Research Article

Journal of Orthoptera Research 2018, 27(1): 23-33

Effects of grazing on orthopteran assemblages of Central-European sand

grasslands

ZOLTAN KENYERES!

1 Acrida Conservational Research L.P., Tapolca 8300, Hungary.

Corresponding author: Zoltan Kenyeres (kenyeres@acridabt.hu)

Academic editor: Corinna S. Bazelet | Received 12 June 2017 | Accepted 14 January 2018 | Published 12 June 2018

http://zoobank.org/931577F0-49AA-4CA9-965A-E2A5FB862ECC

Citation: Kenyeres Z (2018) Effects of grazing on orthopteran assemblages of Central-European sand grasslands. Journal of Orthoptera Research 27(1):

23-33. https://doi.org/10.3897/jor.27.15033

Abstract

The effect of grazing on Orthoptera assemblages has long been the
focus of research worldwide due to the high sensitivity of orthopterans
to changes in vegetation structure. According to previous studies, grazing
has individual, spatially-different effects on orthopteran assemblages. The
current case study was carried out between 2012 and 2016 in a subarea
dominated by open sandy grasslands in the Carpathian Basin. The ~70
ha study area was grazed by 250-300 sheep in 2012. In the beginning
of 2014, the overgrazing pressure was overall reduced, for the most part,
in the examined grassland patches. The study aimed to answer how the
complete abandonment of grazing and moderate grazing influences the
species richness, diversity and density of the orthopteran assemblages. In-
vestigations in Central European sand steppes confirmed that both intense
grazing and the abandonment of grazing have a detrimental effect on the
structure of orthopteran assemblages: (a) the Shannon diversity index was
higher on moderately grazed sites than on grazed and ungrazed ones; (b)
the number of habitat specialists of sandy grasslands was higher on mod-
erately grazed patches than in grazed habitats; and (c) the frequency of
geophilic species was higher on grazed patches than on moderately grazed
and grazing-abandoned ones.

Key words

density, diversity, Hungary, land use intensity, sheep, vegetation structure

Introduction

The structure of habitats and their insect communities exposed
to direct and indirect human impact usually can be considered a
transient state (Pickett et al. 1992). This phenomenon can be well
observed in grazed grasslands where current vegetation structure,
as a background factor to the insect communities, is highly influ-
enced by the intensity, spatial and temporal characteristics and
abandonment of grazing (Carboni et al. 2015, Tork et al. 2016).
Both the vertical and horizontal structure of vegetation, through
the removal of the biomass and through trampling, are changed
by grazing (Asner et al. 2004). Intensive grazing (~overgrazing) in
grasslands usually results in decreased plant diversity, the invasion
of some grazing-tolerant plant species, and the overall degradation

of the habitat-structure (Metera et al. 2010). Nutrient-poor habi-
tats seem to be the most sensitive to grazing intensity (Kruess and
Tscharntke 2002). When the grazing pressure decreases or is aban-
doned in the grasslands of a temperate climate zone, the coverage
of the dominant, narrow-leaved perennial monocotyledons of the
associated plants increases (Critchley et al. 2008), while the cover-
age of annual plant species becomes reduced (Matus et al. 2003).
In addition to the favorable changes for the vegetation-dependent
insect communities, the abandonment of grazing can also cause
the invasion of weeds in the grasslands (Sedlakova and Fiala 2001).
This usually greatly decreases the presence of rare and threatened
Orthoptera (Fonderflick et al. 2014). Therefore, in the habitats
grazed traditionally, the maintenance of extensive grazing is a pre-
requisite for preserving biodiversity (Dolek and Geyer 2002) and
seems to be far more suitable than mowing (Weiss et al. 2013).
The high sensitivity of orthopterans to a change in vegeta-
tion structure is well known (Baldi and Kisbenedek 1997, Sam-
ways 1997, Gardiner et al. 2002, Bazelet and Samways 2011, Weiss
et al. 2013). Based on this, the effect of grazing on orthopteran
assemblages has been the focus of research worldwide and for a
long time (e.g. White 1975, Jepson-Innes and Bock 1989, Prendi-
ni et al. 1996, Kruess and Tscharntke 2002, Jauregui et al. 2008,
Gardiner and Haines 2008, Zhu et al. 2015). Grazing results in
individual, spatially different effects on the orthopterans. Some
studies, for example, revealed lower orthopteran density in grazed
grasslands than in ungrazed ones (Welch et al. 1991, Onsager
2000), while in other cases the opposite was found (Wingerden
et al. 1991). These differences are likely to be due to not only the
habitat-dependent effects of the grazing, but also to species-specif-
ic responses (O'Neill et al. 2003, Jauregui et al. 2008) of orthopter-
ans to intensity (Kruess and Tscharntke 2002, Cease et al. 2012),
timing (Fonderflick et al. 2014) and livestock type (Dolek and
Geyer 2002) of grazing. Most of the results of the assessment of
the impact of grazing suggest that, according to the intermediate-
disturbance hypothesis of Connell (1978), orthopteran assem-
blages show the largest diversity in moderately grazed grasslands
(Batary et al. 2007, Fabriciusova et al. 2011, Jerrentrup et al. 2014).
My study area is situated between the Danube and the Tisza
rivers in the Carpathian Basin in the eastern half of Central-Europe,

JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(1)

24

where the dominant vegetation type is the Pannonian sand steppe
occurring in Europe only in the Pannonian biogeographical region
(6260 Pannonian sand steppes, 2002/83/EC Habitat Directive).
Due to the conditions of the almost humus-less bedrock (sand), the
structure and species composition of the habitats could remain po-
tentially unchanged even for centuries in the absence of any inter-
ventions. However, grazing has been present in the area since, most
likely, the Neolithic (Molnar et al. 2008), so for several thousand
years. Thus, large herbivores played a role in the development of
the actual state of the grasslands (Maté 2014). In the 16-17th cen-
tury, extensive grazing was carried out on a large area of the region
— mainly with cattle and, to a lesser extent, with sheep (Frisnyak et
al. 2015). After the beginning of the 18th century, afforestation was
carried out for the purpose of impeding erosion and for utilizing
areas unsuitable for agricultural use (mainly Pinus sylvestris and Ro-
binia pseudo-acacia were planted). As a result, the extent of areas cov-
ered by forests increased from 3.5% to 60% (Molnar 2003). How-
ever, some hills covered by open sandy grasslands remained, and
until the middle of the 20th century they were grazed by the sheep
of small farmers (Kun 1998). In the past decades and even now,
grazing, especially sheep-grazing, was concentrated in small areas.
The dominance of sheep grazing affected, among other things, the
Fabaceae plant species of the sandstone pastures and their associat-
ed animal species, resulting in the decline or extinction of a number
of plant species at the beginning of the 19th century (Maté 2014).

In addition to the above historical landscape characteristics,
responses of the orthopteran assemblages associated with the typi-
cal habitats of the Central European sand steppes were affected
by several local and global factors. From a local point of view, it
can be said that the orthopteran assemblages of Central European
habitats grazed with different intensity have not yet been suffi-
ciently investigated. Certainly, in regards to the long-term effects
of the various types of grazing, and the long-term effects of the
abandonment of grazing, there are several questions to be an-
swered. Several further questions remain to be answered both lo-
cally and globally with regards to quality and intensity of grazing
being adequate for the most diverse and dense orthopteran assem-
blages (Jerrentrup et al. 2014).

The main questions of the present investigation were the fol-
lowing: 1) How does the complete abandonment of grazing or
moderate grazing influence the species richness, diversity and
density of orthopteran assemblages in grasslands that were heav-
ily overgrazed at the beginning of the study? 2) Is the span of the
study (from 2012 to 2016) enough to detect changes in the struc-
ture of orthopteran assemblages occurring in sandy grasslands
characterized by low plant production? 3) How does the drastic
decrease of grazing pressure impact the density of local popula-
tions of habitat-specific species?

Material and methods

Study area.—The study area is part of the Natura 2000 site Kék-
hegyi lotér (HUKN22037; southern Hungary) (Fig. 1). It is located
at an altitude of ~160 m a.s.l. and is characterized by flat, low
sandy hills and flatlands. The average total duration of annual in-
solation in the region is 2,055 hours. Mean annual temperature is
around 10.6°C. The mean values of absolute maximum and mini-
mum temperatures are 34.7°C and -16.4°C. The average annual
precipitation is 570 mm (330 mm in the growing season) (Dévé-
nyi 2010). The study area is characterized by sandy soils. Based
on analyses of Szilard Szab6 (pers. comm.), mean of the main
soil parameters of the local grasslands are the following: CaCO,

Z. KENYERES

content: 8.5%; humus content: 2.9%; pH-H,O: 7.2; percentages of
the soil fractions: rough: 0.2%; middle-class: 9.7%; small: 80.6%;
mud: 7.1%; loam: 2.3%.

Land use.—Based on the first known detailed map of the study area,
in the 18th century the typical land use of the open sandy grasslands
studied was grazing (http://mapire.eu/hu/map/firstsurvey). In the
19th and 20th centuries some places in the area were afforested,
but the main form of land use on the grasslands was still graz-
ing (see http://mapire.eu/hu/map/secondsurvey; http://mapire.eu/
hu/map/hkf_25e; http://mapire.eu/hu/map/hungary1941). After
World War II the area was used as a closed military base, but the
grazing of sheep continued. When military operations ceased in
1990, the grazing of sheep became increasingly intense. The ~70 ha
study area was grazed by 250-300 sheep in 2012. The overgrazing
pressure was overall reduced in the majority of the study area in the
beginning of 2014 in order to conserve nature.

Experimental design.—Six sampling sites were established, as 50x50
m sized quadrats. Data collection was carried out from 2012 to
2016. One site was on a place ungrazed during the study (Un-
grazed — U-G), three sites were located in places on which grazing
pressure was reduced to zero at the beginning of 2014 (Grazing
Abandoned - G-A), two sites were located in places on which graz-
ing pressure was reduced to a moderate level (Moderately Grazed
—~ M-G) (Fig. 1). Sites were considered to be “grazed” (Grazed - G)
prior to 2014 and the reduction of grazing pressure.

Vegetation and landscape structure.—Measurements of the vegeta-
tion parameters were carried out on 3 plots in each sampling site
of each orthopteran sampling. The following parameters were re-
corded: total vegetation cover (%), average height of the vegetation
(cm), and bare soil (%). Height of the vegetation was measured in
cm with the use of a 30 cm wide and 100 cm high white card. Total
cover of the vegetation was measured in a square meter quadrat
occurring around the spot. Related to each orthopteran sampling,
percentage cover of plant species was estimated. Annual means of
the measured parameters per sampling sites were calculated.

Orthoptera.—During each of the five study years, sampling of the
Orthoptera took place in June, July, and August. In every period, 2
samplings were carried out by sweep-netting on random patches
of each sampling site (altogether 180 samples). Within the 50 x
50 m sized sites, the samplings took place at least 30 m from each
other. Densities were recorded in 10 x 10 m quadrats with 300
sweeps per sample and were completed by direct observations. To
the samples collected by sweep netting I added a simple count
of the number of adult specimens which were detected by direct
observation/collection. Sweep-netted samples were identified to
species level using the works of Harz (1969, 1975). Acrida un-
garica (Herbst), Acrotylus insubricus (Scopoli), Acrotylus longipes
(Charpentier), Calliptamus barbarus (Costa), Celes variabilis (Pal-
las), Dociostaurus brevicollis (Eversmann), Euchorthippus pulvina-
tus (Fischer de Waldheim), Gampsocleis glabra (Herbst), Montana
montana (Kollar), Myrmeleotettix maculatus (Thunberg), Oedaleus
decorus (Germar), Omocestus minutus (Brullé), Platycleis affinis Fie-
ber, Sphingonotus caerulans (Linnaeus), Stenobothrus fischeri (Ever-
smann) were classified as habitat specialists of local sandy grass-
lands. Scientific nomenclature follows Cigliano et al. (2017).
Categories of Uvarov (1977), Ingrisch and Kohler (1998) and
Kristin et al. (2009) were used for classification of life forms (arbori-
col: species found in habitats dominated by tree-sized elements; ar-

JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(1)

Z. KENYERES

25

- iG—>G-A

* .
’ 4 oo

’

Fig. 1. Location and land use-changes of the studied sites (G: grazed; G-A: grazing-abandoned; M-G: moderately grazed; U-G: un-
grazed). Letters to the left of the arrow indicate land use in 2012 and 2013; letters to the right of the arrow indicate land use in 2014,

2015 and 2016.

busticol: species found in habitats dominated by shrub-sized items;
silvicol: species found in forest habitats with a grass understory; pra-
tinicol: species found in grasslands of tall grass; graminicol: species
found in grasslands of short grass; geophilic: species found in grass-
lands characterized by a high percentage of bare soil; psammophilic:
species found in sandy grasslands; pseudo-psammophilic: species
found mainly in sandy grasslands, but also in grasslands occurring
on soil very similar to sand, e.g. dolomite rendzina).
Characterization of climatic requirements of the species as
thermophilic, moderately-thermophilic, mesophilic, moderately-

hygrophilic, and hygrophilic were assigned based on the works of
Varga (1997), Racz (1998), and Ingrisch and Kohler (1998).

Statistical analysis. —Samples collected in the same sampling sites
in the same year were pooled (number of pooled samples was
30: Ungrazed - U-G n = 5; Grazing Abandoned - G-A n = 9;
Moderately Grazed - M-G n = 6; Grazed - Gn = 10). Pooled
samples were used for calculating assemblage variables and sta-
tistical analyses. Shannon diversity index, species richness, den-
sity (individual/10 m2), species number and relative frequencies

JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(1)

26

of habitat specialist species, relative frequencies of geophilic
and graminicol/pratinicol species were calculated and used as
Orthoptera response variables in statistical analyses. Mean val-
ues (4SE) of Orthoptera response variables were calculated for
comparison of structure of orthopteran assemblages exposed
to different grazing pressure. Mann-Whitney U test was used to
evaluate statistical differences among the derived orthopteran
variables. Orthoptera samples were ordered by PCoA (similarity
index: correlation, relative frequency data of species were used
and subtract mean transformed). Generalized linear models
(Poisson distribution; response variables: relative frequencies of
geophilic and graminicol/pratinicol species; predictor variables:
total vegetation cover, average height of the grass, bare soil) were
performed. CCA ordination based on Orthoptera species data
and environmental parameters (total vegetation cover, percent-
age of bare soil, height of the vegetation) were also compiled. All
statistical analyses were performed by using Past 3.14. software
package (Hammer et al. 2001).

Results

Orthoptera diversity and density.—Thirty-five Orthoptera species
(Appendix 1) comprising 2,655 individuals were recorded on 6
sampling sites. The most prevalent species was Calliptamus barbarus
with 580 individuals (24%), followed by Acrida ungarica with 514
individuals (19%), Euchorthippus declivus (Brisout de Barneville)
with 302 individuals (11%), Oedaleus decorus with 279 individu-
als (10%), Oedipoda caerulescens (Linnaeus) with 210 individuals
(8%), Myrmeleotettix maculatus with 185 individuals (7%), Docio-
staurus brevicollis with 117 individuals (4%), Euchorthippus pulvi-
natus with 54 individuals (2%) and Omocestus petraeus (Brisout
de Barneville) with 52 individuals (2%). Shannon diversity, more
sensitive to rare species (Magurran 2004), was highest on mod-
erately grazed sampling sites, and this parameter was lower and
similar to each other on grazed, grazing-abandoned and ungrazed
sampling sites (Mann-Whitney test: U.,,. = 7, Pp = 0.014; Uv ouc =
0, p = 0.008; Fig. 2). Species richness was non-significantly higher
in moderately grazed and grazing-abandoned grasslands, than on
grazed and ungrazed patches (Fig. 2). Overall, Orthoptera density
was higher on the ungrazed patches than on moderately grazed
and grazing-abandoned ones (U,,.4¢ = 1, p = 0.013; Ug, ye = 5) P=
0.023) and non-significantly higher than on grazed ones (Fig. 2).

Composition of Orthoptera assemblages.—PCoA ordination showed
separation of orthopteran assemblages under different grazing
pressure. Sites grouped according to whether they were ungrazed,
grazing-abandoned/moderately-grazed or grazed (Fig. 3). The
number of habitat specialist species was significantly higher on
moderately-grazed patches than on grazed ones (U,,.., = 13.5, p =
0.05) and that parameter was non-significantly higher on grazing-
abandoned and ungrazed patches than on grazed sites (Fig. 4).
The number of habitat specialist species was as high on grazing-
abandoned sites than on the ungrazed site. The frequency of habi-
tat specialist species did not show decided differences related to
grazing pressure (Fig. 4). The frequency of geophilic species was
significantly higher on grazed sites than on moderately-grazed
and grazing-abandoned ones (U,,,, = 9, p = 0.026; U,,, = 15, p=
0.015). Furthermore, the above mentioned parameter was signifi-
cantly higher on ungrazed sites than on moderately-grazed ones
(Uncmc = 4 P = 0.050) (Fig. 5). Parallel to the latter results, the fre-
quency of graminicol/pratinicol species was significantly lower on
grazed sites than on moderately-grazed and grazing-abandoned

Z. KENYERES

a) Species richness
a

No. species

Grazed Moderately Grazing-
grazed abandoned

Ungrazed

b) Shannon overly

-

Diversity

Grazed Moderately Grazing-
grazed abandoned

Ungrazed

Density (individuals/10m7)

Grazed Moderately Grazing-

Ungrazed

grazed abandoned

Fig. 2. Mean values (min-max and +SE) of main parameters of
orthopteran assemblages under different grazing pressure. Signifi-
cant (p<0.05) differences detected by Mann-Whitney U test are
indicated by different letters.

JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(1)

Z. KENYERES

Oca

®., Ungrazed
@c.p

Coordinate ?

27

Grazed

Coordinate 1

Fig. 3. PCoA ordination (sum of all eigenvalues: 20.008, similarity index: correlation) based on Orthoptera data. The different years are
marked by A, B, C, D and E (A: 2012, B: 2013, C: 2014, D: 2015 and E: 2016; e.g. 1-A: site 1 in 2012, 2-B: site 2 in 2013, C-C: control

site in 2014).

Table 1. Main vegetation characteristics of the sampling sites (mean values (+SE) of the measured data in June, July and August).

Grazed Grazing-abandoned Moderately grazed Ungrazed

(June-August) (June-August) (June-August) (June-August)
Vegetation height (cm) 3.340.4 11.741.3 1:8. 32525 26.0+1.8
Vegetation cover (%) 44.0+4.8 78.841.8 82.541.1 72.043.4

ones (U,,,, = 9, p = 0.026; U,.,, = 15, p = 0.015) and the relative
frequency of graminicol/pratinicol species was significantly lower
on ungrazed sites than on moderately grazed sites (Uc. = 4 P
= 0.050) (Fig. 5).

Effects of environmental parameters.—Based on the results of gen-
eralized linear models, total vegetation cover (VCOV), vegetation
height (VH) and percentage of bare soil (BSOIL) were found as
significant predictors of the frequency of geophilic species and,
parallel to this, the frequency of graminicol/pratinicol species
(VCOV/Geo_freq: -0.0055; SE: 0.125; p = 0.002; VCOV/Gra_prat_
freq: 0.0055; SE: 0.018; p = 0.002; VH/Geo_freq: -0.0097; SE:
0.003; p = 0.012; VH/Gra_prat_freq: 0.0097; SE: 0.003; p = 0.012;
BSOIL/Geo_freq: 0.0056; SE: 0.001; p = 0.001; BSOIL/Gra_prat_
freq: -0.0056; SE: 0.001; p = 0.001). Total vegetation cover (VCOV)
and vegetation height (VH) (Table 1, Fig. 6) were negatively re-
lated to the frequency of geophilic species, while the percentage of
bare soil (BSOIL) was positively related. The total vegetation cover
(VCOV) and vegetation height (VH) were positively related to the

frequency of graminicol/pratinicol species, and the percentage of
bare soil (BSOIL) was negatively related.

Three predictor variables contributed significantly to the CCA
ordination. Relative frequency of geophilic species, such as Acrida
ungarica, Acrotylus insubricus, Aiolopus thalassinus (Fabricius), Cal-
liptamus barbarus, Calliptamus italicus (Linnaeus), Celes variabilis,
Oedaleus decorus, and Oedipoda caerulescens, was positively corre-
lated with a high percentage of bare soil (BSOIL) (Fig. 7). On the
other hand, relative frequency of graminicol/pratinicol species,
such as Gampsocleis glabra, Montana montana, Platycleis albopuncta-
ta (Goeze), Stenobothrus fischeri, and Stenobothrus lineatus (Panzer)
was positively related to a high total cover of vegetation and verti-
cally structured grasslands (Fig. 7).

Discussion
Grazing intensity or abandonment of grazing has a detrimen-

tal effect on the structure of orthopteran assemblages (Kruess and
Tscharntke 2002, WallisDeVries et al. 2007, Eschen et al. 2012).

JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(1)

28

10

No. species
o)
ia)

Grazed Moderately Ungrazed

grazed

Grazing-
abandoned

Frequency (%)

Grazed

Moderately Ungrazed

grazed

Grazing-
abandoned

Fig. 4. Mean values (min-max and +SE) of species number and
frequency of habitat specialist species under different grazing pres-
sure. Significant (p<0.05) differences detected by Mann-Whitney
U test are indicated by different letters.

Studied grasshopper assemblages of the Central-European sand
grasslands showed the greatest diversity on patches affected by ex-
tensive grazing (Quinn and Walgenbach 1990, Enyedi et al. 2008).
This result is similar to those of Batary et al. (2007) and Fabriciusova
et al. (2011) from other grassland types of the Carpathian Basin. In
addition to the species with a positive correlation to open soil surfac-
es, these patches provide for the conservation of xero-thermophilic
habitat specialists (Fonderflick et al. 2014) related to heterogene-
ous vegetation (Batary et al. 2007) such as the European Red-Listed
Gampsocleis glabra and Montana montana (Hochkirch et al. 2016).
Extensive grazing therefore results in higher species richness of habi-
tat specialist orthopterans in the moderately grazed patches than in
grazed ones (Fig. 4). Overall density of the orthopteran assemblages

Z. KENYERES

a) Geophilic species

a

=
o
c
oO
=)
oy
2
LL
Grazed Moderately § Grazing- Ungrazed
a8 grazed abandoned
, b) Graminicol and pratinicol species
b
be
3 a zl ac
: =
c
®
=)
oO
2
LL

Grazing-
abandoned

Grazed Moderately

grazed

Ungrazed

Fig. 5. Mean values (min-max and +SE) of relative frequency of
geophilic and vegetation structure-dependent species under differ-
ent grazing pressure. Significant (p<0.05) differences detected by
Mann-Whitney U test are indicated by different letters.

was high on several grazed patches. This is probably explained by
the species-specific response of the orthopterans to grazing (O'Neill
et al. 2003, Jauregui et al. 2008). For example, Chorthippus biguttu-
lus (Linnaeus) (Fonderflick et al. 2014) and Stenobothrus stigmaticus
(Rambur) (Jauregui et al. 2008) reach high density in intensively
grazed grasslands. In the present study, Acrida ungarica, Calliptamus
barbarus and Oedaleus decorus have proved to be species preferring
habitat-structure transformed by grazing. The latter species, ac-
cording to their energy requirements (Fielding and Brusven 1995),
reached much higher abundance on the grazed patches character-
ized by short vegetation and fragmented by open sandy surfaces,
than on the grazing-abandoned habitats. The decline in prevalence
of the geophilic lifestyle due to the abandonment of grazing (Fig.

JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(1)

Z. KENYERES

a) Vegetation cover

Vegetation cover (%)

Grazed

Moderately Grazing-
grazed abandoned

Ungrazed

b) Vegetation height

bc

Vegetation height (cm)

Grazed

Moderately Grazing-
grazed abandoned

Ungrazed

Fig. 6. Mean values (min-max and +SE) of vegetation cover and veg-
etation height on the studied sites. Significant (p<0.05) differences
detected by Mann-Whitney U test are indicated by different letters.

5) has also been proven by revealing a sharp difference between the
habitat requirements of species related to open patches with short
grasses and to closed patches with structured spatial composition
(Fig. 7). The fact that the density of pratinicol and graminicol spe-
cies, as an inverse trend, was significantly low until the abandon-
ment of grazing, is explained by the fact that the vegetation structure
was simplified due to the effect of grazing, which is unfavourable
for pratinicol and graminicol species in terms of a nutritional base,
microclimate, egg-laying possibilities and exposure to predators
(Kruess and Tscharntke 2002, Gardiner and Haines 2008).
Following from the results and suggestions by Jerrentrup et al.
(2014) and Joubert et al. (2016), the conservation of species rich-

29

ness and diversity of the studied orthopteran assemblages can best
be ensured by moderate grazing. The definition of moderate vs.
extensive grazing is not possible in general, but only in relation to
the given habitats. Jerrentrup et al. (2014) estimated that grazing
intensity of ~ 1 Livestock Units/ha can still be considered moder-
ate, resulting in habitat structure rich in microhabitats that ensure
the presence of diverse orthopteran assemblages. However, the as-
semblages of dry steppe grasslands are more sensitive to grazing
pressure. According to the results of Fonderflick et al. (2014), in
the case of extensive grazing of dry steppe grasslands with sheep
(0.24 Livestock Units/ha), the abundance of orthopterans is sig-
nificantly higher in the ungrazed spots than in the post-grazed ar-
eas. According to Gardiner and Haines (2008), in the case of graz-
ing by horses, reduction of intensity from 3.5 horse/ha to 2 horse/
ha can lead to an increase in the diversity and abundance of the
orthopteran assemblage. The grasslands examined in the current
study belong to the extremely low grass-producing habitats that
are highly sensitive to treading. Thus, for the largest species rich-
ness, diversity and density of the local orthopteran assemblages,
the extensive grazing system proposed by Maté (2014) should be
used: 1) Grazing pressure should be set between 0.1-0.2 Livestock
Units/ha; 2) Ungrazed patches should be left in the grasslands
concerned each year. The desirable extent of the latter should be
set as 10-50% of the grazed parcels. The greater the precipitation
in a year, the smaller the percentage of the grasslands that has to be
spared from grazing. Ungrazed areas should be designated at sev-
eral isolated spots, and the grazed and ungrazed patches should be
changed from year to year (on large pastures designated subareas
should be used in every 1st, 3rd, 6th, and 10th year); 3) the land
use must be controlled by the shepherd, ensuring the spread of the
grazing animals.

In the examined habitats the annual grazing schedule is also
important. In this respect, it has to be taken into account that the
changing of vegetation structure affects orthopterans the most dra-
matically in the period when the number of adults in the assem-
blages is at its highest (August in Europe) (Fonderflick et al. 2014).
During this period, the concentration of grazing on smaller, less
natural, weedy patches is also acceptable in order to protect the
valuable vegetation patches and their orthopteran assemblages.
The conservation potential is strongly influenced by habitat con-
ditions (Weiss et al. 2013). This is particularly true for habitats as
sensitive as the Central European sandy steppes, where the grazing
method should always be chosen by taking the weather conditions
into consideration: during dry periods grazing must be moder-
ated, or abandoned completely on the patches most sensitive to
treading (Maté 2014).

On a global or historical scale, it is odd that we consider the
development and use of different grazing systems for habitats that
have been under continuous grazing pressure for hundreds or
thousands of years, in order to preserve their biodiversity. This is
not unjustified, however, given that from the end of the nineteenth
century, in a significant part of Europe, habitats resulting from
moderately disturbing and selective effects of extensive land man-
agement have disappeared, degraded and fragmented to the great-
est extent (Bakker and Berendse 1999). As a result, the remaining
habitat fragments became biodiversity hotspots (Steffan-Dewenter
and Tscharntke 2002), the possible disappearance of which would
be an irreversible loss. The preservation of the latter must there-
fore be given priority. As demonstrated by the present case study,
changing the grazing patterns toward nature conservation-based
land use can have positive results in terms of the protection of di-
verse orthopteran assemblages. However, for successful conserva-

JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(1)

30

@Ste lin

. VCOV
@ Ste fis Myr mace

Axis 2

Graminicol and pratinicol

@Omo min

Omo ruf -5

Z. KENYERES

Geophilic °Cal ita

species

e
BSOIL Oed dec

®
Ail tha

@ @
Agr ung Of caeAcr ins®

Cel var

: -2 e
@ :
Species Pla alb Ste nig
. e@ Gam gla
Pla mon -3
Cd]
Cho bru

-4

Fig. 7. CCA ordination based on Orthoptera data and environmental parameters (VCOV: total vegetation cover; BSOIL: percentage of
bare soil; VH: height of the vegetation). Abbreviations of species names: Acr ins: Acrotylus insubricus; Act ung: Acrida ungarica; Ail tha:
Aiolopus thalassinus; Cal bar: Calliptamus barbarus; Cal ita: Calliptamus italicus; Cel var: Celes variabilis; Cho apr: Chorthippus apricarius;
Cho big: Chorthippus biguttulus; Cho bru: Chorthippus brunneus; Cho mol: Chorthippus mollis; Doc bre: Dociostaurus brevicollis; Euc dec:
Euchorthippus declivus; Euc pul: Euchorthippus pulvinatus; Gam gla: Gampsocleis glabra; Mon mon: Montana montana; Myr mac: Myrmele-
otettix maculatus; Oed cae: Oedipoda caerulescens; Oed dec: Oedaleus decorus; Omo hae: Omocestus haemorrhoidalis; Omo min: Omocestus
minutus; Omo pet: Omocestus petraeus; Omo ruf: Omocestus rufipes; Pla alb: Platycleis albopunctata; Sph cae: Sphingonotus caerulans; Ste fis:
Stenobothrus fischeri; Ste lin: Stenobothrus lineatus; Ste nig: Stenobothrus nigromaculatus.

tion strategies for sensitive communities requiring moderate dis-
turbance, it is important to conduct further, preferably long-term,
studies on the response of orthopteran assemblages as bioindica-
tors (Bazelet and Samways 2011) to the direct and indirect effects
of grazing systems linked to fragmented habitats.

Acknowledgements

The author would like to express his gratitude to Csaba Szine-
tar and to Kiskunsag National Park Directorate for the assistance
during the research. I thank both reviewers for their remarks. Great
thanks from the author go to Corinna S. Bazelet, managing editor
of JOR, for her work with the manuscript and to Nancy Morris, for
her help in repairing stylistic issues of the text.

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Species composition and abundance of the pooled samples of different grazing pressure (LF: life form; EF: ecotype form; G: grazed,
MG: moderately grazed, GA: grazing-abandoned, UG: ungrazed; arbu: arbusticol; geo: geophilic; gra: graminicol; pra: pratinicol; ps:
psammophilic; psps: pseudo-psammophilic; mes: mesophilic; m-ther: moderately-thermophilic; ther: thermophilic)

Taxon
Caelifera
Acridoidea
Acridomorpha
Acrididae
Acridinae
Acrida ungarica (Herbst, 1786)
Calliptaminae
Calliptamus barbarus (Costa, 1836)
Calliptamus italicus (Linnaeus, 1758)
Gomphocerinae
Euchorthippus declivus (Brisout de Barneville, 1848)
Euchorthippus pulvinatus (Fischer de Waldheim, 1846)
Euthystira brachyptera (Ocskay, 1826)
Dociostaurus brevicollis (Eversmann, 1848)
Chorthippus apricarius (Linnaeus, 1758)
Chorthippus biguttulus (Linnaeus, 1758)
Chorthippus brunneus (Thunberg, 1815)
Chorthippus dichrous (Eversmann, 1859)
Chorthippus mollis (Charpentier, 1825)
Myrmeleotettix maculatus (Thunberg, 1815)
Pseudochorthippus parallelus (Zetterstedt, 1821)
Omocestus haemorrhoidalis (Charpentier, 1825)
Omocestus minutus (Brullé, 1832)
Omocestus petraeus (Brisout de Barneville, 1856)
Omocestus rufipes (Zetterstedt, 1821)
Stenobothrus fischeri (Eversmann, 1848)
Stenobothrus lineatus (Panzer, 1796)
Stenobothrus nigromaculatus (Herrich-Schaffer, 1840)
Stenobothrus stigmaticus (Rambur, 1838)

LF EF G MG GA uG
psps ther 172 186 77 fa)
ps ther 203 116 69 241
gra ther 6 0 4 0
gra ther 18 89 60 135
gra ther 0 8 18 28
pra mes 2 0
psps ther 32 48 15 22.
pra mes 1 1 0
pra m-ther 6 12 0
pra m-ther 10 17 11 iL
pra mes 0 0 6 0
pra mes 11 11 13 0
gra ther 22 99 37 27
pra mes 0 0 0
pra ther 1 0
psps ther 4 0 0
gra ther 30 16 4 2
pra mes 1 3 1 0
pra ther 0 9 a3 2
pra m-ther 0 0 0
gra ther 31 0 0
pra m-ther il 0 0 0

JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(1)

Z. KENYERES
Taxon LF
Oedipodinae
Acrotylus insubricus (Scopoli, 1786) ps
Acrotylus longipes (Charpentier, 1845) psps
Aiolopus thalassinus (Fabricius, 1781) gra
Celes variabilis (Pallas, 1771) gra
Oedaleus decorus (Germar, 1826) psps
Oedipoda caerulescens (Linnaeus, 1758) geo
Sphingonotus caerulans (Linnaeus, 1767) psps
Pezotettiginae
Pezotettix giornae (Rossi, 1794) gra
Ensifera
Tettigonioidea
Tettigoniidae
Tettigoniinae
Gampsocleis glabra (Herbst, 1786) psps
Montana montana (Kollar, 1833) psps
Platycleis affinis Fieber, 1853 psps
Platycleis albopunctata (Goeze, 1778) pra
Phaneropteridae
Phaneropterinae
Leptophyes albovittata (Kollar, 1833) arbu

EF

ther
ther

m-ther

ther
ther
ther
ther

ther

ther
ther
ther
ther

ther

JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(1)

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