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67
Journal of Mediterranean Ecology vol. 16, 2018
Effects of changing grazing systems on the threatened genus Peripodisma
(Orthoptera: Acrididae: Melanoplinae) in the Mediterranean mountains of
the southern Balkans
Michèle Lemonnier-Darcemont1, Vassiliki Kati2, Luc Willemse3, Christian Darcemont4
1 Corresponding author. G.E.E.M., P.O. Box 60, 48060 Parga, Greece. Email: Lemonniergeem@free.fr
2 Depart. of Biological Applications and Technology, University of Ioannina, 45110 Ioannina, Greece. E-mail: vkati@uoi.gr
3 Naturalis Biodiversity Center, Darwinweg 2, Postbus 9517, 2300 RA Leiden, The Netherlands. Email: luc.willemse@
naturalis.nl
4 G.E.E.M., P.O. Box 60, 48060 Parga, Greece. Email: Darcemont@free.fr
Keywords: Albanian mountain grasshopper, biodiversity, calcareous grassland, cattle grazing, Cika mountain
grasshopper, endemic, IUCN Red List, land-use intensification, linear indices of abundance, species specializa-
tion index, Tymphi mountain grasshopper.
Abstract
This study examined the effects of pastoralism, including cattle grazing, on populations of three species of locally endemic
and rare Peripodisma grasshoppers in calcareous grassland mountain habitats of northwestern Greece and southern Albania.
The three Peripodisma species are on the IUCN Red List as near threatened, endangered, and critically endangered species,
and cattle grazing had been identified as a key threat to the species. The study sites represented 70% of the known locations
of Peripodisma genus. The region was historically grazed by local breeds of nomadic sheep and goats, but grazing practices
had recently shifted to cattle grazing from non-local cattle breeds. We found a clear relationship between local abundance
of Peripodisma and overall richness of Orthoptera communities. Orthoptera richness decreased at sites with medium to
high impacts of livestock grazing. Cattle grazing had significant adverse effects on overall Orthoptera species richness
and on Peripodisma abundance. Further studies are urgently needed to gather more data and information to guide grazing
management and conservation planning that will provide a more balanced coexistence between livestock and Orthoptera,
especially for the rare Peripodisma species that are in dire need of conservation management.
Introduction
Pastoral farming in the Mediterranean mountains of
Greece and Albania has existed since antiquity. Over
time, pastoralism has shaped mountain landscapes
and vegetation structure (Kizos, 2008) and, as a con-
sequence, the invertebrate biodiversity. Over the last
century, nomadic herding has drastically decreased,
evolving into movements of livestock across elevation
gradients, with the purpose of ensuring forage avail-
ability during summer (Blanc, 1963; Hadjigeorgiou,
2011). Grazing is usually conducted at a low intensity,
as sheep and goats are secured at night and shepherded
on mountain pastures during the day. Additional
changes in pastoralism have emerged in recent decades,
particularly in Greece, due to links with the European
Economic Community. Non-local cattle breeds have
been introduced, resulting in only 0.64% of native cat-
tle breeds (Legakis & Kapari, 2013), due to the new
requirement for production of cattle meat (Hadjigeor-
giou, 2011). Over this period, there has been a strong
decrease in the number of farm holdings, with only a
slight decline of overall livestock numbers, resulting in
concentrations of herds (Eurostats, 2012). The effects
Journal of Mediterranean Ecology vol. 16, 2018: 67-79
© Firma Effe Publisher, Reggio Emilia, Italy
68
Journal of Mediterranean Ecology vol. 16, 2018
of changes from both overgrazing and undergrazing
on vegetation and biodiversity have been object of
investigation (Gkoltsiou, 2011; Hadjigeorgiou, 2011).
The relationships between vegetation and Orthoptera
diversity have previously been demonstrated (Ander-
son 1964, Kati 2012), and the positive or negative im-
pacts of grazing on Orthoptera species and populations
have been studied across a variety of ecosystems and
grazing systems (Smith, 1940; Morris, 1969; Holmes
et al., 1979; Capinera & Sechrist, 1982; Shiyomi et
al., 1982; Grayson & Hassall, 1985; Voisin, 1986;
Gueguen-Genest & Gueguen, 1987; Louveaux et al.,
1996; Kruess & Tscharntke, 2002; Gonseth, 2010).
Our study investigates the effects of grazing on Or-
thoptera. Compared to other animal groups, sampling
of Orthoptera is relatively simple and identification of
most species is easy. Orthoptera are distributed among
various trophic levels of the food webs (carnivores, om-
nivores, herbivores and scavengers), and such trophic
relationships are highly dependent upon environmental
conditions (e.g., elevation, microclimate, vegetation
structure). Orthoptera provide excellent indicators of
ecological conditions (Guegen, 1989, 1990, 1996; Kati
et al., 2004).
We focused on grasshoppers of the genus Peripo-
disma, which are restricted to the Pindos Mountain
Range of Greece and Albania, and were reported to be
threatened by domestic livestock grazing (Hochkirch et
al., 2016). All three species of Peripodisma (P. tymphii,
P. llofizii, P. ceraunii) are included in the IUCN Red
List (Hochkirch et al., 2016). The Tymphi mountain
grasshopper Peripodisma tymphii Willemse, 1972 is
an endangered species (Willemse et al., 2016a) due to
its fragmented distribution pattern on seven mountain
ranges of south-eastern Europe. The distribution of
the Tymphi mountain grasshopper covers the northern
Pindos massif in Greece (Willemse, 1972, 1984; Wil-
lemse & Willemse, 2008), and the adjacent mountains
of southern Albania (Lemonnier-Darcemont et al.,
2015; Lemonnier-Darcemont & Darcemont, 2016). The
Albanian mountain grasshopper Peripodisma llofizii
Lemonnier-Darcemont & Darcemont, 2015 is listed as
Near Threatened (Chobanov et al. 2016), and is known
of only four locations in the mountains of the southern
Albania region of Gjirokaster (Lemonnier-Darcemont
& Darcemont, 2015a; Lemonnier-Darcemont et al.,
2015). The Cika mountain grasshopper Peripodisma
ceraunii Lemonnier-Darcemont & Darcemont, 2015
is a Critically Endangered species (Chobanov et al.,
2016) with a very restricted distribution, only known
from the Cika mountain in Vlore district of Albania
(Lemonnier-Darcemont & Darcemont, 2015b; Lem-
onnier-Darcemont et al., 2015).
All three species are montane specialists that oc-
cur above 1100 m elevation, and have similar habitat
preferences (Figure 1). Peripodisma inhabit grasslands
of Mediterranean montane and alti-Mediterranean
vegetation zones, with Cika and Albanian mountain
grasshoppers known to favour rock or scree slopes
(Lemonnier-Darcemont & Darcemont, 2015a, 2015b).
They share similar life cycle characteristics, with egg
hatching occurring at the beginning of May, and adults
appearing in July through mid-October. These species
have been reported to be threatened by domestic live-
stock grazing, particularly cattle grazing, resulting in
declining population trends for the Cika and Tymphi
mountain grasshoppers (Chobanov et al., 2016; Wil-
lemse et al., 2016). The population trend of the Alba-
nian mountain grasshopper is unknown (Chobanov et
al., 2016).
Given that domestic livestock grazing manage-
ment has recently changed across the region, and the
general impression of a strong and quick reduction of
biodiversity in impacted areas, the aim of this study is
1. To assess whether the local overall score charac-
terising the richness of orthopteran communities
provides a reliable indication of the abundance
status of Peripodisma populations on these southern
Balkan mountains.
2. To identify parameters of pastoralism that have
negative ecological effects on the endemic genus
Peripodisma.
Methods
The taxonomic names and classifications used cor-
respond to the updated Orthoptera Species File Online
(Cigliano et al. 2018).
Sites and sampling
Sampling of orthopteran communities, including
Peripodisma species, was conducted in Greece and
Albania at nine of the thirteen locations where Peripo-
disma species are known to occur (Figure 2), which
represent 70 % of the known localities for the genus.
Table 1 provides locality and environmental informa-
tion for each of the nine sampling sites. Our sampling
sites included each of the three type localities where
the three species were discovered and described from,
two additional sites on the border between Greece and
Albania, and four additional sites that were chosen
randomly among the remaining eight known sites.
69
Journal of Mediterranean Ecology vol. 16, 2018
Figure 1. Example of Peripodisma habitat of subalpine calcareous grassland (Albania, study site 6).
Figure 2. Map of known localities and studied sites. Light green 0-600 m, light brown 600-1200 m, brown 1200-1800 m, dark brown
> 1800 m. Markers 1 to 9: studied sites. Triangles: known sites of P. llofizii not studied. Circles: known sites of P. tymphii not studied.
70
Journal of Mediterranean Ecology vol. 16, 2018
Pastoralism was present in all known locations were
Peripodisma species have been found, however there
were differences in pastoralism practices between sites.
We sampled Orthoptera during a three-year re-
search period (2014-2016) from the end of July to
the beginning of September in each year. This time
range matched the main activity period of the adult
Orthoptera in the study area. At each sampling site, we
studied a large section with homogeneous vegetation
structure, exposition, soil, etc. Sampling was always
performed during the late morning hours, and only
when weather conditions were optimal for diurnal
Orthoptera activity: on sunny days with no clouds and
reduced wind, and temperature close to the monthly
mean temperature (no sampling during heat waves).
It was assumed that pastoralism practices did not vary
over the three-year study.
Environmental and pastoral parameters
All localities had approximately the same envi-
ronmental characteristics: calcareous mountain under
Mediterranean climate, open grassland landscape
above 1500 m elevation. In order to relate variation
in Orthoptera species compositions and abundance’s
to variation in environmental features across the study
sites, we focused on parameters which tend to vary
considerably across sites. Within each site, the vegeta-
tion structure was homogeneous on a very large surface
with no detected changes throughout the three years of
investigation. At each site, we ascertained vegetation
structure and grazing parameters. We recorded the fol-
lowing parameters of vegetation structure: cover (%)
of tree layer (> 4 m), shrub layer (1-4 m), dwarf-shrub
layer (< 1m), herbaceous vegetation or field layer, as
well as bare soil and stones (Prodon, 1988).
Pastoral data were ascertained by direct observation
(characteristic of herds (species, breeds) and number
of individuals) measured on site directly or by using a
drone, and supplemented by personal communications
with local shepherds, and using literature data (Cabal-
lero et al., 2009; Gkoltsiou, 2011; Hadjigeorgiou, 2011)
in order to estimate period and duration of presence
of herds on the site, and type of grazing regime. The
breeds of sheep, goats and cattle were the same across
all sites. We assumed that pastoral conditions were
identical from one year to another throughout our study
Table 1: Information on sites sampled for this study.
Locality Country Latitude Longitude Elevation Habitat Peripodisma
species
1 Mt Llofiz, Gjirokastër county Albania 40°12’56.6” 020°09’52.8” 1720 m
Calcareous dry grassland of the
Mediterranean mountane stage with
Juniperus sp. and Astragalus sp.
P. llof izii
2 Mt Lunxhërisë, Gjirokastër county Albania 40°10’26.6” 020°12’24.2” 1800 m
Calcareous dry grassland (on rocky
ground) of the Mediterranean mountane
stage with Arctostaphylos uva-ursi and
Astragalus sp.
P. llof izii
3 Mt Qores, Vlorë county Albania 40°12’48.4” 019°36’17.1” 1830 m
Calcareous dry grassland (on scree) of the
Oro-Mediterranean stage with Juniperus
sp. and Astragalus sp., in edge of open
pine forest composed of Pinus heldreichii.
P. ceraunii
4 W. Mt Nemërçkës Albania 40°06’19.0” 020°25’19.0” 1750 m
Calcareous dry grassland of the
Mediterranean mountane stage with
Arctostaphylos uva-ursi and Eryngium sp.
P. t ymphi i
5 Mt Buretos, Gjirokastër county Albania 40°01’53.0” 020°18’09.7” 1700 m Calcareous dry meadow of the
Mediterranean mountane stage. P. t ymphi i
6 Mt Silvit, Gjirokastër county Albania 40°03’51.2” 020°29’23.0” 1823 m Subalpine calcareous grassland. P. t ymphi i
7 Mt Murganës, Gjirokastër county Albania 39°47’17.8” 020°23’22.7” 179 0 m
Calcareous dry meadow of the
Mediterranean mountane stage with
Juniperus sp., Astragalus sp. and
Echinops sp.
P. t ymphi i
8 Mt Khionistra, Epirus Greece 39°31’10.0” 020°30’58.3” 1571 m
Calcareous dry grassland with Juniperus
oxycedrus and Satureja montana of the
Mediterranean mountane stage.
P. t ymphi i
9Megalo Papigo, Epirus Greece 39 °58’44. 2” 020°46’07.7” 1905 m Calcareous dry meadow of the Oro-
Mediterranean stage. P. t ymphi i
71
Journal of Mediterranean Ecology vol. 16, 2018
period. We determined a coefficient of relative grazing
impact on Peripodisma by multiplying the number
of individual livestock (all types of livestock; sheep,
goats, cattle) per hectare by the number of months
of pasture on site, but including different weighting
depending on the month of pasture (same weighting
for different grazing animals). We considered that the
impact of grazing in spring was greater than graz-
ing in summer or autumn because our observations
conducted in captivity and in nature showed that the
diet of Peripodisma consisted mainly of forbs (Figure
3), and the availability of flowers for Peripodisma in
their early stages appeared to have been a key factor.
Preferred forbs were mainly Scrophulariaceae, such
as Verbascum spp., and also Fabaceae, such as Astra-
galus spp.. When the livestock herds were composed
of cattle, they were generally moved to high elevation
pastures (habitat of Peripodisma) earlier in the season
than sheep herds. The cattle consumed the tender plant
parts first, such as flowers, which put cattle herds in
direct competition for food with Peripodisma.
Peripodisma sampling method to measure population
density
We used the Linear Indices of Abundance (LIA)
method developed by Voisin (1980) to measure the
density of Peripodisma grasshoppers. Sampling at each
site was restricted to an observer walking ten linear
10m transects (1m wide), without overlap, using a
10m-long rope held by the observer. Each transect was
randomly sampled (by counting without netting) within
the whole pasture area as defined above. The index
obtained by the sum of ten transects, resulted in the
number of individuals of Peripodisma per 100m2. This
sampling was executed first, before any other sampling
aimed to measure the overall richness of Orthoptera
community, which could have caused movement of
Peripodisma grasshoppers, and affected subsequent
sampling for those species.
Sampling method to characterize the richness of the
Orthoptera community
We walked randomly through each sampling area
in all directions, capturing with an aerial insect net all
encountered adult Orthoptera until reaching a sample
of 100 individuals, then recorded it on a data form to
the species rank, and all individuals were then released
unharmed. The surface area of any studied site was
between five hundred to one thousand square meters.
In rare cases where Orthoptera densities were low
at a given site, we ceased sampling after two hours
regardless of the number of captured individuals (i.e.
the total number sampled may not have reached 100).
Sampling without removal has the advantage of
greatly limiting disturbance to the orthopteran com-
munities; nevertheless the observer must be quite
experienced in order to identify individual orthopteran
specimens in the field, and this method has been widely
used by others (Dreux, 1962, 1972; Marty, 1968; De-
faut, 1978; Luquet, 1985; Voisin 1979, 1980, 1986).
This sampling protocol, applicable in open habitats,
was appropriate for the sampled sites.
Data collected were used to calculate four ecologi-
cal indicators of individual abundance and species di-
versity per locality, which were then used to calculate
an overall score characterising the richness of the
Orthoptera community at each site, called « overall
richness of Orthoptera community » (Lemonnier-
Darcemont et al., 2011). The four indices were:
1. Nsc (Number of species capped): Species richness
on each site but with a maximum threshold value
(maximum 30).
2. Rv (Richness value): Quality of richness of the com-
munity. It is the ratio of species with low ecological
valence (i.e. narrow ecological amplitude) found
on the site to the total number of species recorded,
multiplied by 25. This Rv index highlights the lo-
cal interest of species which are more vulnerable
because they are less eclectic in their choices of
elective habitats, it presupposes the knowledge of
the Species Specialization Index (SSI) as defined
Figure 3. Peripodisma diet is mainly forbs.
72
Journal of Mediterranean Ecology vol. 16, 2018
by Julliard et al (2006) of each recorded species.
A given species present in h habitat classes among
H possible habitat classes has a SSI equal to the
square root of (H-h)/h. The SSI reflects the ability
of a given species to occupy various habitats, the
higher the SSI, the lower the ecological valence of
the species. By setting a threshold to the SSI (H/4),
we ranked the species in two classes: low ecologi-
cal valence (high SSI number), and high ecological
valence (low SSI number). We used EUNIS biotope
codes (Louvel et al., 2013) at first levels to define
habitats for SSI computation.
3. En (Equitability indicator): predominance or not
of some species. It is the Equitability index (Es) as
proposed by Barbault (1992), multiplied by 25. The
Equitability index is issued from the index of diver-
sity of Simpson-Weaver (Is = Simpson reciprocal
index = 1/D), as Es = (Is-1)/(Ns-1), Ns being species
richness. When all species are equally abundant, the
indice “En” (= 25 * Es) reaches the maximum value
(25), but decreases when the relative abundance of
the species is not equally distributed.
4. Rn (Richness of vulnerable species): relative abun-
dance of low ecological valence species. The Rn
index is the ratio of number of samples between
species with low ecological values and all species.
Rn = 20 * vn / (Vn + vn), with vn = Σ ni for spe-
cies over or equal to the SSI threshold, and Vn = Σ
ni for species under the SSI threshold, “ni” being
the number of observations of species “i” in the
quantitative sampling.
The overall score ranging from 1 to 100, of the
richness of the Orthoptera community considered is
calculated by summing the four parameters: Nsc +
Rv + En + Rn.
Analysis method
For the first objective of the study, the analysis of
the correlation between overall richness of Orthoptera
community and Peripodisma density was performed
by a simple Pearson’s correlation coefficient com-
puted with the richness per locality and the number of
Peripodisma sampled per 100 m²; and we provided a
plot of the regression analysis by using standard em-
bedded functions of EXCEL. Using XLSAT software,
we performed a principal component analysis (PCA)
to examine the relationships between environmental
parameters, pastoral parameters and Orthoptera spe-
cies richness, and aimed to provide an ordination of
the relationships between key variables.
Results
The richness of the Orthoptera communities (based
on relative abundance) and Peripodisma abundance
(based on quantitative transects) is provided in Table
2. The Pearson correlation analysis between Peripo-
disma densities and the overall richness of Orthoptera
community resulted in a significant and high correla-
tion coefficient of r=0.93 (p=0.0004). A scatter plot
from the regression analysis is provided in Figure
4, R²=0.856. These results demonstrate the overall
richness of Orthoptera community is correlated with
Peripodisma densities across the study site pastures,
and therefore this score may be used as an indicator of
Peripodisma density. 86% of variation in Peripodisma
density is predicted by the score providing the overall
richness of Orthoptera community.
Refer to Appendix A for the list of Orthoptera spe-
cies recorded in the 9 localities visited.
Table 2. Overall score characterising the richness of the Orthoptera community among the study sites, along with population density per
100 m² for Peripodisma species.
Nsc = Number of species; Rv = Richness value; En = Equitability among species; Rn = Richness of vulnerable species.
Orthoptera community Peripodisma
Locality Nsc Rv En Rn Richness Population density
113 9.6 15.5 10.4 49 135
2812.5 18 12.2 51 204
386.2 12 2.4 29 26
4911.1 14.5 10.6 45 127
513 1.9 17.7 2.4 35 85
612 12.5 21.5 10.4 57 230
711 2.3 15 432 37
83 8.3 1.2 0.8 13 9
9412.5 12.2 3.6 32 28
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Journal of Mediterranean Ecology vol. 16, 2018
Figure 4. Relationship between measures of abundance of Peripodisma and overall index of Orthoptera community richness.
Table 3. Vegetation structure and ground cover along with pastoralism parameters across the 9 sites.
NL = non local breed. C = Crossbreed native with non-local breed.
Locality Vegetation structure and ground cover (% cover) Pastoralism parameters
Shrub layer Dwarf-shrub
layer Field layer Bare soil and
stones
Herd Composition %
Period Estimated relative
impact
Sheep Cattle
1 _ 25 50 25 100 July – mid August 125
2 _ 20 30 50 100 July - September 160
3 5 20 20 55 100? ?1June – mid August 525
4 1 20 54 25 1002 June - July 460
5 - 15 55 30 100 July - August 355
6 - 15 55 30 100 July – September 80
7 - 15 55 30 50 50 (NL+C) June – August 550
8 - 30 35 35 20 80 (N L+C) May - September 1360
9 - 20 30 40 100 (N L+C)3May - September 680
The vegetation structure and pastoralism parameters
are provided in Table 3.
The plot of the two main factors of all data recorded
is provided in Figure 5. Results of PCA showed a poor
correlation between sheep pastoralism and Orthoptera
richness and Peripodisma abundance, but a clear nega-
tive relationship when cattle were present.
Locality 3 is an outlier which can be explained by
1 Observed below the study area but no information on their upper elevational limit.
2 Probably also a small unit of goats in their upper elevational limit (50-100 individuals).
3 Since 2016 the area occupied by the Peripodisma is no longer grazed by cattle.
74
Journal of Mediterranean Ecology vol. 16, 2018
grazing over the past decades. We assume that during
past several decades, this locality had been overgrazed.
Although the current grazing pressure seems appropri-
ate, the biotope had not yet recovered its potential full
biodiversity.
We found a positive relationship between the rela-
tive coverage of the herbaceous field layer and the
richness and diversity of the Orthoptera communi-
ties. However more precise parameters (plant species
composition) would probably be necessary to refine
correlation parameters.
The results showed that at sites used only by small
ruminants (sheep and/or goats), the richness of the Or-
thoptera communities was high, along with the highest
densities of Peripodisma, a probable consequence of
a current moderate pastoral pressure. In contrast, we
measured sharp decreases of Peripodisma in areas pas-
tured by cattle of non-indigenous or crossbred breeds
(mainly issued from Brown Swiss cows).
Discussion
We found that the densest populations of Peripo-
disma occurred at sites where cattle grazing was absent.
Moderate sheep grazing pressure did not seem to be a
factor of degradation in those environments, contrary
to what we observed when cattle were introduced, at
least non-local or crossbreeds (Figure 6), that were
not adapted to these Mediterranean mountains. Cattle
generally arrived earlier in the season (due to their
wider foraging habits) and competed early and directly
with phytophagous species of Orthoptera, unlike sheep
which were present only in summer on these pastures.
This was particularly the case with Peripodisma,
which prefer to feed on the first buds, young flowers
and leaves (unpublished data, Lemonnier-Darcemont),
which are also highly desired by cattle.
It is possible that smaller, more rustic local cattle
breeds may have less impact on these natural environ-
ments (due to smaller body size with less consumption,
less soil and vegetation trampling, and a different for-
aging behaviour). Unfortunately we lack comparative
measures to support this assumption because the transi-
tion to non-local cattle breeds was completed mainly
between 1960 and 1970 (Zervas & Boyazoglou, 1977).
After the introduction of breeds of cattle into inappro-
priate lands (lands very different from the original land
of the breed), the process of degradation occurs rapidly
through the decrease of the herbaceous field layer. It
therefore seems important to be able to measure and
monitor grazing impacts as soon as possible, in order
to plan for actions that will help to stop this negative
trend. Such measures have already been practiced in
other ecological regions and environments of northern
Greece (Karatassiou et al., 2009).
Figure 5. PCA plot on results.
75
Journal of Mediterranean Ecology vol. 16, 2018
Figure 6. Crossbreed cattle (Brown Swiss), site number 8
Livestock parasite prevention and other potential
treatments for livestock studied for Coleoptera (Lu-
maret, 2010; Cornille, 2010) could also be considered
as a threat to Orthoptera.
Due to its conservation status and its relationship
with Orthoptera richness, the use of the genus Peripo-
disma as an indicator genus for the environmental
health of grazed montane vegetation formations, seems
an appropriate choice. The measure of overall richness
of Orthoptera community can be performed more easily
and quickly than an accurate measure of the density
of Peripodisma, therefore it could be used for regular
surveys, according to time constraints. Conservation
planning and implementation of such plans for these
threatened and high heritage value taxa would also
provide an effective way to manage the overall biodi-
versity of montane grasslands of the region.
Our findings presented here should be considered
an initial approach for understanding the effects of
livestock grazing on Orthoptera species richness and
Peripodisma species, a snapshot of different localities
more or less impacted by pastoralism. We emphasize
the importance of planning a comprehensive moni-
toring protocol targeted at these sites where newly
introduced breeds of cattle are apparently adversely
impacting Peripodisma species. The objective of such
research would be to better understand the environmen-
tal impacts of cattle and to use that information to plan
conservation management strategies for Peripodisma
and other species that utilize the same montane grass-
land habitats. The future experimental studies could be
performed with different types of grazing treatments,
each paired with non-grazed control studies.
76
Journal of Mediterranean Ecology vol. 16, 2018
Appendix A: Sampled Orthoptera species for each locality.
123456789
Poecilimon gracilioides X X X
Poecilimon jonicus jonicus X X
Poecilimon zimmeri X X X
Tettigonia viridissima X
Decticus verrucivorus X X X X X
Platycleis affinis affinis X
Platycleis grisea X X X
Platycleis intermedia intermedia X
Modestana ebneri ebneri X
Parnassiana sp. X X X
Parnassiana tymphiensis X
Vichetia oblongicollis X X
Sepiana sepium X
Pholidoptera femorata X X X
Pholidoptera macedonica X
Eupholidoptera schmidti X X X X
Bucephaloptera bucephala X
Psorodonotus macedonicus X
Saga hellenica X X X
Gryllus campestris X
Peripodisma ceraunii X
Peripodisma llofizii XX
Peripodisma tymphii X X X X X X
Celes variabilis X X X X X X
Oedipoda germanica X X
Arcyptera microptera XX
Stenobothrus fischeri X
Stenobothrus lineatus X
Stenobothrus nigromaculatus X
Stenobothrus rubicundulus X X X X X
Gomphocerus sibiricus X
Myrmeleotettix maculatus X
Stauroderus scalaris X X X X X X
Chorthippus biguttulus euhedickei X X X
Chorthippus mollis mollis X
77
Journal of Mediterranean Ecology vol. 16, 2018
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