Commentary

Journal of Orthoptera Research 2022, 31(2): 163-172

Orthoptera in the early stages of post-arable rewilding in south-east England

Tim GARDINER!, DOROTHY CASEY?

1 Fisheries, Biodiversity and Geomorphology, Environment Agency, Iceni House, Cobham Road, Ipswich, Suffolk, IP3 9JD, UK.
2 Former Head of Conservation c/o Suffolk Wildlife Trust, Brooke House, Ashbocking, Ipswich, IP6 9JY, UK.

Corresponding author: Tim Gardiner (tim.gardiner@ environment-agency.gov.uk)

Academic editor: Michel Lecoq | Received 15 February 2022 | Accepted 4 April 2022 | Published 30 September 2022

https://zoobank. org/DA579A7 D-6313-4EB2-ADF1-E7BC3B03EA31

Citation: Gardiner T, Casey D (2022) Orthoptera in the early stages of post-arable rewilding in south-east England. Journal of Orthoptera Research

31(2): 163-172. https://doi.org/10.3897/jor.31.82317

Abstract

The ideal aim of rewilding is to restore natural processes to create ‘self-
willed’ ecosystems involving the creation of large areas of habitat subject
to stochastic disturbance, connected by favorable corridors for species to
disperse along. Reversion of arable farmland to grassland and scrub habi-
tats on Black Bourn Valley nature reserve in Suffolk (south-east England)
through non-intervention allowed succession to occur largely unman-
aged. Fields in the early stages of rewilding (4-14 years) are found at Black
Bourn Valley, while pond creation has been extensive since 2010, creat-
ing water edge habitat and heterogeneity to the re-establishing grassland.
Monitoring of Orthoptera revealed statistical evidence that species diver-
sity/richness and field grasshopper Chorthippus brunneus (Thunberg, 1815),
meadow grasshopper Pseudochorthippus parallelus (Zetterstedt, 1821), com-
mon groundhopper Tetrix undulata (Sowerby, 1806) and slender ground-
hopper Tetrix subulata (Linnaeus, 1758) were in higher abundance in fields
>8 years since arable cropping ceased compared to those 4 years post re-
version. Fields =8 years old were probably favorable due to the presence of
microhabitats for basking and egg-laying orthopterans that included ant
hills, sparsely vegetated pond edge and open swards with an abundance of
fine-leaved grasses (Agrostis and Festuca spp.) and a low abundance of leaf
litter. Lagomorph grazing by wild brown hare Lepus europaeus and rabbit
Oryctoloagus cuniculus was critical in maintaining exposed soil for Orthop-
tera in the older fields, while deer paths appeared to create microhabitats
that may be utilized by Orthoptera. We postulate that rewilding schemes
on arable farmland should use a Rewilding Max approach and avoid the
frequent usage of domestic livestock, relying on wild lagomorph and un-
gulate grazers to maintain an open mosaic habitat structure with only in-
termittent cattle, horse, or sheep grazing.

Keywords

Acrididae, biodiversity, bush-cricket, conservation, deer, grasshopper,
Tettigoniidae, ungulate, wilding

Introduction

Orthoptera form an important part of grassland ecosystems
across Europe (Kohler et al. 1987, Ingrisch and Kohler 1998,
Humbert et al. 2009). Orders such as Orthoptera (grasshoppers,

bush-crickets, and crickets) are an important component of in-
vertebrate assemblages in agricultural ecosystems, particularly as
prey for bird and spider species (Joern 1986, Belovsky and Slade
1993, Oedekoven and Joern 1998). Gardiner et al. (2002) suggest
that intensively managed farmland habitats such as arable fields,
heavily grazed improved pastures, and hay meadows have a low
abundance of orthopteran species such as meadow grasshopper
Pseudochorthippus parallelus (Zetterstedt, 1821) and field grasshop-
per Chorthippus brunneus (Thunberg, 1815). To reverse the decline
of insects such as grasshoppers and bush crickets, rewilding of ar-
able farmland may be highly beneficial.

The term rewilding was first used in North America in the
1980s (Noss 1985). Soulé and Noss (1998) proposed three key
components of rewilding: large core protected areas, ecological
connectivity, and keystone species that translated to the 3Cs of
cores, corridors, and carnivores. Over time, Soulé and Noss’s origi-
nal concept has shifted into local interpretations but still incorpo-
rates self-regulatory ecosystems with minimal or no anthropogenic
influence where wild grazers have a critical role (Dempsey 2021).

The role of wild herbivore grazers, such as lagomorphs (e.g.,
rabbit Oryctoloagus cuniculus) and ungulates (e.g., deer), in manag-
ing rewilded grasslands has not been studied in any depth with
the aim of rewilding to recreate ‘natural’ ecosystems (Gordon et
al. 2021b). Deer laydown areas and paths represent variation in
sward height and could therefore be an influence on sward struc-
ture important for Orthoptera. Mixed approaches (e.g., relying on
natural processes and introducing domestic grazers) with only
minimal conservation intervention can be successful and may be
necessary in some situations where non-intervention is not suit-
able (e.g., urban edge).

In recent times, the aim of rewilding in the UK has focused on
restoring natural processes by creating large areas of habitat sub-
ject to stochastic disturbance connected by favorable corridors for
species to disperse along (Carver and Convery 2021, Gordon et
al. 2021a, b). Such ecological restoration on agricultural land that
allows habitats to regenerate with a lack of active farmland (e.g.,
fertiliser application) or conservation management such as con-
trolled livestock grazing is known as Rewilding Max (Gordon et al.

JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(2)

164

2021a,b). Domestic livestock (cattle, sheep, and ponies) are often
used to graze rewilded farmland sites (e.g., Knepp Wildland in West
Sussex, UK (Dempsey (2021)) after the initial establishment phase
and grassland re-establishment (Casey et al. 2020). This form of
conservation intervention without sole reliance on natural grazers
is known as Rewilding Lite (Gordon et al. 2021b). However, intro-
duced livestock can have detrimental impacts on Orthoptera where
stocking density is too high and resultant sward height too short
(Gardiner and Haines 2008). Across Europe, homogenously short
swards established by overgrazing is the greatest threat to Orthop-
tera (affecting 262 species; Hochkirch et al. 2016). The consequenc-
es of livestock grazing are largely influenced by the intensity of graz-
ing, type of grazer, and rotational or seasonal aspects of the regime,
which in turn have an impact on characteristics of grasslands, such
as leaf litter development, plant species presence, sward height, and
vegetation biomass (Marini et al. 2008, Fabriciusova et al. 2011,
Fonderflick et al. 2014, Rada et al. 2014, Kurtogullari et al. 2020).

The abundance of Orthoptera, particularly grasshoppers, is
strongly influenced by sward height, biomass, and the composi-
tion and the availability of bare earth (Clarke 1948, Gardiner et al.
2002). Increased herbage biomass and sward height through aban-
donment of grassland management on rewilded sites could lead
to a concomitant decrease in sward temperatures and extinction of
light near the soil surface, particularly where leaf litter is allowed to
develop (Gardiner et al. 2005). This is an important factor in Or-
thoptera distribution, as Van Wingerden et al. (1992) suggest that
the number of grasshopper species is reduced in such ‘cold’ grass-
lands due to slow egg development and delayed hatching, particu-
larly where there is a dense leaf litter layer (Gardiner et al. 2005). In
these tall and dense grasslands, the availability of bare earth for the
basking and oviposition needs of Orthoptera is often provided by
ant hills where eggs are laid and nymphs are found in spring (Rich-
ards and Waloff 1954). In European grasslands on abandoned ar-
able land, some authors (e.g., King 2006, 2020, 2021) consider the
yellow-meadow ant Lasius flavus to be a keystone species due to the
bare earth habitat provided by its large mounds of earth.

Orthoptera are ideal for monitoring the effects of rewilding
at sites due to established, easily repeated monitoring methods,
speed of response to habitat change, and range of species with dif-
fering habitat requirements (e.g., bare earth, short grass, tall grass,
and scrub species) (Gardiner et al. 2005). Using a simple acoustic
and visual transect methodology accompanied by habitat meas-
urements (e.g., sward height and bare earth), we would expect
there to be a strong species-specific response to rewilding features
(e.g., ant hills) and wild grazing. For example, C. brunneus may re-
quire the bare earth of ant hills for basking and oviposition, while
common green grasshopper Omocestus viridulus (Linnaeus, 1758)
is a species of tall grassland that may benefit from an absence of
lagomorph grazing (Marshall and Haes 1988).

The aim of this paper is to report on a study of the orthopteran
assemblage of rewilded grassland on former arable farmland in
Black Bourn Valley in Suffolk, UK. Results are discussed in relation
to natural grazing pressure (brown hare, rabbit, and deer), grassland
age since reversion commenced, and other factors such as sward
height and bare earth habitat provided by ant hills and pond banks.

Materials and methods

Study site.—Black Bourn Valley has been owned and managed by
Suffolk Wildlife Trust (SWT) since 1995 (it was formerly known as
Grove Farm) in Suffolk, UK (52°15'0.4644"N, 0°51'1.422"E). The
reserve is 119 ha in area and was previously intensively cropped

T. GARDINER, D. CASEY

for agriculture with nitrogen (N) fertilizer applied to a range of
annual crops, including winter wheat. The Black Bourn River runs
along the eastern edge of the reserve and is adjoined by riverside
meadows (outside the scope of this study). The soil is a lime-rich
loam and clay with slightly impeded drainage and moderate fertil-
ity. The farm had 11 farm ponds extant when SWT acquired the
site in 1995. Nine new ponds have been created since 2010 as part
of the Freshwater Habitats Trust’s (FHT) Million Ponds Project.
These ponds have several notably scarce and rare plants (e.g., tas-
sel stonewort Tolypella intricata) for which the reserve has been des-
ignated a Flagship Pond Site by the FHT.

A total of eight fields were selected for this study due to their
differing ages since reversion. Five fields were last plowed and
cropped in 2017, two fields were last cropped in 2013, and one
was cropped in 2007 (Fig. 1, Table 1). Once cropping ceased,
all fields were allowed to naturally revert to grassland and scrub
through succession with minimal intervention apart from mow-
ing of footpaths and occasional light grazing (not on an annual
basis). Throughout the duration of the 2021 summer study period,
there was no active conservation management of any of the fields.
A green lane ran through the site, and most fields were surrounded
by scrubs or dense hedgerows. There were 10 ponds in the fields
last cropped in 2013 or earlier (combined pond area 953 m7’)
and six in the fields last cropped in 2017 (combined pond area
562 m?’). Most of the ponds had gently shelving banks (25-50°
angle) and were sparsely vegetated.

Transect surveys.—A 1-m wide x 400-m long transect was estab-
lished in all eight fields. Transects were arranged in a W shape
(each arm 100 m) to ensure even coverage of each field and avoid
any edge habitat effects. The transect method closely followed the
methodology of Gardiner et al. (2005), Gardiner and Hill (2006),
and Gardiner (2021). Each transect was walked at a slow, strolling
pace (2 km/hr) on 3 occasions from May-August 2021. Adult indi-
viduals of all Orthoptera species along all transects were recorded
acoustically and visually to determine assemblage composition
and species diversity and richness.

Nymphs flushed from a 1-m wide band in front of the observer
were recorded along all transects. As it is difficult to distinguish be-
tween species in the early instars (though not impossible, see Thom-
men 2021), numbers for nymphs of all species were lumped together
for recording purposes. The surveys were undertaken in vegetation
sufficiently short (<50 cm) to minimize the possibility of overlook-
ing nymphs in tall grass or non-stridulating species such as ground-
hoppers (Tetrigidae) (Gardiner et al. 2005). With practice, it was
relatively easy to ascertain the species of adults without capture (Gar-
diner and Hill 2006), although some species, such as long-winged
conehead Conocephalus fuscus (Fabricius, 1793) and Roesel’s bush-
cricket Roeseliana roeselii (Hagenbach, 1822), are significantly under-
recorded using visual transects (Gardiner and Hill 2006). A dual
visual and acoustic monitoring method has been used by Weiss et al.
(2013) to ensure complete coverage of the orthopteran fauna of sites.
In the current study, a stridulation monitoring technique was used to
record adult males of species that stridulated along the transects at
the same time as visual monitoring by flushing. Stridulation moni-
toring has been used to record cryptic species in Essex and has been
found to be effective compared to visual sighting transects and pitfall
traps (Harvey and Gardiner 2006, Gardiner et al. 2010). Bat detectors
were not required in the current study as the first author (TG) was
able to reliably detect stridulating males up to 20 m away either side
of the transect. The weather conditions on survey days were favorable
for insect activity, being largely sunny and warm (>17°C).

JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(2)

T. GARDINER, D. CASEY

Black Bourn River

‘
Secs 4, Valley meadows
\
.

“Further ee
Butchers

oe.
Further “*--..,

eoee tent

Kiln Ground ye. ze

Green lane

100
. Slades

e CC eee
Pond

Railway line

Fig. 1. Layout of the eight experimental fields at Black Bourn
Valley with ponds and corridor habitats highlighted.

Table 1. Characteristics of the eight fields that had been taken out
of arable cropping.

Field name Area Year since Habitat No.ponds Pond areas
(ha) cropping on field (m7)
ceased perimeter

Miller's Field 2.0 2007 H, GL 3 50, 80, 260
Further Thurston Pil 2013 H, GL, M, W 4 80, 72, 98, 138
Kiln Meadow 1.6 2013 H, GL, W 3 32, 50, 140
Tostock Ley 4.4 2017 H 2 85, 90
Kiln Ground 14.7 2017 H, GL, M 2 35,50
First Butchers 3.0 2017 H, GL 2 125, 130
Further Butchers 5.4 2017 H 0
Slades 5:7 2017 H, GL, R 0

Key: H = hedge, GL = green lane, M = marsh, R = railway bank, W = woodland

Natural grazing pressure and habitat characteristics surveys.—A total
of 40 sward heights were recorded at random positions along the
Orthoptera transects using a 1-m rule for each of the eight fields in
early September 2021. In each field, ant hills were counted along
the 400 m long Orthoptera transects in a 1-m wide band; addi-
tionally, the number of individual wild lagomorph (differences
between brown hare and rabbit were not determined, so drop-
pings pooled for both species) droppings (dung balls) were re-
corded in the same 1-m band to ascertain the level of grazing pres-
sure in the fields (Wood 1988, Gibb and Fitzgerald 1998, Millett
and Edmondson 2013).

Four of the six British ungulate species have been recorded at
Black Bourn Valley: fallow deer Dama dama, muntjac Muntiacus
reevesi, red deer Cervus elaphus, and roe deer Capreolus capreolus.
Due to the known presence of ungulates on site, individual deer
pellets (not classified to species) were counted along a 5-m wide
x 200-m long transect in each field in October 2021 when the
vegetation had died back, allowing easy sighting of fecal matter.
The counting of deer droppings (droppings of all species lumped
together for analysis) followed the methods of Marques et al.
(2001). In addition to the dropping counts, deer laydown areas
(flattened areas of grass at least 2 x 2-m in area) and deer paths
(parted grass through vegetation) were recorded where they were
observed within the 5 x 200-m transect in each field.

In the fields where arable cropping ceased in 2017 and in 2013
or 2007, 2 x 2-m quadrats were randomly surveyed (11 and 7

165

quadrats, respectively) to collect vegetation data. The number of
quadrats surveyed in both field types was approximately propor-
tionate to the number of fields studied in each (i.e., 5/8 fields for
2013 or 2007 = 63%, 11/18 quadrats = 61%). Each plant species
recorded in a quadrat, along with bare earth and leaf litter (dead
and decaying vegetative material, typically grass in this study),
was given a DOMIN value in accordance with National Vegeta-
tion Classification survey methods (Rodwell 2006). The DOMIN
scale assigns percentage cover of species or habitat features to a
numeric grade using the following scale: 1, <4% few individuals;
2, <4% several individuals; 3, <4% many individuals; 4, 4-10%;
5, 11-25%; 6, 26-33%; 7, 34-50%; 8, 51-75%; 9, 76-90%; 10,
91-100% (Rodwell 2006).

Statistical analysis

Orthoptera.—For data analysis, two age classes for the fields were
used: 1) fields where arable cropping ceased in 2017 (5 fields) or
2) in 2013 or 2007 (3 fields). This represented either 4-year-old
reversion or >8 years, since arable cropping terminated. All detec-
tions of Orthoptera (visual or acoustic) were summed for each
field for the survey period (3 surveys) to determine the relative
abundance of adults of each species and nymphs in accordance
with previous studies (notably, Weiss et al. 2013). Independence
of transects was assumed, and data was pooled for each one in a
similar way to data analysis in other monitoring studies (Nur et
al. 1999).

Species richness was calculated for each field. Assemblage di-
versity estimates were also calculated using Version 4.1.2. Species
Diversity and Richness software (Pisces Conservation Ltd, IRC
House, The Square, Pennington, Lymington, Hampshire) from
data collated from each of the two methods. The Shannon-Wiener
Diversity Index (H’, Kent and Coker 1992) was calculated using
the total number of adult individuals recorded for each Orthop-
tera species in each field. All data were square-root transformed to
correct for non-normality before analysis (Heath 1995).

To determine whether species richness and diversity and the
abundance of adults (of all species) and nymphs differed be-
tween fields 4 years and 8 or more years since arable reversion
began, a Student's t-test was used for all comparisons. The mean
abundance of the most abundant species/field was also compared
between the two field types. Corrections were made for unequal
variance where necessary using Satterthwaite’s approximate t-test,
which is a method in the Behrens-Welch family (Armitage and
Berry 1994, Heath 1995). Significance was accepted as evidence
based on the following scale in accordance with Muff et al. in
press: p-value >0.1, little or no evidence; 0.05-0.1, weak evidence;
<0.05, moderate evidence; <0.01, strong evidence; or <0.001, very
strong evidence.

Natural grazing pressure and habitat characteristics surveys.—The
counts of ant hills, lagomorph droppings (brown hare and rab-
bit pooled), deer droppings (all species combined), laydown
areas, and paths were individually summed for each field, and
along with DOMIN values for bare earth, leaf litter, the five most
abundant plant species, bryophytes and plant species richness,
data were square-root transformed to correct for non-normality
(Heath 1995). To determine whether all of these variables differed
between fields 4 years old and those 8 or more years post-arable
reversion, a Student's t-test was used in each case with corrections
made for unequal variance where necessary using Satterthwaite’s
approximate t-test (Armitage and Berry 1994, Heath 1995).

JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(2)

166

Results

Nine species of Orthoptera were recorded on the Black
Bourn Valley reserve, including three species uncommon in this
area of Suffolk (Table 2). The most commonly recorded species
were P. parallelus (21% of adult detections) and R. roeselii (20%),
followed by C. fuscus (19%) and C. brunneus (12%). Less common
species included O. viridulus and slender groundhopper Tetrix
subulata (Linnaeus, 1758) (10% each). The common groundhopper
Tetrix undulata (Sowerby, 1806) and lesser marsh grasshopper
Chorthippus albomarginatus (De Geer, 1773) were found in much
lower abundance (3%), while the dark bush cricket Pholidoptera
griseoaptera (De Geer, 1773) was a rarity (1%).

The abundance of 6 of the 9 species (C. albomarginatus,
C. brunneus, O. viridulus, P. parallelus, T. subulata, and T: undulata)
was highest in one field (Kiln Meadow, cropping ceased 2013; Figs
1, 7) where 30% of the total detections of adult Orthoptera were
recorded. For O. viridulus, T: subulata, and T: undulata, a very high
percentage of the total recorded adults for each species were from
Kiln Meadow and Further Thurston combined (82%, 89%, and
73%, respectively). For P. griseoaptera, 83% of adults were recorded
from Miller's Field (cropping ceased in 2007).

There was very strong evidence (p < 0.001) that species richness
and diversity and the abundance of T. undulata were significantly high-
er in fields >8 years since cropping cessation compared to those only
4 years since cropping ceased (Table 2). There was moderate evidence
(p < 0.05) that C. brunneus and P. parallelus were significantly more
abundant in fields =>8 years old and only weak evidence (p < 0.10)
for T. subulata (Table 2). Very strong evidence (p < 0.001) indicated
that the most abundant species (% of total number) in each field was
significantly higher in the younger fields (c. 43%) compared to the
older fields (c. 22%). In 4 out of the 5 fields 4 years post-cropping,
C. fuscus was the most abundant orthopteran, comprising c. 43% of
the total number of Orthoptera overall across the 5 younger fields. In
2 out of the 3 older fields (=8 years post-cropping), P. parallelus was
the most abundant species, forming 20.6% of overall sightings. In
these older fields, C. fuscus comprised only 11.3% of adults.

Table 2. Mean number of Orthoptera nymphs and adults of each
species and species diversity and richness in fields 4 and =8 years
since cropping cessation, significance evidence shown (Student's
t-test).

Species 4 years >8 years tvalue p Evidence
Pseudochorthippus 12.88 +19 360 #11.0 -3.32 0.02 Moderate
parallelus
Chorthippus brunneus 8.2 +418 23.3 43.9 -3.59 0.01 Moderate
Roeseliana roeselii 11.2 28.0 +11.0 -1.28 0.33 None
Chorthippus 1.6 +0.5 4.7 +3.2 -1.23 0.26 None
albomarginatus
Conocephalus fuscus 28.4 +5. 19.7 + 1.07 0.33 None
Pholidoptera 0.2 +02 3.7 43.2 -1.24 0.34 None
griseoaptera
Omocestus viridulus 2.0 26.0 +11.0 -2.74 0.11 None
Tetrix subulata 0.8 +0. 26.00 +11.6 -3.05 0.09 Weak
Tetrix undulata 0.0 +0.0 7.3 +09 -15.12 <0.001 Very

strong
Nymphs (all species) 38.6 +163 61.0 +204 -0.94 0.38 None
Most abundant 42.9 438 22.2 +18 4.40 <0.001 Very
species (%) strong
Species richness 6.2 +04 8.7 +03 -4.44 <0.001 Very

strong
Species diversity 15 +01 1.9 +00 -6.32 <0.001 Very

strong

T. GARDINER, D. CASEY

There was moderate evidence (p < 0.05) that ant hill density,
lagomorph droppings, and the number of ponds were signifi-
cantly higher in fields =>8 years since cropping cessation (Table 3).
Contrastingly, there was very strong evidence (p < 0.001) that leaf
litter was significantly less abundant in fields >8 years since crop-
ping cessation. Only weak evidence was obtained that there was a
higher number of deer paths in the older fields (p < 0.10).

Of the five most abundant plant species, two were significantly
more numerous in fields >8 years since cropping cessation: creep-
ing bent Agrostis stolonifera (moderate evidence) and red fescue
Festuca rubra (very strong evidence; Table 4). Overall plant species
richness was also significantly higher (very strong evidence) in the
older fields (Table 4). In contrast, soft brome Bromus hordeaceus
was more abundant in fields 4 years post-cropping (strong evi-
dence). Two species, Yorkshire fog Holcus lanatus and bristly ox-
tongue Helminthotheca echioides, had similar abundance in both
types of field, as did bryophytes (Table 4).

Discussion
Species richness and abundance in a regional context

The total of 9 species recorded at Black Bourn Valley is
supplemented by species not observed on the transects, which
include the oak bush-cricket Meconema thalassinum (De Geer,
1773) and speckled bush-cricket Leptophyes punctatissima (Bosc,
1792). Omocestus viridulus was relatively abundant, but is a
scarce species in central Suffolk (Ling 2000), while the two Tetrix
groundhoppers are local to the county. On intensively managed
arable farmland in south-east England, O. viridulus is almost never
found (Gardiner 2010b), while groundhoppers such as T. subulata
are occasionally observed in low numbers along field edge
footpaths (Gardiner 2007). The presence of O. viridulus in fields 4
and >8 years since cessation of arable cropping suggests that this
species was quick to establish populations despite an allegedly
slow colonization rate of new grasslands and apparent restriction
to unimproved grassland (Gardiner 2010b).

The overall density of grasshoppers (234 adults/ha) compares
favourably with intensively managed farmland (<100 adults/ha;
Gardiner et al. 2002), indicating that rewilding arable land can
successfully promote large insect populations in the early stages of
reversion to grassland and scrub.

Factors influencing the colonization of rewilded fields

Pond edge.—Pond edge habitat with bryophytes (moss cover) and
bare earth established by digging and grazing lagomorphs may
be important for T: undulata and to a lesser extent T. subulata in
fields =>8 years since cropping cessation where there were small
farm ponds, particularly those created since 2010, or restored old
waterbodies (Fig. 2). Both Tetrix groundhoppers were found to
occur around the ponds in this study, although this may cause
issues with reproductive interference (Hochkirch et al. 2008).
Both groundhoppers can be abundant in river valleys such as the
Bourn, where they require both sparsely vegetated ground and tall
vegetation in close proximity to prevent individuals being washed
away during floods (Musiolek and Koéarek 2017). Groundhoppers
typically utilize micro-habitats in close proximity and can match
their color morph to the substrate for camouflage (Forsman 2000,
Ahnesj6 and Forsman 2006, Forsman et al. 2011). In this way,
melanic forms can be abundant on burned ground or where soil
is darker around silty river valley ponds.

JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(2)

T. GARDINER, D. CASEY

Fig. 2. Pond edge habitat with exposed soil in rewilded fields at

Black Bourn Valley, valuable for groundhoppers (Tetrix spp.) and
field grasshopper Chorthippus brunneus. Photo credit: Tim Gardiner.

The importance of restoring old ponds by vegetation clearance
and creating new ones should be recognized in proposals for re-
wilding at new sites to benefit orthopterans that require bare earth
edges clear of vegetation by lagomorph grazing (Fig. 2).

Ant hills.—To determine why arable reversion to grassland affects
Orthoptera, the habitat preferences of species should be exam-
ined. Habitat preferences of Orthoptera may relate to the choice
of oviposition site, food preferences, vegetation height, and grass-
land management regimes (Clarke 1948, Gardiner 2006, 2009).
Waloff (1950) stated that C. brunneus and P. parallelus lay their
ege pods in the superficial layers of the soil. Bare earth, often on
ant hills, is the usual egg-laying site for P. parallelus, although this
species and O. viridulus have been found to oviposit into grass-
covered soil (Waloff 1950). Exposed soil on ant hills may offer
other benefits for grasshoppers by providing sites where they can
bask (Key 2000), as it is often much warmer than surrounding
vegetation (Fig. 3). Although bare earth overall did not vary be-
tween field types, there were greater numbers of ant hills in fields
>8 years since cropping cessation, which indicates that micro-het-
erogeneity in unvegetated habitats may be important for egg-lay-
ing and basking P. parallelus and C. brunneus, which were in higher
abundance in the older fields (Table 2). Nymphs were evenly
distributed between field types, perhaps reflecting the need for a
diversity of resources including bare earth, but also heterogeneity
in sward structure, which was similar in both. Early instar grass-
hopper nymphs of C. brunneus and P. parallelus are often found
in short grassland near oviposition sites before moving to taller
swards (10-20 cm height) as they mature (Gardiner et al. 2002).
Adults may then return to bare earth and sparse sward patches,
such as those established by ant hills or rabbit grazing.

Ant hills in the older fields may have acted as hot ‘sun traps’
favorable for Orthoptera (Gardiner and Dover 2008, Voisin 1990),
particularly species such as C. brunneus and P. Parallelus, which
were in high abundance in those field types where overall species
diversity and richness were also high (Table 2).

As succession progresses on rewilded sites, the prevalence of
ant hills should be monitored to ensure that this valuable resource
for invertebrates persists. If there is a loss of bare earth on ant hills

eaI A) wiog Se ap) ‘ fs

Fig. 3. Ant hill (Lasius spp.) in rewilded grassland at Black Bourn
Valley, probably used for egg-laying and basking by grasshoppers.
Photo credit: Tim Gardiner.

over time, then site managers could consider periodic rotovation
of strips through fields for early successional, disturbance-depend-
ent species (e.g., C. brunneus and groundhoppers), similar to the
management of nearby Breckland heaths (Gardiner 2020). Any
rotovation should be irregularly undertaken to avoid too much in-
tervention in natural processes, akin to Rewilding Max. However,
a brownfield-type open mosaic habitat with patchy bare earth may
only be possible on poor or moderately fertile soils.

Lagomorph grazing.—The aim of Rewilding Max to reinstate natural
processes ideally means an absence of introduced livestock, such
as cattle, horses, or sheep, relying on wild grazing animals where
possible to manage succession. However, at several rewilded sites,
livestock has been introduced to control natural processes (Rewil-
ding Lite), while wild grazers, such as lagomorphs and ungulates,
have a reduced influence. There is little information on how suc-
cessful wild grazing animals are in maintaining grasslands in the
absence of domestic livestock on rewilded sites.

Wild grazing animals play a significant part in reducing veg-
etation height and cover (Gardiner 2018, Fargeaud and Gardiner
2018). On sea wall pollinator strips, wild rabbit grazing had a sig-
nificant impact on sward height and the density of R. roeselii adults
and orthopteran nymphs (Gardiner and Fargeaud 2020). Rabbits
grazed the closed grassland, reducing grass growth and creating
patches of exposed soil though their burrowing activities that were
favorable for basking nymphs (Gardiner et al. 2002).

JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(2)

168

Clarke (1948) suggested that excessive grazing by rabbits pro-
motes sparser vegetation comprised of less vigorous grass species
(such as sheep’s fescue Festuca ovina) which is consequently more
favorable to grasshoppers, perhaps because of the open sward
structure and warmer microclimate (Gardiner and Hassall 2009).
It is important to remember that microclimate may be crucial in
providing warm temperatures for the development of large Or-
thoptera populations (Marshall and Haes 1988), particularly in
rewilded fields where leaf litter is reduced >8 years since crop-
ping cessation allowing greater penetration of solar radiation to
the soil surface.

Ant hills and pond edges may assume greater importance in
a sward managed only by wild grazers (e.g., brown hares and
rabbits). The lagomorph grazing at Black Bourn Valley is mainly
assumed to be by brown hares, although some rabbit droppings
were located around the ponds where there were bare earth
patches (Fig. 2).

Short sward patches established by lagomorph grazing, particu-
larly when allied to sloping pond banks and ant hills, will have
excessively hot temperatures (>40°C), similar to hay meadows af-
ter cutting (Gardiner and Hassall 2009), which are unlikely to be
favored by grasshoppers in the absence of ‘cool’ tussocks in close
proximity. Both field types had mean sward heights >30 cm, which
could provide grasshoppers with numerous sheltered ‘cool’ areas of
tall vegetation away from ant hills and pond edges where tempera-
tures may be excessively hot during summer. This behavioral ther-
moregulation may account for the persistence of species such as C.
brunneus, P. parallelus, and T. undulata in fields >8 years since crop-
ping cessation where ant hills and sparsely vegetated pond edges
(basking and egg-laying sites) were frequently in close proximity to
cooler tall vegetation for shade-seeking orthopterans (Fig. 2).

Intensive grazing by unmanaged wild rabbit populations in
Epping Forest in the UK led to the extirpation of O. viridulus, a
grasshopper with a preference for tall grassland (Gardiner 2010b).
Rabbits also reduce the abundance of O. viridulus through the crea-
tion of uniformly short swards with little cover from avian preda-
tion and excessively hot microclimates (Gardiner 2021 ). Therefore,
uncontrolled rabbit grazing could pose a threat to O. viridulus on
heavily grazed areas of rewilded sites. The extent to which rabbits
influence the favorability of rewilded grasslands for insect orders
such as Orthoptera should be a focus of further research on arable
reversion sites.

Table 3. Natural grazing and habitat variables for fields 4 and =8
years since cropping cessation; Student's t values and significance
evidence shown for differences between means in each row.

Variable 4 years >8 years tvalue p Evidence
Ant hills/field fd BE DNT 40.3 +4188 -2.53 0.04 Moderate
Bare earth (mean 45 +08 3.4 +08 0.74 0.47 None
DOMIN)/quadrat
Deer droppings/field 4.4 3.7 : 0.20 0.85 None
Deer laydown areas/ 16.2 +6.3 8.7 +39 0.95 0.38 None
field
Deer paths/field 33.2 435 453 +3. -2.29 0.06 Weak
Lagomorph 05 +04 3.5 +09 -3.60 0.01 Moderate
droppings/m7?
Leaf litter (mean 4.3 +08 10 +08 5.29 0.001 Very
DOMIN)/quadrat strong
No. ponds/field 12. : 3.3 -2.69 0.04 Moderate
Sward height (cm) 83.6 +96 85.0 + -0.24 0.82 None
range/field
Sward height (cm)/ 44.1 +456 314 435 1.64 0.15 None
field

T. GARDINER, D. CASEY

The main source of error in this study may be the accuracy of
the lagomorph dropping counts. Droppings may have been easier
to locate in shorter, lagomorph-grazed vegetation and would have
dried and been less likely to decay in such situations compared to
the taller and moister vegetation present in some fields. Therefore,
the lagomorph dropping counts must be viewed with some cau-
tion, and further studies should be undertaken.

Plant species changes and sward height.—Changes in plant species
composition may have been a key influence on the abundance of
C. brunneus and P. parallelus in fields >8 years since cropping ces-
sation where fine-leaved grasses, A. stolonifera and FE rubra, were
in higher abundance (Table 4, Fig. 4). The development of open
sward structure with Festuca and Agrostis grasses in the older field
types benefits these two grasshopper species that require a shorter
sward composed of fine-leaved grass species (Gardiner et al. 2002).
While neither field type offered the optimal mean sward height for
C. brunneus and P. parallelus of 10-20 cm, the presence of diversity
in sward structure in fields 4 and =8 years since cropping cessation
indicates that patchy habitat was present with shorter swards for
grasshoppers (Fig. 4). It also suggests that microhabitats, such as ant
hills and pond edges, may assume greater importance in a sward
managed only by wild grazers (e.g., brown hares and rabbits). The
lagomorph grazing at Black Bourn Valley is mainly assumed to
be by brown hares, although some rabbit droppings were located
around the ponds where there were bare earth patches (Fig. 2).
Short ‘hot’ sward patches established by lagomorph grazing
are unlikely to be favorable for grasshoppers in the absence of
‘cool’ tussocks in close proximity. Both field types had a mean
sward height >30 cm, which will have provided grasshoppers with
numerous sheltered ‘cool’ areas of tall vegetation away from ant
hills and pond edges where temperatures may be excessively hot.

Ungulate grazing.—Wild ungulates are browsers that consume
grasses, sedges, shrubs, and trees (Uresk and Dietz 2018). How-
ever, the influence of deer grazing on swards was assumed to be
minimal at Black Bourn Valley (Table 3), despite the positive
(Adler and Proud 2021) and negative (Gardiner 2011) influence it
can have elsewhere.

In subalpine pastures in the Swiss Alps, Spalinger et al. (2012)
found no direct effect of wild ungulate grazing (red deer and
chamois). However, they did observe the small-scale alteration of
habitats and plant N content by ungulates, which in turn affected
Orthoptera abundance and diversity. Gardiner (2011) noted a re-
duced species richness of Orthoptera in red deer-grazed enclosures
near Broadway Tower in the Cotswolds, UK, where sward height
was mainly 5-10 cm with only occasional ungrazed taller vegeta-
tion (Gardiner 2011).

Table 4. Mean DOMIN abundance data/field for five most abun-
dant plant species, bryophytes (mosses, liverworts and hornworts)
and species richness for fields 4 and =8 years since cropping cessa-
tion, significance evidence shown (Student's t-test).

Species/family 4 years >8 years tvalue p Evidence
Helminthotheca 49 +1.0 23 +405 1.43 0.17 None
echioides
Agrostis stolonifera 10 +0.8 3.4 +05 -2.77 0.02 Moderate
Bromus hordeaceus 3.8 +0.7 0.6 +03 3.65 0.002 Strong
Holcus lanatus 2.9 +09 19 +04 -0.02 0.98 None
Festuca rubra 0.6 +05 3.9 +04 -7.05 <0.001 Very strong
Bryophytes 48 +04 40 +08 1.16 0.29 None
Species richness 13.7 +10 283 +1.1 -4.93 <0.001 Very strong

JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(2)

T. GARDINER, D. CASEY

Fig. 4. Rewilded grassland at 4 years (right) and =8 years (left)
since cropping cessation. Note the greener vegetation of the soft
brome Bromus hordeaceus-dominated field (right) compared to the
red and brown hues of the creeping bent Agrostis stolonifera and red
fescue Festuca rubra of the older fields. Photo credit: Tim Gardiner.

Deer droppings and laydown areas were not significantly dif-
ferent between field types in the current study, indicating that un-
gulate grazing was evenly distributed across Black Bourn Valley
(Fig. 5). There was weak evidence of greater deer path frequency
in older fields (Fig. 6), indicating that ungulate passage may have
had localized benefits for grasshoppers that need heterogeneity
in sward structure. However, the shorter grass of deer paths was
not numerous enough to make a difference to either mean sward
height or structural range (Table 4), despite the micro-habitat mo-
saic and structural diversity that has been observed in other studies
(Adler and Proud 2021). Deer may control the regrowth of scrub
in the long term to maintain the mosaic of sparse grassland and
woody vegetation, while ants and lagomorphs create the necessary
bare earth for disturbance-dependent invertebrates. Seed dispersal
can also occur due to ingested seeds being deposited across fields
in fecal matter from grazing deer and lagomorphs, which may lead
to the establishment of A. stolonifera swards (Eycott et al. 2007). It
is possible that A. stolonifera swards favorable for C. brunneus and
P. parallelus have developed along this successional trajectory at
Black Bourn Valley.

Unstudied factors influencing colonization

Corridor linkage.—The presence of an ancient green lane corridor
with hedgerows and grass margins may have allowed O. viridu-
lus to spread quickly into the rewilded fields (Fig. 7). The main
population of O. viridulus appeared to be centered in two of the
older fields where cropping ceased in 2013, from which individu-
als probably colonized the fields taken out of cropping in 2017.
Both fields with high concentrations of O. viridulus are located
next to a species-rich unimproved meadow and the Black Bourn
River Valley, which may have been sources of colonizing grasshop-
pers and likely explains the fast colonization into rewilded fields
(it was present in 7 out of the 8 fields). In the field nearest to Black
Bourn River (Kiln Meadow; 100 m), 5 out of 8 of the remaining
species were in highest abundance, further highlighting the ben-
efits of connectivity between rewilded fields and corridors to aid
quick colonization.

Fig. 5. Deer laydown areas with shorter vegetation (and drop-
pings), creating localized variation in sward structure in both field
types. Photo credit: Tim Gardiner.

Range-expanding species such as C. albomarginatus, C. fuscus,
and R. roeselii, which are spreading rapidly due to climate change
(Gardiner et al. 2002, Gardiner 2009), were early colonizers of
fields, being in similar abundance in fields 4 and =8 years since
cropping cessation (Table 2). These pioneer species benefit from
unmanaged grasslands and can build up large populations in such
habitats on farmland in the UK (Gardiner et al. 2002, Gardiner
2009). Conocephalus fuscus formed just over 40% of the orthopter-
ans counted in fields 4 years since cropping cessation, falling to c.
11% in those =8 years since cropping cessation, where P. parallelus
was the most abundant species.

Scrub development.—In the oldest field (Miller’s Field, 2007), which
had well-developed scrub patches, P. griseoaptera was in its high-
est abundance, reflecting a well-documented preference for woody
habitats (Marshall and Haes 1988). It should also be remembered
that patches of scrub are desirable for red-list declining birds such
as turtle dove Streptopelia turtur and common nightingale Luscinia
megarhynchos, both of which were recorded in the two rewilded fields
where cropping ceased in 2013 (Further Thurston and Kiln Meadow)
at Black Bourn Valley. Therefore, light scrub encroachment is desira-
ble from a conservation viewpoint, particularly as it also supports or-
thopterans such as L. punctatissima and M. thalassinum, both of which
can spread from green lanes into adjacent habitats (Gardiner 2010a).
A beating survey of the hedgerows and woodland would reveal the
distribution of arboreal bush crickets across the rewilded area.

JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(2)

Fig. 6. Deer paths through tall grassland, creating localized varia-
tion in sward structure in older fields. Photo credit: Tim Gardiner.

Soil fertility. —On soils with high fertility (e.g., clay) farmed inten-
sively for years, a low diversity, tussocky sward less favorable to
Orthoptera may develop (Gardiner 2006, 2009). Natural regen-
eration on highly fertile arable soils often produces grassy swards
of low diversity with minimal benefits to Orthoptera due to unfa-
vorable tall sward height and lack of structural diversity (Gardiner
2009) or to butterflies that are restricted because of the dearth of
nectar sources (Field et al. 2005, 2006, 2007). Therefore, farmland
suitable for the maximal rewilding benefits must be carefully cho-
sen. Farmers could remove fields from arable production and re-
wild them on the infertile or difficult-to-crop parts of their farms,
perhaps rewilding 10-15% of total crop area. Rewilding Max may
be the preferred option in these circumstances for Orthoptera, in
that active conservation grazing is probably unnecessary where
wild grazers such as lagomorphs can have a significant beneficial
impact for invertebrates.

Conclusion: Can rewilding aid Orthoptera conservation on
farmland?

Intensive agriculture has significantly reduced the diversity and
abundance of Orthoptera on UK farmland (Gardiner et al. 2002,
Cherrill 2010, 2015). Taking less agriculturally productive land out
of arable cropping and allowing it to naturally regenerate (Rewild-
ing Max) with minimal domestic livestock grazing and only wild
grazing by lagomorphs and ungulates can produce swards that are
quickly colonized (c. 4-8 years) by Orthoptera species not usu-

T. GARDINER, D. CASEY

Black Bourn River

Railway line

Fig. 7. Layout of the eight experimental fields at Black Bourn
Valley (year since cropping cessation shown) showing the likely
dispersal corridors onto rewilded farmland and the main green
lane corridor, which may have acted as a green highway to fields at
greater distance from the river valley corridor.

ally found on farmland, such as O. viridulus or T. undulata. Rewil-
ded fields should incorporate features such as old and restored
ponds to provide bare earth bank habitat for groundhoppers (e.g.,
T: subulata), while ant hills develop as the grassland matures, pro-
viding exposed soil and shorter swards for basking and oviposit-
ing grasshoppers (e.g., C. brunneus). Lagomorph grazing in and
around ponds and ant hills enhances the value of these bare earth
features by maintaining them at an early successional stage. Of
course, the response is species specific, and here the heterogeneity
of micro-habitat is important to cater for short (e.g., T; undulata)
and tall grassland (e.g., O. viridulus) species. A patchwork of dif-
ferent swards may also help to promote the highest diversity of
insects (Kruess and Tscharntke 2002).

Acknowledgements

The authors would like to thank Suffolk Wildlife Trust for
supporting the project and giving permission to survey the fields.
We would also like to extend our gratitude to the Suffolk Natural-
ists’ Society for funding the research through their Morley Grant
Scheme. Professor Rob Fuller kindly helped with the discussion
and review of the draft manuscript, while Anna Saltmarsh and
Joanna Hodgson assisted with the fieldwork. Prof. Axel Hochkirch
and an anonymous reviewer provided very constructive reviews of
this paper.

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