
Bical
z
z
| HR | 0.75
Lycaena helle ((SCHIFFERMULLER], 1775)
Erebia melas (HERBST, 1796)
Tomares ballus (Fasrictus, 1787)
Pyrgus sidae (ESPER, 1782)
Fipparchia volgensis (MAZOCHIN-PORSHNYAKOV, 1952)
R
R
alle
R
R
R
R
R
R
1K
R
R
Ie
R
HR
R
R
R
R
R
R
R
R
R
R
R
R
R
R
On
;
On
XS)
On
HHR 0.69
Zerynthia cerisyt (GODART, 1822)
Leptidea duponcheh (SYAUDINGER, 1871)
Polyommatus admetus (EsreR, 1785)
IP 0.69
R
R
0.67
| HER |
0.69
RS]
| TI |
Plkebejus sephirus (FRIVALDSZKY, 1835) R 0.67
Pieris mannit (MAYER, 1851)
Carcharodus lavatherae (ESPER, 1783)
Coenonympha oedippus (FABRICIUS, 1787)
Coenonympha leander (ESPER, 1784)
Polyommatus dolus (HUBNER, 1823)
94 | Lycaena ottomana (LEFEBVRE, 1830)
216 | Callophrys avis CHAPMAN, 1909
412 | Exphydryas cynthia (|SCHIFFERMULLER], 1775)
100
602
560
416
De
tat
da
P
P
0.72
Scare
P
P
iS
Ro
Jal
R
Jal
R
R
H
H
R
R
Jel
IR
R
H
R
jel
Jal
R
R
R
ix
z
ils
ie
I
lle
lle
ils
is
i
i
iB
le
Ny
ee 0.64
, R 0.63
R
Ie
Ie
ls
R
Is
R
L
I
R
R
R
let
H
a
eal
H
lel
H
H
z
Gegenes pumilio (HOFFMANSEGG, 1804) 0.62
Fipparchia senthes (FRUHSTORFER, 1908) | HHR | 0.61
Erebia oeme (EsPer, 1805) eae
Euphydryas maturna (LINNAEUS, 1758) Te
Erebia styx (FREYER, 1834)
Erebia casstoides (REINER & HOHENWARTH, 1792)
(species complex)
R
R
0.61
ily 0.60
H
lal
On
ra =P oOTHR ITA] oO
oe) N | oo
H
R
|
338
oe
Polyommatus damon (|SCHIFFERMULLER], 1775)
=
Ss
S
S
ise
S
>
x.
S
is
a
to
fy
—
~I
Qo
em
I
>)
644 Climatic Risk Atlas of European Butterflies
4
Cupido alcetas (HOFFMANSEGG, 1804)
Papilio alexanor Esper, 1799
Gonepteryx farinosa ZELLER, 1847
Melanargia russiae (ESPER, 1784)
Euchloe belemia (EsPER, 1798)
Boloria titania (Esrer, 1793)
Mehtaea varia (MEYER-DOR, 1851)
Table D.1.4: 36 European butterflies with the highest percentage of false presences (> 0.39):
(incl. overall risk categories with full and without dispersal)
i Risk
p disp
144
150 | Prerts rapae (LINNAEUS, 1758)
154 | Prerts napi (LINNAEUS, 1758)
466
388
494
248
504 | Maniola jurtina (LINNAEUS, 1758)
Papilio machaon LINNAEUS, 1758
Callophrys rabi (LINNAEUS, 1758)
Abporia crataegi (LINNAEUS, 1758)
fi
390 | Vanessa cardui (LINNAEUS, 1758)
396
296 | Ania agestis ([SCHIFFERMULLER], 1775)
402
Leptidea sinapis (LINNAEUS, 1758) & Leptidea real
REISSINGER, 1990 (species complex)
i |
iS
Oe
66 | Boloria euphrosyne (LINNAEUS, 1758)
76 | Plebejus argus (LINNAEUS, 1758)
168 | Cokas croceus (GEOFFROY, 1785)
262 | Glaucopsyche alexis (PODA, 1761)
Thymehcus lineola (QCHSENHEIMER, 1806)
0.47
[rr [045
[045
PR
By
BR
PR
PR:
PR
PR
PR
PR
Pare
PR
PR
PR
PR
PR
ArAre
13
PR
PR
PR
ded Re
Bae
Pre
LR
PR
Pit
=
D. Discussion of methodological limitations 645
Ta | Gonpes a epwcs 5) | pe [pk [048 —
| 352 | Argynnis niobe Linnavus, 1758) | PR LPR 043
D.2 Interaction of taxonomic status and modelling results
A special problem in niche modelling is presented by sister species or sibling species.
They typically occupy very similar niches because they originated from the same
ancestors, but are normally allopatric in distribution (no overlap in their areas). If we
take the present distribution of just one of such a pair, we are bound to experience
problems in the modelling, as the niche model will encompass the entire distribution
of the climate space which may be occupied by the sister species. ‘This leads to rather
poor modelling results. If the distribution of these species is pooled, the model fits
are normally much better.
Within this atlas we repeatedly came across this problem. ‘The selection of some
species was already affected, for example when some of the sister species had a very
limited distribution (below our threshold of 20 UTM grids) and thus have been
excluded from our analysis. However, their climate space might have been modelled
through the other sister or sibling species.
We have selected the following examples to illustrate some of the effects:
First we have a more detailed look at the Ewchloe ausonia complex. The maps and
results presented in part C.2 include Ewchloe ausonta (s.str.), Euchloe cramert, and Euchloe
simplonia. lf we differentiate between the results for the complex and the two three
included here, we can see substantial differences in the output. ‘These are summarized
in table D.2.1 and also presented as maps (Figures D.2.1-4).
The results suggest that E. ausonia (s.stt.) would suffer much less from
climate change and under all scenarios would gain in climate space assuming
full dispersal. In contrast, the climate space of FE. crameri and E. simplonia would
decrease under nearly all scenarios. The models for the three species together lies
in between these results. Although there are large areas of false presences, these
are compensated by right presences and right absences in E. crameriand E. ausonia
(LEr8-) 9bb- | (Yocr'rb-) SET- | (%98°S6-) LOOP- | (ohE'OL-) 9E6T-]| (%H6'TE-) LBL-| (YbB'STL) ILLT | CHbT'SS-) TLHTZ- | (%16'S-) S9Z- SVUD
(%66°SL>) ZOP- | (ETT) ELI- | CESSES) BLbE-| oL'6S>) ZobZ- | (%HLT9Z-) 9Z9-| (PSL) ELT | (LG'Sb-) BS0Z- | (%67'L-) LZE- | nave
(ples) ebb | (SESE) LBI- | ColF'69-) LORZ-| (66'S) Sho] (ELI) 6It-] BSE) O9LL| (LTE) otI- |r Bee | OCS]
(%19'L9-) 85¢- | (%6ELI) Z6- | (obeLS-) 68Ez-| (H9ELZ) ZHII- | (%LETI-) 96z-| (68TH) 9ZOI | (%S6'7-) 6ZOT- (aLoret) 98s | SVED.
(moos) oe- [meet 19 — | ores) toze-| Carre) 9er1- | Corre) zee-]_ @oes'ze) 8LL| (o'r) O801-] (oe07) 16- | aavE|
(%61'79-) 67E- (68°) O1- | @8S'LP-) 9861- | (%89'E1-) LLS- | (Es'tI-) es] (% 19°28) P08 | (68 LI) ZO8- (ngs) tL | Saas
essodstp oN, | yessodsip Jn | essodstp on, | [essodstp qn | [essodstp on | yessodstp Jn | Jestodstp on, | [essodstp NY [oo]
s]J9o
67S bLib 76ET 8th pis pordnos0
Apussoig
piuojduts aojqgony LLIUD AD 90]GINT “S'S DIUOSND 30]GINT xajduros pruosnp aojqony Sse
Ajayeredas eiuojduis aojyonyg
pue VWaWeID ao/YONF ‘(@Suas }OUjS Ou} Ul) BUOSNe BojyONF pue x9jdwod e/uOsNe BsojYyONF du} JO} Huljjapow syewjo Jo syNsey :1°S'G sIGeL
Climatic Risk Atlas of European Butterflies
646
D. Discussion of methodological imitations 647
(s.str.). In the EF. ausonia complex and in E. simplonia there are less false presences
but more false absences. This leads to the effect that the AUC values are rather
similar in all cases.
Few false absences
In many instances we come across sister species that can be modelled quite well
independently from each other, with little overlap between the derived climate niches.
A good example 1s Scohtantides baton and Scolitantides vicrama, (Yable D.2.2. and Figures
D.2.5 & 6). Here the models show differences in climate requirements which suggests
different evolutionary histories for the two species.
Similar in terms of output statistics are the sister species Lycaena hippothoe and Lycaena
candens (Yable D.2.3; Figures D.2.7-8). However the map of the absolute distribution
is very different between the two species and the distribution points of L. candens are
to a large extent included 1n the climate space of L. hippothoe. ‘This contributes to the
large number of false presences which in general lead to the low AUC. The climate
space model of L. candens 1s relatively poor if we look at right presences as well as false
absences and the high AUC 1s due to the many right absences across Europe. Here
a combination of both species would clearly lead to an improvement of the climate
models and highlights the “disadvantage” of the separate modelling of sister species
which we can observe in several other cases. ‘Two other examples of sister species ate
shown below: a) Spzaka sertorius and Spiaha orbifer (Lable D.2.4; Figures D.2.9-10); and
b) Hipparchia semele and Hipparchia senthes (Yable D.2.5; Figures D.2.11-12).
Table D.2.2: Results of climate modelling for the sister species Scolitantides baton and
Scolitantides vicrama
a Scolitantides baton Scolitantides vicrama
Presently
occupied grid
el Full baa No AEBS Full soe No sreeens
648
Climatic Risk Atlas of European Butterflies
Figure D.2.1: Actual distribution
and modelled present climatic niche
of the Euchloe ausonia complex
7
Figure D.2.3: Actual distribution
and modelled present climatic
niche of Euchloe crameri
Figure D.2.5: Actual distribution
and modelled present climatic
niche of Scolitantides baton
=
Figure D.2.2: Actual distribution and
modelled present climatic niche of
Euchloe ausonia (in the strict sense)
Figure D.2.4: Actual distribution
and modelled present climatic
niche of Euchloe simplonia
Figure D.2.6: Actual distribution
and modelled present climatic
niche of Scolitantides vicrama
D. Discussion of methodological limitations
649
Table D.2.3: Results of climate modelling for the sister species Lycaena hippothoe and
Lycaena candens
[_ ES Lycaena hippothoe Lycaena candens
Presently
occupied grid 14109
cells
| Fulldispersal_| No dispersal | Full dispersal
Table D.2.4: Results of climate modelling for the sister species Spialia sertorius and Spialia
orbifer
P| Spicatia sertorius | Spialia orbifer
occupied grid 6768 2728
cells
| Fullidispersal_| No dispersal_| Full dispersal_| No dispersal_|
Table D.2.5: Results of climate modelling for the sister species Hipparchia semele and
Hipparchia senthes
a Hipparchia semele Hipparchia senthes
Presently
occupied grid 13833 840
cells
73
:
a | Full dispersal No dispersal Full dispersal No dispersal
650
Climatic Risk Atlas of European Butterflies
Figure D.2.7: Actual distribution
and modelled present climatic
niche of Lycaena hippothoe
Figure D.2.9: Actual distribution
and modelled present climatic
niche of Spialia sertorius
FigureD.2.11:Actualdistribution
and modelled present climatic
niche of Hipparchia semele
Figure D.2.8: Actual distribution
and modelled present climatic
niche of Lycaena candens
FigureD.2.10:Actualdistribution
and modelled present climatic
niche of Spialia orbifer
FigureD.2.12: Actualdistribution
and modelled present climatic
niche of Hipparchia senthes
Jk;
——
OUTLOOK: CLIMATE CHANGE AND
BUTTERFLY CONSERVATION
652 Climatic Risk Atlas of European Butterflies
E.1 Direct and indirect climate change impacts on
butterflies and biodiversity
Climate change affects biodiversity and ecosystem processes both directly and
indirectly. An example of a direct effect is when critical climate limits are altered,
which directly affect species physiological mechanisms (e.g. winter temperatures or
annual or seasonal precipitation patterns). Indirect effects can be through complex
interactions with other processes, such as invasion of exotics, loss of pollinators and
interactions with environmental chemicals. Further responses to climate are due to
interactions with land use and land use change.
Habitats and trophic interactions
Biodiversity responds to the distribution of habitats as well as to climate factors (and
other pressures), so understanding habitat change is also critical. Climate change
may alter: (1) the distribution of clmatically-suitable areas for a given species, and
thus change the geographic match between otherwise suitable habitats and climate
(potentially endangering the species), (11) the distribution of suitable habitats, for
example if vegetation structure 1s affected by climate, (11) the distribution of land-use,
and (tv) what is currently recognised as suitable habitat for a given species may change
if habitat choice 1s at least partly determined by thermal environment. In addition, (v)
land-use may change for reasons unconnected to climate.
Although we do not expect particular habitats to move with climate change as
intact collections of species, climate-driven changes to the boundaries of structural
elements of the vegetation (e.g., of evergreen and deciduous forest; tree lines) may
be as predictable as the boundaries of individual species. Such changes would have
major implications for the community structure of all taxonomic groups. A map of
the changes of the potential natural vegetation (PNV) under the BAMBU scenario is
shown if Figure E.1.1 for the presence and the conditions around 2080.
Many of the other habitat/trophic interactions mentioned above have been
researched in recent years with the project ALARM (Settele et al. 2005, 2007,
2008). In the context of butterflies, analyses have been started to study effects
of climate change in combination with other factors. An example is the study
on Boloria titania (see page 638ff). Here the model was not able to predict the
occurrence of this species in the Baltic States, but the model became far more
accurate when the constraints of the larval host plant Pojgonum bistorta were
included (Schweiger et al. in press). This shows that including other essential abiotic
and biotic environmental factors can help to improve model accuracy, but in most
E. Outlook: Climate change and butterfly conservation 653
Anchclalpie desert
ATCEC/alpepe burda
* Boreal/alpine mixed woodland
® Borealalping conifer forest
Heriboreal mixed forms
= Temperate beech and mixed beech forest
® Temperate mixed broadleaved fonast
Thermophitous mined broad-leaved forest
© Mediterancan sclerophyiiceus foresiwoodland
© Mediterancan sclemphyfous shrubland
® Steppe woodland
Sleppe
Fig. E.1.1: Modelled current (averaged for 1961-1990) and future (2071-2100) potential
natural vegetation (PNV) in Europe under the BAMBU (IPCC A2; HadCM3) emission
scenario; from Hickler et al. (in press, modified).
cases we lack crucial information about the multitude of species interactions that
may add to the effects of climate.
Climate envelopes for European butterflies as a starting point for future
research and conservation
“Pure” climate change models as presented in this atlas are only a first, but very
important step. Until more sophisticated methods and models become available, we
have to rely on what we have in order to draw conclusions for conservation actions.
Moreover, until we determine exactly how climate change (in concert with other
factors) will affect biodiversity, it is sensible to consider worst case scenarios under the
cautionary principle. Consequently, we have selected the worst case result under all the
given scenarios until 2080 for our classification of climate risk.
However, even under a very conservative classification of risk, the vast majority of
European butterfly species has to be regarded to be at risk through climate change. An
important next step will thus be to test model outputs against field data for butterflies,
as these ate a group of species that are expected to track climate changes quickly due
to their sensitivity and short generation time. In particular, we can expect the habitat
generalists and mobile species to be the first and quickest to respond.
The literature already contains many examples of apparent responses to climate
change (e.g, Walther et al. 2002, Parmesan & Yohe 2003), but genuine tests against
model predictions are largely lacking. If we have a closer look at our simulation results,
which are based on the distribution of species from roughly 1980 to 2000, we can see
that some predictions seem to have become true already. Good examples are
° Cacyreus marshalh, which has already spread across Italy;
° Plebejus argiades, which has moved northward e.g. in Southern Germany;
654 Climatic Risk Atlas of European Butterflies
° Pieris mannit, which 1n 2008 has suddenly expanded its Northern range limit in
Switzerland and in August 2008 has reached Germany;
° Brenthis daphne, which has extended its range from Alsace to Southwest
Germany since the mid 1990s.
In future studies considerable attention will have to be paid to the development
of a global climate change envelope model and model testing, because statistically
appropriate and ecologically plausible predictive models are a key prerequisite for
the assessment of the biodiversity risks resulting from the projected climate change.
Models will also have to be improved to assess impacts of climate and landscape
change scenarios on biodiversity at the European scale, assessing the role of species
characteristics in projected susceptibility. Physiological traits (e.g., temperature
thresholds for development; evapotranspiration) will have to be considered as well as
functional characteristics, such as the types of habitats occupied and dispersal capacity
(Warren et al. 2001, Luoto & Hetkkinnen 2008).
Climate change and evolution
Recent reviews and meta-analyses of aquatic and terrestrial studies from throughout
the world have revealed consistent evidence of climate change effects on species. These
include the advancement of the timing of life-cycles (phenology), northward expansions,
southern retractions, movement up mountain slopes, and abundance changes such that
species with southerly distributions have tended to increase in the northern hemisphere,
whilst northerly distributed species have declined (e.g. Parmesan 1996, Parmesan et al.
1999, Walther et al. 2001, Parmesan & Yohe 2003, Wilson et al. 2005). Furthermore,
recent evolutionary shifts in phenology, habitat choice and migration direction are also
consistent with a response to climate warming (Berthold et al. 1992, Berthold & Pulido
1994, Berthold 1998, Bradshaw & Holzapfel 2001, Thomas et al. 2001).
At the receding edge of a species range, local populations are forced “outside the
niche”. This raises the question whether sufficient evolutionary speed 1s available to
rescue such a species which clearly faces extinction (Pease et al. 1989, Holt 1990, 2008,
Gomulkiewicz & Holt 1995)? Here we need deeper insight to understand both niche
conservatism (species seem to have much the same niche limits over a broad geographical
range or over long periods of evolutionary history; Bradshaw 1991, Holt & Gaines
1992, Wiens & Graham 2005) and rapid niche evolution (Reznick & Ghalambor 2001).
Butterflies can evolve quite rapidly in ways that could influence their distributional
limits. There are examples of species shifting host preferences over periods of less
than ten years (Singer et al. 1993), and if these host species have different geographical
distributions, a large shift in the butterfly’s range can easily be expected. Rapid evolution
of dispersal abilities is likewise known from butterflies (Mill et al. 1999), with spreading
species developing both a larger thorax and a greater ability to fly in newly colonised
patches. Geographical ranges reflect both niches and dispersal abilities, which can each
E. Outlook: Climate change and butterfly conservation
evolve and lead to range shifts. In other cases, butterfly niches can be maintained,
even over very large ranges (Crozier & Dwyer 2006). In a study of a North-American
skipper, one basic aspect of the species’ niche — the thermal environments in which it
can persist, versus where it declines toward extinction - appears to have been conserved
ovet an enormous distance (Holt 2008).
Understanding the reasons and mechanisms of niche conservatism, and anticipating
when to expect rapid niche evolution instead, is of crucial importance. However, our
current level of understanding is extremely limited. Much of the literature on niche
conservatism 1s essentially phenomenological in nature, reporting correlations between
species distributions and various environmental attributes (“ecological niche models”).
This provides an essential starting point but, according to Holt (2008), what we really
need is a deeper mechanistic understanding of the factors that either constrain or
facilitate niche evolution. He states that “this understanding requires one to take a highly
integrative approach to science, as explanations for niche conservatism can reflect a
wide spectrum of forces and constraints, from limitations on genetic variation (Blows &
Hoffmann 2005), to tradeoffs emerging from how organisms are engineered from the
gene to the whole phenotype (Hansen & Houle 2004), to the details of demography and
spatial movement patterns (Holt & Gaines 1992, Holt 1996, Kawecki 1995), to the nexus
of interspecific interactions (Ackerly 2003, Case et al. 2005).” (Holt 2008, p. 3).
We sincerely hope that this atlas also serves as a tool to encourage research into these
topics, as butterflies seem to be an ideal group of organisms to study. We know a lot about
the species, they can be raised in the laboratory (genetic and functional studies of traits),
they have short generations, they are popular and diurnal, and they are ectotherms (hence
sensitive to thermal conditions). They also typically have close interactions with other
species (host plants, predators, parasitoids; e.g, van Nouhuys & Hanski 2004, Anton
et al. 2007, Schweiger et al. in press), which makes the interspecific dimensions of their
niches amenable to quantitative modelling (e.g., Mouquet et al. 2005, Johst et al. 2006).
Thus Holt (2008) suggests that these traits collectively make butterflies potentially highly
useful in developing a deeper understanding of the phenomenon of niche conservatism,
which not only would be a satisfying intellectual endeavour tn its own right, but also of
vital importance for conservation (see chapter E.3, page 658f.).
Biodiversity Risk Assessment
Our study has also shown that biodiversity risk assessment to environmental
changes is heavily based on species modelling for current and future conditions
(e.g. Segurado & Araujo 2004, Thuiller et al. 2003) and conservation planning for
biodiversity (e.g. Araujo & Williams 2000, Araujo & Williams 2001, Araujo 2002). The
first vulnerability assessments of European biodiversity were undertaken in a global
change context, but the challenge is now turning these broad-scale assessments of
vulnerability into probabilistic assessments of risk at the regional (e.g. national) and
local scales at which mitigation strategies must be developed.
655
656 Climatic Risk Atlas of European Butterflies
E.2 Butterflies as indicators of environmental change
Butterfly indicator developments
Government representatives at the 2002 World Summit of Sustainable Development
pledged ‘a significant reduction in the current rate of biodiversity loss by 2010’. ‘The
commitment of the EU to protecting biodiversity is even stronger by aiming at halting
biodiversity loss by 2010 (Balmford et al. 2005, Gregory et al. 2005). Butterflies may be
useful as biodiversity indicators for reporting on the development towards the EU 2010
target. Unlike most other groups of insects, butterflies have considerable resonance with
both the general public and decision-makers (Kiihn et al. 2008a). Butterflies are also
relatively easy to recognize and data on butterflies have been collected for a long time and
by many voluntary observers. The method of monitoring butterflies is well described,
extensively tested and scientifically sound (Pollard 1977, Pollard & Yates 1993).
As a result butterflies are the only invertebrate taxon for which it is currently possible
to estimate rates of decline in many parts of the world (de Heer et al. 2005, Thomas 2005).
However, butterflies can only be regarded as good biodiversity indicators if it is possible to
generalise thetr trends to a broader set of species groups (Gregory et al. 2005). There is still
some debate on how well butterflies meet this criterion. Hambler & Speight (1996, 2004)
claimed that this group 1s likely to experience greater declines than other organisms due to
their herbivorous life strategies and thermophily, but Thomas & Clarke (2004) convincingly
rejected both arguments. Based on a comprehensive review of studies into their life-history
traits, relative sensitivity to climate change, and adjusted extinction rates, Thomas (2005)
concluded that butterflies may be considered representative indicators of trends observed
in most other terrestrial insects, which together form a major part of biodiversity.
Trends per butterfly species can be combined into a unified measure of biodiversity.
To achieve this, Van Swaay et al. (in press) followed Gregory et al. (2005) 1n averaging
indices of species to give each species an equal weight 1n the resulting indicators. When
positive and negative changes of indices are in balance, we would expect their mean to
remain stable. If more species decline than increase, the mean should go down and vice
versa. Thus, the mean of such an index 1s considered a measure of biodiversity change.
The results of national butterfly monitoring schemes may be combined to create
an indicator at a supra-national level (see also Henry et al. in press). Based on the
procedure described for European birds (see Gregory et al. 2005), a preliminary
erassland butterfly indicator has been developed (Van Swaay & Van Strien 2005). The
countties covered were mainly from Western Europe. The results showed that average
erassland butterfly abundance has declined by almost 50% during the last 15 years,
which is most probably linked with the agricultural intensification in Western Europe
(Van Swaay & Warren 1999, Gregory et al. 2005). The decline is much stronger than
E. Outlook: Climate change and butterfly conservation
the decline of the farmland bird indicator, which has fallen by 19% in the same period
(Gregory et al. 2008). This corresponds with the findings in the UK where butterflies
have experienced greater losses than birds (Thomas et al. 2004).
The European Environmental Agency has already recommended the development
of European butterfly indicators (EEA 2007a), and these may lead to indicators that
are comparable to the farmland bird indicator, which has been adopted as a biodiversity
indicator by the EU (Gregory et al. 2005). ‘The addition of a butterfly indicator would
be valuable to complement the bird indicator to report against political targets (e.g. the
EU’s 2010 target), because butterflies operate at smaller spatial scales and more closely
represent insects.
The grassland butterfly indicator offers the possibility to detect large scale effects
of either abandonment of agricultural land (especially occurring in Eastern and
Southern Europe) or intensification of agricultural practices, a process that has slowed
in parts of Western Europe, but is ongoing in many other European regions.
Butterflies as climate change indicators
Recent climate change has already affected the distributions of many species (Hill et al.
2001, Walther et al. 2002, Parmesan & Yohe 2003, Wilson et al. 2005, Franco et al. 2006,
Fickling et al. 2006) but future changes are likely to have even more severe impacts
(Sala et al. 2000, Thomas et al. 2004, Uhuiller et al. 2005, Araujo et al. 2006, Broenniman
et al. 2006). These impacts are often assessed with bioclimatic envelope models which
relate the current distribution of species to climatic variables to derive projected future
distributions under climate change (e.g. Huntley et al. 2004, Heikkinen et al. 2006).
The restriction of studies just to climatic variables has been criticized by some
authors (e.g. Davis et al. 1998, Pearson & Dawson 2003) and there have been calls
for the consideration of other factors that determine species distributions such as
dispersal, land cover, and biotic interactions (Guisan & Thuiller 2005, Ohlemuller et
al. 2006, Ibanez et al. 2006, Heikkinen et al. 2006, Schweiger et al. in press). However,
we decided to base the current atlas entirely on climatic variables in order to provide
a complete overview of potential climate change impacts for a group of ectothermic
insects which are expected to react relatively quickly on climatic changes.
In addition to the grassland butterfly indicator, there is a good possibility of
producing a climate change indicator for European butterflies that would summarise
trends in species whose distributions may be most affected by climate change. Similar
indicators are also in progress for European birds (Gregory et al. 2007).
The development of the indicator can, among others, be based on the results of the
present atlas, which highlights species for which we might expect changes within longer time
periods. As the relationship between abundance and distribution of species ts well supported
scientifically, we can test whether species that are expected to gain niche space actually
increase in abundance within their core areas of distribution, and where there ate decreases
in abundance towards the trailing (mostly southerly or low altitude) edge of distribution.
657
658 Climatic Risk Atlas of European Butterflies
E.3 Climate change and butterfly conservation
The results of this atlas show that climate change 1s likely to have a profound effect
on European butterflies. Although some aspects may seem unstoppable, there are still
some ways to mitigate some of the negative impacts.
1) Maintain large populations in diverse habitats
In order to allow species to adapt and give them more time to evolve, we should
adjust land use and management practices to maintain large populations of
butterflies and create or maintain diverse microclimatic conditions which could
mitigate climate change effects on the larger scale. There are many examples of
how microclimatic conditions are able to support populations of butterflies in
areas that are by macroclimatic standards far out of their climatic niche. Good
examples are species like Lycaena helle, Conenonympha oedippus, Boloria eunomia and
Boloria aquilonanis. We need to ensure effective conservation of existing protected
areas and important habitats and manage them to maintain large, diverse
populations. The deliberate creation of habitat heterogeneity within such sites
may also give species scope to shift within their habitats and move to cooler
microclimatic conditions. ‘To achieve this on the ground we need well resourced
and targeted agri-environment schemes, and place biodiversity at the heart of
forestry and land use policies.
2) Encourage mobility across the landscape
We need to ensure that we reduce, and 1f possible remove, any barriers to dispersal
in the landscape. Many butterfly habitats are now highly fragmented and we should
place far more effort on habitat restoration and improving the links between
habitat patches to give butterflies a chance to move and respond to climate change.
We need to place far more emphasis on the conservation of whole landscapes
and to build ecosystem resilience and connectivity — 1n particular in connection
with the Natura 2000 network and its coherence. Well resourced and targeted
agti-environment schemes are essential in delivering on the ground, together with
enlightened planning policies.
3) Reduce emissions of greenhouse gasses
There is a great opportunity to influence the general policy on reducing
ereenhouse gas emissions (as generally described in the SEDG scenario; see
chapter A.2, page 13f.). As we have mentioned in the methods, the results of
our scenarios are not actual predictions and we should use them for what they
ate most suited: to compare different overall futures. If we translate the overall
E. Outlook: Climate change and butterfly conservation 659
pictures presented in chapter C.4 (page 629ff.) into action for conservation, the
vast majority of species would have a more sustainable future under the SEDG
scenario compared to the futures under GRAS.
4) Allow maximum time for species adaptation
The results show us that the climate change risks for butterflies increase much
faster after a certain lag phase. This could give us hope that if we take immediate
action there is a chance to avoid some of the worst effects on the majority of our
butterfly species — and most probably on other aspects of biodiversity! Our hope
is that we can direct environmental change to encourage as much adaptation by
natural selection as possible, which might contribute to the evolutionary rescue of
species. Given the pace of environmental change we are inflicting upon the rest
of the diversity on the planet, the living world needs all the help it can get (Holt,
2008) — and we hope that we can contribute to this through this atlas, particularly
as a tisk communication tool.
5) Conduct further research on climate change and its impacts on biodiversity
Further research is needed so that we continue to improve our understanding
of climate change and its impacts on biodiversity. The findings will be vital to
improve our adaptation strategies in the future. This study is an important step in
understanding the impact on butterflies, but highlights the limitations of current
models. In particular, we need to develop models for the approx. 150 species that
have such restricted distributions that they could not be reasonably modelled using
the current method.
F
—————
APPENDICES, REFERENCES
AND INDEX
662 Climatic Risk Atlas of European Butterflies
Appendix 1 — Model performance
663
Appendix 1
Poros [ero [avo [eso [aro | a0 | seo 100 | RR Ca RC SIND I / STE SHAT RHA
Se] NS | 5OOE | | GN | Ste | (NS 00] For NT || Ssh || NS" | 00 Ay
1G) BEG) S| ALG) AUG) NS NS 3 | INS 5 ENS NS eee abs | bene ea CO] 0
Climatic Risk Atlas of European Butterflies
664
s]J29 potdnss0
Eu
co
redex
Tro
(xoqduros) (9Q8] “ANAOEL) vewosup sojqouTy
(Q6LI “Uadsy) vemajog aojqonrzy
(GOST ‘asdsy) aaagdua suusozy
[SQ] ‘aH VHOS-HORIMAL] MaUMasd SUBGIOGIUTZ eer
698] AAONIAAV.LS Sapeouag dua suupq20qjupy oer
(QGL] ‘SQUVNNI'Y) Saueuppins supqooqjuEy Rea
(L881 ‘NOINA}) Zasvom vopydrT | 9Z1
(LL8L UAONIANV.LS) Yaqouodup vapudry | pZL
(xofduros) 66] WAONISsRY Had / (BGLT ‘SAAVNNIT) sedouss vapydry | TI
66L] “Uaasy sounxan oyrdey | OTI
QG/] ‘SAHVNNYY “ovgovu oyrdrg | 811
(SLT ‘SOaVNNIT) sumqopod sopyaq dt | OLT
(QGL] SSAAVNNI'Y) o7odp suissvusng | pT
(COLI ‘snioriavy)) sugaogd sumssvusvy | TY
oO
loo)
—
(QG/] ‘SQUVNNI'Y) audsomoum suissvusny | OTL
(ZZ8I “IAVAOD) asusa vaqgu4a7 | BOT
(SLL ‘[waTiowaainos)) vuaxtyod viqqud4a7 | QT
(LOLI SSANVNNI'Y) Duan viqqud4a7 | FO
(COLT ‘snIoniavy) sumppomsou sauasory | TOT
(POST ‘DDASNVIWIOP) oymund souadary | (KOT
(LLL “taasy) suunyds sapozgaQ | 36
(QGLT ‘SQHVNNI‘]) Ducdoo vuadsaty
oO
nN
(GLLT ‘AHOUNEINALLOY) Hoan snoyauigy.|
(LOLT ‘vaog) sagsauts snoyaulg | 76
(9081 “USINISHNASHOQ) yoauy suoyaud | 0
(67ST ‘NSO Supornyes sruypgqo04sajAvy | QQ
(LLLI ‘sVTIVg) “oman suvyqar01gavy) | 98
Pav SSSSSCSC*i
665
Appendix 1
(QGL] ‘SQUVNNI'T) avaimvouns vuenal T
(COST “HIMoMvy) odsip puanat]
(GLLL ‘ea TiIonesetHos)) aaq vuavaT
(TOLL ‘SQaVNNI']) svarygd vuanal T
(LOLT ‘SQaWNNI'T) ayodog xtaagqou0y
Lygy aria, vsoutavl xilsagqauory
(GL ‘SQUVNNI‘]) ampgs X4ayPauory
GOGT “AMA Seswaunavfy svyoy
(8GLT ‘SANVNNI'T) yolg svyoD
(OSL “WHasy) aacaqzosuga spy 07D
(OSLI Wadsy) aopiuslu svyory
OCR] “ANACHAA] 0770q SVY0D
(SQL SAOWNHOND) Suar0u9 spyOy
(SORT SUAS) aynua svyoy
(QGL] ‘SAHVNNI‘Y) ouarynd svyoy
(OSL “adasy) auomongd svy0D
(xojdusos) (27. ] ‘snioniavy) esmpa / (GLI ‘SQaVNNI’]) ampydop vyuog
(OO8T WANAOED) apyyr) yuo
(L6OLT “MANaNE) aweuolag seer
(8GL] ‘SANVNNI'‘]) 20 Susaig
(R781 UAAAD) aera seat
(QGL ‘SASVNNI'T) aga sada
(TGQT “UMAVIN) Zape Svat
(NOT AHONIGAVLS) aeadamsy Setar
(QG/] ‘SQAVNNI') anoussndg Siar
(QGL] ‘SQUVNNI'T) soarynus vuodE7
(POST MANINED) S207 aojqoury
| cro | sso | coo | oor | 860 | ogo | 980 | (PPR WaVHOS-HORTaAp]) swapuns ouenat T | z0z._ |
TEETER | CESS ERE |S SE
+
—_
N
Climatic Risk Atlas of European Butterflies
666
667
Appendix 1
Sigs pentane | esses) |iezal tory issess enas eee ony
(QZ6] ‘ALIN A) SesuafMoU DID | 00¢ |
(C6L] ‘SQIORIAVY) SAUANDLLD DiI | 867. |
(CLL) ‘faa rinweastatHos)) sasaap peau | 967. |
(LZQT “ZIOHOSHOSY]) Déauin4s DID Ena
(O8LT ‘Uadsy) wopamna vest Becca
(SE8l AMZSCTVARLD) seq das suloqape
(Q6L] SMANNON) symjiquo sulaqa J
(ZEegy “tvanasiog) yabn suloqa
(Q6L] ANNAN) “opunys suloqag
(I8L1 SHOONS) waydo sulagag
(6LLI SNASSVULSOUAG) “omoudoudauv sulaqaf J
(LOL] ‘SQNVNNI'T) Sope sulaqarg.
(QGLI ‘SQUVNNI'T) svouv sulogag
(CLL ‘a iIgWetHos]) woyp suvsued
(6LLI ‘WASSYULSOUAG) SwoqpsnDU suDsUaq
(GLLI ‘WASSYULSOUAG)) SHzaap SUUDTUAY J
(QSL ‘SQAVNNI‘Y) “oun suviuegg
(Q1Q] SXHNISHNASHOQ) S202 DULIOT
(Qz78q “IvAnasiog) sdourjau agatsdoourpy
(LOLT ‘vaog) saxap agasdomnjy
(CIQ] ANaAED) sardound sapyurjyoos
(LLLI SSVTIV q) Homo sapyunjyoas
(TERT SNNVINSUH AG) SHadDg sapyUviyors
(LEST “LAMM g) SMinLLaoUagD sepuyUDjy os
(c 98 “AMOOW) DULDAINA SAPLUDIYOIS
(6LLI ‘XASSVMLSOUAG) wojng sapyurjyors
(QGL] ‘SQUVNNI'Y) SHOtbuD DULSLIAD
06C
88C
98C
V8C
C8C
O8C
SLC
OLS
VLE
CLC
OLE
89C
99¢
VIC
C9C
09C
8SC
9SZ
vVSC
N
Ke)
N
OSC
8h
I8tg
Climatic Risk Atlas of European Butterflies
668
LOV
s]J20 potdnss0
Eu
sae
eldest
Tro
Tro
(LLLT ‘Svtive) apony stuuiay | $Se |
(QG/] ‘SAUVNNI'T) agom seuudoapy | zse |
(SLL ‘watiawasianos)) adgpp seuudiupy | ose |
(8G/1 ‘SASVNNI‘T) VAyaD seuudouEy | Bre |
(GLLI ‘uarrignasnos)) aopund seuuliup7 | OE |
(QGL] ‘SQAVNNIT) Legged seuul op | +re |
(Z8L SONMAVHOIV’]) S47a0 vaqggr]
(QG/ | ‘SQHUVNNI‘]) DULY StUDaUDET
(CLL ‘(aa rianeattHos]) “omvp snjouuolog
(CTBT ANALY) sop sujouuolgog
(OEQ] AAU) wend supwuuowgog
(GOL Wade) SHpaMpy suoUULo[g Og
(LSQ] “AaeTVHOS-HORTAAPY) sung suppuluolyog
(TSQ] ‘wattyHOS-HonMapy) swundsig supouuoog
(LOLT ‘vaog) wopuo suppuuolog
(SLL ‘DUNANALLOY) MoU aq SHIDO] OT
(SLL ‘[eaTioawesatnos)) seagdop suppuuotjog
(QO8] SAHWISHNASHOQ) Sada supouuolyog
(G/L SOUNAWALLOY) smn supouuMolog
(PEST AANAINV)) Saasuagy SuppuLuoly og
(Z6LI MAAIANHOS) Supuvuly snjouuoljog
(LSQ]. ‘NINLSUAAAS)) Swarsat SUpDULULON Og
(SLL ‘[aarianeaettHos]) seyaop sujoumog og
(CZQ] SMANAAED) eaqasa suppuuolyog
(CLL SDUNANALLOY) saoeuwuuas sumun’y
(6Z8T ‘NHOIAYN) S72 DiI | r0e |
(QCQ] AATM) Sosauy DIU | Zoe |
pov
669
Appendix 1
Cor
Toro
Tro
Tro
i
(GLI ‘SQUVNNI']) vunia vIuGosDAp
(OBLT ‘waasey) wnqiry syoqdutn’ | 90% |
(IQLT ‘aasy) seyaudoqzuox syog funy | 0b |
(QL. ‘SAAVNNI'T) SoxorgaGod syvqdual yy | ZOb |
(8GL1 ‘SASVNNI'T) Vdoyun syoqg dug | OOF |
(GLLT ‘watrwva) 7000 syrq dun’ | BoE |
(BSL ‘SaavNnry) aqpr-0 syoqduant | 96€ |
(GL ] ‘SQHVNNI'Y) a0gjun Seva | 76e |
(GLI ‘SQNVNNI‘T) 02 S70/0f7 | ZOE |
(QGL] ‘SAAVNNI'Y) “upavo vssauv | 06S
(QSL ‘SAAVNNI'Y) Yyunyoyp vsseuvA | QQ¢
(OL8T ‘XHONTAAV.LS) Manus vuojog | 98¢
(8061 “THHOLLS) seepucpinbp vuuojog | 8¢
(SLL ‘uatTiawaaimnos)) syed vuojog | 78¢
(LOLL ‘NImoag) radu/f ruojog | OBE
(9081 WANED) oq7 vuopog | BLE
(LOL ‘SOAVNNI'T) mp Puopog | OLE
(LOLI ‘Nitoug) haf ruopog | pL¢
(POLI SNHCINNHOS) vanuengs vuojog | TLE
(SLL ‘[WariaWwasttHos]) ages puojog | OLS
(COLI “eldsyy) PHIY DUOjOT | BOE
(GLI ‘SQUVNNI'T) amtsouqdua vuojog | 99¢
(66L1 “tHasyy) rouns ruojog | +9¢
(GLLI ‘[WaTiaNesttHos]) yoray saqquasg | TOS
(OSL SWassvULsoung) aqdop seqyuasg
(GLLI SONNANALLOY) 042 seqguasg | 8G¢
(QGL] ‘SAAVNNI'Y) YeMOgsN] LOSST | OSE
S
No)
loo)
Climatic Risk Atlas of European Butterflies
670
671
Appendix 1
(SLL ‘[WarraWweasetHos]) ofavum viqasry
No)
—
Leo
(O¢QT WAAAM]) alg dua viqassyT
(GOST ‘waasy) youn viqary
(QSL ] ‘SQAVNNI']) 2904 DigadT]
(9¢g] ‘vison) vusdy opaydouod( ty
(FLL ‘NHOS) woraG sag douod(yy
(9G/] ‘SQaVNNI']) vagal roupyy | +s |
(QG/] ‘SQUVNNI'T) saquruodg sndozuvgdg |
(C6LT ‘sntoriavy) vgasqing vinowt | OOS
(P68 ‘NLLNVTIV A) 2972709 DUO | 86h
(LLLT ‘SQavNNI']) sauoqay voit | 96h
(GL SSASVNNI'Y) supeqdud vyduduoussy | $6
(PEL ‘Uaasy) sapuny vgduauousoy | TEF
(LOLI ‘SQaVNNI']) ovag vydukuousvy | O6F
(Z8LI “UHasyy) smsop vydudtuousoy | ggy
(Q6L] SXANNON) “apne vg duuous0> | 9Rh%
(987. [ ‘NASAVHSMOg) womans nqduduous0’y | pRp
(LOLI ‘SQAVNNI']) veavup vyduuoussy | Zr
ONG] ‘SHMTH Sesuadopoqu vyduluous0y | 8p
(18L] ‘satortavy) saddipa0 vg duuous0y | Qi
(POLL aTIAWN) ey vgduduous0y | 914
(COLT ‘SQUVNNIT) aueqon vIudoT | bly
(QGL] ‘SANVNNI'Y) Manus DIDULULOLSU I. \ TL
(L8L] ‘snioriavy) vunpyodomed vpuuloisvy|) Oly
(LOL ‘SQHVNNI‘]) (ada DIDULULOLSD T
ice)
x2)
(QC). ‘SQHVNNI'T) Pusadon aunt | 99h
(LLL SMHINVUD) PUraxod DuUTY | POF
ezceal oripil| ised srry ington Ova ikea
| ss | sso | sro | 100 | 860 | Loo | cso | 860 | (Q6L1 WANNA) md vigasyy | OCG
seo pationa | sara oda ody | wae | Ba [wrx | ow | i
Climatic Risk Atlas of European Butterflies
672
673
Appendix 1
674 Climatic Risk Atlas of European Butterflies
Appendix 2 — Scenario results
675
Appendix 2
%9T'8L-)
ISOI-
%68'TS-)
LLb€-
%69'S8-)
%10'69-)
%9L'S6-)
98ZI-
%67'S9-)
G8Tb-
%16'66-)
8SOI-
%EP'88-)
ZhTE-
(PL L9-)
8r8I-
(16 16-)
ZS¢-
(L5°69-)
88Z-
%89'SC-)
9FET-
%G8'EL-)
60bC-
%1.7 68°)
LO8-
%8r LP-)
%88'86-)
8ZEI-
%9L'b8-)
c9G¢-
%76'96-)
€SS¢-
%MLEEL-)
000C-
%bG'89-)
6¢9t-
%8b'66-)
18¢-
%LL'OL-)
€67-
%8S'06-)
OLLI-
%Cb'69-)
SZ6C-
%6S'S8-)
Z6LT-
%€C8'St-)
6£19-
%GE'S6-)
Z98-
%65'09-)
6ZLOI-
%ST EL)
8LI-
%LO'9T-)
1SOI-
%T9'TS-)
6261"
%LG' LE)
SZOL
%L1'8-)
ESg-
(°S0'L)
iG
)
)
(°49°SZ
901
(HL'0Z.
GZ
%91'09-)
808-
WET LE)
L8LI-
%6T LL-)
WIT TS)
%98'PI-)
900T-
(41489)
Z9T-
(4EE'€¢)
Sel
(A1LE'SE-)
S¢P-
(419'LS)
LTbT
(41L'8)
$87
(4L7'€9-)
CLS-
(0499 '0¢-)
%6S'8T-)
%88'19-)
1¢8-
WbL'6C-)
ZS6I1-
%LL9L-)
908-
%GL'6-)
ZOST-
%LT'9T-)
LLLI-
%16'0S-)
S6I-
%EL'LS-)
6£7-
% 65 6t-)
L19-
%SE LT-)
SrLI-
WEP'SP-)
Z8bI-
%E9'LS-)
VZS-
%60'81-)
cOTE-
%87'89-)
LI6-
%/S'88-)
8o6-
%88'hS-)
ZIOC-
%8T9E-)
L86-
%€6'ST-)
SSLI-
(4€5°6S-)
8ZT-
(4879S)
€€T-
(417 Sb-)
+6S-
(486 8b-)
BLSI-
(486 '6-)
SSZI-
(46h +9-)
€8s-
(4G9' 17>)
PERE-
%S0'EL-)
186-
%TE LP-)
ZLLT-
%59'06-)
096-
%TE'19-)
SPCC"
%BL'SE-)
9L6-
%ER'SC-)
STbC-
(468° L9-)
097-
(%1E'SS-)
6TC-
(414'9S-)
S69-
(4¢F'6E-)
1991-
(€0'8S-)
C68I-
(“PLT T-)
9LbI-
(048°69-)
1¢9-
(1L'T8)
6ZCC
%60'0I-)
98LI-
%GG°9T-)
OFT-
MLV ET-)
G8CT-
Ltb- ZS¢-
%L9'8T-)
ZI8I- 8Z6I-
L¢¢- tILI-
(ATVI) | (%8S'0)
891
6II-
%TEOS) |(%G6E'ET)
CCC"
%EL'LI-)
0/9)
€0¢- S
(%SE61L) | (6¢'rh)
8ZS LIZI
(%86'C-) | (ZL'8T-)
Gy
%GI6C) |(oLb'C)
SZC FOL
OFT
(7791-)
%W6L'TE-) | (%1Z'9Z-)
OLL-
%6L'0S-) | (%6S'CS-)
(68'r-) |(9r'9T-)
88- OLI-
%99'6-)
Pag (79°11)
Col 6LE-
(%9G'be-)
€L8T-
(CELT WAaSy) apes sHouy
(CLQT ANaQEY) zvgpsve sub
(COST MANAAED Mapassay smyqouls
(QO8T SUHWISHNASHOQ) o7oug smjqous
(CZQ] ANaNE) vafiqso vywids
(POST ‘DOASNVINIOFY) swaopias byes
(GQ WadHYHOS-HORMIAEY)
sipuaggd vyords
(OPST UNANVY) saoyarg suposyAV’?)
(/€9'L-)
6- (CIOL ‘NIGUHAMY) SYL[UaLLO SUPOADYIAV)
(LpQl SATIN) smsafeoopf suposwqouv’y
(SQL UMaSy) emuaqyniny supoMDYUD’D
(OL8T SUAdSy) aayy supovyavy
(Peg “IvANasIOg) aque seuulary
(9G 1 ‘SQAVNNI‘Y) S450 seuuary
N SE ‘oO ioe) cS ian! + ‘oO oO =, N + No) ice)
ica) foo) a + Re) 09 =) A + S aS ne EA e
S Ne) Nad Nad Ns = Ss ~ ~ ~ oo oO oO
o
S
iam
677
Appendix 2
%69'PS-)
%€8'L9-)
%Gr'Sb-)
€lrr-
%LLYL-)
LL9T-
(F0'°T8-)
SOOT-
(%6'1t-)
OZT-
(4GL'€T-)
(0 TS-)
%6S'99-)
LLU
%G6 LS")
Gr9-
%78'89-)
6979-
%L9'TI-)
le
cro)
OG 1S:
%b6'L8-)
8LO¢-
CT £67)
CrlI-
(“%r1'S9-)
cre
PSEC)
£689-
%8L'19-)
vOCL-
%66'CS-)
€€S9-
%19'Ob-)
168L-
%18'89-)
%G8'SL-)
8SLr-
PT CL)
FO8-
Vee wais)
SZOL-
%9TTI-)
lias
%96'6L-)
L16¢-
%6T L6-)
SOrE-
%/T'66-)
9IZI-
%~6b'SP-)
6r<6-
%6L'9L-)
9906-
%b8'SS-)
€L¥S-
%GT'S9-)
Gr08-
%1S'P8-)
ThOr-
%19°SE-)
tooe-
% 97 Lb-)
OZG—
%G6'T1S)
OL6Z
%/.8'¥8)
8SIr
%L6'bb-)
vLSI-
(19 '6S-)
L€L-
(PT LD
(49 LT-)
STPE-
%16'TS-)
61¢E-
%b8'LT-)
OLGg-
%€6 TTS)
ZZOE
%90'0L)
ZEPE
%bS'09-)
611Z-
(4Gh'bL-)
Z16-
(Eh €T-)
€ZI-
(ATL 8T-)
Ogss-
%TE'€9-)
ZLOS-
%LGTS-)
vLS-
%GL'ET-)
€91T-
%SP' COL)
€LOF
% CG 'F8)
Lvl
%b6 Ob-)
89GT-
88 O-)
SSr-
%PE'ST-)
80€T-
%PL'VI-)
adi
%LL'TS-)
L¥8I-
%LL'S9-)
SO8-
%PLLE-)
G6I-
%b6'SI-)
OFeG:
% 61" LI-)
vbLi-
%8S'CT-)
L0G6C-
%GE9I-)
OFLC
%bO'EP-)
POCT-
%€8'6b-)
9ZIE-
%98'FC-)
B8e-
%TL'TE-)
0867-
(046° L9-)
8Z8-
(467 8E-)
LOZ-
(1'91-)
60€¢-
(418° 8T-)
80b¢-
(LE 1)
680C-
(47E'8T-)
Z6b-
(%0'81-)
90SE-
(48¢ Lb)
€LTC-
%9P'SS-)
6LPE-
%G8'Cr-)
S8r-
%LESE-)
ae
%bL'T9-)
9617-
(429° L-)
L@6-
(%7S'St-)
6£7-
(PL '6I-)
8S0b-
(416'€€-)
SO0+-
(LOC)
6SET-
(ET EC)
L6O+-
(%Z8'0C-)
ShOb-
(S0'0+-)
00Z7-
(°480'0€-)
L88I-
%TZO' 162)
S891
OGLE 1S>)
8Z9-
%GE'SI-)
800T-
(469 Tb-)
609T-
(46L'LI-)
861-
(°489°6I1-)
€6LI-
(%L0°6S¢)
6L0Z
(°469'S9)
%LC'6E-)
%99'61-)
Igoe
%G7'SI-)
O88I-
%bT E1-)
Clee.
%EL'9T-)
616-
%LY OP)
Gl6or
%1T'8T-)
v1e-
%LL'TT-)
€86I-
% FT OSE)
0807
% PT C9)
860€
%16'EE-)
L8II-
(19'8S-)
SIL:
(Er L-)
6¢-
(8'9-)
86E1-
(L9' FT")
€16T-
(rst)
Sbb-
(/7¢9°81-)
€87C-
(19 F1-)
6€8T-
(QG/ | ‘SQNVNNI'Y) O7odp suisspusngy
(C6LI ‘snlontav 4) sagaogd suisspusngy
(QSL ‘SQUVNNI'Y) eM(SOMaUu SHISSDUADT
(ZZQT “Iavaon) aSasas viqguaey
(GLLL ‘[uarrianerstos))
puaxiyog viygudsay
(LOLI ‘SQUVNNI']) vusns viqguisezy
(C6LI ‘SMIORIAV]) SHMDpOLSOU Sauadar)
(POST SDOUSNVNIIOP)) oyeaind sauaiary
(LLLI Has) suuny ls saporgacQ
ee)
nN
(QC | ‘SQHVNNI‘]) Dudion Duadsapy
No)
nN
(CLL HOUNENALLOY)
uoagav snayauly J,
nN
(LOLI ‘vaog) saugsaapds snayaulG |,
(9081 HNIHHNASHOQ)
nyoauy suouaudy [,
S N
— nN
(678 (NADINAD Supoonmes sry Pa004ag4v’D
ioe)
ioe)
| oaas | neva
[svi |
yessodstp | yessodstp | [essodstp | [essodsip
ou ou
ou ou Wy
yessodstp | pessodsrp | [essodstp | [essodszp | [essodstp | tessodsrp | ,essodsrtp | tessodstp
Tey Tey oe ou. Ty
i)
S
ay
Climatic Risk Atlas of European Butterflies
678
%TL'69-)
S69-
%PbL'LL-)
60€7-
%99°€T-)
%GP'9L-)
(%6L'€-)
608-
(AEF TI-)
98II-
[essodstp
ou
(LTE)
Cr19-
(9G PL-)
OSET-
(16'S-)
8S0T-
(%L0'06-)
9Z9-
(SS L6-)
LE8-
%.0°68-)
888-
%L0'88-)
P19T-
%8L OE")
9LbL-
%GL'P8-)
8ZEI-
%EE06-)
600I-
%GT I¢-)
$99G-
WELLL)
veG-
%b6 LT-)
LYST-
%99'0¢-)
Z8IE-
[naive
Jessodstp
ou
%MTL'EP-)
LOTS-
%Z0'L8-)
€vLT-
%PL'SS-)
CLYC-
%56'86-)
6+8-
%6E'L6-)
LL6-
%1TL6-)
L88T-
%MTH'ES-)
OFOOT-
%9€ 06-)
9LPI-
%EP'S6-)
990I-
%89'6¢-)
Z6LL-
(4L 198°)
Z6S-
(4€S'61-)
LOLb-
(LT Ob-)
69I+-
[svi |
Jessodstp | jessadstp
ou Ty
(9 8b-)
Bce-
(/91'68-)
S9L-
LPC)
vCT-
LT Ph)
€L€l-
% 6h LT-)
6SIZ-
(9 TD)
LE9T
(7'Th)
O8¢r
(41 Sb-)
G¢SI-
(4b L8-)
L.09-
(465°L6-)
LE8-
(0466°S9-)
8h6I-
%GE'ST-)
S9Lb-
WIG TL)
961
%TT9S-)
8Z9-
(476 60)
SOTE
[naive
Jessodstp
Tey
%69'LI-)
%~6r'SS-)
%BEC6-)
6+9-
%G6'86-)
6+8-
%EGLL-)
%98°S9-)
(0467 L9-)
ZOb-
(°490'T)
ae
(480° €9)
Core
sv
[essodstp
Tey
%L6TI-)
9TH"
%9L'8t-)
LESI-
%68'LI-)
ZOS-
WPT HS)
LLE-
%19'09-)
0zS-
%8h'19-)
€19-
((TZL'8S-)
trLI-
(498'91-)
89I¢-
(°466'°9S-)
668°
(prL'€9-)
ZLL-
(417 €T-)
$6€C-
(/18°8S-)
bOr-
(0469'b-)
€8b-
essodstp
ou
%GTLI-)
LZO+-
%G0'€T-)
LSbZ-
%TS' Lb)
LOSI-
%91'Z9-)
ZCb-
(045°S9-)
Z9G-
(°466'49-)
8+9-
(049 6S-)
C9LI-
%TL'6I1-)
LOLE-
%G0'€9-)
886-
%69°TL-)
ZI8-
CCLC-
(049% €9-)
9¢h-
(%10°T-)
8tb-
(417'8-)
ZS8-
[naive
Jessodstp
ou
%80°ST-)
(/78'81-)
6Ibt-
(Y6¢L1-)
997¢-
(89°ES-)
Z691-
(LV49-)
OFb-
(49° L9-)
Ogs-
%18'69-)
969-
%bL'L9-)
ZLOC-
%LY'ST-)
L8Lb-
%b9'EL-)
vSII-
%€9°9-)
9¢8-
%TL'6I-)
GLSE-
(%80°99-)
bSb-
(8r'¢-)
ChL-
(418° TT-)
9TZI-
[svi |
essodstp
ou
(41 1'8-)
(0489°97-)
99T-
(46G"L-)
OTbI-
(%19'€7)
OLE
(/6L'61
IZ
)
(46001)
6781
(°481'8)
trLl
(46°69)
660+
(“EE 1-)
E€1¢-
(AP L'€-)
(0469 'T¢-)
(4G L'01-)
1ZOC-
(0496 '8h)
19L
(%9°9T-)
98I-
(%6L'9¢)
8I8E
[essodstp
Ty
yessodsrtp | [essodsip
Tey Tey
(48°ET-)
IbZ€-
(b1'8-)
O€SI-
(%461'ST-)
$6L-
(469° LS-)
O0b-
(AST'ES-)
9Gb-
%LL'SE-)
9G¢-
%9T 9F-)
PLel-
%8S°ST-)
8Z67-
(Lb 0F-)
8LT-
(L460)
BS0E
Svud
%TL)
881
(161)
Zo
(rS'0)
LG
(QGL] ‘SQUVNNI‘]) sanjnu vuogEy
ian!
—
(POST ANAAED) S257 aopqoury
(xodusoo)
(9081 ANAAED) vewosnp aojqoury
(Q6LI “Waasy) vameag aoqoury
(GOgT “asasy) aay dua suse
ISS] “Weel VHOS-HORRIa PY
LUIUNAS SUDGYIOYIULT
698] HONICAV.LS
SOptou ay: qua SUDYIOUIU
(QSL ‘SQHVNNI'T)
SOULMD PADI SUDIOUI UY
a)
—
(L881 ‘NOINay]) sasuom vapy da]
‘oO
N
—
(LL8T SHONTAaNV.IS)
yoqouodup vapydr T
N
—
(xo[dusos) N66] AHONISSHR
you | (BGL] SSAuVNNI'Y) sedouss vapydaT
66LI “Wiasy sounxap oyrdog
QC/] ‘SANVNNI'T “ovqovu oysdoy
ioe)
al
al
(SSL ‘SQAVNNI'Y) saequpod sapyoug dy
o
S
iam
679
Appendix 2
%69'68-)
%b8'96-)
6+6-
%CP'9OE-)
968S-
%10'98-)
6S0C-
%GL'9P-)
Z9GP-
(88'FS-)
I8L-
%8T LE-)
80LS-
%EL'LS-)
60S-
OGL OSC?)
SS9-
%16'TC-)
£909-
%/b'8L-)
P89I-
%68'PC-)
8009-
%80'LL-)
SI¢I-
%6'89-)
006-
%80'9€-)
%/€'86-)
+96-
%60'9F-)
O9FL-
%96'PS-)
€SPS-
Lb LL)
ETO;
%b8'Sr-)
SLPL-
% SOL)
789-
%bELL-)
vrs-
%6LTE-)
€S78-
%97'88-)
+681-
%LEEE-)
€r08-
%PS'L8-)
L6rI-
%19'8L-)
€SOI-
%+0'0S-)
[Sots
%~6b'P8-)
%6b'6E-)
Z9S-
(%7Z0°9¢)
919
(6 EE)
Ob
=)
“bo EL)
nee
%b6'6C-)
9TP-
(ACF TE-)
O8Z-
(04699 T-)
LG6I-
(46601)
G20
(HLELE) |(%8S Tr)
69 LIL
(%EL'01-)
ItI-
%1T'86-)
%GL'9T-)
%19'ST-)
COG
/%99'8T-)
€8SL-
% RLFC)
%C6'91-)
%18'OI-)
OSLI-
%€8'E1-)
Ieee
%/Y'TE-)
O1ZE-
%G0'9¢-)
cls
%G6'8C-)
LYS-
%96'Cb-)
0S
%TSTI-)
GrOe-
%€9'9C-)
98L-
%GH'EC-)
CES
LVL)
619-
Geer)
STTE-
(475° 19°)
919T-
(41208)
16L-
(49°S1-)
S7ST-
(%80°0S°)
6611-
(0486 0€-)
©90¢-
(“zo TE")
OSb-
(47 EE-)
967-
(4CE'GE-)
Sib
(Th LL-)
ZCOE-
(4h Sb-)
ZE6-
(77 L-)
ZrLI-
(418 Lb-)
9IL-
(4L6'SP-)
+09-
%69'1S-)
9617-
%PLTS-)
SO8-
%TTSI-)
1€67-
MLE TH)
686-
%b6 CH)
119-
(YEG IF-)
OL¢-
(417'0S-)
$6S-
(ACF F1-)
618E-
((9'8h-)
erOI-
(SEO)
8S7C-
(0496 9b-)
€08-
(AZ. ES-)
869-
(PL L-)
6ZE-
(149)
LEOL
(48 9T 1)
LOLZ
%65'S¢-)
Icp-
%8E'h)
+6
%PE 0)
Z8
CA wad)
b1Z
(“br OL)
91Z
(467 TS-)
I8IZ-
(%9L'0-)
€ZI-
(4F0'FE)
(469'F1-)
607-
(68°71)
€L61
%T6'TL)
1%
%b8'0)
if
%F6'SE-)
87SI-
%08-)
PSL-
0%65°C-)
Otr-
%6L'T8)
7861
%TETE-)
S60E-
%9E'0E-)
Zr
(478'TO)
CGE
(89°€T-)
LIZ-
(b0'FF-)
1ZS-
(%8ETI1-)
GLZE-
(69'¢-)
168-
%6'T-)
GZ
OCR] “AAMT Yay spy0D
(G8LT SAOWMOMD) swarou svyoy
(GOST “WHase) aznda svy0D
(GL ‘SQUVNNI'Y) ovanjnd spyoD
(OSL ‘uHasy) auomongd svyoD
(xojdusos) (1/1 sn1ontav,)
peupa | (SSL ‘sOaVvNNe) ampydop ryuog
(OO8T ANNE) apyyv vyuog
(16LT SWANANEY) aveuolag sear
(SSL ‘SQHVNNI'D) 2@0u susan
(QZ7Q] AAAS) emia susan
(SSL ‘SQAVNNI'D) adie seat
(LGQT UHAVIN) MUUDU Seuatg
(%86°Z1)
CCC (OORT SAHONIAAVLS) sagomsy Sedat]
(
(QGL] ‘SQUVNNI'Y) apousstug Stat
ou , ou ,
yessodstp | jessodstp | [essodstp | [essodstp | tessodstp | ,essodsrp | ,essodstp | tessodsrp | tessodstp | [essodszp | [essodstp | testodstp
ou ou Ty Wy ou ou Wy 1m.
)
S
ay
Climatic Risk Atlas of European Butterflies
680
(419 9b-)
0@Z-
(ET Tb)
1609-
%66'Cr-)
%6T'8T-)
(4€4'88-)
S99-
(0460°SL-)
€61¢-
Jessodstp
ou
(496'°€L-)
OFL9-
(%10'19-)
9189-
(AS1'EL-)
OFE-
(ER LS-)
LLS8-
(91 8¢-)
LLO¢-
%69'6¢-)
896-
%8L Tr)
9800I-
%/%69'19-)
1Z9Z-
%IT LS)
69¢-
%60'PS-)
ZCOL-
%90'9S-)
60%S-
%ZO'P6-)
LOL-
%TH 16-)
L88¢-
[naive
Jessodstp
ou
%EG'E8-)
ZLOL-
%G8'EL-)
0SZ8-
%T79'S8-)
SO+-
%66'TL-)
LL90I-
%bO'TS-)
88It-
PT 6t-)
LOZI-
%LL'SS-)
LLI€I-
%6b'9L-)
OSTZE-
%Lb'08-)
61S-
%G0'94-)
98b6-
%76'89-)
6+99-
(490'°FL-)
$876-
(414 L6-)
€€L-
(%7'S6-)
Sb 0r-
[sv |
Jessodstp | [essoadstp
ou Ty
(HEh'Sh)
9S9¢
(7Z0'69-)
Gigs
(4b '6T-)
8LTE-
(4bS°9¢-)
OTPS-
(04€0'79)
Z66F
(0/,99°S¢-)
9ZS8-
(p8°ET-)
CLOI-
(A61'b-)
LT-
(481 'F1-)
LLLI-
(9 1'S9-)
O6b-
[naive
Jessodstp
Ty
(47 Lb)
%CH OE-)
€6CI-
%EOTC-)
trl-
%ET'OC-)
OF Lt-
%1S'ST-)
(417 FTC-)
(%7L'T9-)
L997-
essodstp
ou
[essodstp
Tey
(%€6'LT-)
SPST-
(/9¢'F7-)
IZLZ-
(48h'8¢-)
Z8I-
(€9'8T-)
ZOLT-
%EV6L)
6€SI-
%80'€T-)
C9¢-
OHSE CC)
LIvI-
%L9'ST-)
8Z7E-
%BL'IT-)
LOIZ-
%8T CC)
€6LT-
(04€0'S9-)
68b-
(LL'TE-)
IS¢I-
(4E'P¢-)
6¢8r-
(46°C)
GS8¢-
(AEF OE-)
O0rE-
(481 Sb)
vOT-
(4G197-)
9L.8¢-
%89'°ST-)
Z9TI-
%16'61-)
L8b-
%GT'6I-)
ZO9F-
%b6'SE-)
LZSI-
%GL'6C-)
88I-
(0490'T€-)
L667-
(479 '€T-)
196Z-
(AER GE)
S9S-
[nana
[essodstp
ou
(L9°TH-)
0Z09-
(Lb)
ZGCr-
(%GS'9¢-)
€80+-
(AZZ TS)
Lye-
(8TTE-)
ZZ9b-
%T6'IC-)
P9LI-
%ET'8ST-)
989-
%1.6'61-)
9LLY-
%18'SC-)
L€z-
%1.8°6I-)
Z60+-
(0486 '€¢-)
8LTZE-
(L0°LT-)
€6EE-
(PO FL-)
6SS-
oo
Jessodstp
ou
(PT 6C-) | (%9TET-) | (HBO'FE>)
LLIt-
(4L0'T)
611
(8L IZ)
COL
(97'S-)
(7 LT)
ZO1Z
%6LOT)
£86
%Gh'E-)
Leb-
%TE'SS)
Ob
[essodstp
Tey
(77ST)
Ze
(0486 'LI-)
S99T-
(47S bb)
E8SE
(/6¢'6)
677
(4€6'91-)
(69 TD)
Gy
( )
(AT L'TE-)
9FT-
inane
essodstp
Tey
808+-
(LY TES)
6S6C-
(9G TZ)
ZOL
(LL'07-)
8L0¢-
%EO'LE)
0867
0/89" T-)
It-
WTP 9T-)
€T6E-
%EL'0-)
I¢-
%TZO'ET)
$8
%8S'°6-)
€L6I-
(/6r'T-)
trI-
(L79-)
98L-
(41519)
Cor
Svud
[essodstp
(GLLI SOUNANALLOY) Yoqdiyy vuavatT
ee)
nN
ol
(LOLT ‘vaog) smgy vuavat]
(O€QT “AMACAIA]) Durucoyo DuaDIeT
(8G/] ‘SQHVNNI‘]) apaunvesne DuavIeT
(CORT ‘HINOMW])
ioe)
=
Lygy aaa, vsoumvf xiaqouory
ign
ioe)
—
(86/1 ‘SQHVNNIY) euaogqu xiuaq@auor
ioe)
=
COG] “AAANY Seswauvonfy svyoy
(8GL] ‘SQUVNNI'D) yoy svyoD
(OSL Udase) aaaqgostiga spyoy
ie
Si
(OSL Udasy) avopuusi spyory
N
i
=!
o
S
iam
681
Appendix 2
%EET8-)
SrbTC-
%8T'6E-)
+9Tb-
%ET OS)
06Tt-
%L0'8¢-)
LL9b-
%EE 6Y-)
LLYS-
WET H6-)
80b-
(49G'PT-)
0SZ¢-
(%81'06>)
Bc¢-
(4€9'EL-)
(466 Tb)
%6E €6-)
LI8Z-
%GLIS-)
819S-
%1.0'69-)
668S-
“br 6b-)
9909-
(%€¢'09-)
8699-
(b8'S6-)
SIb-
(7 0r-)
G6S8-
(HL'76-)
89¢-
%88'T8-)
GECE-
MLL ES)
99bL-
%GS'ES-)
Z169-
%S1'69-)
CrLe-
%b8'S8-)
b9G-
%G6'8S-)
%EL'S9-)
9€IL-
%TS'8L-)
90L9-
%1.6'LL-)
LS98-
%S1'86-)
STb-
MLL ES-)
LObIT-
%bT 667)
$6¢-
%LU'T6-)
60L¢-
%bL'99-)
$816-
%G8'EL-)
ZES6-
%EC08-)
1S9¢-
%86'F6-)
$Z9-
VE LS
% 9G ¢r-)
9¢8b-
(71)
(% 69°01
OL
)
>)
%9E'EL-)
%G0'6C-)
%€8'0S-)
bb9S-
%86'69-)
CO¢-
(41Y'S¢-)
OLSL-
(o9'S8-)
OFE-
(466 bb)
ZT9-
(E971)
DLS
(4156S)
%16'99-)
PL0C-
%16'6E-)
CCE-
%LO'TC-)
G88I-
%67'8I-)
tbTC-
%/b'99-)
O8¢L-
%TL'9G-)
(49L' Lb)
(415°89-)
LSLT-
%Lb'8S-)
O9LI-
W6V'TC-)
Ihbc-
%68'LT-)
Z8ET-
%LTST-)
LOTE-
%6L'€T-)
Lr97-
%G1'68-)
98¢-
%16'ST-)
LOFE-
%E8'LL-)
60¢-
%bO'8T-)
619¢-
%99°9]-)
LSL-
%66'Cr-)
687-
%69'6¢-)
009S-
(“6° LE-)
€S6T-
(46¢°49-)
8c6I-
%€6'9T-)
-ZG6C-
%16'Ob-)
b6r¢-
%19°9T-)
SS67-
(4°L8-)
BL¢-
(°466'0C-)
L8bb-
(46¢°6L>)
SI¢-
(4€0'19-)
9GbC-
(ET LT)
LOLE-
(46'F7-)
bIT¢-
(ITC?)
IZrI-
(4€T LS-)
9L¢-
%L6'Cr-)
OFE¢-
%9F'LL-)
ISIz-
%bO'TE-)
OLES-
% 6G Lt-)
ZGSE-
%LE6C-)
96S¢-
%89'TE-)
6Z9¢-
(4L.0'06-)
06¢-
(469'°CT-)
ThSt-
(68'S8-)
Ir¢-
(%)€1'S9-)
1Z9Z-
(€0'0¢-)
OLIt-
(LG F¢-)
Z9bb-
(0468'F¢-)
b8SI-
(10°¢9-)
bib-
(“8h'ST)
SOTI
H/T LI)
LI6I-
%9¢'S9-)
€87-
%6L'TL-)
G87-
%99°TC-)
Z16-
%L6'V1-)
6L0C-
%8ELI-)
CVCT-
%LL'b9)
1h67Z
%TC 6)
Z61
(%7¢'9-)
(0469'C-)
€19-
%9G'PL-)
967-
%TEOF-)
P9O8I-
%L0'0C-)
SLLT-
%90'0I-)
867Z1-
% 16'S)
ZE9OL
WLS PC)
19I-
%TTST-)
L¥SI-
%9P'ST-)
L@8T-
(1¢°¢9-)
GLT-
(b6'81-)
6b 0t-
(9'°6L-)
9T¢-
(pb 8C-)
1SSI-
(9 TT")
SOTE-
(499'TT-)
SZ6C-
(“bb Ob)
Sel
(Tr LT-)
Si-
(POST WINANED) yse anos
(OLLI “Waasy) 82042 anus
(L8L] ‘sotortavy) 2aeds anugos
(GL ‘SANVNNV'Y) umd mnuyvs
(ZBL | SHOONS]) Mugqjv- Un S
GOGT ‘NVINAVHD sep saqdoyv)
(SLT ‘SQaVNNIT) sqm slag doo)
(L8LT ‘sororav,]) sjjvq saunuioy,
(COLT “Uaasy) sacoqos sedosoar
(QSL 1 ‘SAHWNNIY) Smuanb sumo.
(QSL ‘SAHVNNI']) ayujag v720q |.
(PELL UHasy) woupssaqy vuaval T
(PPS WakvHOS-HoRtsyY)
suapuvs DUDE
(LOLT ‘SASVNNI‘]) eoqjoddig vuavatT
ou , ou ,
Jessodstp | yessodsrp | [essodstp | [essodstp | tessodstp | ,essodsrp | [essodstp | tessodsrp | tessodstp | [essodszp | [essodstp | testodstp
ou ou Ty Wy ou ou Wy Wy
i)
S
ay
Climatic Risk Atlas of European Butterflies
682
(410'89-)
CZ8I-
(470° LT-)
(419'bb-)
%EL'98-)
isi
%8h'9T-)
6L6¢-
%18'68-)
SOOT-
%LSOb-)
%CL'98-)
(°490°9T-)
€99I-
(49'0S-)
THOS"
[essodstp
ou
(67 68-)
Z6E€T-
(%S1'SS-)
bL6T-
(7T'FS-)
eebl-
(b8°9C-)
VCSL-
(b1'09-)
8ZEI-
%8E'86-)
LOOC-
% C8" LE-)
G89S-
%69'Gb-)
ZL99-
%96'96-)
L6L-
GLEEG)
7CG6C-
LP CP)
OLLT-
%€T'89-)
Bless
[nave
Jessodstp
ou
%LS'L6-)
v19C-
%EL'8S-)
CCLe-
%8G'C8-)
6077C-
PLO)
9¢€00I-
%P8'08-)
G8LI-
%9S'66-)
Le0g-
%9b'TS-)
€88L-
%99'86-)
TOES
%9L'P9-)
ZOLS-
%S1'66-)
S18-
“Lv OL)
GS8¢-
(°/001-)
008-
(4G LS-)
999¢-
(41 L'8L-)
LZ19-
[svi |
(6% 90'¢-)
1-
)
(88° LI-
Lte-
(16 TL)
6851
%16'1E-)
1S9-
(46 '69-)
Gis-
(L8°E1)
S88
(O%€L'0)
ES
Jessodstp | [essodstp
ey Tey
(%68°S8-) | (%8r'F6>)
L€Sz-
(ELS) | (%Z9'8h)
60¢
(EEC)
98-
(
%86'PC-)
LOIS-
(41 L'08)
Z8LI
(460°TE-)
(97 8¢-)
L&b-
(b5°T6-)
6SL-
TEST)
BCR
%79'8L-)
679-
%LE LI)
Clic
%PE'8T-)
90TT-
%/.8'6C-)
890I-
PT 6C-)
LEOI-
%6S'91-)
L8E¢-
PETE)
bIL-
%80'9F-)
OFG-
(/EVTI-)
ZZ8I1-
(48G°TL-)
O1g8-
(049G°CC-)
ZEOE-
(%8'0L>)
Z8S-
(981)
6SL-
(79 FS")
LEb-
(PL 9T-)
O¢OI-
(465 '6C-)
CO€T-
(47ST)
Z8I-
(0466 '0S-)
99¢1-
(0469 '6C-)
66SI-
(414 9¢-)
+96-
(416 LI-)
OL9E-
%LY'TE-)
LIL
%98'9L-)
89SI-
“be FI-)
6€1T-
(4LEFT-)
GLZE-
(477 BL-)
€+9-
(0458°ST-)
898-
(%7T'8T-)
9STI-
jnanval svuo | Saas | na
yessodstp | jessadstp
my my
| osoc_ | osoc_|
[naval
essodstp ea essodstp
ou
ees
(L0°6+-)
OIZ-
(LT FS)
vSbI-
(8'LT-)
Z96-
(97 Iz")
Iper-
%9T TH)
€£6-
%GG'LL-)
Z8SI-
%76'LI-)
Z69T-
%LL'98-)
196-
%/L'O€-)
yelr-
%ZO'T8-)
999-
(49 0C-)
OZLI-
(%79'69-)
EGS
(Er 6I-)
OrzI-
[svi |
eld
% BLES)
nal
%9L'TT-)
%GESS)
S867
SFL)
Ize
)
(
(HT 81)
866
"Gz>)
(“0° 07)
6LZ1
[essodstp
my
(148)
(40° 8b-)
L8ZI-
(0460'9¢)
%bE'SP-)
90¢-
%6L'91-)
9G7C-
%EL'Y)
6S7
)
(%T79'FE-
LLt-
(LLTD)
SI8
[nave
Jessodstp
my
(67 LI)
bl
(°460'0+-)
PLOI-
(6b 9)
8961
(/LVST-)
LOb-
(469'€1-)
G6LT-
%TH LT)
CL
)
%TL'0S-)
99¢-
((S1'b7-)
Gree"
(56 TL)
+59
(/SL'8¢-)
OL¢-
(68°41)
0S6
oo
[essodstp
my
(LEQ “LAAT g)
SUSPLAAOUAGD SapyUvIYOIS
(GOQT “AMOOP) PMRUIA sepyUvIyOIS
LLL Sassy uIsoutg)
UOJDG SaptUvyyoI Ss
>
Wa)
N
ee)
+
(GL ‘SQUVNNI'Y) S7odtuy vuLsHIaD
(F081 SDOSNVIMOPY) seyayp opduy
x2)
—f)
N
(O88 ] XHONIAAV.LS) smyruojovep opedu’y
(LLL ‘svTIVg) sappiiuv opiduy
(6781 ‘NADIA Saleso opid’y
(GLLI ‘XISSANY) saummuu opidiry
(ZOQ] ‘NAINA, vusiny DUads~7
(LOLI ‘SAAVNNI‘T) svogyeued sagordr]
(R681 ALLAg) yoqsanu snaswy
(LOLI ‘SAAVNNI'Y) svo1700q saprduir]
(/8/] ‘saiontavy) annw unos
o
S
iam
683
Appendix 2
%8E'SP-)
O8SPr-
%18'9G-)
SLIE-
%8G'EC-)
Z619-
%6T'9T-)
¥c0S-
%90'Th-)
¥6LT-
%CO'EL-)
€F9T-
(46h 19-)
P86¢-
(“6 Ob-)
(486'49-)
Wwva
a
ou
%LL'OS-)
LSOS-
%ELTL)
+90b-
%~TH CP)
9008-
%G9'8E-)
CoGus
%90'€S-)
GzGe-
%8S'P8-)
L90¢-
%G8TL-)
€TLP-
%€8' 6b)
6659-
%P8'88-)
8C6C-
% 68° 16-)
Este:
%TH'SL-)
L16¢-
PEO LL-)
%TS'19-)
6079-
%Lb'S8-)
9LLY-
%9E'PS-)
€ZOOI-
%€T'0S-)
86S6-
%TLSEL-)
€88r-
%19'S6-)
09rE-
%F9'S9-)
7698-
%bE96-)
981¢-
(%75'86-)
the
%8T CP)
G8L9-
%G1'88-)
LZSS-
%98'F6-)
90F-
[sv |
ed
%/ TCP)
L9¢F-
%66'TE-)
€809-
%GT'TI-)
IvL-
(4G¢°ST-)
800I-
(HEL TL)
LOC
%b0'6r-)
6b6r-
%LEEP-)
986L-
%66'S1-)
ZIOI-
“TH LY)
9TLI-
(48h bT-)
L8SI-
(480°€T-)
ZELI-
(4€8'49-)
61€C-
(1b Ch)
6S9C-
ae
%6L'09-)
SE19-
CAC)
GS9-
%61'PS-)
7666"
% 8S TT-)
Cake
Gas)
%6b'F9-)
(%G°8L-)
96S7-
(418°9S-)
ZEOT-
(4ST OL)
16SI
(°/001-)
€6bI-
(4ST Th)
6F9T-
(49¢9T-)
OL-
(49 L'0€-)
POle-
%/9'1T-)
966¢-
%6F'9I-)
LSS
%TLOE-)
100c-
%bTe'SS-)
6661-
%69'€¢-)
P8IT-
(/€7'ET-)
9LOE-
IV TY)
LEI.
%h8'LS-)
690C-
%F0'S9-)
1L6-
%LT6C-)
ered i
%TL'6E-)
OLI-
ou
(Lb'vr-)
681-
%G6'9¢-)
S90T-
%LG6'ET-)
61hr-
% 6S LI-)
L9¢¢-
(491 FS")
096I-
(467 OF)
ZL9C-
(486 1T-)
8S8T-
(%EL'LS-)
6061-
(L9°79-)
SETC-
(bT'6-)
Srrl-
(069°SL-)
O€TI-
CA 4 ae)
€99C-
| svuo | oaas | na
ro)
yessodstp | jessodstp | [essodstp | [essodstp | Tesrodsip eee Jessodstp | [essodsrp | [essodstp
Tey
[naval
eid
%9L'SS-)
LEG-
%bT LE-)
€SI¢-
%86'LY-)
1897-
%6L'8T-)
60€S-
%LL IC)
Sblb-
%8h Ob-)
689T-
(41 1'69-)
LOSZ-
(HEL 6r-)
VCTE-
AIS TS)
ZLIb-
(416°SS-)
6F8I-
(88'89-)
b9OPT-
(ALVET-)
S90T-
(448'08-)
LOZI-
(S0'¢+-)
669T-
bo
eid
%8h'6C-)
GL6C-
% 18 LL)
G66
P)
%10'8S-)
L98-
| oaas |nanva| svuo_
yessodsrtp | jessodstp | [essodstp
Tey
%TL'TO)
ae)
€08T-
%G0' TZ)
OL
%/8'€T-)
LOpr-
%6S'0C-)
16zI-
my
(%L4'0-)
Gu
(ALE 0¢-)
S90E-
%TL'8T-)
967¢-
%6VIT)
SCIZ-
%68'0T-)
B8eI-
(460047)
ISrI-
(%1L'€e-)
LeSt-
(%99°9T-)
907Z-
(Sb 6c")
bL6-
(ALT LE)
€CEI-
(/9L9L-)
OFLI-
(Er b7-)
ZESI-
(T8L1 HOON) arapypdo sulaqag
(6LLI WissyuIsoUg)
UOUOUTOAMGAD SULoGA J
(LOL ‘SQUVNNI']) supe suloqag.
(QGL] ‘SQAVNNI']) sou sulogag
(CLL ‘[eearrranaeettHos))
UOYD SUUDEUAY J
(6LLT MASSyUIsOUg)
SHOGSNDU SUDSUAY
(LLL MASSVULSOMAG) Seajay SuUBsUad J
(9G ] ‘SQAVNNI']) “ou suvsuayg
(OTST ANIAHNASHOQ) s7or DULOT
(ZI “IvAnasiog)
sdourjau ayatsdoonnj
(LOLI ‘vaog) sexap agatsdoonny
(CIQT SWHNaNP) sazdound sapyuviyoos
(LLL ‘SVIVg) wouo sapyuvjyors
(ZEQT SNNVINSUAAG) S70IDg SopryUnyy0s
oO
S
ay
Climatic Risk Atlas of European Butterflies
684
(0418'6S-)
90LT-
(0490'°98-)
886I-
(1b PC)
ScTr-
(416'€T-)
(0499'9G-)
%9'CS-)
981S-
PETE)
€S6r-
%LTT8-)
Jessodstp
ou
LESLD
ZIT
EGiGe-)
9919-
MELTS)
EG
%/1'88-)
Gtr-
“bE 'S6-)
618-
%LL'SL-)
Orel
%6r'Sr-)
6£69-
%b6'F6-)
OVET-
%97'€S-)
eee
%6L'S8-)
OZOT-
%LE'6E-)
68r-
%9G'86-)
G89-
%6S'99-)
inane
Jessodstp
ou
%LP'TS-)
Cea
%81'86-)
89TT-
%G6' LY)
€TE8-
%€0'19-)
GZ9Z-
%Z0'L6-)
S8r-
%6L'L6-)
Org-
%EL'E8-)
LE08-
PLS)
SIL8-
%L0'66>)
SPPT-
%99'8S-)
¥S8L-
%8L'C6-)
SLI-
(47S 6b-)
S19-
(%001-)
G69-
(%17'S8-)
€9¢-
[sv
Jessodstp | [essadstp
sh Tey
(18)
LOG
CAR AKA)
Cll
/8'89-)
16S-
%EL'TS-)
Cr6r-
%F6'F1-)
GiGes
%Lb'89-)
Z691-
%€9'6C-)
L96¢-
% GL LT)
x9
(%7¢-)
1Zz-
(4¢4 TS")
S8II-
(49 L¢-)
L19I-
(449 '6S-)
00¢-
%EP'OL-)
S09-
%bS'EL-)
OLIL-
%TL'E8-)
¥S0C-
(%G'8r-)
€6+9-
(46¢'6C-)
G9¢-
(466'°C6-)
9+9-
(488° TT-)
€6-
nana
Jessodstp
my
(Th)
(0485°0L-)
GG¢-
% COPS")
b9r-
%LS'18-)
988L-
%BETE-)
L8LP-
%C6'8L")
0S6I-
%Tb'ES-)
eye
%69'6C-)
€6r-
%CL'86>)
Z89-
%67'SS-)
CEz-
po
[essodstp
my
(416 bb-)
O¢0T-
(79 L9-)
ZOSI-
(4€8'F1-)
vLST-
(/€6'81-)
bI8-
(%1'0S-)
ZSC-
(46 TL-)
819-
(HSE'8¢-)
80LE-
%8E'9I-)
66%C-
%1L609>)
GOSI-
%G8'9C-)
9L9-
%99'8T-)
9G¢-
(ZV 9L-)
67S
(0488°€S-)
6CC-
essodstp
ou
(4£6'76-)
ver-
(4° 1S-)
STET-
(469'0L-)
ec9I-
(46¢'F1-)
86hC-
(466°S¢-)
SZSI-
(69'6S-)
00¢-
(415°6L>)
€89-
%9G'TS-)
S86b-
% 8h OT)
vCIE-
/%69'99-)
8r9I-
Lb LE-)
L10S-
(476 '€9-)
09L-
(418°9T-)
eee-
(%L7 LL)
LeS-
nana
[essodstp
ou
(466'S6-)
Sbb-
(8¢°6S-)
$89T-
(80° T8-)
€L8I-
(89 1T-)
€OLE-
(ALIFE)
LOVI-
% 68" L8-)
GSL-
% 6S LY-)
LO9F-
%18'€T-)
1¢9¢-
%6S'°PL-)
CrSI-
%6b'SP-)
1609-
(°490'0L-)
€€8-
(96 1¢-)
L6¢-
(L5°6L-)
€Ss-
Boe
essodstp
ou
(46 '08-)
8L¢-
(%79'T1
©9
)
*)
(466° F¢-)
OFES-
(°%96'1)
667
(ATS TS-)
€LZI-
(45° 8T-)
OZ8¢-
(6479)
bl
(b8'81-)
bET-
(4L4°99-)
Z9b-
[essodstp
Ty
(%€6'C6-)
ber-
(49 L'€¢-)
(46'SF-)
(4650S)
168b-
(476'89-)
6Lb-
laanva
yessodstp | [essodstp
Tey Tey
(SLLL ‘RerrianwamrttHos))
SILO SHIDULUMOM OT
(%5°S6-)
OFb-
(/16'6¢-)
POSt-
(H18°ES-)
A
a)
—
co)
(CZQI WANANED) Magosa sujpumoGog
(SLL SOUNAWALLOY) Saoupuumas sumuv’)
NS)
=)
isa)
2S
(673 SNHDTAIA)) SDL DIOL
(/81'67-)
SGala
%76'E1-)
OL-
%£0'69-)
€6S-
(QCQ] WHAM) Sosazuy vidupr
(QZ6T ‘ALIMAA) S2MazUOU DIMEZ
(C6LI SSMIORIAV ]) SANUANDIUD DIME
(GLLI ‘[eaTinweatHos)) sysai veoup
(IZQT. SZVIOHOSHOS}]) MAWIDAI DID
(/6¢'F9-)
16SI-
(p66 Th)
919S-
(S70)
€ (R6L1 WANNA) smnjequo suleqafg
(L0°F7-)
667-
(AISLE)
LGt-
BoE
(OSL Uddse) “opauna vim
(Gegy ‘AxNZSCrIVARY]) smueqdas suleqa
(zogy “IvAnasiog) yubv sulogarg
(R6LL ANNO) “opunys sulaqae,
o
S
iam
685
Appendix 2
(41b'6C-)
LO9¢-
(499°SL-)
69LI-
(HET F6-)
eCb-
(67 88-)
LSEI-
(6001-)
0S6-
(H1E°€6-)
6¢L-
(%7Z0'6S-)
L61¢-
(88 rb-)
96%S-
LLL)
Bee9-
%TC 6S")
OLSI-
%TP'99-)
ZC9b-
%EG'E8-)
%€6'S8-)
600C-
%EL'66-)
9GP-
%GT'S6-)
b9rI-
%L'66-)
L8L-
%79'EL-)
€8b9-
%L6'SS-)
$S89-
%LL'TS-)
9TOL-
%1.6'69-)
GS8I-
%TS'18-)
€L9S-
%99°9¢-)
LS8-
%.0'6P)
SOL
%bO'SL-)
(4L4'89-)
1069-
(/6L'66-)
99b-
%99'9F-)
1601-
WES SP)
LG6-
%9T'88-)
%TE'S9-)
(485 °66-)
9+6-
(%Z0' LZ)
b1Z
CATS)
SO+-
(%Z0'Fb-)
LOLT-
(LS 07-)
80ZS-
(AZL'01-)
OFL-
(0498'T8-)
%/T'09-)
60bI-
%76'9¢-)
6L-
%LU'L6-)
LYb-
%EG'6S-)
S16-
%G0'L6-)
TCO"
% 10°61)
ISI
(Po LT-)
LtSI-
(049G'87-)
L€ZL-
%6L'88-)
06I-
%bO'EL-)
9¢¢-
%8L'bS-)
Zh8-
%66'C9-)
86S-
%ECOL-)
is¢
(499'°8T-)
vCST-
(A7L'91-)
LvOT-
“by 9T-)
CvCT-
WbS'TI-)
ZSTE-
%GL'0E¢-)
OFIZ-
%7G'SE-)
€8S¢-
%~b6'E8-)
Z6E-
%T8'PE-)
L9IZ-
%T9'PS-)
LLTI-
%TL'68-)
Z61-
%/8'08-)
ZLE-
(049¢°99-)
OZOI-
(89°€L-)
00L-
(486° 0L-)
6SS-
(40 0€-)
SP9T-
(/€T ITZ-)
009T-
(PL Th)
GLSE-
(49 'Th-)
O¢II-
(4S1'ST-)
9¢8¢-
(476 9¢-)
69S7-
(4L7'SS-)
GSS
%LULY-)
8767"
%LTS9-)
9ZSI-
%6E'V6-)
ZOC-
%Bb'Es-)
P8¢-
%LO'OL-)
9LOI-
“CY PL)
LOL-
(4LL'9L-)
809-
(HEL OF-)
L8S¢-
(LT Tb)
L8b¢-
(L8'6+-)
CCEI-
(ALT LI-)
bLor-
(0486 O-)
tZ8T-
(49+ IZ)
OLI
%Gh'¢)
POE
WPL TL)
09S
90" L-)
66S-
%T'EE-)
088-
%96'S-)
80SI-
“TE TT)
L8L
%b9'8T-)
6887-
%LP'6T-)
ZSb-
PLL)
169-
%/.0°€S-)
Ese
(460° TT-)
989-
(9G Lt-)
ZLII-
(0468°S1-)
(%L8'SL-)
6FE-
(4Th'TS-)
86b-
(HIE 8D)
Srl
(14 01-)
LI6-
(H€8'0¢-)
S19T-
(/€8'Ct-)
ZOLI-
(90°T1-)
(10'C-)
OFI-
(99 b+-)
GOSb-
(SLLI ‘[errignerentnos))
Moulop suIvUUUO Og
(CT8T MANANEY) Sop supoumoljoy
(OST MAAdMY) wna suouuoljog
(SQL I ‘Uaasy) supaupy supouuolog
(1SQ]. SaeTVHOS-HORAAEY)
SUDIIG] D SULDUM oy Od
(GQ. aetVHOS-HORRIAEY)
snuvdsiy supouuoljoy
(LOLI ‘VaOg) “opus sujouuolgog
(GLLI ‘OUNANALLOY)
SUBLDI[ AG SHJDUUIUOL Of
(SZLI ‘[warrianasentHos])
seugqop snpouuo]og
(Q08] XHINISHNASHOQ) Soda smpvUuuolgoy
(SLL SOUNAWALLOY) Si090 SuoUULUOL OT
(POST AHNHINV)) Sapereaqy supouluolg og
(Z6L] NACISNHOS) supupiuv suypuiuogog
(GST ‘NISISWLLES]) SUarsane SHpDUMULO] OT
ou Tay Ty
yessodstp | jessodstp | [essodstp | [essodstp | [essodszp | ,essodsrp | ,essodstp | tessodsrp | tessodstp | [essodszp | [essodstp | testodstp
ou Ty Wy ou ou ou
oO
S
ay
Climatic Risk Atlas of European Butterflies
686
(465° LT-)
6+9t-
(08'87T-)
(0469'99-)
%L6'Sb-)
€9+9-
(16° TS-)
CCCE-
[essodstp
ou
(%90°€¢-)
SILL:
(bS'P¢-)
L¥LT-
(%€0'S8-)
Cree
(%%90°7S-)
8987-
(9¢ 19-)
09b8-
%PL'TI-)
€80¢-
%b0'Sb-)
9P6L-
%GELS-)
Cr18-
%ST'TS-)
ZISOI-
%6L'9L-)
ZLIF-
%9L'SP-)
€S78-
% 87 €8-)
890¢-
%1G'P9-)
800+F-
inane
Jessodstp
ou
%80'TH-)
6S06-
%6T EP)
erre-
%GT'06-)
crre-
%€L'99-)
9L9¢-
%G6'SL-)
ZLYOI-
HSV TL?)
£000I-
% 6G LL-)
6r8¢-
%1TT9-)
68Z01-
%89'PL-)
r090I-
%LEE9-)
VCO I-
%TEL8-)
ad hae
%49'(09-)
€80LI-
%bS'T6-)
60r¢-
%b6'T8-)
Sr1s-
[sva |
(91 '8h)
€S97
(476 67")
OTT F-
%LL'O€-)
9TCP-
%60'0C-)
TCE
(47 0C-)
ZOLb-
(L0°TC-)
€18-
Jessodstp | [essadstp
ve Tey
(E496)
LOOT
(0485°S¢-)
90Gb-
%Z0'9F-)
LL+9-
%GE'6L)
096
(494 '9¢-)
S6bL-
(0499°9T-)
ZL8P-
(48¢' rb)
SCOI-
nana
Jessodstp
my
(Mb Lb)
(48° €T-)
O8rI-
no
[essodstp
Ty
%6L'6I-)
ZIP
%TH'TC-)
€8LI-
%69'SP-)
lites
%6G'EC-)
1¢9b-
%LL'OC-)
9¢8T-
%CS PL)
607C-
%b6'LI-)
L96T-
%9G'0T-)
616C-
(61 9T-)
STEE-
(4€6'6C-)
9Z9I-
(PE8T-)
1S€¢-
(LS bb-)
ZH9OI-
(%LESE-)
Z61C-
Jessodstp
ou
(46% LT-)
SZS-
(0499'9T-)
L8S¢-
(469° LT-)
66€1-
(0486°SS-)
SEIZ-
(BL vc")
S9CI-
(“bb b¢-)
SPLb-
(°480°67-)
880b-
%IL9E-)
v6LI-
%18'0C-)
Ibre-
%F0'ST-)
9GG¢-
%BETC-)
66Sb-
(0460 Lb)
9GST-
(419'0C-)
99L¢-
(%78'7S-)
9FG6I-
inanvar
[essodstp
ou
(49L'SP-)
bL8-
(98°97-)
Z8LS-
(HEV'CT-)
09LI-
(°60°09-)
ZGCC-
(Y67'TE-)
6LLI-
%G0'8T-)
bhos-
%61'SP-)
CPCC-
%/8'€T-)
8h6C-
%9T HE)
G98r-
%LO'VC-)
L¥Gb-
(ET Lb)
99G7-
(/16'€T-)
69¢¢-
(ALL'8S-)
Core
fo
Jessodstp
ou
%PL'61-)
eke
%LL'SI-)
TrOP-
%86'61-)
68S1-
%CS FL)
+SS
(49'ST-)
LSIC-
(469'€1-)
6L9-
(€8"L-)
S6ZI-
(%8°S1-)
Z8S-
[essodstp
Ty
(%16'L-)
IST-
(G°91-)
(EST -)
(Ab LPC)
O8cl-
fnanvar
(ALE 0¢-)
Ogs-
(415°9T-)
LOLS-
(996 1-)
b9SI-
(%80°L>)
OLZ-
(S8'E1)
COL
%19'6I-)
vOLT-
~SrPL)
1€07-
%G6b'S1-)
ZOGT-
(7'9T-)
OZLE-
(F0'FT-)
G88T-
(8°01)
L8S-
(141)
LLST-
(479 TE-)
S9TI-
no
yessodstp | [essodsip
Tey Tey
(LLY ‘Ra TIONAIHOS]) aoay seqyuang
(OSLT “UussyuLsoung) eaqdop sijuarg
NS)
isa)
(QSL ] ‘SQaVNNI'Y) au(Souqdua vuopog
(66LI ‘wads ey) vemouna DuofOg
N
Ne)
isa)
(CLL SOUNAWALLOY) O2 Sayjuadg
(QC). ‘SQAVNNI'Y) PeuogzH] DuLOSST
(LLLL ‘sv TIve) anpory stuuddapy
(QGLI ‘SAHVNNI'Y) aqoe seuudiapr
(SLLI ‘[earrianenttos))
addipy siuulstpy
(8G ‘SQAVNNI']) PAyoD seuuliapy
(GLLI ‘[uarrianetttos))
vaopuvd stuudcapy
(QGL] ‘SaHVNNI') pegged seuulsapy
(ZL SONLIAVHOIW’]) 52720 vaqglgr
(QGL] ‘SQAVNNI']) BZN SuDaUUDL]
o
S
iam
687
Appendix 2
%€0'LT-)
868S-
%L0'ST-)
OCG:
%G9'LT-)
%76'6C-)
(480°€€-)
LLL9-
“cr bb)
7696-
%F0'9E-)
€8EL-
%~bS'SP-)
OLI6-
%66'SP-)
18¢01-
%TC 6b)
S7Ob-
%8G'1S-)
Z8L-
%GT'SP-)
0697-
%16'9T-)
Bey:
%LL'T9-)
6cc5>
%G9'()9-)
Oree-
%G6'CP-)
9668-
%8h'Sb-)
%TS'SS-)
tha
%€8'0S-)
Z1POI-
%€9'8S-)
GOST I-
%Z9'09-)
P89EI-
%Z0'06-)
CCre
%8E'L9-)
610S-
%/%60'99-)
ZOOI-
%L8'EC-)
6S"
%69'9L-)
+6£9-
%1T9L-)
L61b-
(/6E'ES-)
LZ601-
(481 9-)
IST I-
%EL‘OC-)
L6¢r-
%8L'TC-)
800S-
96 'Ob-)
L6I-
%80'LP-)
6SEE-
(%69'S¢)
(PL 0¢-)
16Z9-
(%0°9)
OLT
%b0'9C-)
€98L-
%89'F1-)
%€E0'SE-)
%19'89-)
0c:
%9b'SP-)
EEGs
(415° LT-)
(468°C)
OSSZ-
(LL Th)
6FS8-
(8°60)
OLS
%8T Ob-)
%G0'ST-)
(%L0°9L-)
681b-
(°/001-)
8zS-
(470 1S-)
errOl-
(49L'S7)
a
(z'S1-)
Se
(%96°ST-)
69eE-
%8T OL)
€88C-
%IV LI)
bLee-
%6E EL)
ESE
%EITE)
899C-
%CG TE)
c6r-
%8E TE)
6161
%BEVE)
96¢-
%60'8c-)
es
%S0'SE-)
O€6I-
%G0'66-)
€ZS-
%19PT)
LEOS-
%L Or)
99L-
(PS 8T-)
o¢e-
(16ST)
bOre-
%€0'61-)
€SIt-
% 6S LI)
ZO9E-
%61'ST-)
€LOS-
04S Ves)
LS8b-
(417 LT-)
O€@C-
(4S 0E-)
€Ob-
(19 PZ)
LSbI-
(%8L'TC-)
OL¢-
(48¢'6C-)
OSbC-
(%979C-)
OFbl-
(41.9'86-)
1ZS-
%9T'€T-)
vir-
%T9'1T-)
SILb-
%66'61-)
+60+-
Aer Gas)
O81S-
%EV'TC-)
©90S-
Onrces)
GZ9T-
(479 I4-)
1¢9-
(%8L'TE-)
6+GI-
(97 8¢-)
0G61¢-
(/€6'8T-)
€6S1-
(47 66>)
VES-
(ALY IE-)
Zrbh9-
% 9G TI-)
TCSC-
(49€'L1)
60€
(49°ST-)
bOCE-
(%H1S°8-)
bbLi-
(86 '07-)
VCC
%TO'KC-)
+961-
%L881-)
98T-
%6E'TC-)
[ols
%6S'1-)
Regs
%TT LI)
OCP]
%F0'9T-)
vorl-
%L1'86-)
SIS-
66°C")
VIS
(45°61)
Bre (QGL] ‘SQAVNNIY) avoqtan save
(8GL] ‘SAHVNNI‘Y) 07 SeS 7
(89° LT-)
GESC-
(/>6'6-)
SCOT-
(r9'91-)
OS¢eE-
(r0'81-)
€LOv-
(/68'87-)
ZIET-
(HEVE-)
ozs
(86 0¢-)
ZhSI-
(/8h'TC-)
S9¢-
(QSL 1 ‘SAAVNNI‘]) “puro vssauvn
(9SLI ‘SOaVNNIT) Puurynyv pssouny
(OL8T HONIANAV.LS) Mans puojog
(06[ “IMHOLLS) sapugynby vuojog
(SLLT ‘[uarrianaannos)) sand puojog
(I6LT ‘NITsOUg) v30u/ puojog
(9081 SWANSQH) og) vecopog,
(LOLI ‘SAAVNNI']) Mp Duopog
(L6LT ‘NiTsoug) vlad puopog
(%1L'8C>)
18ST-
(%11'86>)
8IlS-
(ALL'6T-)
£609-
(POLL WHCINNHOS) vaprunqs vuogjog
(GLLY ‘[Mariqnaastnos)) avaas puojog
(COLT Wasy]) Peuryy Duco
(4
ou iy Tay
yessodstp | jessodstp | [essodstp | [essodstp | tessodszp | ,essodsrp | ,essodstp | tessodsrp | pessodstp | [essodszp | [essodstp | testodstp
ou ou Ty 1. ou ou ou Wy
i)
S
ay
(486 bb)
SC6S-
(%€°6T-)
806I-
(47067)
I8FI-
(419° LT-)
LLG6I-
%SUTL-)
G87-
%59'T6-)
(%LT7'SS-)
€6CL-
(H1E9¢-)
S9CT-
(8'86-)
ZZ9I-
(bl bE-)
OFbTC-
(GLTL-)
€6I-
%b0'€9-)
6+T-
%1.6'96-)
%91'L9-)
1468-
%8L'9G-)
6S9¢-
%b6'66-)
OF9T-
%98'Lt-)
67S"
%79'98-)
€€TC-
%8L'T8-)
LTE-
%79'66-)
(%Z1'LI-)
SILI-
(0/4G9°SS)
L86€
(%EL'9T-)
Sr-
%LL' TY)
S9I-
%1.9°96-)
(497 TS")
%9IT'ES-)
OIT-
%79'66-)
OCP IZ)
L8LT-
%66'P1-)
9L6-
%86'89-)
ZELI-
%GO'PE-)
6¢bC-
WIT LY)
LTI-
%TC LE-)
LYI-
(/Sr'08-)
ZOOI-
(466 9b-)
P8I-
(4G6¢'87-)
L81Z-
(486° LT-)
Z69E-
(%L0°T1-)
98/-
(bS°9L-)
9GZI-
%69'€1-)
186-
%TL'SP-)
€TZI-
(LT7'¢9-)
8bC-
(S1'7E-)
€LbT-
(Z0'0¢-)
196¢-
(b8'°61-)
ZOCI-
(L0°08-)
bI¢I-
YSPC)
vSLI-
%79'LS-)
(414 TT-)
(048699)
19Z
(LEST)
(%TL'61-)
(4654S)
bIZ
(L491)
LOTT
(%90°07-)
LY9T-
(bE'8-)
obS-
(G7'EL-)
ZOTI-
%9T'SE)
9TSTZ
%G9'ST-)
69-
(9G ] ‘SQAVNNI'Y) Vexun verIya\
(GLLI SOUNAWALLOY) vedunv solaply dey
(IT “avaoy) wuazuofsap solaptqdury
(QG/. 1 ‘SOMVNNI'Y) DUMYDUL SD
(6S81 ‘SSRLLANAAD
pipausague solaphy dary
(GLLI ‘[uarianentos))
pigguds soapy uy
S
Sorel
se
GAs ‘NVINIVC) 2uupl sD
%8P'Sr-)
L8L¢-
%9L'0E-)
O8ZI-
%90'LS-)
LOTL-
%8E br)
PCETC-
SIEI-
%6L'EL-)
06£6-
%LO'SS-)
$887-
9LTI-
%G0'€T-)
L99I-
%1.6'€9)
OSE
SIEI-
%EL'ET-)
CH6T-
%G6'LS)
SCE
SSI-
%LS'SP-)
O8I-
%G6'T8-)
S60I-
(76'8-)
¥C-
)
(H6r'61-
LL
(%7E'6L>)
LvOI-
THES)
ZEI-
%LTTS-)
980I-
(467 0C-)
08-
(%ZE'6L-)
L¥OI-
%TL'LE-)
6+ I-
%80'08-)
LSOI-
(4€6'€T-)
ShOE-
(4€9'€¢-)
LOLI-
%1.5°9T-)
801TZ-
%TC'E9)
LIC€
(G/] ‘SAHVNNI']) Cuma DiMqosDAE™
(OSL IT Waasyy) wngyr-7 syvy daly
%65'68-)
€R9¢-
“bb 9C-)
ZS68-
WSL LE)
6L09-
%bO'br-)
%G0'86-)
L¢0b-
(46L'6b-)
LLO8-
(%77'S9-)
%9L'€C-)
O1Zz-
%90' T¢-)
9ZGb-
(of 0S -)
S08-
(9'TS)
%8T CP)
8ELI-
(4LG°CC-)
GC9E-
(04€8'6)
%18°SS-)
LOCC"
%E8'TS-)
078°
(419°TE-)
1SZS-
(4S LL)
(75° EF-)
68LI-
“%bT OC)
OLZ¢-
“EP LI-)
9087-
%1.9'0C-)
g9¢-
%~TH ET)
0867-
%ET9T-)
0Ss-
(461 8b)
1861-
(41 L'9T-)
9¢TP-
(LE°6T-)
LEL¢-
(%90'S¢-)
9€8I-
(OM 1°€S-)
€8IT-
(9L'LT-)
CObrr-
(416'8)
vIST
(ALLE)
SSI-
(%79'9-)
ZLT-
(%90'°8T-)
+98T-
(ISL Udase) sepamoqyuor syvg Gung
(8GL] ‘SANVNNIY) sosogaqod syvq qual py
(96/1 ‘SQaVNNI']) vdoyuv syog duly
.
+
Climatic Risk Atlas of European Butterflies
688
nN
(sa)
v8L- [Bee le L¢6 60L O8¢T (GLLI SIVA) ada syog ual py
%GE-) | (zr 9Or-) (/L99T-) |(%60' FZ | (HTL'ET) | ob9° LTD | MIL bZ7) | S'S-) (r6'T1-)
68hL- $£66- L9GE¢- vSIS- 9¢6C- OP8z- 6LG6¢- 988- €807-
Jessodstp | jessadstp | [essodstp | [essodszp | tessodszp | ,essodsrp | ,essodstp | tessodsrp | tessodstp | [essodszp | [essodstp | testodstp
ou ou ou Wy Ty Wy ou ou ou Wy Wy Wy
(GL ] ‘SQHVNNI‘T) auqyv-9 syog (uy
o
S
iam
689
Appendix 2
%18' Lb)
06€7-
%16' LP)
Goer
%G0'9S-)
ZSE9-
%6S'8E-)
Gags
%GG LB")
lg tke
%BE'ES-)
L9€T-
%6L'LL-)
EGre-
(91 0L°)
cc¢-
%LES8-)
LLOZ-
GEG Dis)
SIL¢-
%bS'68-)
1S¢-
%LL'PS-)
%/.0°LS-)
ZITE-
%6L'OL-)
€S79-
%BEEL-)
SIE8-
%8T'0S-)
Coil
%8TF6-)
S¢Or-
PE VL)
88zC-
(487 €6-)
6967-
(LL'S8°)
ver-
(ATT 16>)
a Waa
(46¢0L-)
€L09-
%EG'8S-)
SCG
%8h'TS-)
G8Tr-
%Lb'86>)
98¢-
%1T7'69-)
PERG:
%80° TT-
69'S T)
168
) | (%78'T-)
L9I-
%68'O1-)
%9'8E-)
9F91-
%E LHL)
G79
%b6'E1-)
%78'SP-)
1961-
%ZO'TS)
ZLEL
(%L'€r-)
16€I-
(41 'FT-)
€TI-
(479 '0T-)
O8LI-
(%€0'S¢-)
OZ8I-
(469'7£9)
O8bT
%bb'h)
vST
%8E9L-)
%GT'61-)
0867-
%6S°TS-)
bSece
%80' TL)
(°49°07-)
OLOT-
(816011)
Src
(0496 '€@)
CHS
%E0'TC-)
6SZI-
“bh'6C-)
0097-
%BE8C-)
6ber-
%1T9I-)
OLSze
%€6'0E-)
VECls
%TSHC-)
Leal
%ES'SS-)
187-
%TH'S9-)
FASS)
%6S'LT-)
Z8ET-
%€ESTI-)
%GELT-)
IZrl-
%G8'PS-)
GIc
%EGOT-)
Gi Gls
%EL'TS-)
aa
%78'6E-)
8OI-
%CF'6L-)
9TLI-
(“oer 0¢-)
Srre-
(%€9'LT-)
O€LT-
(456'8r-)
G607-
(%S7'Sc")
LUI-
(0499 'Tb-)
9ZEI-
(048°SP-)
ZEC-
(418°EL-)
Z6LI-
(60°61)
8r9I-
(41E'81-)
OLZz-
(%90'¢r-)
LETC"
%G1'S9-)
16€1-
%8T'ES-)
68EI1-
%89'0E-)
GS9-
60" ES)
Pel
%1L9'PE-)
6€L-
%/68'€8)
L8IZ
%89'0S-)
Z801-
% BCP)
LSI
(LOGL “UHONTANV.IS) Myrmpas sopsMauT
(POL] ‘SQAVNNI'Y) Yyeuvo symaurT
(QGL1 ‘SAAVNNID) yudod sapuaury
%GO'LT-) [(%L0°9) | (%98'7T) | (%78"Z)
OFS I- LYE SEL 191
%66'SE-) |(tP'TT-) 18001) | (%S'ST-)
6LIE- 6001- 068 69€1-
%EG TH) (L778) |(%B8L'ET-)
L¢6- G69T-
(GLLI ‘DYNAWALLOY) Lyng varIYIN
618b- (Lp8 1 ‘NNVINSSY) syuniuojuug vartyay\y
%€9'ET-) | (L'0)
8S9¢- 601
(OS8] “REISOIN) vyainp verjyayy
(ACF OF)
L861-
(VLE)
L91-
(ALIS)
9GLI-
(0468'8S-)
867-
(61'S L-)
veSI-
(~z'Ze-)
O8LT-
(0499 'TT-)
989T-
(Or Th)
vSIT-
(£962)
89T1
(ALS L)
SEE
%9E'S)
€9C-
%G/'E-)
6I-
%EPE-)
ZE8-
%bS'E1-)
6911-
% 8h OT)
OrST
%Lb' TE)
SEO
(%ZL'0)
188
(19°09)
%/9'F1-)
L9v-
%9L'€1-)
Chis
(H€ES81)
C6L (LGQ]. ‘NIMISUATES]) Saprouaqpind vary ayy
(1T81)
108
(67'ST-)
SO8-
(b0'ET-)
99- (6821 SONV’]) vunuip vevjyay
(67 0S-)
O@ZI-
(o/ 0S MAN =)
TIESI=
(80 TL)
(LSQT MAC-IsANN)) vane varyyayy
(ZEQ]T AAMC) auotap vavjya
(6LLI ‘taasyy) vuaGpap vorzyayy
(SLLI ‘[uarianenttos))
DIAL DODIY INE
(OZR ANANEL) eaaqzan verjyayy
(6LLI SAZIOD) aqaogd varyayy
| oaas | naiva |
yessodstp | yessodsrp | [essodstp | [essodstp | tessodszp | ,essodsrp | ,essodstp | tessodsrp | tessodstp | [essodszp | [essodstp | testodstp
ou ou Ty Wy ou ou Wy Wy
ou , ou ,
i)
S
ay
Climatic Risk Atlas of European Butterflies
690
(0499 Lb-)
97Z9-
(4 18'S¢-)
Z6E€T-
(%L'0¢-)
O@rb-
(45'6¢-)
1Z9¢-
%GEVE-)
%GT91-)
%PL'T9-)
HRS Se
(0498'L9-)
69LI-
yessodstp
ou
(%65°SP-)
L8r-
(%L6°9S-)
Ibrl-
(“bh LE-)
LOSZ-
(81 Tr-)
+619-
(GF 1S-)
LLL
%LP'6b-)
LOI8-
%61'9T-)
€66¢-
%LL'SS-)
SPSS
“bb 18-)
COLI-
%LL'L8-)
EGE
%6L'TL-)
O7SI-
%9F'98-)
VSTC-
nana
Jessodstp
ou
%8S°L9-)
Clue
%/.6'99-)
LYL8-
%bG'9C-)
Dyke
%TH'TS-)
869L-
(484'79-)
Raine
%S1'6S-)
EiehGe
%LESE-)
0808-
%Z9'EL-)
Z88-
%SL 167)
L008-
%TO'EL-)
OLTL-
%S1'06-)
G88I-
(76'96-)
SI6I-
(85°78)
€9LI-
(b'€6-)
Gupc-
[sv |
(8¢69)
Cer
(G7 Ze")
CIZb-
(I'S)
O8L
(
%G9'9T-)
Orre-
%9F OL)
ZrOL
%E'C6)
1S61
(LV OF-)
S6L-
%/b'S)
vl
)
(
(%9'C€
918
Jessodstp | [essadstp
ou Wy
(1 LTO)
(4€9'L¢-)
S1Gb-
(%F6'9-)
610I-
(LETH)
6L8E-
%€8'9C-)
L809-
(0487'S)
9TS
(FT OL)
“AWG
(49'9G-)
IZEI-
(4G LT-)
8SP-
(LL'LS)
90ST
inanvar
Jessodstp
my
%EL'SH-)
8SS/-
%79'E1-)
EVILS
%86' OP)
% 6b CL)
TELE
EG
% 49" LE)
L8L
(PL 1S)
(467° €9)
6F91
| svuo | oaas |
Jessodstp | jessodstp
Wy ou
|_osoc_|_osoc_|
%b6 LEZ) | (%S'ST-)
697-
(1 E9¢-)
er Ly-
%99'TE-)
Z8IT-
%G LI-)
69SC
ON Se>)
Ghee
%SL'6I-)
S9I¢-
%LEBE-)
6Sr-
%6E'PP-)
668¢-
(A7E'6S-)
bLII-
(L8°0S-)
980I-
(AC 6E-)
8Z0I-
(41G°FT-)
61€¢-
(EC OL-)
60I-
(478 °TE-)
L8@b-
(LEE1-)
£68-
%PLSI-)
£99T-
%86'LE-)
LL¥S-
%CH'ST-)
LOcP-
%9G°9¢-)
8or-
%8E'Ob-)
LYSE¢-
(“6 '0-)
S80C-
(LT 8T-)
68S-
(4S '09-)
861 I-
fnanvar
[essodstp
ou
(498° LT-)
bLL¢-
(4S¢'6L-)
€09-
(ALE TS)
L€¢-
(AL T6¢-)
601S-
(b0'EE-)
LOCT-
%86'€T-)
1ZS¢-
%€6'6E-)
9G9¢-
%LL'9T-)
SIbb-
%€9'LI-)
Z99T-
%99'Cr-)
€7S-
(97 F¢-)
ZIP:
(pP'€T-)
88b-
(°480°69-)
LOGT-
[sv |
essodstp
ou
(%L8°T-)
(PL Ch)
(°%98'0-)
9ZI-
(%LS°¢-)
06S-
(°%6r'0)
(as
(4S4 IZ)
LSE
)
(HET OC-
LLLI-
(H€7'¢)
GE
(
PL PPL)
PLOE
(b'8E-)
09L-
| oaas_| nana
yessodstp | [essodstp
Ty ey
[essodstp
Ty
(S18)
vOll-
(L491)
Cal
(46 OLL)
OLLI
(4€€°€T-)
(ACh bb)
(4€S'F1-)
LOvZ-
(LG 1-)
6SE¢-
(ALL'FL)
LLY
(46h TL)
(“cr'rr-)
618°
(80° TT-)
LOST-
(%96'S7)
L161
(%76'S1)
891
(1167)
ZO8E-
(/tC'P-)
Z8T-
%59'01-)
v9OSI-
PT VE-)
CULG
(L8/] ‘satoniav yy) saddypeo vy duuous0)
(POLL CATION) Py vgduious0’y
(COLT SSQHVNNIY) aueqov vsurdoT
N
>
(QGL] ‘SQUVNNI'Y) Quan vpUMOISD TT
(L8L] ‘sniortavy)
vuvjyodomad VIDULMOISD
(LOL ‘SQUVNNI']) Madam vIvUMOISD TT
(9G ‘SQHVNNI']) Pa&asav azununy
(LLLI UHINVUD) DUDIANOd DINED
%/%69'¢-)
Grg-
%8T LI)
LOZ (GLLI
%TS'8T-)
v0ST-
(9¢'9-)
€¢9-
(56 TET)
6SLZ
((Sr'tr-)
OF8-
SVaD
(SGT ‘SQaVNNVY) sue rungodg
“ATIQNAIHOS]) vye rangodr
(673 UAAMMY) sya vanjody
(LOLI ‘SQAVNNIY) sev saxvuvy’y
(COLT ‘TIOdODS) suamae sodang
(TLLI ‘svrivg) 0gddos sudan
i)
S
iam
691
Appendix 2
%19'69-)
LECH)
O€ZL-
Ce Ms)
€L9T-
%6L'SL-)
%G9'LC-)
(/91'68-)
8E19-
(699 ' T-)
0606-
(10° 16-)
OST¢-
(668°98-)
ZhOT-
%TE LP)
6b9T-
%ELPE-)
vI9OL-
%1T7'06-)
OPL-
%6L'S9-)
Eeoze
%€6'68-)
L0SZ-
%8h'0S-)
GLY-
%€0'Th-)
G6CG-
%BL'SS-)
bLbL-
%68'T6-)
%90'€6-)
9069-
%TEHG-)
ZS8II-
%LO'TL-)
GO0TI-
%GE'86-)
vOre-
%Z0'L6-)
O87C-
%78'09-)
668¢-
%6L'9F-)
SEr0l-
%6F'96-)
86L-
%LT€6-)
Tee=
%69'96-)
689C-
%GEOL-)
Z99-
%L1'09-)
lis
%€0'69-)
0S€6-
%€0'TS-)
Z8S¢-
%€6'61-)
Over
%96'TC-)
8T8E-
%TS'9P-)
OL9T-
%16'LE-)
168-
%LO'VI-)
(469° LI-)
LLET-
(7 9b)
IS¢
%LE'ER-)
6€LS-
%LE'OE-)
L7Z99-
%TL'OE-)
10s
%€6'T9-)
9 We
%TETS-)
90ZI-
%LT8I-)
S9OTI-
(41 8'8T-)
(414'69-)
(0497 '9T-)
9G¢¢-
(%76'9S)
woh
%EE'G6L-)
L9b¢-
%69'TH-)
(ET LS-)
PZ0C-
(Th 'Th)
TCE
(°490'0€-)
6907-
(68'S T-)
ard om
OnClice)
06L¢-
%9F'9C-)
+S61-
%18'8S-)
ZSel-
%€9'TC-)
[Stile
%BSSI-)
OLbE-
(68 1S)
rae
(“Sb'rS-)
676I-
(£0 L9-)
PO8I-
%9T'EE-)
ian
%78'8I1-)
GIPC-
% 9b 0T-)
Wiz
%TL'L9-)
v1S-
pho ee)
L8I¢-
%89'FS-)
L¥SI-
%99' 19°)
ShTb-
%6L'6I-)
S8Ib-
%9'ST-)
bLtb-
%18'6S-)
OLOT-
(04€8'6S-)
90FI-
(1b TC)
L¢vl-
(8 LT-)
BL8E-
(61 TL)
L6S-
(0468°€T-)
Grs-
(%L5°89-)
LOGI-
(0469 '6C-)
6LT-
(p66 '07-)
889T-
%LO'8E-)
LELt-
%bL'6S-)
0691-
%/T'0S-)
bSPE-
%/LG'OT-)
68bb-
%€0'6C-)
OFSb-
%€6'S9-)
Z8TC-
(449'89-)
€19I-
(8¢9T-)
1691-
(6'61-)
6¢bb-
(44069)
Lis:
(19°E+-)
SPSI-
(%79°9L-)
SZIZ-
(479 L€-)
bS¢-
PL’ ET)
8S67C-
%GLET-)
68¢-
%69'TI-)
S08-
%1T LT)
GEE
%16'6E-)
OIFI-
%LT8I1-)
80S-
%GL'LI-)
LOI-
%EL'S)
Go);
%8C'61-)
Gites
“TP Cb)
87CI-
%EL'LS-)
VL6¢-
OHA Fas
%GT'LI-)
%TT TI)
Ee
%GT'Sr-)
66¢-
%9G'9Z)
146
“TH LP)
CSlls
%LEOL-)
L6-
% 9 T-)
vLI-
(PLL ‘NEAS) wovay ayaqdouodty 1
%89'PE-)
186-
%6T'6E-)
869T-
%89'C1-)
986C-
(%r0'r1-)
Oree-
(97 €¢-)
IST I- (LLL ‘SQNVNNI'Y) Sauoqyzy viUOd
(%96°6¢-) (GL ‘SQaVNNI'T)
6¢6- smpiy dung vy duuouso’y
(%L6'L>)
L1S-
(%60°¢T-)
OC6C-
(8G/ | ‘SQAVNNI']) vuqunl omy
(9GLT ‘SQaVNNI']) squcuady sudoyuvy dp
(COLI ‘sntoniavy)) vgasqyog vinod
(FORT ‘NELNVTIV A) 272000 VIMO T
(P8L] dase) sapuvy vyduluousoy
(LOLI ‘saavNNI’D) avaq vg dudious0)
(Z8LI SUAdsy) smlop vyduiuous0’)
(6S F1-)
LGe
(8L'0¢-)
9¢8-
(77°)
LOT-
(%L'0)
06
(Q6L1 ANNA) Byapavs vq fuuous0’y
(98/1 ‘NasSAVHYOg)
nouns vq uuous0’)
‘SQHVNNI'T) veuvup vq duuouso’)
(LOLI
ONG6L ‘SHMTY sesuadopoqu vy duuous0D
sags |nawva| svao | oaas |nawva| svao | oaas |nawva] svao | oaas |nawva| svao |
yessodstp | jessodstp | [essodstp | [essodstp | tessodstp | ,essodsrp | ,essodstp | tessodsrp | tessodstp | [essodszp | [essodstp | testodstp
ou ou ou Ty Wy ou ou ou Wy Wy my
oO
S
ay
Climatic Risk Atlas of European Butterflies
692
(10° LS-)
(%Z0'SL-)
LL8-
%66'1b-)
6681-
%80'TS-)
Orl-
%10'0S-)
6L9-
%19'0b-)
%LE6-)
(497 T€-)
G68¢-
(%)77'S9-)
SrSI-
[essodstp
ou
(71 Lb)
€8r-
(%G'P9-)
0997-
(499 '66-)
OLFI-
(b8°TS-)
OL0T-
(9F'TL-)
LYg-
%GT TS")
SIET-
%8T 8b-)
989-
%TL'6b-)
999-
%90'Lb-)
9ZSI-
%G8'8r-)
199-
%61'S9-)
1Sc-
%TT 8S)
SIZI-
“be Eh")
88EC-
%EL'L8-)
Z8PT-
nana
Jessodstp
ou
(04€8'99-)
S89-
(%EL'6L-)
88Z¢-
%PL'89-)
Z6SC-
%19'08-)
€h6-
%6S'€9-)
9L8T-
%CS'P9-)
L16-
%9E'19-)
ZE8-
%G9'9G-)
LE8I-
%78'T9-)
0S8-
%96'8L-)
vOE-
(0499°€L-)
IvSl-
(oS 1S")
SIr9-
(451967)
OCLT-
no
(LEL)
bO¢
(19 Tb-)
€8SI-
(18 1Z-)
GST-
%98' OL)
lor
YAS)
66r-
% 9G 1¢-)
STP-
%9L'9L-)
oS:
Oren)
B9¢-
((Gh'TC-)
86Lc-
Jessodstp | tessadstp
ou Wy
(4LS'C)
901
(%TC'TG6-)
LYEI-
(A61'Ch-)
SO9I-
(461 '0T-)
9¢T-
(°%66°9)
91¢
(4L9'9T-)
6LE-
(AL L'CC-)
LUL-
(0468°91-)
87C-
(461 €T-)
GLT-
(0465°S¢-)
S¢hb-
Jnana] svuo |
Jessodstp
my
(460'0S-)
e1s-
%GT7'S6-)
SOPI-
%€8'8S-)
8CTC-
%8EHC-)
GRC-
%TVTL)
8h
%8E Th)
8gc-
%CH TE-)
9TP-
%F6'0C-)
6L9-
%TL'6C-)
ZOP-
%CV TE-)
(%6'€T-)
(417 €8-)
¥S¢T-
[essodstp
Ty
be OF)
LOG6I-
%66'6C-)
Ivl-
%/b'0S-)
06S-
%€9'LP-)
€88I-
(4G7 1 ¢-)
brr-
%1L6'SE-)
L8r-
%C6'TE-)
8901-
bv VE)
99¢-
“br 6C-)
SZ8-
(60° T¢-)
GL8E-
(67 9b-)
80€T-
essodstp
ou
(GS TE-)
Bcs-
(40° T€-)
bLT-
(4L1'ST-)
8S7-
(0469 '6¢-)
Se9I-
(460'°6L-)
99LT-
%1L0'CS-)
809-
%60'FE-)
CHSIe
%BELT-)
68¢-
% 49 TC-)
6Cr-
%6S°TE-)
LSOI-
(471 6C-)
v6E-
(0479'9¢-)
Ivl-
(49L'9¢-)
69L-
nav
Jessodstp
ou
(460°C)
Ale
(86'6¢-)
eS¢-
(PE re-)
Ate
(97'SS-)
6LTC-
(89'F8-)
6h71-
%ELTE-)
LOTI-
%ZO0'6S-)
069-
%TTC OY)
9TTT-
%65'9¢-)
0zs-
%L6'6E-)
ZhS-
%HEL Ob)
1zel-
(%S OF)
SrS-
(89 b+-)
ZLI-
(%8S°TS-)
OOTT-
no
essodstp
ou
(464 9T-)
ZSb-
(467 0C-)
80C-
(466 1Z-)
S¢8-
(Mb 1T-)
(AE €T-)
(04S0'S7-)
vCS-
[essodstp
my
(486 1T-)
(HLS'8-)
€¢-
(46L'0C-)
S¢r-
nave
yessodstp | [essodstp
my Tey
(67'S¢-)
Z09-
(L0°LT-)
6£7-
(bore)
9IbI-
(/6+'69-)
SZOI-
(461 TT>)
908-
(b8'0¢-)
S6EI-
(°490'6C-)
Clr
(68'F7-)
LE¢-
(477 O€-)
086-
(/6¢°9T-)
LS¢-
(/1L'ST-)
66-
(418° L€-)
I6L-
no
(Q6L1 SUANNOAUG) SHUDRIOGID DIGIT]
isa)
(L8L] ‘snIoniavy]) espouse vIqaLry
(L6LT ‘NIDIONg) Bp viqaszy
S
Sa)
Ww
(L6LI ‘NIDIONg) yqua viqadTy
oe)
(Q6LT SUANNOAUG) Puouy DIqgary
(LLLI Haase) sdoryjan vigour]
(SLL ‘VIssany) sudunjau viqary
(Post “HANANED) méogd veqadry
(EL SHOONS) wouqdida viqauy
(GLLL ‘[uarrianerstaos))
OJUDUL DIGIATT
(OCQT WAAR) ydqdua viqary
(SOQ “Uadsy) ynine viqauy
(QL ‘SOAVNNI'Y) D004 DeqasTy
Sat
—
wa)
(9¢QI ‘vson) vuadig apaq douodyy
o
S
iam
693
Appendix 2
%LTT9-)
L891-
%CL6b)
BEC
%69'ST-)
%TL'T3-)
%bL'S9-)
C18I-
%9L' Lb-)
€7S-
%/L'99-)
vor-
%LP'P9-)
60r-
%17'S9-)
O87-
%/.0'9P-)
vLL-
(LS°ES-)
0¢9-
%EG'79-)
GLG
%GL'TS-)
006-
%68'TS-)
%G8'99-)
Zi:
%TE9L-)
96+-
%9€'68-)
S09-
%€9'08-)
ace
%SLP8-)
19¢-
(481 '09-)
LLOI-
(475°€9-)
LYL-
(4€6'LL-)
GEC
(bb'SL-)
OLP-
%10'98-)
Ivc-
%T1'99-)
SZII-
%6E'EL-)
8r9-
%GO'€1-)
09¢-
PL’ €T-)
09T-
9)
Or
% GETS)
617
%96'PS-)
6bS-
%8G°LS-)
LYT-
(SL'8T-)
cies
(%8L'SS-)
9S9-
(65'0©)
C61
CAtaca)
L8S-
%GE'CT-)
br9-
%9LPI-)
Sols
%€6'€T-)
Z9I-
(467 6C-)
98I-
(%66°81-)
GLE
(“ob9'vb-)
STS
(440'ST-)
8SI-
(469'bT-)
69-
(HE 1E-)
PES-
(487° TE)
G87-
%TS Lb)
SrlI-
Cae)
Cire
%9T TH)
icin
%/.6'Ob-)
O€II-
%LEVT-)
8SI-
%/.8'69-)
€Ly-
LY bb)
CSG
%16'TS-)
Ee
%C6'EE-)
OES
%~SEVE)
vOr-
MET LY)
GOT-
(/€0'8¢-)
OrT-
%61'F6-)
Z9I-
/88'€C-)
Oca
%G9'PEC-)
90¢-
(491 LL>)
691I-
(414 81-)
0¢s-
(/66'6¢-)
COLI-
(417'S7-)
9LT-
(4LL'9€-)
6€T-
(480°TL>)
88b-
(%87'S¢-)
VCC"
(470 8b-)
90Z-
(1'8T-)
ZLb-
(PL 'TE-)
SLE-
(48L'9¢-)
09I-
(L0° L-)
L6T-
(°460' L6-)
LOI-
(67 VT")
89-
%9LP8-)
FTA Be
%b0'LT-)
SIZI-
%9€'0S-)
68E1-
%bS'SC-)
1€7-
%96'9L-)
1ZS-
(ATL Lb-)
COE-
(bS'19-)
b9T-
(6 T Th)
Z69-
(0498'8¢-)
LSb-
(6'9b-)
bOT-
(“818P-)
bO¢-
(48'86-)
OLI-
(LG'8¢-)
80I-
%GG'1S-)
I8L-
%PLTT-)
0S6-
%ET €-)
68-
% 18 LI-)
Cie
% PLT)
6LI
% 8h T-)
Ol-
%EVTI-)
fee
)
(79° 9b-
007-
% ET HC?)
LOF-
%99'8T-)
LEG-
%7S'S-)
vC-
%F6'01-)
(41L'SC)
rib
(4) 17'S9-)
(46'S-)
ESi-
%TL'TL)
IS
-)
%€6'S6-)
S9I-
%PT'LS)
O91
(%478'F9-)
Z86-
(86°9T-)
ZIZI-
%ITLT-)
867-
%79'81)
IZ1
99°)
al
(%9'LT-)
ZII-
(Abr 6S-)
SSZ-
(/88're-)
98S-
(pb8'Ee-)
(090°S1-)
S6-
(%€6'S6-)
S9I-
(6L'9@)
a
ou , ou , 104 ,
yessodstp | jessodsrp | [essodstp | [essodsrp | tessodstp | ,essodsrp | ,essodstp | tessodsrp | tessodstp | [essodszp | [essodstp | testodstp
ou. oe Tey Ty og OU, Tey Tey
(Q6L1 SAHNNOAN) SUNj0IM DIGAAFT
(GOST “sddsy) aaa0 vigour]
(O6LI “ISHUME]) Seam vigarTy
(9781. “IVANASIOg) sypuoau Diqary
(Q6L1 WANNA) VUrUOM DIqaLy
(PEQy “UAAMIY) MGS vigour]
(OSLI “Uaasy) aouoad viqauy
(xojdusos) (761 | “HIVANNAHOLT
2 UANLAY]) Saprowssw9 vigadry
(LQL] waa) smenpug viqaary
(LQ Walt VHOS-HORMAEY)
DUDULOLJO DIGALT]
(61ST ANANEY) aalgsrda viqasry
(SOT lHaSe) Bysaum vIqadT]
(GOT “AHaSH) atlas vegas]
(Q6L1 WANNA) 9779 PIgadsy]
o
S
ay
Climatic Risk Atlas of European Butterflies
694
(46S 9b-)
£68-
(419°9L-)
G¢SZ-
(F0'8E-)
L8bT-
(486° 19-)
L19Z-
(4L0°F6-)
ZCSI-
bb LL-)
90LI-
%8L'SL-)
€¢8-
%EL'S6-)
Srel-
%G9'ES-)
MLE TCS)
(“bT'68>)
ZGEI-
(479 'FT-)
90LT-
[essodstp
ou
(S1'+9-)
L€ZI-
(669'68-)
8967-
AIS TL)
990¢-
(65° L6-)
6LSI-
%6L'€8-)
9F8I-
%1.6'16-)
610I-
%6L'66-)
ZOPI-
%67'99-)
60b-
%88'LL-)
€L8-
%1.S'86-)
PL€l-
%TG LP-)
€¢99-
%TI'S6-)
Ir l-
%G1'8T-)
S9TI-
[naive
Jessodstp
ou
%T9'bL-)
ZCbI-
%Cb'96-)
16L€-
%69'L9-)
STrb-
%19'18-)
COre-
%1.S'66-)
LI9I-
(ATH 'CG-)
9¢0T-
%b8'96-)
€LOI-
GPCL?)
Le
%T76'68-)
800I-
%1.T'66-)
bOSI-
WIP PE)
OFSI-
[svi |
(86'€9)
ZS9
(%8T9L)
O€ZE
(67 9F-)
6bL-
%91'LI-)
BL¢-
%99' LL-)
b6L-
%b6'9L-)
L801-
%LL'STI)
OLL
(“6 17)
OFT
(%89°SL-)
SSOI-
(9'¢)
8b
(“9 6b-)
ZSL-
Jessodstp
Ty
Jessodstp | [essodstp
ou Ty
(LY 0I-)
LOZ-
(461'€9-)
1607-
(°498'68)
OLE
(46L'79-)
9TOI-
(487 LT-)
109-
(416 16>)
610I-
(LEFT)
(4L5°9¢-)
OLb-
(491 '0-)
bu
(%TL'L9-)
9ZOI-
(461 ET)
(F0'€9-)
980C-
LPT)
Peel
%€S' LOL)
SOC
%EL'T9-)
SIOI-
%b9'8T-)
1¢9-
%b8'96-)
€LOI-
MEVEL)
r10I-
% 9901)
WIS CL)
%b9'Gb-)
ZSL-
WPL TE)
trrl-
[essodstp
Ty
%LO'TE-)
G6S-
%68'6t-)
1S9I-
%b6'ET-)
G9SI-
%9L9F-)
LS6I-
(LET)
LLEI-
(4b LFS")
90ZI-
(ALE 6F-)
LYS-
(4869)
9LOI-
(bS'GP-)
18Z-
(bor)
LGt-
(TE TSL-)
LYOr-
(/17'61-)
1897-
(br 9L-)
8ST I-
(46¢°CC-)
900T-
essodstp
ou
(0,96'SS-)
6+S-
(“1 6C>)
L€0b-
(0499 '€¢-)
9+9-
(0484'89-)
99TC-
(491 97°)
OILI-
(4 19°SP-)
P<6I-
%TS'09-)
CCEI-
%8b'6S-)
6S9-
%£6'08-)
LELI-
%9T Lb)
16Z-
(449 Lb-)
veS-
(ET 18>)
L€LI-
(49° 17")
S1OE-
[naive
[essodstp
ou
(419'8S-)
Gis-
(8 TE-)
SObb-
(87 0F-)
€LL-
(/66°7L-)
bIbe-
(S0'F¢-)
9TCT-
%98'h8-)
€LEI-
% 9 TL-)
ZLSI-
%LTT9I-)
069-
%89'88-)
9FCI-
“CP 6)
SO¢-
(04G¢°€S-)
86S-
(8E'88-)
C&TI-
(414'9T-)
989¢-
[sv |
essodstp
ou
(4¢ Ch)
(04G¢°€S)
ZITT
(416 T€-)
€0L-
(47S Tb)
09b-
(°460'6L)
88h
(416 19-)
€98-
(%EE9)
¥88
[essodstp
oo
(%6L'6>)
96-
(4€9'€T-)
697E-
(%L0'SZ)
8b
(HL FS-)
OI8I-
%1S'S9-)
090I-
%GS'08)
LGb
2)
inane
essodstp
Tey
(48'S)
Goh
(/9L'ST-)
b9SE-
(4€8' LZ)
bES
(91 0F-)
6ZEI-
(/L°T1)
0¢8
%10°9S)
GLET
%GE LP)
699-
LL TP)
Lb6-
(LL8 1 SWHONIAAV.LS) Moves viqowwddapy
N
ive)
(POLL SSQHVNNI'Y) auordsay vigouwd day]
(COLT ‘rIodons) sof rrqasvd dit]
Ne)
ice)
Te)
(COLL ‘tIOdO0S) sap seourpy
©
(OSLI “uHasy) vavjw suas
(COLT ‘sniortavy) wmsal snags
©
(POST SODUSNVWAHOPL) Sau2 vidavuvjayy
(COLI UNAS) 22UIILII0 DIGADUDJAINT
(OLLI ‘AZ ING) akin viIRuuma
(17 18°)
IWIl-
(S911)
(P8L] Udase) vssany visuvuvjayyy
(O6L1 MINANP) Seaqoyy viduvuvpayny
(pb)
$7
(96'FL-)
ShOI-
(°460'0-)
ZI-
oo
[essodstp
Tey
) DayIvIVE DIRIDUIIATY
(P8LI UNAS) avessms vIRADUDIITNT
Ne)
ive)
(QQ ] ‘NNSAVHMNOG) aoupund viqaary
o
S
iam
695
Appendix 2
%97'99-)
Clee
%LS Pb)
OL9C-
%6b'89-)
%16' 16>)
%L0 18>)
ve-
%8S'1S-)
060¢-
0 88'9G-)
65>
%bL'L6-)
O¢0T-
(%8'8S-)
18r-
(%%Z0'8S-)
L¢cp-
(69¢°99-)
8L0S-
(MSL 18-)
S6Sr-
%TL'SL-)
1¢9-
%97'96-)
OLIZ-
89" L9-)
£69-
%19°S9-)
L1OF-
%76'VL-)
Go
%10'9S-)
%9G'68-)
69¢-
%88'P9-)
L88¢-
%PEOL-)
OIL:
%~Tr'66-)
S90C-
%9L'9L-)
€79-
%GYPL-)
6tbS-
%L0'T8-)
€819-
%EL'S6-)
18¢s-
%LE 16)
LOL-
%Gr'66-)
O81Z-
%9L'08-)
LZ8-
%1L8'P8-)
88 1S-
%9T'L8-)
9S8-
% Lb OL-)
Sr L6-
%96'LI-)
vL-
)
%BEOE-
O78 I-
“bo OF)
O¢r-
%CF'16-)
6681-
6° €T)
£61
%1S°S-)
ZOP-
(Sr'7)
G8
(696 01-)
9T9-
(b9' FZ)
LOZ
(€L'0S-)
ZIII-
(6°70)
GUE
(%79'I-)
667
%EC LY)
S6I-
%66'PC-)
9607-
%GT'6C-)
ZL
SLO”)
9ZOT-
PT BT-)
L€Z-
%F0'9I-)
OLTI-
(4G L'T1-)
196-
(a Cz-)
ZEEI-
(HECEs)
O87-
(°4G7'69-)
SISI-
(/€6'€T-)
SbT-
(LT'8-)
90¢-
(4b0'Fb-)
Zr
(0496 'Lb-)
$¢99-
%LT 6b-)
€0C-
%6E Lb-)
6£8C-
%19'TS-)
O8r-
%TH 667)
S90T-
%9€'6E-)
Gee
%€O'TE-)
(“PT 0C-)
(0498°TS-)
tbb-
%€G'S9-)
Ong
%68'6C-)
Vals
%LS'6C-)
89¢-
%~TH'18-)
1691-
%LO'P-)
09¢-
%80'9T-)
ZOG6T-
%19'0E-)
80€T-
((S I4-)
OLce
%19' Lb)
OS¢-
%bO'PL-)
CZOile
%L0'LY-)
Z8r-
%€8'LE-)
L¥6I-
%b6'8C-)
Z8¢-
Oh eee)
G60¢-
(0465°€9-)
Z9C-
(4EE'FE-)
LSOZ-
(0465°€9-)
Z9T-
(4EE'FE-)
LSOC-
(EL FE-)
€TE-
(477 18>)
L891-
(465 Ob-)
TEC"
(“6 0E-)
9S7C-
(466° LE-)
+98T-
(4¢4 Lb)
999C-
(488'FE-)
€67-
(419° Th)
6¢b-
%6L'EL-)
vOS-
%6L'EL>-)
vOS-
%LO'EP-)
O0b-
%€9'C8-)
LELI-
(410° Lb-)
G8¢-
(pb LE-)
LELT-
(%69 It)
Cvle-
(L7TCS-)
8C6T-
(%S6'St-)
98¢-
(/bS'S8-)
SL8I-
(41'0S-)
Ore:
(9 Et-)
OF9T-
(415 6b-)
%1S'6b-)
¥0C-
%GPr'6I-)
S9OLI-
%G9'0T-)
Z61-
%€0'T8-)
€891-
%6E TT-)
Sy he
0/)G8'8)
Gr9
%TO'FL)
LSOT
% 61)
8901
% BE L-)
Z9-
AGE -CC-)
bese
/69'8-)
68-
“TP OL)
POOL
(GS Tb-) }(H€L0S-)
%PLYC-)
OFFI-
%EL'0S-)
607-
%PLYC-)
OFFI-
(9¢°9T-)
Lve-
(467 €8-)
6TLI-
(HEV'C-)
I8I-
(/6L'b-)
6rS-
(%9S'0-)
cr
(99°)
L8b
(ZV 01-)
S8-
%€0'6S-) | Sr Lr)
$67I- OFOT-
M6) | (%86'8-)
9F- Z6-
%76'0-) | (%L7'8)
9¢- 90
(QSLI ‘SQaVNNI']) sadqsilqa supuncgy
(9081 WANIAED eynl srauac
(G8L1 “TION sym0IF stoUaK
(L6LT ‘NIIIONG) Pusou srouac_)
(LES AGH —) vyaqjun vavsDqyopnasy
(POL ‘SQAVNNI']) seasmg vuvsoyD
(GLLI ‘SQIOMtaV ]) ane vesajueg
(GLLT ‘[uarrianerstanos))
DING JaID DUDSHUJalL7
(QO6T WAIOISHA]) sagzuas vequvd gay]
(LOLT ‘SAHWNNI']) vepif viqauwddapy
(Gpgy UaAaay) omyvl rqouvddipy
(99L] “HOVNaNyY) suugyeys viqoavdday
(ZG6]. ‘AONVANHSUOd
-NIHOOZVYN) Sesuacpon viqaswod apy
(9G ‘SAAVNNI'T) was viqounddapy
ou , ou ,
yessodstp | jessodsrp | [essodstp | [essodstp | tessodszp | ,essodszp | ,essodstp | tessodsrp | tessodstp | [essodszp | [essodstp | estodstp
ou ou Ty Wy ou ou Wy Wy
oO
S
ay
696 Climatic Risk Atlas of European Butterflies
Appendix 3: Risk category statistics of European butterflies
under different scenarios
Table App. 3.1: Number of European butterfly species in different risk categories under
different scenarios
HHHR = extremely high risk (loss >95%; AUC > 0.75)
HHR = very high risk (loss >85%; AUC > 0.75)
HR = high risk (loss >70%; AUC > 0.75)
R = risk (loss >50%; AUC > 0.75)
LR = lower risk (loss < 51%; AUC > 0.75)
PR = potential risk (any loss or gain; AUC < 0.76)
LR with incr = lower risk with gain under full dispersal (AUC > 0.75)
climate change risk categories
5
a=
a “ S <=
a0 — & 5s apie
2050 no dispersal
2080 full dispersal
2080 no dispersal
Appendix 3 697
Table App. 3.2: Percentage of European butterfly species in different risk categories under
different scenarios (excluding the PR-category)
HHHR = extremely high risk (loss >95%; AUC > 0.75)
HHR = very high risk (loss >85%; AUC > 0.75)
HR = high risk (loss >70%; AUC > 0.75)
R = risk (loss >50%; AUC > 0.75)
LR = lower risk (loss < 51%; AUC > 0.75)
LR with incr = lower risk with gain under full dispersal(AUC > 0.75)
climate | climate change risk categories =| risk categories
aon CS) EY
(Yo species)
RAS % Se eee
HHHR
LR total
LR with incr
Chmpeies) [BAMBI i)
(o species)
GRasy | 12| 41| tor] a] oz] oo
(0 species)
GRAS 1.6 6.1 63.9 17.6
2080 no dispersal
(0 species)
698 Climatic Risk Atlas of European Butterflies
References
Ackerly DD (2003) Community assembly, niche conservatism, and adaptive evolution in
changing environments. International Journal of Plant Sciences 164 (Supplement): 165-184.
Albre J, Gers C, Legal L (2008) Molecular phylogeny of the Erebia tyndarus species group
combining CoxII and ND5 mitochondrial genes: A case of a recent radiation. Molecular
Plylogenetics and Evolution 47: 196-210.
Anton C, Zeisset I, Musche M, Durka W, Boomsma JJ, Settele J (2007) Population structure
of a large blue butterfly and its specialist parasitoid in a fragmented landscape. Mokcular
Ecohgy 16: 3828-3838.
Araujo MB (2002) Biodiversity hotspots and zones of ecological transition. Conservation Biology
6: 1662-1663.
Araujo MB, New M (2007) Ensemble forecasting of species distributions. Trends in Ecology ¢™
Evolution 22: 42-47.
Araujo MB, Williams PH (2000) Selecting areas for species persistence using occurrence data.
Biological Conservation 96: 331-451.
Araujo MB, Williams PH (2001) The bias of complementary hotspots toward marginal
populations. Conservation Biology 15: 1710-1720.
Araujo MB, Thuiller W, Pearson RG (2006) Climate warming and the decline of amphibians
and reptiles in Europe. Journal of Biogeography 33: 1712-1728.
Balmford A, Bennun L, Ten Brink B, Cooper D, Coté IM, Crane P, Dobson AP, Dudley N,
Dutton I, Green RE, Gregory RD, Harrison J, Kennedy ET, Kremen C, Leader Williams
N, Lovejoy TE, Mace G, May RM, Mayaux P, Morling P, Phillips J, Redford K, Ricketts
TH, Rodriguez JP, Sanjayan MA, Schei PJ, Van Jaarsveld AS, Walther BA (2005a) The
Convention on Biological Diversity‘s 2010 target. Sczence 307: 212-213.
Balmford A, Crane P, Dobson AP, Green RE, Mace G (2005b) The 2010 challenge: data
availability, information needs and extraterrestrial insights. Philosophical Transactions of the
Royal Society of London Series B-Biological Sciences 360: 221-228.
Barry S, Elith J (2006) Error and uncertainty in habitat models. Journal of Apphed Ecology 43:
413-423.
Berthold P (1998): Bird migration: genetic programs with high adaptability. Zoology — Analysis
of Complex System 101: 235-245.
Berthold P, Pulido F (1994) Heritability of migratory activity in a natural bird population.
Proceedings of the Royal Society of London, Serres B 257: 311-315.
Berthold P, Helbig AJ, Mohr G, Querner U (1992) Rapid microevolution of migratory behaviour
in a wild bird species. Nature 360: 668-670.
Blows MW, Hoffmann AA (2005) A reassessment of genetic limits to evolutionary change.
Ecology 86: 1371-1384.
Braby MF (2005) Provisional checklist of genera of the Pieridae. Zootaxa 832:1-16.
Bradshaw AD (1991) Genostasis and the limits to evolution. Phzlsophical Transactions of the Royal
Society of London B 333: 289-305.
Bradshaw WE, Holzapfel CM (2001) Genetic shift in photoperiodic response correlated with
global warming. Proceedings of the National Academy of Sciences, USA 98: 14509-14511.
References 699
Broenniman O, Thuiller W, Hughes G, Mideley GF, Alkemade JMR, Guisan A (2006) Do
geographic distribution, niche property and life form explain plants’ vulnerability to
global change? Global Change Biology 12: 1079-1093.
Case TJ, Holt RD, McPeek M (2005) The community context of species borders: ecological
and evolutionary perspectives. Ozkos 108: 28-46.
Cohen J (1960) A coefficient of agreement for nominal scales. Educational and Psychological
Measurement 20: 37-46.
Crozier L, Dwyer G (2006) Combining population-dynamic and ecophysiological models to
predict climate-induced insect range shifts. The Amenvan Naturahst 167: 853-860.
Davis AJ, Jenkinson LS, Lawton JH, Shorrocks B, Wood S (1998) Making mistakes when
predicting shifts in species range in response to global warming. Nafure 391: 783-7806.
de Heer M, Kapos V, ten Brink BJE (2005) Biodiversity trends in Europe: development and
testing of a species trend indicator for evaluating progress towards the 2010 target.
Philosophical Transactions of the Royal Socety B-Biological Sciences 360: 297-308.
de Jong, R (1972) Systematics and geographic history of the of the genus Pyrgwsin the Palaearctic
region. [ydschrift voor Entomologie 115: 1-121.
Descimon H, Mallet J (in press) Bad species — How ecology and evolution confuse butterfly
taxonomy. In: Settele J, Shreeve TG, Konvicka M, Van Dyck H (eds.): Ecology of Butterflies
in Europe. Cambridge Untversity Press.
Dobson A (2005) Monitoring global rates of biodiversity change: challenges that arise in meeting
the Convention on Biological Diversity (CBD) 2010 goals. Philosophical Transactions of the
Royal Soctety B-Brological Sctences 360: 229-241.
EC [European Commission] (2001) A sustainable Europe for a better world: a European
union strategy for sustainable development. COM(2001)264 final, Brussels. http://www.
europa.eu.int/eur-lex/en/com/cnc/2001/com2001_0264en01.pdf.
EEA [European Environment Agency] (2005) The European environment — State and outlook:
http://reports.eea.europa.eu/state_of_environment_report_2005_1/en.
EEA [European Environment Agency] (2007a) Halting the loss of biodiversity by 2010: proposal
for a first set of indicators to monitor progress in Europe. Office for Official Publications
of the European Communities. http://reports.eea.europa.eu. Cited 6 Dec 2007.
EEA [European Environment Agency] (2007b) Europe’s environment — the fourth assessment.
Elith J, Graham CH, Anderson RP, Dudik M, Ferrier S$, Guisan A, Hymans RJ, Huettmann F,
Leathwick JR, Lehmann A, LiJ, Lohmann LG, Loiselle BA, Manion G, Moritz C, Nakamura
M, Nakazawa Y, Overton JM, Peterson AT, Phillips SJ, Richardson K, Scachetti-Pereira R,
Schapire RE, Soberon J, Williams S, Wisz MS, Zimmermann NE (2006) Novel methods
improve prediction of species’ distributions from occurrence data. Ecography 29: 129-151.
Erhardt A (1985) Diurnal lepidoptera: sensitive indicators of cultivated and abandoned
grassland. Journal of Appled Ecology 22: 849-861.
Erhardt A, Thomas JA (1991) Lepidoptera as indicators of change in the semi-natural grasslands
of lowland and upland Europe. In: Collins M, Thomas JA (eds.): The Conservation of Insects
and their Habitats. Academic Press, London.
ESRI ArcGIS [9.2] (2006) Redlands, California, Environ. Systems Res. Inst.
Fiedler K (2006) Ant-associates of Palaearctic lycaenid butterfly larvae (Hymenoptera:
Formicidae; Lepidoptera: Lycaenidae) — a review. Myrmecologische Nachrichten 9: 77-87 (plus
, digital supplementary material“ at: http://myrmecologicalnews.org/cms/images/pdf/
volume9/mn9_77-87_supplement.pdf).
700 Climatic Risk Atlas of European Butterflies
Fielding AH, Bell JF (1997) A review of methods for the assessment of prediction errors in
conservation presence/absence models. Ezironmental Conservation 24: 38-49.
Franco AMA, Hill JK, Kitschke C, Collingham YC, Roy DB, Fox R, Huntley B, Thomas CD
(2006) Impacts of climate warming and habitat loss on extinctions at species’ low-latitude
range boundaries. Global Change Biology 12: 1545-1553.
Geiger H, Scholl A (1982) Ponta daphdwe in Sideuropa — eine Gruppe von zwei Arten.
Mitteclungen der Schweixerischen Entomologeschen Gesellschaft 55: 107-114.
Gomulkiewicz R, Holt RD (1995) When does evolution by natural-selection prevent extinction?
Evolution 49: 201-207.
Graham MH (2003) Confronting multicollinearity 1n ecological multiple regression. Evohgy 84:
2809-2815.
Gregory RD, Van Strien AJ, Vorisek P, Gmelig Meyling AW, Noble DG, Foppen RPB, Gibbons
DW (2005) Developing indicators for European birds. Philosophical Transactions of the Royal
Society B-Biological Sciences 360: 269-288.
Gregory RD, Vorisek P, Van Strien AJ, Gmelig Meyling AW, Jiguet F, Fornasari L, Reif J,
Chylarecki P, Burfield TJ (2007) Population trends of widespread woodland birds in
Europe. [bis 49: 78-97.
Gregory RD, Vorisek P, Noble DG, Van Strien AJ, Pazderova A, Eaton ME, Gmelig Meyling
AW, Joys A, Foppen RPB, Burfield TJ (2008) The generation and use of bird population
indicators in Europe. Bird Conservation International 18: 223-244.
Guisan A, Thuiller W (2005) Predicting species distribution: offering more than simple habitat
models. Ecology Letters 8: 993-1009.
Guisan A, Zimmermann NE (2000) Predictive habitat distribution models in ecology. Ecological
Modeling 135: 147-186.
Hambler C, Speight MR (1996) Extinction rates in British non-marine invertebrates since 1900.
Conservation Biology 10: 892-896.
Hambler C, Speight MR (2004) Extinction rates and butterflies. Sczence 305: 1563.
Hanley JA, McNeil BJ (1982) The meaning and use of the area under a ROC curve. Radiology
143: 29-36.
Hansen TF, Houle D (2004) Evolvability, stabilizing selection, and the problem of stasis. In:
Pigliucci M, Preston K (eds.) Phenotypic integration: Studying the ecology and evolution of complex
phenotypes. Oxford University Press, Oxford, pp. 130-150.
Harte J, Ostling A, Green JL, Kinzig A (2004) Biodiversity conservation — Climate change and
extinction risk. Nature 430 DOT: 10.1038 /nature02718.
Heikkinen RK, Luoto M, Araujo MB, Virkkala R, Thuiller W, Sykes MT (2006) Methods and
uncertainties in bioclimatic envelope modeling under climate change. Progress in Physical
Geography 30: 751-777.
Heikkinen RK, Luoto M, Virkkala R, Pearson RG, Korber JH (2007) Biotic interactions improve pre-
diction of boreal bird distributions at macro-scales. Ghbal Ecohgy ¢~ Biogeography 16: 754-763.
Henry P-Y, Lengyel 8, Nowickt P, Julliard R, Clobert J, Celik ‘IT, Gruber B, Schmeller DS, Babi
V, Henle K (in press) Integrating biodiversity monitoring initiatives: potential benefits
and methods. Bzodiversity and Conservation.
Hernandez PA, Graham CH, Master LL, Albert DL (2006) The effect of sample size and
species characteristics on performance of different species distribution modeling
methods. Ecography 29: 773-785.
Hickler ‘T, Vohland K, Costa L, Cramer W, Miller PA, Smith B, Feehan J, Kuhn I, Sykes M (in
press) Vegetation on the move — where do conservation strategies have to be re-defined?
References 701
In: Settele J, Penev L, Georgiev T, Grabaum R, Grobelnik V, Hammen V, Klotz 8, Kuhn
I (eds.) Atlas of Biodiversity Risk. Pensoft, Sofia.
Hickling R, Roy DB, Hill JK, Fox R, Thomas CD (2006) The distributions of a wide range of
taxonomic groups are expanding polewards. Global Change Biology 12: 450-455.
Higgins LG (1976) The classification of European butterflies. Collins, London (1975).
Hill JK, Thomas CD, Lewis OT (1999) Flight morphology in fragmented populations of a rare
British butterfly, Hesperia comma. Biological Conservatoin 87: 277-284.
Hill JK, Collingham YC, Thomas CD, Blakeley DS, Fox R, Moss D, Huntley B (2001) Impacts
of landscape structure on butterfly range expansion. Evohgy Letters 4: 313-321.
Holt RD (1990) The microevolutionary consequences of climate change. Trends in Ecology and
Evolution 5: 311-315.
Holt RD (1996) Demographic constraints in evolution: Towards unifying the evolutionary
theories of senescence and niche conservatism. Evolutionary Ecology 10: 1-11.
Holt RD (2008) IJEE Soapbox: Charismatic mesofauna: Butterflies as inspiration and test for
theory that integrates ecology and evolution. [srae/ Journal of Ecology and Evolution 54: 1-5.
Holt RD, Gaines M (1992) The analysis of adaptation in heterogeneous landscapes: Implications
for the evolution of fundamental niches. Evo/tionary Ecolgy 6: 433-447.
Honey MR, Scoble MJ (2001) Linnaeus’s butterflies. Zoological Journal of the Linnean Society 132:
277-399.
Huntley B, Green RE, Collingham YC, Hill JI<, Willis SG, Bartlein PJ, Cramer W, Hagemetyer
WJM, Thomas CJ (2004) The performance of models relating species geographical
distributions to climate is independent of trophic level. Ecology Letters 7: 417-420.
Ibanez I, Clark JS, Dietze MC, Feeley K, Hersh M, LaDeau 8, McBride A, Welch NE, Wolosin
MS (2006) Predicting biodiversity change: Outside the climate envelope, beyond the
species-area curve. Evolgy 87: 1896-1906.
IPCC (Intergovernmental Panel on Climate Change; 2001) Clamate Change 2001: The Scientific
Basis. Contribution of Working Group I to the Third Assessment Report of the Inter-
governmental Panel on Climate Change. Cambridge University Press, Cambridge, UK.
Johst K, Drechsler M, Thomas J, Settele J (2006) Influence of mowing on the persistence of
two endangered large blue butterfly species. Journal of Applhed Ecology 43: 333-342.
Kadmon R, Farber O, Danin A (2003) A systematic analysis of factors affecting the performance
of climatic envelope models. Ecological Applications 13: 853-867.
Kawecki TJ] (1995) Demography of source-sink populations and the evolution of ecological
niches. Evolutionary Ecology 9: 38-44.
Klots AB (1933) A generic classification of the Pieridae together with a study of the male
genitalia. Entomologica Americana 12: 139-242.
Kudrna O (1977) A revision of the genus Hipparchia Fabricius, 1507. Classey, Faringdon.
Kudrna O (1984) On the taxonomy of the genus Hipparclia Fabrictus, 1807, with descriptions
of two new species from Italy. Fragmenta Entomologica 17: 229-243.
Kudrna O (2002) The distribution atlas of Euroopean butterflies. Oedippus 20: 1-342.
Kudrna O, Belicek J (2005) On the ‘Wiener Verzeichnis’, its authorship and the butterflies
named therein. Oedippus 23: 1-32.
Kuhn E, Feldmann R, Harpke A, Hirneisen N, Musche M, Leopold P, Settele, J (2008a) Getting
the public involved into butterfly conservation — Lessons learned from a new monitoring
scheme 1n Germany. [srae/ Journal of Ecology and Evolution 54: 89-103.
Kuhn I, Sykes MT, Berry PM, Thuiller W, Piper JM, Nigmann U, Aratjo MB, Balletto E,
Bonelli S$, Cabeza M, Guisan A, Hickler T, Klotz $, Metzger M, Midgley G, Musche M,
702 Climatic Risk Atlas of European Butterflies
Olofsson J, Paterson JS, Penev L, Rickebusch S, Rounsevell MDAR, Schweiger O, Wilson
FE, Settele J (2008b). MACIS: Minimisation of and Adaptation to Climate change Impacts
on biodiverSity. GAIA — Ecological Perspectives for Science and Society 17/4: 393-395.
Kuhn I, B6hnine-Gaese K, Cramer W, Klotz S (2008c). Macroecology meets global change
reseatch. Global Ecology ¢~ Biogeography 17: 3-4.
Kuhn I, Vohland K, Badeck F, Hanspach J, Pompe S, Klotz S (in press) Aktuelle Ansatze zur
Modellierung der Auswirkungen von Klimaveranderungen auf die biologische Vielfalt.
Natur ¢ Landschaft 84(1).
Lobo JM, Jimenez-Valverde A, Real R (2008) AUC: a misleading measure of the performance
of predictive distribution models. Global Ecology ¢» Biogeography 17: 145-151.
Luoto M, Heikkinen RK (2008) Disregarding topographical heterogeneity biases species
turnover assessments based on bioclimatic models. Géobal Change Biology 14: 483-494.
Luoto M, Virkkala R, Heikkinen RK (2007) The role of land cover in bioclimatic models
depends on spatial resolution. G/obal Ecology ¢~ Biogeography 16: 34-42.
Manel S, Williams HC, Ormerod SJ (2001) Evaluating presence-absence models in ecology: the
need to account for prevalence. Journal of Apphed Ecology 38: 921-931.
McPherson JM, Jetz W (2007) Effects of species’ ecology on the accuracy of distribution
models. Ecography 30: 135-151.
Miller LD (1968) The higher classification, phylogeny and zoogeography of the Satyridae. Mem.
Am. Ent. Soc. 24: 1-174.
Mitchell TD, Carter TR, Jones PD, Hulme M, New M (2004) A comprehensive set of high-resolution
grids of monthly chmate for Europe and the globe: the observed record (1901-2000) and 16 scenarios
(2001-2100). Tyndall Centre for for Climate Change Research, University of East Anglia,
Norwich, UK.
Mouguet N, Belrose V, Thomas JA, Elmes GW, Clarke RT, Hochberg ME (2005) Conserving
community modules: a case study of the endangered lycaenid butterfly Maculinea akon.
Ecolgy 86: 3160-3173.
New M, Hulme M, Jones J (2000) Representing twentieth-century space-time climate variability.
Part I: Development of 1901-96 monthly erids of terrestrial surface climate. Journal of
Chmate 13: 2217-2238.
Ohlemuller R, Grittt ES, Sykes MT, Thomas CD (2006) Quantifying components of risk for
European woody species under climate change. Global Change Biology 12: 1788-1799.
Office of International Epizootics (OIE) (2000): International Animal Health Code.
Parmesan C (1996) Climate and species’ range. Nature 382: 765-766.
Parmesan C, Yohe G (2003) A globally coherent fingerprint of climate change impacts across
natural systems. Nature 421: 37-42.
Parmesan C, Ryrholm N, Stefanescu C, Hill JK, Thomas CD, Descimon H, Huntley B, Kaila
L, Kullberg J, Tammaru T, Tennent WJ, Thomas JA, Warren MS (1999) Poleward shifts
in geographical ranges of butterfly species associated with regional warming. Na/ure 399:
579-583.
Pearson RG, Dawson TP (2003) Predicting the impacts of climate change on the distribution of
species: ate bioclimate envelope models useful? Gébal Evolgy and Biogeography 12: 361-371.
Pearson RG, Thuiller W, Araujo MB, Martinez-Meyer E, Brotons L, McClean C, Miles L,
Segurado P, Dawson TP, Lees DC (2006) Model-based uncertainty in species range
prediction. Journal of Biogeography 33: 1704-1711.
Pease CM, Lande R, Bull JJ (1989) A model of population-growth, dispersal and evolution in
a changing environment. Ecology 70: 1657-1664.
References 703
PechP, Fric Z, Konvicka M, Zrzavy J (2004) Phylogeny of Maculineablues based on morphological
and ecological characters: evolution of parasitic myrmecophily. Cladistics 20: 362-375.
Pena C, Wahlberg N, Weingartner E, Kodandaramaiah U, Nylin S, Freitas AVL, Brower VZ
(2006) Higher level phylogeny of Satyrinae butterflies based on DNA sequence data.
Mokecular Phylogenetics and Evolutin 40: 29-49.
Pollard E (1977) A method for assessing changes in the abundance of butterflies. Bzo/ogial
Conservation 12: 115-134.
Pollard E (1988) Temperature, rainfall and butterfly numbers. Journal of Apphed Ecology 25:
819-828.
Pollard E, Yates TJ (1993) Monitoring butterflies for ecology and conservation: the British butterfly monitoring
scheme, 1st edn. London, Chapman, Hall.
Pompe 8, Hanspach J, Badeck F, Thuiller W, Kthn I (2008) Climate and land use change
impacts on plant distributions in Germany. Bzology Letters 4: 564-567.
Porter AH, Schneider RW, Price BA (1995) Wing pattern and allozyme relationships in the
Coenonympha arcania group, emphasising the C. gardetta-darwiniana contact area at Bellwald,
Switzerland. Nota kpidopterologica 17: 155-174.
Porter AH, Wenger R, Geiger H, Scholl A, Shapiro AM (1997) The Pontia daplidice-edusa hybrid
zone in Northwestern Italy. Evolution 51: 1561-1573.
Pulliam HR (2000) On the relationship between niche and distribution. Evolggy Letters 3: 349-361.
Randin CF, Dirnbéck 'T, Dullinger S$, Zimmermann NE, Zappa M, Guisan A (2006) Are niche-
based species distribution models transferable 1n space? Journal of Biogeography 33: 1689-1703.
Reznick DN, Ghalambor CK (2001) The population ecology of contemporary adaptations:
what empirical studies reveal about the conditions that promote adaptive evolution.
Genetica 112-113: 183-198.
Rickebusch S, Thuiller W, Hickler T, Araujo MB, Sykes M, Schweiger O, Lafourcade B (2008)
Incorporating the effects of changes in vegetation functioning and CO2 on water
availability in plant habitat models. Brology Letters 4: 556-559.
Roy DB, Rothery P, Moss D, Pollard E, Thomas JA (2001) Butterfly numbers and weather:
predicting historical trends in abundance and the future effects of climate change. Journal
of Animal Excology 70: 201-217.
Sakamoto Y, Ishiguro M, Kitagawa G (1986) Akaike information criterion statistics. D. Reidel
Publishing Company, Dordrecht, Tokyo.
Sala OF, Stuart Chapin HT F, Armesto JJ, Berlow E, Bloomfield J, Davis F, Dirzo R, Froydis I,
Huber-Sanwald E, Huenneke LF, Jackson R, Kinzig A, Leemans R, Lodge D, Malcolm
J, Mooney HA, Oesterheld M, Poff L, Sykes MT, Walker BH, Walker M, Wall D (2000)
Global biodiversity scenarios for the year 2100. Sczence 287: 1770-1774.
Schweiger O, Settele J, Kudrna O, Klotz S, Kiihn I (in press) Climate change can cause spatial
mismatch of trophically interacting species. Evol/ogy.
Segurado P, Araujo MB (2004) An evaluation of methods for modelling species distributions.
Journal of Biogeography 31: 1555-1568.
Seoane J, Carrascal LM, Alonso CL, Palomino D (2005) Species-specific traits associated to
prediction errors in bird habitat suitability modelling. Evolggecal Modelling 185: 299-308.
Settele J, Hammen V, Hulme PE, Karlson U, Klotz S, Kotarac M, Kunin, Marion G, O’Connor
M, Petanidou T, Peterseon K, Potts S, Pritchard H, Pysek P, Rounsevell M, Spangenberg
J, Steffan-Dewenter I, Sykes MT, Vight M, Zobel M, Kuhn I (2005) ALARM: Assesing
LArge scale environmental Risks for biodiversity with tested Methods. GALA — Ecological
Perspectives in Science, Humanities, and Economics 14: 96-72.
704 Climatic Risk Atlas of European Butterflies
Settele J, Kuhn I, Klotz $, Hammen V, Spangenberg J (2007) Is the EC afraid of its own
visions? Sczence 315: 1220.
Settele J, Spangenberg J, Kiihn I (2008). Large projects can create useful partnerships. Na/ure
453: 850.
Shirozu T, Yamamoto H (1956) A generic revision and phylogeny of the tribe Theclint. Szeboldia
1: 330-421.
Simonsen TJ (2006) Fritillary phylogeny, classification and larval hostplant: reconstructed
mainly on the basis of male and female genitalic morphology. Bzological Journal of the
Linnean Society 89: 627-673.
Singer MC, Thomas CD, Parmesan C (1993) Rapid human-induced evolution of insect host
associations. Nature 366: 681-683.
Smith B, Prentice IC, Sykes MT (2001) Representation of vegetation dynamics in modelling
of terrestrial ecosystems: comparison of two contrasting approaches within European
climate space. Global Ecology and Biogeography 10: 621-638.
Spangenbere JH (2007) Integrated scenarios for assessing biodiversity risks. Swstainabk
Development 15: 343-356.
Sykes MT, Prentice IC, Cramer W (1996) A bioclimatic model for the potential distribution of
northern European tree species under present and future climates. Journal of Biogeography
23: 203-233.
Thomas CD (1994) Extinction, colonization, and metapopulations: Environmental tracking by
rare species. Conservation Biology 8: 373-378.
Thomas JA (2005) Monitoring change in the abundance and distribution of insects using butterflies
and other indicator groups. Philosophical Transactions of the Royal Society B 360: 339-57.
Thomas JA, Clarke RT (2004) Extinction rates and butterflies — Response. Svzenve 305: 1563-1564.
Thomas JA, Morris MG (1994) Patterns, mechanisms and rates of extinction among
invertebrates in the United Kingdom. Philosophical Transactions of the Royal Society of London
Serves B-Biological Sciences 344: 47-54.
Thomas CD, Bodsworth EJ, Wilson RJ, Simmons AD, Davies ZG, Musche M, Conradt L (2001)
Ecological and evolutionary processes at expanding range margins. Nature 411: 577-581.
Thomas JA, Telfer MG, Roy DB, Preston CD, Greenwood JJD, Asher J, Fox R, Clarke RT,
Lawton JH (2004) Comparative losses of British butterflies, birds, and plants and the
global extinction crisis. Sczence 303: 1879-1881.
Thuiller W, Araujo MB, Lavorel S (2003) Generalized models versus classification tree analysis:
a comparative study for predicting spatial distributions of plant species at different scales.
Journal of Vegetation Science 14: 669-680.
Thuiller W, Araujo MB, Lavorel S (2004) Do we need land-cover data to model species
distributions in Europe? Journal of Biogeography 31: 353-361.
Thuiller W, Lavorel S, Araujo MB (2005) Niche properties and geographical extent as predictors
of species sensitivity to climate change. Ghbal Ecology and Biogeography 14: 347-357.
Thuiller W, Albert C, Araujo MB, Berry PM, Cabeza M, Guisan A, Hickler T, Midgley GF,
Paterson J, Schurr FM, Sykes MT, Zimmermann NE (2008) Predicting global change
impacts on plant species’ distributions: Future challenges. Perspectives in Plant Ecology
Evolution and Systematics 9: 137-152.
Tolman T, Lewington R (2008) Coddns Butterfly Guide. HarperCollins. London.
Trivedi MR, Berry PM, Morecroft MD, Dawson TP (2008) Spatial scale affects bioclimate
model projections of climate change impacts on mountain plants. Global Change Biology
14: 1089-1103.
References 705
van Nouhuys S, Hanski I (2004) Natural enemies of checkerspot butterflies. In: Ehrlich PR,
Hanski I (eds.) Ox the Wings of Checkerspots: A Model system for Population Biology. Oxford
University Press, Oxford.
Van Strien AJ, van de Pevert R, Moss D, Yates ‘TJ, Van Swaay C, Vos P (1997) The statistical power
of two butterfly monitoring schemes to detect trends. Journal of Apphed Ecology 34: 817-828.
Van Swaay CAM, Van Strien AJ (2005) Using butterfly monitoring data to develop a European
grassland butterfly indicator. In: Kuhn E, Feldmann R, Thomas JA, Settele J (eds.) Stadies
on the Ecology and Conservation of Butterflies in Europe. Vol. 1, General Concepts and Case
studies. Pensoft, Sofia-Moscow.
Van Swaay CAM, Warren MS (1999) Red Data book of European butterflies (Rhopalocera) Nature and
the Exnvironment, No. 99. Council of Europe Publishing, Strasbourg.
Van Swaay CAM, Warren MS, Lois G (2006) Biotope use and trends of European butterflies.
Journal of Insect Conservation 10: 189-209.
Van Swaay CAM, Nowickt P, Settele J, Van Strien AJ (in press). Butterfly Monitoring in Europe —
methods, applications and perspectives. Bzodiversity and Conservation.
Vodolazhsky DI, Stradomsyk BV (2008a) Phylogenetic analysis of subgenus Po/ommatus (s. str)
Latreille, 1804 (Lepidoptera: Lycaenidae) based on mtDNA markers. Part I. Caucasian
Entomological Bulletin 4(1): 123-130.
Vodolazhsky DI, Stradomsyk BV (2008b) Phylogenetic analysis of subgenus Po/yommatus (s. str)
Latreille, 1804 (Lepidoptera: Lycaenidae) based on mtDNA markers. Part II. Caucasian
Entomological Bulletin 4(2): 237-242.
Wahlberg N, Nylin S (2003) Morphology versus molecules: resolution of the positions of
Nymphals, Polygonia and related genera. Cladistics 19: 213-223.
Walther GR, Post E, Convey P, Menzel A, Parmesan C, Beebee TJC, Fromentin JM, Hoegh-Guldberg
H, Batrlein F (2002) Ecological responses to recent climate change. Nature 416: 389-395.
Walther GR, Burga CA, Edwards PJ (eds.) (2001) “Fingerprints “of Chmate Change — Adapted behaviour
and shifting species ranges. Kluwer Academic/Plenum Publishers, New York / London. 329 pp.
Warren MS, Hill Jk, Thomas JA, Asher J, Fox R, Huntley B, Roy DB, Telfer MG, Jeffcoate S,
Harding P, Jeffcoate G, Willis SG, Greatorex-Davies JN, Moss D, Thomas CD (2001)
Rapid responses of British butterflies to opposing forces of climate and habitat change.
Nature 414: 65-69.
Wiemers M (1998) Coenonympha darniniana. A hybrid taxon? New insights through allozyme
electrophoresis. Mem. Soc. R. Belg. Ent. 38: 41-70.
Wiemers M (2003) Chromosome differentiation and the radiation of the butterfly subgenus Agrodiaetus
(Lepidoptera: Lycaenidae: Polyommatus) — a mokcular phylogenetic approach, Dissertation.
Universitat Bonn.
Wiemers M (2007) Die Gattung Coexonympha Hubner, 1819, in Europa: Systematik, Okologie
und Schutz. Oedippus 25: 1-42.
Wiens JJ, Graham CH (2005) Niche conservatism: integrating evolution, ecology, and
conservation biology. Annual Review of Ecology, Evolution, and Systematics 36: 519-539.
Wilson RJ, Gutiérrez D, Gutierrez J, Martinez D, Agudo R, Monserrat VJ (2005) Changes to
the elevation limits and extent of species ranges associated with climate change. Evohgy
Letters 8: 1138-1146.
Climatic Risk Atlas of European Butterflies
abencerragus 254£
acaciae 228
achine 474£
actaca 582£
acteon 94£
adippe 350
admetus 332£
Admiral 388£, 446ff
Admiral, Poplar 446f
Admiral, Red 388f
Admiral, Southern White 450f
Admiral, White 448f
aegeria 466
aetherte 426£
aethiops S24
agestis 296
Aglais 392 ££
aglaja 348
alberganus 584£
albicans 330£
alveae 36£
alcetas 246£
akiphron 198
alcon 274£
alexanor 120£
alexis 262£
alfacariensis 178£
alveus 18
amandus 314£
andromedae 60£
anteros 802£
anthelea 610£
Anthocharis 128ff
antiopa 400£
Apatura 458 ff
Aphantopus 502£
Apollo 110ff
apollo \14€
Apollo, Clouded 110f
Apollo, Small 112f
Abporia 142£
aquilo 2B6£
aquilonaris 384£
Species index
(of modelled butterflies)
Araschnia 408£
arcania 482£
arethusa 604£
Arethusana 604£
arge STA£
argiades 242
arsiolus 248
argus 216
Argus 292ff
Argus, Blue 802f
Argus, Brown 296f
Argus, Geranium 292f
Argus, Northern Brown 298f
Argus, Scotch 524f
Argus, Silvery 304f
Argus, Southern Mountain 300f
Arrgynnis S44 f£
argyrognomon 280£
Arica 292 ff
arion 268£
armoricanus 16£
artaxerxes 298f
atalanta 388£
athalia 444£
aurelia 440 £
aurinia 420£
ausonia 138£
avis 2166
baeticus 44£
ballus 212£
bathseba 500£
baton 250£
bavius 256£
belemia 136£
bellargus 824£
bellieri BOE
betulae 206£
Blue 230£, 234 £6, 306ff
Blue, Adonis 324£
Blue, African Grass 236f
Blue, Alcon 274£
Blue, Alpine 288f
Blue, Amanda’s 314f
Blue, Anomalous 332f
Blue, Arctic 286f
Blue, Baton 250f
Blue, Bavius 256f
Blue, Black-eyed 264f
Blue, Chalkhill 326£
Blue, Chapman’s 316f
Blue, Chequered 258f
Blue, Common 318f
Blue, Cranberry 282f
Blue, Damon 338f
Blue, Dusky Large 272f
Blue, Eastern Short-tatled 244£
Blue, Eros 320f
Blue, Escher’s 308f
Blue, False Baton 254£
Blue, Furry 336f
Blue, Glandon 284f
Blue, Green-underside 262f
Blue, Holly 248f
Blue, Idas 278f
Blue, Iolas 266f
Blue, Lang’s Short-tailed 234f
Blue, Large 268f
Blue, Long-tailed 230f
Blue, Mazarine 306f
Blue, Meleager’s 322f
Blue, Mother-of-peatrl 312f
Blue, Osiris 240f
Blue, Panoptes 260f
Blue, Provencal Short-
tailed 246f
Blue, Provence Chalkhill 328f
Blue, Reverdin’s 280f
Blue, Ripart’s Anomalous 334f
Blue, Scarce Large 270f
Blue, Short-tailed 242f
Blue, Silver-studded 276£
Blue, Small 238f
Blue, Spanish Chalkhill 330f
Blue, Turquoise 310f
boeticus 230£
Boloria 364 ££
Species index 707
brassicae 144£ centaureae B4£ dorus 488£
Brenthis 358££ cerisyi 108 dorylas 310£
Brimstone 180ff Charaxes 456£ Dryad 584f
Brimstone, Powdered 182f chariclea 372£ dryas 584£
Brintesia 806£ Chazara 608f duponcheli 124
briseis 808f chrysippus 618 edusa 160£
britomartis 442£ chrysotheme 174 egea 398F
Brown 464£, 468f£, 504 €f, cinxia 422 embla 528£
S40ff, SSOfF circe 606£ Emperor 458ff
Brown, Arran 510f cirstt T4£ Emperor, Freyer’s Purple 458f
Brown, Autumn 556f cleopatra 184£ Emperor, Lesser Purple 460f£
Brown, Black 558f Cleopatra 184f Emperor, Purple 462f
Brown, Bright-eyed 560f Coenonympha 476f£ epiphron 518
Brown, Dewy 564f Colias 162£f epistygne S42£
Brown, Dusky Meadow 506f = Comma, False 406f erate 166£
Brown, Large Wall 472f Comma 396ff, 406f Erebia 10££
Brown, Lattice 464£ comma 96£ ergane 192£
Brown, Marbled 554f Comma, Southern 398f eriphyle S14£
Brown, Meadow 504f Copper 186ff eros 820f
Brown, Mnestra’s 540f Copper, Balkan 202f Erynnis 32f£
Brown, Northern Wall 470f Copper, Grecian 194f escheri 308£
Brown, Oriental Meadow 508f Copper, Large 190f esculi 226£
Brown, Ottoman Brassy 544f = Copper, Lesser Fiery 204f Euchloe 136f£
Brown, Piedmont 562f Copper, Purple-edged 200f eumedon 292£
Brown, Spring 542f Copper, Purple-shot 198f ennomia 304£
Brown, Stygian 552f Copper, Scarce 192f eupheme 134£
Brown, Swiss Brassy 546f Copper, Small 186f euphenoides 130£
Brown, Wall 468f Copper, Sooty 196f euphrosyne 366£
Brown, Water 550f£ Copper, Violet 188f Euplhydryas 410
Brown, Woodland 474f coridon 326£ euryale S12£
bryoniae 156£ cramera 294£ asi S86£
cacaliae 82£ crataegi 142£ Jarinosa 182£
Cacyreus 232 croceus 168f fatua 598
c-album 896f Cupido 238 ff Favonius 208£
callidice 158£ Cyaniris 806£ ferula 580£
Callophrys 214 ff cynthia 412£ Festoon 104ff
Camberwell Beauty 400f damon 338£ Festoon, Eastern 108f
camilla 448£ Danaus 618£ Festoon, Southern 106f
candens 202£ daphne 360£ Festoon, Spanish 104f
Carcharodus 36f£ daphnis 322£ Jidia 600£
cardamines 128 daplidice 160£ flocciferus 40£
Cardinal 346f decoloratus 244£ freija 8T4E
cardui 390£ deione 434£ Jrigga 380£
carlinae T2f desfontainii 418f Fritillary 340f, 344f, 34866 410ff
Carterocephalus 86 ££ dia 376£ Fritillary, Aetherie 426f
carthami S6£ diamina 432£ Fritillary, Arctic 372f
cassioides S48 didyma 430£ Fritillary, Asian 414f
cecilia A98£ disa 580£ Fritillary, Assmann’s 442f
Celastrina 248 dispar 190 Fritillary, Balkan 386f
celtis B42 dolus 336£ Fritillary, Bog 364f
Climatic Risk Atlas of European Butterflies
Fritillary, Cranberry 384f
Fritillary, Cynthia’s 412f
Fritillary, Dark Green 348f
Fritillary, Duke of
Burgundy 340f
Fritillary, False Heath 482f
Fritillary, Frejya’s 372f
Fritillary, Frigga’s 380
Fritillary, Glanville 422f
Fritillary, Grisons 486f
Fritillary, Heath 444f
Fritillary, High Brown 350f
Fritillary, Knapweed 424f
Fritillary, Lapland 410f
Fritillary, Lesser Marbled 358f
Fritillary, Lesser Spotted 428f
Fritillary, Marbled 360f
Fritillary, Marsh 420f
Fritillary, Meadow 438f
Fritillary, Nickerl’s 440f
Fritillary, Niobe 352f
Fritillary, Pallas’ 354f
Fritillary, Pearl-bordered 366f
Fritillary, Provencal 434f
Fritillary, Queen of Spain 356f
Fritillary, Scarce 416f
Fritillary, Shepherd’s 382f
Fritillary, Silver-washed 344f
Fritillary, Small Pearl-
bordered 370f
Fritillary, Spanish 418f
Fritillary, Spotted 480f
Fritillary, Thor’s 378f
Fritillary, Titania’s 368f
Fritillary, Twin-spot 362f
Fritillary, Weaver’s 876f
galathea 568£
gardetta 486£
Gatekeeper 496ff
Gatekeeper, Southern 498f
Gatekeeper, Spanish 500f
Gegenes 100
Geranium Bronze 232f
glacials 614£
glandon 284£
Glaucopsyche 262f£
Glider 452ff
Glider, Common 452£
Glider, Hungarian 454f
ghycerion AB4£
Gonepteryx 180ff
gorge 538F
gracca 386
Grayling 586ff, 610ff
Grayling, Alpine 614f
Grayling, Balkan 602f
Grayling, Baltic 616f
Grayling, Delattin’s 594f
Grayling, Eastern Rock 590f
Grayling, False 604f
Grayling, Freyer’s 598f
Grayling, Great Banded 606f
Grayling, Norse 612f
Grayling, Rock 588f
Grayling, Striped 600f
Grayling, Tree 596f
Grayling, White-banded 610f
Grayling, Woodland 586f
gruneri 132
Hairstreak 206ff
Hairstreak, Black 220f
Hairstreak, Blue-spot 222f
Hairstreak, Brown 206f
Hairstreak, Chapman’s
Green 216f
Hairstreak, False Ilex 226f
Hairstreak, Green 214f
Hairstreak, Hex 224f
Hairstreak, Provence 212f
Hairstreak, Purple 208f
Hairstreak, Sloe 228f
Hairstreak, Spanish
Purple 210f
Hairstreak, White-letter 218f
Hamearis 340£
Heath 476f, 480ff
Heath, Alpine 486f
Heath, Chestnut 484£
Heath, Dusky 488f
Heath, Eastern Large 480f
Heath, Large 476f
Heath, Pearly 482f
Heath, Russian 492f
Heath, Scarce 490f
Heath, Small 494f
hecate 362
hecla 170
helle 188
hermione 588£
hero 490£
hersamon 204£
Hesperia 96£
Heteropterus 84£
Hipparchia 586f£
hippothoe 200£
hispanus 828
hyale 176£
hyperantus 502£
Hyponephele 506f£
warus 318£
idas 218£
iduna 410£
ila 460£
ilicis QO4£
ines 518£
ino 308£
intermedia 414£
10 3892£
Tolana 266£
zolas 266£
Iphichdes 116£
iris AB2E
Tssoria 856£
jasius A56£
jurtina S04£
jutta 816£
Kirinia 464£
knysna 236
krueperi 146£
lachesis 570£
Laeosopis 210£
L-album 406£
Lampides 230E
laodice 3854£
larissa 57 2£
Lasiommata 468f£
lathonia 356£
lavatherae 388£
leander 492£
Leptidea 122ff
Leptotes 234£
levana 408£
Libythea 342£
ligea 910£
Limenitis 446 ££
Lineola 9O£
Lopinga 474£
/ucina 840£
/upina 508
Lycaena 186f£
Jycaon S06E
machaon 118£
maera 472£
malvae 66£
malvoides 66£
Maniola 504£
mannit 148£
manto 516£
Map 408f
marloyi 34£
marshall 2326
maturna 416£
medusa D82f
megera 468
melampus S22
Melanargia 566 ££
melanops 264£
melas 558£
Mebhtaea 422 ff
meolans 562£
metis 458£
minimus 238f
Minois 584£
mnemosyne 110£
muestra DA0£
montana D54£
montensis 800£
morpheus B4£
morset 126£
myrmidone 172£
napi 194
nausithous 21 2f
neoridas 556£
Neptis 45266
Nettle-tree Butterfly 342f
nictas 804£
niobe 352£
nivescens B12£
norna 612£
nostrodamus 102£
Nymphalis 396££
occttanica 51T6£
Ochlodes 98£
oedippus 478£
oeme 560£
Oeneis 612 ££
onopordi 10£
optilete 282
Orange-tip 128f, 132
Orange-tip, Gruner’s 132f
Orange-tip, Sooty 134f
orbifer 5O£
orbitulus 2B8£
orientalis 42£
orton 258f
osiris QAOE
ottomana 194£
ottomana D44£
Painted Lady 390f
palaemon 86£
palaeno 164£
pales 382£
pamphilus 494£
pandora 346£
pandrose 564£
panoptes 260£
paphia 844£
Papilio 118f£
Pararge 466£
Parnassius \10f£
parthenoides 438£
Peacock 892f
petropolitana 470
pharte 520£
Phengaris 268f£
phicomone 162£
phlacas 186£
phlomidis 46£
phoebe 424f
phoebus 112£
Pieris 144 ff
pirithous 234£
Plain ‘Tiger 618f
Pkebejus 276££
pluto 5386£
podalirius 116£
pohchloros 402
Polommatus 308
polyxena 106£
Pontia 158f£
populi 446£
pronoe 550
proto 52
prani 220f
Pseudochazara 610£
Species index 709
pumilio \00E
Pyrgus SOfE
Pyronia 496
guercus 208£
rapae 150£
reali 122£
reducta 450£
rhamni 180£
rhodopensis AB0£
Ringlet 478f, 502f, 5126,
526ff, 548f
Ringlet, Almond-eyed 534f
Ringlet, Arctic 530£
Ringlet, Blind 520f
Ringlet, Common Brassy 548f
Ringlet, de Prunner’s 526f
Ringlet, Eriphyle 514f
Ringlet, False 478f
Ringlet, Lapland 528f
Ringlet, Large 512f
Ringlet, Lesser Mountain 522f
Ringlet, Mountain 518f
Ringlet, Silky 538f
Ringlet, Sooty 536f
Ringlet, Woodland 582f
Ringlet, Yellow-spotted 516f
ripartii 334£
rivularis AB4£
roboris 210
roxelana 464£
rubi Q14£
rumina \0O4£
russiae SO6£
sappho 452£
Satyr 580ff
Satyr, Black 582f
Satyr, Great Sooty 580f
Satyrium 218 ff
Satyrus 580f£
Scolitantides 250££
selene 310£
semele 5925
semiargus 806£
senthes 602
sephirus 290
serratulae 68f£
sertorius 48£
sidae 58£
stlricolus 88£
Climatic Risk Atlas of European Butterflies
sinapis 122f
Skipper 32ff
Skipper, Alpine Grizzled 60f
Skipper, Carline 72f
Skipper, Chequered 86f
Skipper, Dingy 32f
Skipper, Dusky Grizzled 62f
Skipper, Essex 90f
Skipper, Foulquier’s
Grizzled 80f
Skipper, Grizzled 66f
Skipper, Inky 34f
Skipper, Large 98f
Skipper, Large Chequered 84f
Skipper, Large Grizzled 78f
Skipper, Lulworth 94f
Skipper, Mallow 36f
Skipper, Marbled 38f
Skipper, Mediterranean 102f
Skipper, Northern
Chequered 88f
Skipper, Northern
Grizzled 64f
Skipper, Oberthtir’s
Grizzled 76f
Skipper, Olive 68f
Skipper, Orbed Red-
underwing 50f
Skipper, Oriental Marbled 42f
Skipper, Persian 46f
Skipper, Pigmy 100f
Skipper, Red-underwing 48f
Skipper, Rosy Grizzled 70f
Skipper, Safflower 56f
Skipper, Sage 52f
Skipper, Silver-spotted 96f
Skipper, Small 92f
Skipper, Southern Marbled 44f
Skipper, Tessellated 54f
Skipper, Tufted Marbled 40f
Skipper, Warren’s 82f
Skipper, Yellow-banded 58f
Speckled Wood 466f
Spialia 46 ££
spini 222
statilinus 596£
styx 552
Swallowtail 116ff
Swallowtail, Scarce 116f
Swallowtail, Southern 120f
syhanus 98
syhestris 92
syriaca 590E
Syrichtus S2fE
tages 82
tagis 140
teleius 210£
tessellum SAE
The Hermit 608f
Thecla 206£
thersites 316£
thore 378£
Thymelicus QOfE
titania 368£
tithonus 496£
tityrus 196£
Tomares 212£
Tortoiseshell 394f, 402ff
‘Tortoiseshell, Large 402f
Tortoiseshell, Small 394£
Tortoiseshell, Yellow-
legged 404f
triaria 526£
trivia 428£
tullia 476£
Two-tailed Pasha 456f
tyndarus SA6£
urticae 894£
Vanessa 888 ££
varia 436£
vicrama def
virgaureae 192£
voleensis S94F
w-album 218f
warrenensis B2£
White 122ff, 136ff, S66ff
White, Balkan Marbled 572f
White, Bath 160f
White, Black-veined 142f
White, Dappled 138f
White, Eastern Wood 124f
White, Esper’s Marbled 566f
White, Fenton’s Wood 126f
White, Green-striped 136f
White, Green-veined 154f
White, Iberian Marbled 570f
White, Italian Marbled 574£
White, Krueper’s Small 146f
White, Large 144f
White, Marbled 568f
White, Mountain Green-
veined 156f
White, Mountain Small 152f
White, Peak 158f
White, Portuguese
Dappled 140f
White, Small 150f
White, Southern Small 148f
White, Spanish Marbled 578f
White, Western Marbled 576f
White, Wood 122f
xanthomelas 404£
Yellow 162ff
Yellow, Berger’s Clouded 178f
Yellow, Clouded 168f
Yellow, Danube Clouded 172£
Yellow, Eastern Pale
Clouded 166£
Yellow, Lesser Clouded 174£
Yellow, Moorland Clouded 164£
Yellow, Mountain Clouded 162f
Yellow, Northern Clouded 170£
Yellow, Pale Clouded 176f
Zegris 134£
Zerynthia 104 ff
Zizeeria 236