Active Management of Fruit Orchard Meadows Under The Influence of Suburbanization is Important for Insect Diversity

Abstract

Fruit orchards under different types of management represent the most common agroforestry practice in central Europe. Traditional fruit orchards with trees usually planted in meadows are at a surplus, providing suitable habitats for many plant and animal species. We examined the influence of different management and biotope types on three insect groups. This study was conducted in thirty orchards across the capital city of the Czech Republic – Prague (496 km²). We investigated the diversities of butterflies, hymenopterans and beetles. Their species richnesses mainly benefitted from orchard management and were partly higher at xerothermic sites than at mesic sites. Red-listed species did not show any clear patterns. Open-landscape specialists were influenced by management, while forest species were influenced by habitat type. Generally, orchard abandonment led to insect biodiversity loss. Therefore, active agricultural management appears to be essential for insect biodiversity conservation in orchards, and different management and biotope types provide suitable conditions for specific species. Mowing and maintaining orchards are two important biodiversity management actions in highly human-populated landscapes.

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Active Management of Fruit Orchard Meadows
Under The Inuence of Suburbanization is
Important for Insect Diversity
Patrik Rada
University of Hradec Králové https://orcid.org/0000-0001-8039-5193
Petr Bogusch
University of Hradec Kralove: Univerzita Hradec Kralove
Pavel Pech
University of Hradec Kralove: Univerzita Hradec Kralove
Jan Pavlíček
Lesák
Jiří Rom
Environmental Protection Department
Jakub Horák (  jakub.sruby@gmail.com )
University of Hradec Kralove: Univerzita Hradec Kralove
Research Article
Keywords: biodiversity, traditional fruit orchard, Lepidoptera, Hymenoptera, Coleoptera, agroforestry
DOI: https://doi.org/10.21203/rs.3.rs-299688/v1
License:   This work is licensed under a Creative Commons Attribution 4.0 International License. 
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Abstract
Fruit orchards under different types of management represent the most common agroforestry practice in
central Europe. Traditional fruit orchards with trees usually planted in meadows are at a surplus,
providing suitable habitats for many plant and animal species. We examined the inuence of different
management and biotope types on three insect groups. This study was conducted in thirty orchards
across the capital city of the Czech Republic – Prague (496 km²). We investigated the diversities of
butteries, hymenopterans and beetles. Their species richnesses mainly benetted from orchard
management and were partly higher at xerothermic sites than at mesic sites. Red-listed species did not
show any clear patterns. Open-landscape specialists were inuenced by management, while forest
species were inuenced by habitat type. Generally, orchard abandonment led to insect biodiversity loss.
Therefore, active agricultural management appears to be essential for insect biodiversity conservation in
orchards, and different management and biotope types provide suitable conditions for specic species.
Mowing and maintaining orchards are two important biodiversity management actions in highly human-
populated landscapes.
Introduction
European agroforestry practices have a long history starting with slash-and-burn agriculture and
continuing with forest pastures or fruit orchards (King 1987). Traditional fruit orchards are the most
abundant type of agroforestry in central Europe. The majority of fruit trees are planted in mowed or
pastured grasslands (Herzog 2000; Mosquera-Losada et al. 2009). Agroforestry as such is usually
considered to have a positive effect (e.g., Bignal and McCracken 1996; Myczko et al. 2013; Horak 2014a)
on biodiversity. However, different types of agroforestry have different impacts (Varah et al. 2013).
Schroth et al. (2004) summarized three major positive effects of agroforestry. First, agroforestry provides
secondary habitats for species that can tolerate some level of disturbance. Second, it connects natural
habitat fragments better than less tree-dominated land use, partly because traditional fruit orchards
combine the conditions of forest and grassland habitats in one place (Horak 2014b). Third, in some
cases, agroforestry can also reduce the rate of conversion to more intensive practices. This effect may be
related to higher biodiversity, which can, for example, successfully control pests without the need for
pesticides (Simon et al. 2011).
The habitat area or landscape context of the orchard may be another important factor (Steffan-Dewenter
2003). In this study, we are investigating traditional fruit orchards in the large area of a million-person city
that includes more than one hundred freely accessible orchard meadows (Janeček et al. 2019).
Orchard meadow biological diversity depends on many factors (Steffan-Dewenter 2003; Steffan‐
Dewenter and Leschke 2003; Čejka et al. 2018). Vegetation cover management seems to be very
important. However, groups of animals differ in their preferences. While birds or butteries prefer
abandoned or moderately managed orchards (Myczko et al. 2013; Horák et al. 2018), it appears that
carabids have the opposite preference for tilled or even herbicide-treated ground cover (Miñarro and

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Dapena 2013). In contrast, wasps are indifferent to management practices, according to another study
(Steffan‐Dewenter and Leschke 2003).
The aims of this paper were to determine whether different types of management, in the context of
habitat type, can inuence insect biological diversity in traditional fruit orchards and, if so, which
management type is the most suitable for current orchards in the city of Prague.
Methods
Study area
We studied traditional fruit orchards in Prague, the capital city of the Czech Republic. The area of Prague
is 496 km2 and is densely populated (1.3million citizens in 100 thousand houses), but it offers
biodiversity hotspots (Kadlec et al. 2008). The topography is diversied, with diverse bedrock, and divided
by the Vltava River as well as its river and stream tributaries. This region includes undeveloped areas with
habitats ranging from humid (e.g., wetlands close to streams) to very dry (steppes on southern hill
slopes) and from seminatural (e.g., natural reserves) to articial (e.g., brownelds). The mean elevation is
235 m a.s.l. (with a range from 177 to 399 m a.s.l.). The mean annual temperature is approximately 9°C,
and the mean annual precipitation is approximately 525 mm.
This area includes more than 100 freely accessible traditional fruit orchards representing artifacts of
agroforestry that supplied the city with fruits from its former rural peripheries (Horák and Trombik 2016).
We selected 30 of these orchards that were widely distributed throughout the territory of the city for our
research (Fig.1; Table S1).
Sampling methods
The collection of insect diversity data was focused on the three most species-rich insect orders:
Lepidoptera, Hymenoptera and Coleoptera. We selected one group for each insect order: day-active
butteries (clade Rhopalocera and family Zygaenidae; hereafter butteries), Aculeata (hereafter
hymenopterans) and click beetles (Elateridae; hereafter beetles).
The taxa studied as biodiversity indicators should meet certain requirements, such as having a well-
known taxonomy and natural history and/or being widely geographically distributed with the presence of
ecologically specialized species in the study region (Gayubo et al. 2005).
Day-active butteries are one of the most studied insect groups worldwide, mainly because of their easy
identication, well-known niches and ability to colonize new habitats (Thomas 2005). Aculeate
hymenopterans are also well studied (Gayubo et al. 2005; de Souza et al. 2010; Heneberg et al. 2013;
Heneberg and Bogusch 2020), with a special contribution in the form of social species – mainly bees and
ants (de Souza et al. 2010). Elateridae is one of the most diverse beetle families, with a wide range of
habitats and economic importance as pests (Traugott et al. 2015). These beetles can also serve as good

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bioindicator species (Dušánek 2011). All of these reasons led to the selection of these insect groups as
our indicator species.
We used the most suitable sampling method for every group studied during 2017. Butteries were
recorded using a time-limited survey (15 minutes). During the active season (April to August), seven
surveys were performed under ideal weather conditions (> 17°C, from sunny to partly cloudy skies, from
calm to mild wind). We surveyed all currently available major sources of nectar where more buttery
species were expected in similar habitats of central Europe (e.g., Kadlec et al. 2012).
Hymenopterans were collected using colored Moericke traps (150 x 150 x 45 mm length/width/depth)
(e.g., Heneberg and Bogusch 2014; Aranda 2018). We improved the traps by using half-yellow and half-
white pan traps. Ten traps were installed in every orchard in May, June, July and August. They were
always recollected after 72 hours. A colored stick was placed near every trap to ensure that the traps were
installed in similar places during the sampling periods. Traps were installed to cover all major present
conditions of orchards; therefore, ve of them were placed under the tree canopy, and ve were placed
between crowns. The traps were lled with saturated saline solution with a small amount of detergent to
reduce the surface tension of the liquid. This solution preserves insects.
Beetles were collected using passive window trunk tree traps (i.e., placed on tree trunks) from April until
the end of August. Each trap consisted of crossed transparent plastic panes (400 × 500 mm), a dark top
cover for protection and a plastic funnel leading down into a container with saturated saline solution and
a small amount of detergent. One trap was always placed in every orchard at breast height
(approximately 1.3 m above the ground) facing south (Horák et al. 2013a). The traps were recollected
every two weeks.
The collected hymenopterans and beetles were then transported in plastic bottles to the laboratory for
sorting and identication. Recorded or collected individuals of all studied orders were identied to the
species level. The only exception was buttery species, whose identication was based on DNA
sequencing or genital preparation (Friberg and Wiklund 2009). These species were identied as different
species sensu lato (e.g.,
Leptidea
spp.).
All identied species were divided according to the recent literature (Laibner 2000; Beneš et al. 2002;
Macek et al. 2010, 2015) into generalists and species with a preference for forest or open landscapes.
This distribution was then adjusted according to our expert knowledge. All identied species were also
compared with the Red List of threatened species of the Czech Republic (Hejda et al. 2017). For every
orchard, the threatened species index (Harcourt and Parks 2003) was calculated. This index consisted of
a threat category number (near threatened (1), vulnerable (2), endangered (3) and critically endangered
(4)) multiplied by the number of individuals from every category in the orchard.
Study variables
Three environmental variables were used in this study. The rst factor was the intensity of management.
The intensity of management often inuences biodiversity in different ecosystems (Bengtsson et al.

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2000; Rook and Tallowin 2003; Kampmann et al. 2008). Thus, we were interested in whether it has an
impact on the biodiversity of traditional fruit orchards. The intensity of management was recorded on an
ordinal scale. The rst level was abandoned orchard with no management (coded as 1). Management in
these orchards usually ended in the early 1990s, and such orchards have since become overgrown with
shrubs and young trees (Horák and Trombik 2016; Fig. S1). Moderately managed orchards were those
that were mowed a maximum of twice per season, often with unmowed patches left behind (coded as 2),
and intensively managed orchards were those that were mowed more than twice per season (coded as 3)
but not periodically as an urban lawn. Overall, we studied nine abandoned, 14 moderately managed and
seven intensively managed orchards.
The second studied variable was biotope type, which might be essential for the presence of particular
species (van Swaay et al. 2006). Biotopes were divided based on the type of vegetation according to
Chytrý et al. (2010). Two types of biotopes were recorded: xerothermic and mesic. Xerothermic vegetation
consisted of species such as
Festuca
spp.,
Stipa
spp.,
Dianthus carthusianorum
,
Centaurea stoebe
and
Eryngium campestre
. Mesic vegetation included species such as
Arrhenatherum elatius
,
Poa pratensis
,
Plantago lanceolata
and
Trifolium pratense
. In total, we studied eight orchards with xerothermic biotopes
and 22 orchards with mesic biotopes.
The third variable was orchard area (MEAN = 17538.75 (m²); SE = 4063.25; MIN = 2011.10; MAX =
90512.60). We vectorized all orchards from aerial photographs of Prague from 2015. Then, we adjusted
their borders according to a terrain survey and calculated their areas using ArcGIS 10.6.1. Orchard area
might inuence total biodiversity (Cornelis and Hermy 2004); hence, we used area as one of the predictors
to generalize our models.
Statistical analyses
Statistical analyses were performed using R version 3.6.1. Dependent variable normality was tested using
the Shapiro-Wilk test, and if necessary and possible, transformation was used to achieve a normal
distribution. If the distribution was signicantly different from normal or the values were close to
signicant, we used the packages MASS (Venables and Ripley 2002) and pscl (Jackman 2017) to test for
adherence to a Poisson distribution using the OdTest function. If the distribution was signicantly
different from a Poisson distribution, a negative binomial distribution was used. Theta was calculated for
every model using a tted negative binomial generalized linear model (glm.nb function).
Generalized linear mixed-effect models (GLMMs) were t using the packages nlme (Pinheiro et al. 2020)
and MASS with every dependent variable (glmmPQL function). Intensity of management and type of
biotope were included as xed independent variables. Orchard area was included as a random factor in
every model to generalize the results while accounting for the variation in this factor. The results were
visualized using the packages ggplot2 (Wickham 2016) and visreg (Breheny and Burchett 2017), which
offer visualization of variable dependency on multiple independent variables while accounting for
random factors.

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The threshold for running dependent variable analysis was set to the presence of at least eight species of
a dened category, and cases with only a few observations in interaction with two independent variables
were excluded from testing.
Canonical correspondence analyses (CCAs) were performed for every order (Lepidoptera, Hymenoptera
and Beetles) using the lattice (Sarkar 2008), permute (Simpson 2019) and vegan (Oksanen et al. 2019)
packages to obtain interactions between every single species of the selected orders and independent
variables. These analyses revealed the dependency of species composition on independent variables.
The results were visualized with the ggplot2 package.
Results
In total, 50 species of butteries represented by 2 219 individuals, 165 species of hymenopterans
represented by 4 610 individuals and 25 species of beetles represented by 242 individuals were observed
and trapped (Table S2).
Species richness
There was a signicant positive inuence of moderate management (t = 4.50; P < 0.001) and intensive
management (t = 2.07; P < 0.05; Fig.2) in fruit orchards on the observed species richness of butteries in
comparison with that in abandoned orchards. There was no signicant difference between moderate and
intensive management (Table1). Buttery species richness was signicantly higher in xerothermic than
in mesic (t = 2.84; P < 0.01; Fig.2) orchards.
Moderate (t = 2.14; P < 0.05) and intensive management (t = 2.58; P < 0.05) were signicantly preferred by
hymenopterans according to species number (Fig.2). There was no signicant difference between
moderately and intensively managed orchards. There was no signicant difference between the types of
biotopes (Fig.2; Table1).
Beetle species richness was not signicantly inuenced by the intensity of management or the type of
biotope (Fig.2; Table1).
Response of red-listed species
We observed 11 red-listed buttery species, eight red-listed hymenopterans and two red-listed beetle
species.
Red-listed buttery species were not signicantly affected by the intensity of management (Table1).
Signicantly more species were recorded in xerothermic biotopes than in mesic biotopes (t = 2.57; P <
0.05).
There was no signicant impact of intensity of management or type of biotope in the case of
hymenopteran species (Table1).

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Beetles were not analyzed due to the low number of species.
Open and forest landscape specialists
Open landscape-related butteries exhibited signicantly higher species richness in moderately managed
orchards (t = 3.88; P < 0.001) and intensively managed orchards (t = 2.12; P < 0.05) than in abandoned
orchards. There was no signicant difference between moderately and intensively managed orchards
(Table1). No signicant difference between types of biotopes was observed (Table1).
Open landscape-related hymenopteran species richness was signicantly higher in moderately (t = 3.19;
P < 0.01) and intensively managed orchards (t = 3.63; P < 0.01) than in abandoned orchards. There was no
signicant difference between moderately and intensively managed orchards. There was a signicant
impact of the type of biotope. There were more species present at xerothermic sites than at mesic sites (t
= 2.44; P < 0.05).
In the case of open landscape-related beetle species richness, there were no observed signicant impacts
(Table1).
Forest-related buttery species richness was not signicantly inuenced by the intensity of management
(Table1). There were signicantly more species in xerothermic biotopes (t = 3.45; P < 0.01) than in mesic
biotopes.
Forest-related hymenopterans were signicantly dependent on neither the intensity of management nor
the type of biotope (Table1).
Forest-associated beetles were not signicantly dependent on the intensity of management or the type of
biotope (Table1).
Species composition
We did not observe a signicant relation between buttery species composition and intensity of
management or type of biotope (Table2).
Our analysis indicated a signicant impact of the environment on hymenopterans (Table2). Specically,
there were fewer species in abandoned orchards. There were mainly
Andrena carantonica
and
Myrmica
rubra
.
Andrena ovatula
,
Andrena strohmella
and
Andrena oricola
had a preference for intensive
management of orchards (Fig.3).
Andrena polita
,
Bombus pascuorum, Formica pratensis
and
Lasioglossum lativentre
preferred moderately managed orchards.
Polistes nimpha
,
Andrena bicolor
and
Andrena dorsata
preferred xerothermic orchards, and
Halictus tumulorum
,
Nomada avoguttata
and
Andrena varians
were present in mesic biotopes (Fig.3).
There was a signicant inuence of orchard management and biotope type on the species composition
of beetles (Table2).
Agriotes ustulatus
preferred abandoned orchards, while
Selatosomus gravidus
occurred in intensively managed orchards. Moderate management was preferred by the species
Ampedus

Page 8/24
glycereus
and
Ampedus pomorum
.
Agriotes pilosellus
and
Melanotus villosus
preferred xerothermic
biotopes. Mesic biotopes were preferred by
Agrypnus murinus
(Fig.3).
Discussion
Our results indicate a mainly positive impact of the management of traditional fruit orchards on the
studied insect taxa. We also found that orchards planted in xeric environments tended to have higher
species richness.
Traditional fruit orchards might be depicted as grasslands with scattered trees (Plieninger et al. 2015) or
forests with very sparse canopies (Horak 2014a). This depends on the density of planted trees and the
stamina of the fruit tree species (Janeček et al. 2019). Fruit orchards provide suitable conditions for
species associated with open landscapes that were formerly pastured (Mosquera-Losada et al. 2012).
The abandonment of pastured grasslands and forests, from a long-term perspective, usually leads to a
decline in biological diversity (Benes et al. 2006; Queiroz et al. 2014; Uchida and Ushimaru 2014).
Therefore, from this perspective, it is not surprising that any type of orchard management has a positive
effect. On the other hand, intensication of mowing regimes is also decreasing biodiversity (Uchida and
Ushimaru 2014). Forest management has changed to more intensive approaches using high forests with
dense canopies (Benes et al. 2006). Thus, orchards might serve as a habitat for many species that
originally lived in traditionally hayed meadows (Cizek et al. 2012) and those associated with open forests
(Horak 2014a).
Managed orchard grasslands with xerothermic biotopes can serve as refugia for many species (Čejka et
al. 2018; Šantrůčková et al. 2020). The area of xerothermic grasslands is declining due to land
abandonment, afforestation, building activities or even conversion due to atmospheric nitrogen input or
fertilization (Tscharntke et al. 2005; Janišová et al. 2011). This has negative consequences for their
biodiversity (Buscardo et al. 2008; Janišová et al. 2011). In addition, orchards appear to be very similar to
dry traditionally managed woodlands (Hédl et al. 2010), where canopy openness is much higher than in
managed forests. This is mainly caused by competition among trees and was even multiplied by wooded
pasture (Konvicka et al. 2008) or coppicing in the past (Altman et al. 2013). These forests, typically with
xerothermic vegetation, have recently declined (Hédl et al. 2010). Traditional fruit orchards might
supplement these natural habitats and serve as habitats for species that originally lived in natural and
seminatural habitats. This may lead to higher biodiversity in xerothermic orchards. In addition, such
orchards can also serve as transitional habitats that facilitate migration between natural habitats
(Steffan-Dewenter 2003).
Butteries
The species richness of butteries in our study was most promoted by the management of orchards.
Furthermore, dry orchards were more species-rich than mesic orchards.

Page 9/24
Abandoned orchards are often akin to shrubland or forests having undergone natural succession for
some years after clear cutting (Prach 1994; Balmer and Erhardt 2000). The remaining grafted fruit trees
are overgrown, and their crowns are withering due to competition (Horák et al. 2018). The majority of day-
active butteries in central Europe prefer open or at least semiopen habitats (Beneš et al. 2002). There are
known examples of gradual declines in buttery fauna with increasing densities of trees and shrubs
(Erhardt 1985; Fartmann et al. 2013). Young tree forests are especially species poor (Balmer and Erhardt
2000). This is probably one of the most important reasons for the preference of butteries for actively
managed sites. This was also conrmed by the preferences of open landscape specialists for managed
orchards. Succession toward forests was not protable even for forest-related species. The main reason
is that forest-associated species mainly prefer sparse forest canopies (Beneš et al. 2002, 2006).
We predicted that moderate management would offer more heterogeneous biotope conditions, which
usually lead to higher species diversity (Noordijk et al. 2009; Cizek et al. 2012). More heterogeneously
managed sites should have more nectar sources and provide a more diverse understory of vascular
plants (Erhardt 1985; Steffan-Dewenter and Leschke 2003). In addition, research has shown that buttery
species richness mainly declines with intensive understory cultivation (Erhard 1985; Ekroos et al. 2010)
and that butteries prefer rather patchy mowing (Varah et al. 2013). Surprisingly, there was no signicant
difference in the effects of moderate and intensive management on the number of buttery species. This
nonsignicant difference can be explained by the complex 3D structure of managed orchards. This
means that, even after mowing, this habitat type still offers owers and shelter in tree crowns (Herzog
1998). In addition, some species can also benet from tree sap, honeydew or rotting fruits (Shreeve 1984;
Ômura and Honda 2003). Nevertheless, this observed issue remains relatively unclear – as abandoned
orchards also often offer the abovementioned complementary resources. One of the possible reasons
could be the temporal emigration of butteries to neighboring habitats followed by regression when the
condition of the intensively mowed orchard improves (Baum et al. 2004; Ouin et al. 2004). The most
surprising nding was that red-listed species were not inuenced by management. However, we observed
one endangered species, two vulnerable species and eight near-threatened species, which illustrated the
importance of orchards in a landscape context. These species might have such a strong biotope
preference that variation in local management has little effect (Fattorini 2010).
Xeric orchards contained more buttery species and of red-listed species. Surprisingly, forest specialists
also preferred xeric orchards. Xeric biotopes, including orchards, are often less accessible than other
biotopes. This is a possible reason why they have stayed relatively untouched by suburbanization
(Ouředníček 2007). Therefore, xeric orchards and similar dry habitats can be suitable for many species –
even those not specialized for dry habitats (Dostálek and Frantík 2008; Kadlec et al. 2008). This is
conrmed by the nding that some urban areas (such as railway verges or brownelds) could mimic
natural xeric habitats (Konvicka and Kadlec 2011). These orchards are still rather marginal. However, they
can facilitate connectivity or mimic natural xeric steppes. In addition, they might also serve as transitional
habitats or stepping stones (Horak 2014a) in fragmented areas of cities (Horák 2016). This is probably
the reason why orchards in xerothermic biotopes contained more species of both butteries and red-listed
butteries.

Page 10/24
Forest specialist species richness was higher in xerothermic biotopes than in mesic biotopes. Xerothermic
orchards might imitate and substitute natural xeric forests due to their similarly high canopy openness
(Chytrý 1997; Hédl et al. 2010). Such openness is needed for forest specialist butteries to nd suitable
basking sites, while forest cover serves as an important food source and shelter (Shreeve 1984; Dennis
and Sparks 2006). In addition, xerothermic forests are currently less common in central Europe because
modern forestry practices shifted them to mesic forests (Hédl et al. 2010). Thus, butteries originally
associated with dry forests had to nd supplementary habitats with similar living conditions, and
traditional fruit orchards provided such conditions.
Surprisingly, the species composition of butteries did not differ among management and biotope types.
For example, we observed some forest-related species (e.g.,
Argynnis paphia
and
Celastrina argiolus
), but
they were present in all types of orchards. Nevertheless, these species were not as common as other
species in the orchards, suggesting that orchards are simply transitional habitats for these species in the
landscape context. On the other hand, transitional habitats might be as ecologically important as natural
habitats (Schmitt and Rákosy 2007), underlining the importance of traditional fruit orchards for insect
biodiversity.
Hymenopterans
Hymenopterans benetted from management in orchards, and they were unaffected by the type of
biotope. The only exceptions were open-landscape specialists, which preferred xerothermic sites. This
might be because ground cover in xerothermic biotopes is often bare soil, which is ideal for nesting in
many hymenopteran species (Heneberg et al. 2014; Fortel et al. 2014; Bogusch et al. 2020). In addition,
active management such as mowing might multiply this effect since disturbed habitat is often preferred
(Fortel et al. 2014). Orchards with old veteran fruit trees also host a high diversity of species that nest in
wood cavities, especially smaller ones (Horák et al. 2013b; Bogusch and Horák 2018).
Many hymenopterans prefer habitats in early successional stages (Heneberg et al. 2013; Taki et al. 2013;
Heneberg and Bogusch 2020). Such stages could be maintained in orchards by intensive management. A
more heterogeneous understory is at a surplus in managed orchards since intermediate disturbance
reduces the dominance of competitive species and increases plant species richness (Curry 1994). This
offers more available food sources (Steffan-Dewenter and Leschke 2003). In this context, it is not
surprising that we observed one critically endangered species (
Andrena similis
), three vulnerable species
and four near-threatened species. Their incidence is another indication that traditional fruit orchards
could provide suitable habitats for insect biodiversity.
Management practices were clearly preferred, and there were fewer species in abandoned orchards.
Nevertheless, some of the species – for example,
Myrmica rubra
and
Andrena carantonica
– preferred
abandonment. An explanation in the case of
M. rubra
might be that this species prefers nesting in leaf
litter or within woody debris (Groden et al. 2005), which is more common in abandoned orchards. In the
case of
A. carantonica
, the only reasonable explanation might be that this species does not require as
high a temperature for pollination as the European honeybee (
Apis mellifera
) (Chansigaud 1975) and

Page 11/24
collects pollen and nectar, usually from tree owers (Macek et al. 2010; Westrich 2018). Thus, it might use
shrub overgrowth as an inhibitor of high temperatures in the understory (Breshears et al. 1998), providing
a competition benet.
Other species found in the abandoned orchards, such as
Temnothorax crassispinus
and
Lasius
fuliginosus
, usually occurred in forest biotopes. Therefore, their preference was rather natural (Macek et
al. 2010). Moderate management was preferred in the case of species such as
Lasioglossum lativentre
and
Formica pratensis
whose natural biotopes are forest steppes. Thus, moderately managed orchards
with patchy mowing and the presence of old-growth trees might serve as articial supplements.
Nevertheless, these orchards were also preferred by habitat generalists such as
Andrena polita
and
Bombus pascuorum
(Macek et al. 2010), which might instead primarily benet from the continuous
presence of nectar (Croxton et al. 2002). Intensively managed orchards were inhabited by species such as
Andrena ovatula
,
A. oricola
and
A. strohmella
, which corresponded to their preference for open biotopes
and oral specialization (Macek et al. 2010; Westrich 2018; Bogusch et al. 2020).
Beetles
Click beetles are known to prefer veteran trees exposed to sunlight (Horák and Rébl 2013). Therefore,
managed orchards appeared to be ideal habitats. Nevertheless, beetles were indifferent to the
management and type of habitat in our study. Species richness was higher in moderately managed
orchards than in abandoned orchards, but this difference was not signicant. From this trend, we might
conclude that moderate management could have a positive impact on beetle diversity.
We trapped species displaying all types of biotope preferences, from a preference for crop elds (
Athous
haemorrhoidalis
and
Agrypnus murinus
) to a preference for high forests (
Melanotus villosus
and
Dalopius marginatus
). This leads to the conclusion that traditional fruit orchards can host a very wide
range of species that can benet from meadow, trees or both within one habitat. This was also conrmed
by the presence of red-listed species – we trapped one vulnerable (
Brachygonus megerlei
) and one near-
threatened species (
Ampedus rupennis
). Both of these species are saproxylic (Zaharia 2006; Brunet and
Isacsson 2009). Thus, old trees present within habitats are essential for them, even in agricultural
habitats.
Species composition differed among the studied orchard categories. Abandoned orchards were preferred
by
Agriotes ustulatus
. This species is usually found in places with lower vegetation cover (Mertlik 2016),
which appears to contradict our ndings. However, this species also prefers soil with a higher humus level
(Čačija et al. 2018), which is found mostly in abandoned orchards due to decaying plant residuals (Horák
et al. 2018). Moderately managed orchards were inhabited by
Ampedus glycereus
and
A. pomorum
,
which are mostly known from forests, and they clearly benetted from the presence of old trees (Laibner
2000). Our studied sites were typical in terms of the presence of old fruit trees, which could mimic the
forest environment for these species. Intensive management of orchards was preferred by
Selatosomus
gravidus
. This species is usually present in steppe biotopes, and intensively managed orchards are
probably also suitable (Laibner 2000), as in the case of some hymenopterans.

Page 12/24
Our results were in some cases opposite to the ndings of current research regarding species
composition between biotopes. In xerothermic biotopes, the species composition included
Agriotes
pilosellus
, which is supposed to prefer mesic biotopes, and
Melanotus villosus
, which is usually described
as a generalist (Laibner 2000). Nevertheless, in mesic biotopes, we trapped species such as
Selatosomus
gravidus
, usually mentioned as having a preference for xerothermic biotopes, and
Agrypnus murinus
,
which is thought to be a generalist or even a pest species (Laibner 2000). We can provide context for this
information with the nding that
Selatosomus gravidus
could also be found at mesic sites (Bulgakova
and Pyatina 2019). Moreover, this species prevailed in intensive orchards, where intensive mowing can
cause even mesic biotopes to be drier than in the case of using less intensive practices (Kobayashi et al.
1997). Thus, all of the abovementioned reasons might explain this preference for mesic sites.
Conclusion
Traditional fruit orchards, as humanmade habitats, appear to be very important for many insect species.
Many red-listed species were present in these habitats, which combine the conditions of grassland and
forest in one place.
Management of orchards was essential for many species and thus very important for biodiversity. There
was a nonsignicant difference in species richness between moderate and intensively managed
orchards. Nevertheless, the species composition was different between them. Thus, we can conclude that
the presence of both approaches is important in terms of maintaining large-scale and long-term species
diversity. Xerothermic biotopes showed, in some cases, higher species diversity, and species composition
differed between xerothermic and mesic biotopes. Thus, the presence of orchards with both biotopes is
important.
Declarations
Acknowledgments
We are grateful to Jana Zemanová and Alena Pacáková for their help with beetle and hymenopteran
sampling and to Petr Boža for his help with the identication of click beetles.
Funding
This study was supported partly by the project of MHMP (OBJ/OCP/54/12/00100/2017) for NGO Lesák
and partly by a UHK-specic research project (2116/2020).
Conict of interest
All authors declare that they have no conict of interest.
Data availability Not applicable.

Page 13/24
Ethics approval Not applicable.
Consent to participate Not applicable.
Consent for publication Not applicable.
Code availability Not applicable
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Tables
Due to technical limitations, table 1,2 is only available as a download in the Supplemental Files section.
Figures

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Figure 1
Map of study sites located in the capital city of Prague in the context of the Czech Republic and Europe.
The black area in the cadastral map of Prague shows the real orchard area, with the orchard numbers in
gray Note: The designations employed and the presentation of the material on this map do not imply the
expression of any opinion whatsoever on the part of Research Square concerning the legal status of any
country, territory, city or area or of its authorities, or concerning the delimitation of its frontiers or
boundaries. This map has been provided by the authors.

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Figure 2
Species richness dependency on the intensity of management for (a) butteries (Lepidoptera), (c)
hymenopterans (Hymenoptera) and (e) beetles (Coleoptera) and the type of biotope for (b) butteries, (d)
hymenopterans and (f) beetles in Prague (Czech Republic) traditional fruit orchards. Note that plots
based on GLMMs indicate relative changes in species richness between categories of study treatments,
the black line is the mean, shaded boxes represent the 95% condence intervals, and black dots represent
individual observations

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Figure 3
Species composition relationships with the intensity of management and type of biotope for (a, b)
hymenopterans (Hymenoptera) and (c, d) beetles (Coleoptera) in Prague (Czech Republic) traditional fruit
orchards. Plots based on CCA on the left show individual species distributions (small dots) in relation to
the treatment categories (large dots), and species abbreviations (two letters from the Latin genus and
three from the species name) are visualized without categories on the right. Abbreviations of species
names in alphabetical order are as follows: (b) An_cin = Andrena cineraria; An_dor = Andrena dorsata;
An_fal = Andrena falsica; An_c = Andrena oricola; An_g = Andrena orivaga; An_ful = Andrena
fulvago; An_gra = Andrena gravida; An_hae = Andrena haemorrhoa; An_hel = Andrena helvola; An_chry =
Andrena chrysosceles; An_nig = Andrena nigroaenea; An_nit = Andrena nitida; An_ova = Andrena ovatula;
An_pol = Andrena polita; An_str = Andrena strohmella; An_var = Andrena varians; An_vir = Andrena
viridescens; Bo_hyp = Bombus hypnorum; Bo_lap = Bombus lapidarius; Bo_pas = Bombus pascuorum;
Ca_fal = Camponotus fallax; Cr_qua = Crossocerus quadrimaculatus; Fo_pra = Formica pratensis; Ha_tum
= Halictus tumulorum; He_nie = Hedychrum niemelai; Hy_ann = Hylaeus annularis; Hy_con = Hylaeus
confusus; La_ema = Lasius emarginatus; La_ful = Lasius fuliginosus; La_ltv = Lasioglossum lativentre;
Me_cen = Megachile centuncularis; Mi_dah = Mimumesa dahlbomi; My_rub = Myrmica rubra; No_a =
Nomada avoguttata; No_gut = Nomada guttulata; Os_bcr = Osmia bicornis; Os_cae = Osmia
caerulescens; Po_nim = Polistes nimpha; Ps_con = Psenulus concolor; Te_cra = Temnothorax

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crassispinus; Tr_att = Trypoxylon attenuatum (d) Ag_bre = Agriotes brevis; Ag_mur = Agrypnus murinus;
Ag_pil = Agriotes pilosellus; Ag_spu = Agriotes sputator; Ag_ust = Agriotes ustulatus; Am_gly = Ampedus
glycereus; Am_nig = Ampedus nigroavus; Am_pom = Ampedus pomorum; Am_ruf = Ampedus rupennis;
At_bic = Athous bicolor; At_hae = Athous haemorrhoidalis; Br_meg = Brachygonus megerlei; Ca_bip =
Calambus bipustulatus; Ca_eri = Cardiophorus erichsoni; Me_vil = Melanotus villosus; Se_gra =
Selatosomus gravidus
Supplementary Files
This is a list of supplementary les associated with this preprint. Click to download.
Table1.jpg
Table1a.jpg
Table2.jpg
Supplementarymaterial.docx