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ORIGINAL PAPER
On the surrogate value of red-listed butterflies for butterflies
and grasshoppers: a case study in Grammos site of Natura
2000, Greece
Konstantina Zografou ÆSpyros Sfenthourakis Æ
Andrew Pullin ÆVassiliki Kati
Received: 19 July 2008 / Accepted: 19 November 2008 / Published online: 9 December 2008
ÓSpringer Science+Business Media B.V. 2008
Abstract We tested the surrogate value of butterflies,
red-listed butterflies and grasshoppers for each other in
terms of diversity patterns congruence and complemen-
tarity at a site in the NATURA 2000 network. Grammos
Mountain is proposed as a new Prime Butterfly Area for
Greece: it supports a total of 56 grasshopper species and
112 butterfly species, 24 of which are of European con-
servation concern (SPEC) and two of Prime Butterflies
Area Project. We found a strong congruence in the species
richness patterns of SPEC butterflies, butterflies and
grasshoppers, because three common ecological factors
influenced them: number of flower heads, altitude, and
cover of low trees or bushes (Redundancy Analysis,
CANOCO). Each complementarity network maintained
quite well the species richness of the other two target
groups (\18% average species loss). SPEC butterflies were
the best surrogate group overall and therefore we propose
that they should be monitored on a permanent basis.
Keywords Complementarity Conservation
Mountain biodiversity Indicators Lepidoptera
Orthoptera Protected area
Introduction
One of the 22 headline indicators to assess progress
towards the 2010 target of halting further biodiversity loss
is the monitoring of selected species (Mace and Baillie
2007). Given the restricted funding, expertise and time
resources to monitor the full spectrum of species diversity,
we need to identify taxa which are well-known, readily
surveyed, and can also reflect the distribution patterns of
other unrelated taxa more difficult to sample or identify
(Gregory 2006; Noss 1990). If such surrogate taxa become
available, conservation practitioners could make good
management decisions and apply monitoring schemes
without needing multi-taxa data. In this vein, several
studies attempted to reveal indicator taxa, in the sense of
cross-taxon surrogacy (e.g., Bilton et al. 2006; Howard
et al. 1998; Jonsson and Jonsell 1999; Kati et al. 2004c;
Lawler and White 2008; Lawton et al. 1998; Prendergast
et al. 1993; Ricketts et al. 2002; Sauberer et al. 2004).
Given that only a small fraction of the global number of
species is known, and that more than half of them are
insects (Thomas 2005), the need to identify robust surro-
gates for insect diversity is even more pronounced.
Butterflies (Lepidoptera, Papilionoidea and Hesperioi-
dea) are among the most popular taxa with well-known
distribution patterns and population trends at least for some
parts of the world, such as Europe (van Swaay and Warren
1999). They are among the most threatened known taxa,
exhibiting declining trends due to severe habitat loss and
climate change (Thomas et al. 2004; Warren et al. 2001).
Because butterflies are among the few invertebrate taxa
that can be monitored cost-effectively at a national or a
European scale (Roy et al. 2007), their surrogate value
towards other target taxa has been often tested and proved
to be either good (e.g., Pearman and Weber 2007) or poor
K. Zografou V. Kati (&)
Department of Environmental & Natural Resources
Management, University of Ioannina, Seferi 2, 30100 Agrinio,
Greece
e-mail: vkati@cc.uoi.gr; kikikati@hotmail.com
S. Sfenthourakis
Section of Animal Biology, Department of Biology,
University of Patras, 26500 Patra, Greece
A. Pullin
School of the Environment and Natural Resources, Bangor,
Gwynedd LL57 2UW, UK
123
J Insect Conserv (2009) 13:505–514
DOI 10.1007/s10841-008-9198-6
(e.g., Gutie
´rrez and Mene
´ndez 2007; Lawton et al. 1998;
Lovell et al. 2007).
On the other hand, grasshoppers (Orthoptera) are a
major component of rural biodiversity (Ryszkowski et al.
1993); they play a central role in food webs, as primary
herbivores and abundant food resource for other taxa.
Some species have been used as indicators of land use
change and disturbance (e.g., Andersen et al. 2001; Baldi
and Kisbenedek 1997; Kati et al. 2006; Kruess and
Tscharntke 2002), but their value as surrogates of other
target taxa is usually poor (e.g., Duelli and Obrist 1998).
Besides, the species-level taxonomy, ecological require-
ments, distribution patterns and vulnerability status of this
group are still poorly known, and high expertise is needed
for their identification (Green 1998). For these reasons,
conservationists consider them only rarely for conservation
management and monitoring schemes (Kati et al. 2004a).
Greece is among the richest countries in Europe in terms
of insect diversity and one of the least studied (Balletto and
Casale 1991; Munguira 1995). In the present study, we
investigated the diversity patterns of both butterflies and
grasshoppers on the Grammos Mountain, a remote area of
NW Greece, which is also a protected site of the European
network NATURA 2000. Apart from general studies cov-
ering the distribution patterns of both groups at a national
scale (Pamperis 1997; Willemse 1984), no research has
ever been conducted in the Grammos area. The main
purposes of the present study are: (a) to explore red-listed
Lepidoptera, Lepidoptera and Orthoptera value as surro-
gates of each other in terms of diversity pattern covariance
and of complementarity and (b) to explore the ecological
factors affecting the relationships found. In doing so, our
study earns also a practical scope, that of exploring the
conservation value of the Grammos site for insect diversity
and interpreting our findings in the context of conservation
management decision for this area.
Methods
Study area and sites
The study area is situated in northwestern Greece (long.
20°500, lat. 40°210). It is a protected site of the European
network NATURA 2000 (GR1320002) and its area covers
350 km
2
. Altitude varies between 600 and 2,520 m, and
climate is of the mountainous type (average annual tem-
perature 8–12°C, average annual rainfall 800–2,200 mm).
The main habitat types encountered in Grammos Mt are
broadleaved forests (41%), grasslands (30%) conifer for-
ests (21%), rocky slopes (5%) and rural mosaics (3%)
(Fig. 1). It is a sparsely populated area, where logging,
periodical livestock grazing and small-scale cultivations
constitute the main, low impact, human activities. We
selected 22 sites, including nine forested, seven grassland
and six agricultural sites, in order to represent the main
vegetation types of the study area (Table 1; Fig. 1).
Sampling
We sampled Lepidoptera in 22 predefined plots of
100 m 9100 m, one per site. We covered each plot with
systematic transects, recording all individuals for a stan-
dard time period of 60 min (time for specimen
identification excluded). We visited each plot three times in
2007 at 25 day intervals: May, June and July (66 plots in
total). We recorded the presence of Orthoptera species in 5
smaller quadrats of 10 m 910 m, randomly located within
each of the above plots, at a distance of at least 10 m away
from each other (110 quadrats in total). Sampling was
conducted once (August) at the peak of Orthoptera adult
activity. We sampled the above groups using a hand held
net and only between temperatures of 19 and 27°Cto
standardize sampling efficiency. We recorded 15 environ-
mental parameters in the 110 quadrats in each sampling
period (440 quadrats in total) (Table 1). We estimated the
cover of trees (T
1
,T
2
) and bushes (Sh
1
,Sh
2
) as the vertical
projection of their crown area (%), the percentage cover of
herbs (He), rocky substrate (R
1
,R
2
) and bare ground (B).
We also recorded the altitude (Alt), the slope (Sl), the
average height of herbs (Hh) by taking four measures (cm)
in each quadrat, the number of flower heads (F) and soil
humidity (using Hobo U12 data logger). Finally, we esti-
mated grazing intensity (G) (minimal: occasional livestock
passage, low: periodical grazing) and human disturbance
(D) in a zone of 500 m (minimal: dirt road presence, low:
asphalted road and human settlements).
Data analysis
We estimated the overall number of species, using the non-
parametric species estimators (1,000 permutations) that
consider the presence of rare species recorded by 1–2
individuals (Chao 1), or present in 1–2 quadrats (Chao 2)
for Lepidoptera and Orthoptera, respectively (Colwell
2006; Magurran 2004). The diversity of Lepidoptera was
calculated in terms of species richness (S) and the Shannon
index (H), considering the total number of individuals
recorded per plot (N1) in all three seasons. We calculated
the Orthoptera species richness (S) and the overall fre-
quency of species presence across the five quadrats per plot
(N2). We tested the surrogate value of Lepidoptera species
of Conservation Concern (SPECs), of all Lepidoptera
species and of Orthoptera by comparing the congruence of
their species richness patterns using Spearman rank cor-
relation coefficient. Considering one group as the surrogate
506 J Insect Conserv (2009) 13:505–514
123
group and the other two as the target groups, we formed the
complementary network after the surrogate group, starting
by the richest-in-species site and adding those sites con-
tributing more new species, until all species were included
in the network. For the SPEC network, we followed the
same procedure, starting with the sites including more
SPEC 2, 3 and 4 species successively. We then calculated
the species proportion of every group included in each step
of the networking procedure and we calculated their
average difference. This was a measure of efficiency of the
surrogate complementary network vis-a
`-vis the target
groups, without the bias of the area selected (different
number of sites forming the network).
We calculated the per plot average values (5 quadrats)
of the environmental parameter for Lepidoptera (mean
values of the 3 seasons) and for Orthoptera. In order to
explore the environmental factors shaping the community
composition, we conducted a Redundancy Analysis
(RDA) using CANOCO software (ter Braak and Smil-
auer 2002). The RDA method extracts the major
gradients in the data that are accounted for by the
measured environmental parameters. The position of a
species in the resulting diagram indicates the degree of
dependence by the closest environmental parameters
(arrows). The diagram shows only the species of con-
servation concern together with the species sufficiently
influenced by the parameters (fit [25%), and only the
significant (P\0.05) environmental variables that did
not suffer from collinearity (1,000 iterations of Monte–
Carlo test).
Fig. 1 Main habitat types and
sampling sites in the study area
(Grammos Mt, Greece)
J Insect Conserv (2009) 13:505–514 507
123
Results
Species richness
We recorded a total of 112 Lepidoptera species (2,115
individuals), 24 of which were of European conservation
concern (SPEC), and 3 more species were protected under
Bern Convention (Appendix 1). Nine of the SPEC species
had a threatened status in Europe with their global popu-
lation concentrated in Europe (SPEC 2) or not (SPEC 3) and
fifteen species had their global population concentrated
mainly in Europe (SPEC 4) (Appendix 1). Two of the SPEC
species sampled (Parnassius apollo and Euphydryas auri-
nia) were also target-species of Prime Butterfly Areas
project (PBAs) (van Swaay and Warren 1999). We sampled
56 Orthoptera species (Appendix 2), including Pholidop-
tera fallax, whose distribution is known to be limited in
eastern Greece (Willemse 1984). The estimated species
number was 118 for Lepidoptera (Chao 1) and 56.7 for
Orthoptera (Chao 2), proving our sampling to be exhaustive
for both groups (95 and 99% species sampled, respectively).
Diversity patterns
A mixed thermophilous forest (F3a) was one of the most
species-rich sites for both Lepidoptera and Orthoptera
(Table 2). It was a low semi-open forest including a stream,
a well-developed shrub undergrowth and herb layer. Simi-
larly, an abandoned rural mosaic (A4a), combining fields
with hedges and thermophilous wood plots supported great
species richness for both groups. Sub-alpine grasslands at
lower (1,300 m) and higher (1,500–1,800 m) altitudes were
also species-rich habitats for Lepidoptera (G3c) and
Orthoptera (G2b, G3a), respectively. On the other hand,
Table 1 Site description on the basis of the average values of the environmental variables recorded in the five quadrates (10 m 910 m)
Site Habitat code Habitat description Topography Vegetation Ground Impact
Alt (m) Sl T
1
T
2
Sh
1
Sh
2
He Hh
(cm)
FR
1
R
2
BHu
(%)
GD
F1a 9,130 Beech forests of Fagus spp. 1,751 2 5 0 0 0 3 2 4 1 1 2 51 ??
F1b 1,839 2 5 0 0 1 3 2 3 1 1 2 44 ?
F2a (924A) Oak forests of Quercus spp. 946 1 3 2 1 1 4 3 2 0 0 3 51 ??
F2b 1,085 2 3 2 2 2 3 3 3 0 0 1 58 ??
F3a (925A) Mixed thermophilous forests 1,100 2 0 3 2 0 3 3 6 2 1 1 52 ?
F3b 932 2 0 4 3 1 2 2 1 0 0 3 40 ?
F4 3,220 Riverine vegetation 690 1 0 2 1 1 1 2 3 3 0 2 48 ???
F5 9,270 Fir forest of A. borisii-regis 1,329 1 5 0 2 1 2 2 3 2 0 2 49
F6 9,530* Pine forest of P. nigra 1,118 1 4 2 0 1 4 3 3 0 0 2 51 ?
G1a 4,090 Grassland-opening in pine forest 1,473 1 0 0 0 0 5 3 8 0 0 0 48 ?? ?
G1b 1,236 2 1 0 0 1 4 3 5 1 1 1 57 ?? ??
G2a Grassland-opening in beechwood 1,914 2 0 0 0 0 5 3 7 0 0 0 55 ?
G2b 1,745 2 0 2 0 3 3 3 7 0 0 1 52 ??
G3a Subalpine grassland 1,672 2 0 0 0 0 4 2 8 1 0 1 59 ??
G3b 2,077 2 0 0 0 4 4 3 4 2 0 2 50 ?
G3c 1,300 2 0 1 0 0 5 3 8 1 0 0 61 ?
A1a 1,020 Rural mosaic (fields with hedges) 1,132 1 0 2 0 0 5 3 5 0 0 0 62 ???
A1b 1,201 1 0 1 1 0 5 3 10 0 0 0 22 ?
A2 (1,020) 9(9,250) Rural mosaic 9Q. trojana trees 798 1 0 2 1 1 5 3 8 1 0 1 58 ??
A3 (1,020) 9(1,050) Rural mosaic 9village ruins 971 2 0 1 1 1 4 2 7 2 0 1 65 ??? ?
A4a (1,020) 9(925A)
95,130
Rural mosaic 9mixed thermophilous
wood X bushes of J. communis
825 2 0 2 2 1 3 2 7 1 0 1 52 ?? ??
A4b 827 1 0 2 2 2 3 3 5 0 0 1 57
Habitat codes refer to Annex 1 of the Directive 92/43/EEC. Additional Hellenic habitat types in parenthesis
Topography-climate, Alt altitude (m), Sl slope (1:0–15%, 2:15–30%). Vegetation, T
1
trees ([10 m height), T
2
trees (2.5–10 m height), Sh
1
shrubs
(0.5–2.5 m height), Sh
2
shrubs (\0.5 m height), He herb cover, Hh mean height of herbs (cm) (1:1–5%, 2:6–25%, 3:26–50%, 4:51–75%,
5[75%). Fflowerheads (1\10, 2:11–20, 3:21–50, 4:51–100, 5:101–200, 6:201–300, 7:301–400, 8:401–500, 9:500–600, 10:[600). Ground,
R
1
rocks (0.5–1.5 m diameter), R
2
rocks ([1.5 m diameter), Bbare ground. Hu humidity (%). Impact, Ggrazing intensity, Dhuman disturbance,
?minimal, ?? low, * priority habitat type
508 J Insect Conserv (2009) 13:505–514
123
closed canopy forests (e.g., F3b, F5) were among the
poorest sites for both groups (Tables 1,2).
Regarding the most threatened Lepidoptera species in
the study area, we recorded Thymelicus acteon (SPEC 2) in
the species-rich thermophilous forest (F3a) and in a rural
mosaic combining patches of fields and oakwoods (A2).
We recorded P. apollo (SPEC 3, PBA) in the sub-alpine
grassland (G2b) and in a beech forest opening (F1b) and
E. aurinia (SPEC 3, PBA) at the species-rich grassland
(G3c).
Surrogate value
We found significant congruence between all species
richness patterns examined. The patterns of Lepidoptera
species of European Conservation Concern (SPEC) were
congruent with those of Lepidoptera (rho =0.796,
P\0.01) and Orthoptera (rho =0.686, P\0.01).
Besides, Lepidoptera and Orthoptera had also congruent
species richness patterns (rho =0.659, P\0.01).
The SPEC Lepidoptera functioned as the best surrogate
group when complementarity is concerned. When com-
paring the average species richness included in each step of
the complementary network selection, SPEC network (10
sites) succeeded to conserve even more species (3%) of
Lepidoptera and lost only 9% of Orthoptera species on
average (Fig. 3). The complementary network of Lepi-
doptera (12 sites) lost on average a comparative proportion
of 11% of SPEC and 18% of Orthoptera species. Finally
the complementary network of Orthoptera (9 sites) lost on
average a comparative proportion of 14% of SPEC and
18% of Lepidoptera species (Fig. 3).
Environmental factors
The number of flower heads, cover by low trees (2.6–10 m)
and altitude influenced significantly the diversity patterns
of Lepidoptera, affecting positively 89, 80 and 20% of
species, respectively (34% of variance explained) (Fig. 2a).
Concerning the threatened butterfly species in Europe,
cover by low trees and flowering affected positively the
presence of T. acteon, Glaucopsyche alexis and Pseudo-
philotes vicrama. Flowering affected also significantly the
presence of E. aurinia and E. medusa, whilst low trees
determined the presence of Scolitantides orion. Altitude
determined significantly the presence of P. apollo,Poly-
ommatus eroides and Erebia medusa. None of the above
mentioned parameters affected the presence of Maculinea
alcon.
The number of flower heads, cover by low bushes
(\0.5 m), altitude and bare soil influenced significantly the
diversity patterns of Orthoptera community, affecting
positively 75, 57, 45, and 18% of species, respectively
(35% of variance explained) (Fig. 2b).
Discussion
Surrogate value
We found a strong congruence in the species richness
patterns of SPEC butterflies, butterflies and grasshoppers,
suggesting the potential use of each taxon as a good sur-
rogate for the other two. Although butterflies are a
Fig. 2 Redundancy analysis diagram (RDA) presenting the signifi-
cant (P\0.05) environmental factors affecting the community of a
butterflies and bgrasshoppers. Only SPEC species and species with a
fit [25% are shown
J Insect Conserv (2009) 13:505–514 509
123
charismatic group that is well-studied, easily identified and
monitored, fulfilling the criteria for a good ‘‘umbrella’’
taxon (New 1997), studies on their surrogate value for
other taxa are contradictory. On one hand, they present
congruent diversity patterns with birds and tiger beetles at
continental scale (Pearson and Cassola 1992), with birds
and vascular plants at national scale (Pearman and Weber
2007) and with plants or bumblebees at local scale (Grill
et al. 2005; Niemela
¨and Baur 1998; Vessby et al. 2002).
On the other hand, several studies have shown their poor
surrogate value towards other taxa (Grill et al. 2005;
Gutie
´rrez and Mene
´ndez 2007; Howard et al. 1998; Lawton
et al. 1998; Lovell et al. 2007; Niemela
¨and Baur 1998;
Osborn et al. 1999; Ricketts et al. 1999,2002). Grass-
hoppers are more difficult to identify and monitor and their
surrogate value has been more rarely tested. They are
known to have a good cross-taxon congruence with bryo-
phytes, vascular plants, gastropods spiders, carabid beetles,
ants, or birds in agricultural landscapes (Sauberer et al.
2004), but also poor surrogate value for a number of other
taxa (Duelli and Obrist 1998; Kati et al. 2004c; Niemela
¨
and Baur 1998). Contrarily to our findings, butterflies and
grasshoppers are known to present incongruent diversity
patterns in temperate grasslands (Niemela
¨and Baur) and
savannas (Lovell et al. 2007).
We also found a strong correlation between red-listed
butterfly and overall butterfly species richness. This rela-
tionship is documented at a broader scale (Pearman and
Weber 2007) but not at local scale (Gutie
´rrez and Mene
´ndez
2007). These discrepancies may be related to the effects of
the different habitat templates in the various studies and
indicate that the use of certain taxa as surrogates of
Table 2 Insect species
diversity in the sites sampled
and species of European
conservation concern
concentrated in Europe
(SPEC2) or not (SPEC3)
Numbers in parenthesis indicate
target-species of prime butterfly
areas (PBAs)
HShannon index, N1 sum of
individuals, N2 sum of
frequencies, Snumber of
species
Site Habitat description Lepidoptera Orthoptera
SN1 HSPEC2 SPEC3 SN2
F1a Beech forests of Fagus spp. 25 81 2.79 1 11 14
F1b 20 55 2.64 1 (1) 7 8
F2a Oakwood of Quercus spp. 20 33 2.83 7 15
F2b 32 116 3.18 3 13 25
F3a Mixed thermophilous forests 53 156 3.79 1 1 19 42
F3b 5 5 1.6 2 2
F4 Riverine vegetation 30 79 3.16 6 10
F5 Fir forest of A. borisii-regis 9 17 1.97 2 3
F6 Pine forest of P. nigra 23 68 2.78 10 17
G1a Grassland-opening in pine forest 34 99 3.29 1 17 35
G1b 26 74 2.96 1 10 22
G2a Grassland-opening in beechwood 18 60 2.67 7 27
G2b 31 101 3.13 3 (1) 22 45
G3a Subalpine grassland 17 52 2.56 1 18 41
G3b 14 49 2.2 2 14 49
G3c 51 196 3.52 2 (1) 15 43
A1a Rural mosaic (fields with hedges) 27 75 3.1 8 21
A1b 40 180 3.45 1 11 27
A2 Rural mosaic 9Q. trojana trees 44 187 3.27 1 2 17 33
A3 Rural mosaic 9village ruins 33 108 3.23 13 33
A4a Rural mosaic 9mixed thermophilous
wood 9bushes of J. communis
51 183 3.51 1 18 31
A4b 35 141 3.22 14 26
Fig. 3 Average proportion of species richness included in each step
of the complementary network formation, designed after the surrogate
group (SPEC butterflies, butterflies and grasshoppers)
510 J Insect Conserv (2009) 13:505–514
123
biodiversity should not be uncritically generalized among
different scales and regions but should be locally evaluated.
Could we attribute the congruence found herein from an
ecological perspective? Butterflies and grasshoppers are
both terrestrial thermophilous taxa sharing similar traits in
their life history and ecology. We found that three common
environmental parameters influenced their patterns: num-
ber of flower heads, altitude, and cover of low trees or
bushes. Butterflies are positively affected by the number of
plants in flower because of the dependence of nectar-
feeding species (Grill et al. 2005;O
¨ckinger and Smith
2006). The response of grasshoppers may be due to the
indirect effect of the flowering season that coincides with
the peak of grasshopper activity at higher altitudes, or also
to their dependence on colourfull substrates as hiding
places against predators (Isely 1938). Both taxa also
respond to vegetation structure and negatively to altitudinal
gradient (Gebeyehu and Samways 2002; Gutie
´rrez and
Mene
´ndez 2007; Kati et al. 2004a; Sawchik et al. 2005;
Wettstein and Schmid 1999).
Testing the surrogate value of the taxa on the basis of
complementarity, we found that each surrogate network
maintained quite well the species richness of the other two
target groups (less than 18% average species loss). The
complementary networks of butterflies and grasshoppers
are known to conserve adequately several other unrelated
taxa in the tropics and Mediterranean regions, respectively
(Howard et al. 1998; Kati et al. 2004b). According to our
results, the complementary network of SPEC butterflies
was the most efficient of all, as it succeeded to conserve
overall butterfly richness (no loss) and grasshopper rich-
ness (\10% average loss).
Our results supported that the red-listed butterfly species
could be a good surrogate group for the conservation and
monitoring of butterflies and grasshoppers, because they
presented congruent diversity patterns with these taxa and
also the greatest efficiency in conserving them in their
complementary network. Butterflies satisfy a number of
criteria for a valuable surrogate for other insect taxa and
they are among the most well-studied invertebrate groups
(New 1997; Thomas 2005). During the last decades the
distribution and abundance of many European butterfly
species have declined seriously (12% of the total diversity)
because of the changes in the European environment (Pullin
1995; van Swaay and Warren 1999). In order to stop this
enormous loss a good database and monitoring network of
butterflies and their trends has been developed in several
countries to provide adequate information such as distri-
bution, threats and conservation measures for threatened
species (SPEC). Several studies also propose butterflies as
valuable indicators for environmental change, focusing on
their quick response to habitat destruction and demon-
strating that they can be used for monitoring trends in
habitat quality (Pywell et al. 2004; van Swaay et al. 2006).
More recently, butterflies have been assessed as one of the
most useful indicator groups of climate change (Forister and
Shapiro 2003; Mene
´ndez et al. 2007; Oostermeijer and van
Swaay 1998; Roy and Sparks 2000; Warren et al. 2001).
Grammos as a prime butterfly area
The area of Mt. Grammos is a site of the European NA-
TURA 2000 ecological network that satisfies also the
criteria of a Prime Butterfly Area (PBA), since it includes
populations of two target-species of the Prime Butterfly
Area project (van Swaay and Warren 1999). Furthermore,
Grammos area hosts great species richness with 120 out of
the 576 European Lepidoptera species, with a high pro-
portion (24%) of species of European Conservation
Concern (SPEC). The area includes also highly restricted
montane species which have a very limited possibility of
adapting to global warming and may be seriously threa-
tened in the future (Dennis 1993; Wilson et al. 2005). We
propose therefore that the Grammos protected area should
be added as an eleventh site in the network of Prime
Butterfly Areas (PBAs) of Greece.
Insect diversity conservation
The proposed selection of this mountain area as a PBA
would require specific conservation measures on the whole
mountain area, in order to secure viability of populations
through the supply of multiple favorable habitat patches.
We found that habitats hosting high diversity of butterflies
and grasshoppers are mostly those of an open structure,
such as subalpine grasslands, woodland clearings and old
rural mosaics. Traditional agricultural practices should be
encouraged (Grill and Cleary 2003), since further aban-
donment of agricultural land and its conversion to early
forested stages could lead to a decline of important butterfly
species such as E. aurinia (van Swaay et al. 2006). Rota-
tional grazing that supports high grass species richness and
abundance and also more grasshopper and butterfly species
should also be encouraged (Gebeyehu and Samways 2003;
Poyry et al. 2005; van Swaay et al. 2006; WallisDeVries
et al. 2002). Furthermore, in view of on-going climatic
change, the role of such mountainous and subalpine habitats
as refugia for lower altitude species can prove crucial in the
near future. According to our results, a monitoring network
of Red-listed butterfly species should be developed in order
to obtain detailed information on their population trends,
well reflecting also overall butterfly and grasshopper
diversity. Finally, the role of these insect groups as surro-
gates of overall biodiversity has to be evaluated further
through a more extensive survey of more taxa and indices of
habitat quality.
J Insect Conserv (2009) 13:505–514 511
123
Acknowledgments We are grateful to ‘‘Proodeutiki Enosi Pyrso-
giannis’’ for financially supporting our field research and to L.
Willemse and A. Grill for their valuable support and help in specimen
identification. We also thank H. Papaioannou, P. Gerofoka, G. The-
odoropoulos, G. Michaelides and G. Adamides for field assistance.
Table 3 Lepidoptera species
(120) in Grammos protected
area (2007) and their
conservation status according to
the Red Data Book for
European Butterflies (van
Swaay and Warren 1999)
The nomenclature used follows
Karsholt and Razowski (1996)
B
Species annexed in Bern
convention;
2
SPEC 2;
3
SPEC
3;
4
SPEC 4; * PBAs; species in
parenthesis are recorded outside
the sites sampled
PAPILIONIDAE Melitaea trivia Lycaena thersamon
Iphiclides podalirius Mellicta athalia Lycaena tityrus
Papilio alexanor
B
(Nymphalis antiopa) Lycaena virgaureae
Papilio machaon Nymphalis polychloros Polyommatus bellargus
Parnassius apollo
3,
*Polygonia c-album (Polyommatus coridon)
4
Parnassius mnemosyne
B
(Polygonia egea) Maculinea alco
3
Zerynthia polyxena
B
Vanessa atalanta Polyommatus daphnis
4
PIERIDAE Vanessa cardui Plebeius argus
Anthocharis cardamines LIBYTHEIDAE Plebeius argyrognomon
Aporia crataegi Libythea celtis Plebeius idas
Pieris ergane RIONIDAE Plebeius pylaon
Pieris mannii Hamearis lucina Polyommatus dorylas
4
Pieris napi HESPERIIDAE Polyommatus eroides
3
Pieris rapae Carcharodus alceae Polyommatus icarus
Colias alfacariensis
4
Erynnis tages
4
Pseudophilotes vicrama
3
Colias aurorina Ochlodes venata (Satyrium acaciae)
4
Colias croceus Pyrgus armoricanus Satyrium ilicis
(Gonepteryx cleopatra) Pyrgus malvae Satyrium spini
Gonepteryx rhamni Pyrgus sidae Scolitantides orion
3
Leptidea duponcheli Spialia orbifer Aricia anteros
4
Leptidea sinapis complex Spialia phlomidis SATYRIDAE
Pieris brassicae Thymelicus acteon
2
Coenonympha arcania
Pontia daplidice complex Thymelicus lineola Coenonympha leander
NYMPHALIDAE Thymelicus sylvestris
4
Coenonympha pamphilus
Aglais urticae LYCAENIDAE Coenonympha rhodopensis
4
Argynnis adippe Polyommatus escheri
4
Erebia medusa
3
Argynnis aglaja Polyommatus thersites (Erebia melas)
4
Argynnis niobe Polyommatus admetus
4
Erebia ottomana
Argynnis Pandora Polyommatus amandus Hipparchia fagi
4
Argynnis paphia (Agrodiaetus damon)
3
Hipparchia syriaca
Boloria graeca
4
Polyommatus ripartii Hipparchia volgensis
4
Brenthis hecate Aricia agestis Hyponephele lupinus
Brenthis daphne Callophrys rubi Brintesia circe
4
Boloria dia Celastrina argiolus Kirinia roxelana
Boloria euphrosyne Cupido minimus Lasiommata maera
Euphydryas aurinia
3,
*Cupido osiris Lasiommata megera
Inachis io Cyaniris semiargus Lasiommata petropolitana
Issoria lathonia Cupido alcetas Maniola jurtina
(Limenitis reducta) Glaucopsyche alexis
3
Melanargia galathea
4
Melitaea arduinna Leptotes pirithous Melanargia larissa
4
Melitaea cinxia Lycaena alciphron Pararge aegeria
Melitaea didyma Lycaena hippothoe Pyronia tithonus
Melitaea phoebe Lycaena phlaeas
Appendix 1
See Table 3.
512 J Insect Conserv (2009) 13:505–514
123
Appendix 2
See Table 4.
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