OR I G I N A L A R T I C L E
Crop type rather than production method determines
functional trait composition of insect communities on arable
land in boreal agricultural landscapes
Marjaana Toivonen1 | Erja Huusela2 | Terho Hyvönen2 | Ari Järvinen3 |
Mikko Kuussaari1
1Finnish Environment Institute (Syke), Helsinki,
Finland
2Natural Resources Institute Finland (Luke),
Jokioinen, Finland
3Natural Resources Institute Finland (Luke),
Jyväskylä, Finland
Correspondence
Marjaana Toivonen, Finnish Environment
Institute (Syke), Latokartanonkaari 11,
FI-00790 Helsinki, Finland.
Email: marjaana.toivonen@syke.fi
Funding information
Strategic Research Council of the Academy of
Finland, Grant/Award Number: 352638
Abstract
1. To understand the potential consequences of arable land use changes for insect
conservation and ecosystem functioning, it is fundamental to know how insect spe-
cies with different functional traits respond to crop choice and production method.
2. This study examined the effects of crop type and production method on functional
traits of butterfly, bumblebee and carabid beetle communities using species abun-
dance data from 78 fields in Southern Finland. Surrounding landscape composition
was also accounted for. The studied traits were associated with dispersal capacity,
habitat or diet specialization and phenology—the key determinants modifying spe-
cies responses to agricultural disturbances and land use changes.
3. Butterfly habitat breadth was narrowest and wingspan shortest in long-term fal-
lows. Fallows also supported the highest share of butterflies overwintering in early
development stages and bumblebees with late-emerging queens. The tongue length
of bumblebees was longest in organic oat fields, probably due to flowering weeds
with long corolla.
4. For carabid beetles, the proportion of poor flyers and carnivores was highest in
perennial crops and fallows. Carabid beetles overwintering as adults were relatively
more abundant in organic than in conventional production, probably due to more
intensive tillage in organic fields.
5. In all insect groups, poor dispersers and/or specialists decreased with increasing
arable land cover in the surrounding landscape.
6. Increasing the area of long-term fallows and perennial crops and enhancing within-
field plant diversity while maintaining landscape heterogeneity would promote
insect species sensitive to agricultural disturbances and land use changes and their
associated ecosystem services in boreal farmland.
K E YWORD S
bumblebees, butterflies, carabid beetles, community-weighted means, dispersal capacity, fallows,
organic production, specialization, species traits
Received: 15 January 2024 Accepted: 1 May 2024
DOI: 10.1111/afe.12629
This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium,
provided the original work is properly cited.
© 2024 The Authors. Agricultural and Forest Entomology published by John Wiley & Sons Ltd on behalf of Royal Entomological Society.
Agr Forest Entomol. 2024;1–12. wileyonlinelibrary.com/journal/afe 1
INTRODUCTION
Agricultural land use change and management intensification are
among the main drivers of insect decline globally (Wagner, 2020). In
agricultural landscapes, insect diversity and abundance have declined
due to the loss of semi-natural habitats and landscape heterogeneity
(Hendrickx et al., 2007; Tscharntke et al., 2005), and the adoption of
high-intensity management practices such as high pesticide and inor-
ganic fertilizer inputs (Geiger et al., 2010; Gossner et al., 2016). While
it is clear that intensive agriculture is damaging insect diversity, the
effects of various arable land use options on insect community com-
position are still not well understood.
Arable land can temporarily provide plenty of resources for some
insects (Holland et al., 2004; Knapp et al., 2019; Martins et al., 2018).
However, in Europe and many other parts of the world, habitats on
arable land are mostly homogeneous and ephemeral: the vegetation is
typically dominated by few plant species, and the land is regularly dis-
turbed by management such as ploughing, mowing and pesticide use.
This leads to insect movement between arable land and adjacent
semi-natural habitats that, compared to crops, provide more heteroge-
neity and temporal continuity in resources and more stable conditions
(Knapp et al., 2019; Martínez-Núñez et al., 2022; Martins et al., 2018).
Among arable land use types, fallow or set-aside land, that is, land
temporarily taken out of production, can host relatively high insect
diversity, including species that are not able to persist in more dis-
turbed crop fields (Kovács-Hostyánszki et al., 2011; Toivonen
et al., 2015, 2022). In many European countries, farmers are paid agri-
environment subsidies for establishing and managing different types
of fallows, of which long-term fallows and fallows sown with diverse
seed mixtures are probably most beneficial to insect diversity (Alanen
et al., 2011; Boetzl et al., 2021; Toivonen et al., 2015).
Insect communities on arable land are constrained by species’
functional response traits (i.e., characteristics of species important for
their response to the environment) that influence their ability to move,
establish, reproduce and tolerate disturbances. Typically, habitat or
resource generalists with strong mobility can cope with intensive agri-
cultural land use and homogenous landscapes better than specialist
species with poor mobility (Börschig et al., 2013; Gámez-Virués
et al., 2015; K}orösi et al., 2022). The trophic levels of the species may
also affect the responses: predators have been suggested to be more
dependent on landscape diversity than herbivores that respond more
strongly to local habitat type (Woodcock et al., 2010). Furthermore,
traits associated with life cycle timing and pace, such as overwintering
stage, voltinism and the length of the activity period, may be linked to
land use intensity and landscape structure (Börschig et al., 2013;
Gámez-Virués et al., 2015; K}orösi et al., 2022). For example, species
with short activity periods have been reported to suffer from intensive
management more than species with longer activity periods (Börschig
et al., 2013; Gámez-Virués et al., 2015). Insects’ resource needs and
vulnerability to disturbances also vary depending on their life cycle
stage (K}orösi et al., 2022). Thus, the ability of an insect species to use
arable land is dependent on the temporal match between insect life
cycle, field operations and resource provision on arable land.
Functional effect traits (i.e., characteristics of species important
for their effect on the ecosystem) of insect communities, in turn,
affect ecosystem functioning and service provision, such as the sup-
pression of pests and weeds on arable land (Greenop et al., 2018;
Guenay-Greunke et al., 2023; Welch & Harwood, 2014) or pollination
success of insect-pollinated crops and wild plants (Bommarco
et al., 2012; Ehlers et al., 2021; Woodcock et al., 2019). Hence, know-
ing how insect species with different functional traits respond to ara-
ble land use is a prerequisite for understanding the consequences of
arable land use change for both insect conservation and insect-driven
ecosystem services.
Crop choice and production method are important determinants
of habitat quality for many taxa on arable land (Stein-Bachinger
et al., 2021; Toivonen et al., 2022). For example, cereal fields, which
dominate agricultural land use in most of Europe (Eurostat, 2016), are
relatively resource-poor habitats for many insects (Toivonen
et al., 2022; Yvoz et al., 2020). The types of crops grown are affected
by the production method (Barbieri et al., 2019), and both production
method and crop type influence management practices, such as tillage
system and agrochemical use. Crop choice and production method are
also of high political interest and under pressure for change. For
example, in the European Union, increasing domestic plant protein
production as well as increasing the share of agricultural land under
organic farming have been set as targets (European
Commission, 2020). Still, functional traits of insect communities have
seldom be compared between various crop types while also account-
ing for production methods (but see Eyre et al., 2013).
This study examined how key functional traits of insect communi-
ties on boreal arable land are influenced by crop type and production
method. The studied traits were associated with dispersal capacity,
habitat or diet specialization and life cycle timing—the key determi-
nants modifying species responses to agricultural disturbances and
land use changes. The crop types included seven common annual
and perennial crop types, as well as long-term fallows as an alternative
land use option. We expected that (1) the proportional abundance of
species with poor dispersal ability is higher in perennial crops and fal-
lows than in annual crops, as persisting in a more disturbed habitat
requires a better dispersal ability; (2) the proportional abundance of
habitat or diet specialists is highest in fallows, which have more
diverse vegetation and a higher share of wild plants than crop fields,
and are thus more likely to provide suitable resources for specialist
insects and (3) traits associated with life cycle timing, including over-
wintering stage and spring emergence time differ between arable land
use types since these traits affect insects’ ability to tolerate distur-
bances and utilize resources at different times of the year.
MATERIALS AND METHODS
Study design
Data on butterflies, bumblebees and carabid beetles were obtained
from a biodiversity study of 78 fields in Southern Finland (60.09936–
2 TOIVONEN ET AL.
 14619563, 0, D
ow
nloaded from
 https://resjournals.onlinelibrary.wiley.com/doi/10.1111/afe.12629 by Duodecim Medical Publications Ltd, W
iley Online Library on [13/09/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on W
iley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
61.01251
 N, 23.40421–25.14284
 E), described in detail by Toivo-
nen et al. (2022). Landscapes in the study region are typically mosaics
of arable land and coniferous and mixed forests. The study fields
represented a range of arable land use types, called as ‘crop types’ in
this paper: (1) spring cereal (oat; n = 10), (2) winter cereal (rye;
n = 10), (3) faba bean (n = 10), (4) spring oilseed crops (n = 10),
(5) cabbage (n = 8), (6) grass ley (n = 10), (7) arable pasture (n = 10)
and (8) long-term fallow (n = 10) (Table 1). Fallows are a popular agri-
environment measure in the study region and they are often retained
in the same place for several years or even decades. The studied fal-
lows had 8–35 years old grassland swards. Five conventionally man-
aged and five organically managed fields were selected for each crop
type, except for oilseeds and cabbage, which are uncommon in
organic farms. For oilseeds, seven conventionally managed and three
organically managed fields were selected. For cabbage, eight conven-
tionally managed fields and no organically managed fields were found.
All organically managed study fields were situated in organic farms
certified according to the EU regulation (Regulation (EU) 2018/848).
Fallows were also selected from organic and conventional farms,
although the production method does not affect the management of
long-term fallows. The study fields were separated by at least 500 m.
For the study fields that represented the same crop type, the mini-
mum distance was 1 km.
Proportions of the areas covered by arable land, forest, built-
up area and waters and wetlands were calculated for four radii
(100, 250, 500 and 1000 m) around the central points of the study
sites using ArcGIS 10.8.1 (Esri, 2020). Data on arable land cover
were obtained from the Finnish agricultural land parcel register
(Finnish Food Authority, 2021). Data on the other land use types
were derived from CORINE Land Cover 2018 dataset (Finnish
Environment Institute, 2018). Fallows were situated in landscapes
with higher forest cover and lower arable land cover than the
other crop types (Table 1). This could not be avoided because
farmers prefer to have long-term fallows in field parcels that are
low productive, irregularly shaped or located far away from the
farm centre and these kinds of parcels are concentrated in for-
ested landscapes.
Insect sampling
Butterflies and bumblebees were counted five times during the sum-
mer of 2020 at approximately 2-week intervals, from 27 May until
7 August, in weather conditions allowing insect activity. Counts were
performed using the standard line transect method (Pollard &
Yates, 1993), in which an observer walks along a fixed route at a
steady speed and counts all the individuals within an imaginary box of
5  5  5 m in front of him or her. In each field, two 50 m-long tran-
sects were placed parallel at 5 and 20 m distances from the field edge
farthest from the forest. Butterflies and bumblebees were identified
as species. An exception was bumblebee individuals belonging to the
Bombus lucorum group (B. lucorum (L.), Bombus terrestris (L.), Bombus
cryptarum (Fabiricius) and Bombus magnus Vogt) that were treated as
one species, B. lucorum, in the analyses.
Carabid beetles were sampled by pitfall trapping between 8 and
25 June 2020, with two consecutive 1-week trapping periods per
field. Three pitfall traps (9 cm in diameter, 8 cm in depth, transparent
plastic) filled with concentrated NaCl liquid were placed in each field:
two traps were placed next to the ends of the pollinator transects, at
5 and 20 m distances from the field edge, and one trap between them,
at 12.5 m distance from the edge. The traps were covered with ply-
wood roofs to prevent flooding during rain. All carabid beetles were
identified at the species level. In two fields, pitfall trapping could not
be conducted, and in one field, the traps were in place only during
T AB L E 1 The descriptions of the crop types, the numbers of study fields per crop type and production method and the covers of arable land
and forest within the 250-m radius around the central points of the study sites.
Crop type Description
Number of study fields
Land use cover (%) within
250 m, mean
Conventional Organic Arable land Forest
Spring cereal Oat Avena sativa L. 5 5 (4) 68 23
Winter cereal Rye Secale cereale L. 5 5 72 18
Faba bean Vicia faba L. 5 5 66 23
Spring oilseeds Annual mass-flowering Brassicaceae: oilseed rape Brassica
napus L. subsp. oleifera, turnip rape Brassica rapa L. subsp.
oleifera and camelina Camelina sativa (L.) Crantz
7 3 66 24
Cabbage Brassica oleracea L. var. capitata 8 (7) 0 58 30
Grass ley Mean age of the sward 3.5 years (median 2.5 years), mowing
2–3 times/year
5 5 69 28
Arable pasture Mean age of the sward 7.5 years (median 4.5 years); various
animal types and grazing intensity
5 5 (4) 62 30
Long-term fallow Mean age of the sward 18.1 years (median 17.5), mowing 0–1
times/year
5 5 30 49
Note: The numbers of study fields in parentheses indicate the fields where full data on carabid beetles were collected, if not collected in all study fields.
INSECT FUNCTIONAL TRAITS ON ARABLE LAND 3
 14619563, 0, D
ow
nloaded from
 https://resjournals.onlinelibrary.wiley.com/doi/10.1111/afe.12629 by Duodecim Medical Publications Ltd, W
iley Online Library on [13/09/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on W
iley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
one trapping period. These fields were excluded from the analyses of
carabid beetle traits.
Species traits
For each insect group, three species traits associated with dispersal
capacity, habitat or diet specialization and life cycle timing were
selected. These traits were selected because they have been shown
to be key determinants modifying insects’ responses to agricultural
land use (Börschig et al., 2013; Gámez-Virués et al., 2015; K}orösi
et al., 2022; Toivonen et al., 2016) and because previously published
measures of these traits were available for all three insect groups.
Butterfly species traits were wingspan (as a measure of dispersal
capacity; Kuussaari et al., 2014; Sekar, 2012), adult habitat breadth
and overwintering stage. The average wingspan of females was based
on Marttila et al. (1990). Adult habitat breadth (an inverse measure of
habitat specialization) was expressed as the number of habitat types
occupied by adult butterflies, as in Komonen et al. (2004). Overwin-
tering stage (Marttila et al., 1990) was an ordinal variable describing
the development stage and ranging from 1 (egg) to 4 (adult).
Bumblebee species traits were body size (a measure of dispersal
capacity; Greenleaf et al., 2007), tongue length (a measure of diet spe-
cialization; Goulson & Darvill, 2004) and spring emergence time. Body
size, measured as the average body length of females (from the front
edge of the head to the tip of the abdomen), was based on Söderman
and Leinonen (2003), supplemented by Scheuchl and Willner (2016)
for some species. Tongue length was according to Ranta and Lund-
berg (1980) and Goulson et al. (2005, 2008). Spring emergence time
of queens (Scheuchl & Willner, 2016; Söderman & Leinonen, 2003)
was expressed as month and it ranged from 4 (April) to 6 (June).
Carabid beetle species traits were flight ability, feeding prefer-
ence and overwintering stage. All three traits were categorical with
two levels. The categorization of species was according to Hanson
et al. (2016), supplemented by information from Lindroth (1985,
1986). Species with no or reduced wings, dimorphic species and those
for which flight observations are missing or rare, were classified as
poor flyers, while the species with fully developed wings and docu-
mented flight ability were classified as good flyers. Based on feeding
preference, carabid beetles were divided into species with mainly car-
nivorous diet and those with omnivorous or herbivorous diets. Fur-
thermore, the species were grouped into those overwintering in adult
stage and those overwintering in larval stage.
Statistical analyses
Prior to statistical analyses, insect observations of each field were pooled
over the season (five transect counts for butterflies and bumblebees and
two trapping periods for carabid beetles) and the locations within the
field (two transects for butterflies and bumblebees and three traps for
carabid beetles). The pooled abundance data were used to calculate
community trait values for each field. For butterflies and bumblebees,
the trait values of individuals observed on a field were averaged to get
the community-weighted mean for the field (following, e.g., Kuussaari
et al., 2021; Toivonen et al., 2016). For carabid beetles, the proportions
of individuals in the groups of poor dispersers, carnivores and adult
hibernators were calculated for each field.
The effects of crop type and production method on the commu-
nity trait values were analysed using generalized linear mixed models.
The response variables were the mean wingspan, adult habitat
breadth and overwintering stage of butterflies; the mean body size,
tongue length and spring emergence time of bumblebees and the
logit-transformed proportions of carabid beetles in the groups of poor
dispersers, carnivores and adult hibernators. The models were fitted
using the glmmTMB function of the R package glmmTMB (Brooks
et al., 2017) with Gaussian error distribution.
Two types of models were fitted. Firstly, to analyse the effects of
crop type on the insect traits, all eight crop types were compared
using models with crop type as a fixed factor, the percentage cover of
arable land in the landscape as a covariate and farm as a random fac-
tor. For the percentage cover of arable land, the radius of 250 m was
used for butterflies and carabid beetles and the radius of 500 m for
bumblebees. The landscape variables were selected by comparing the
correlations of species richness and abundance of the insect groups
with arable land and forest covers within four landscape radii
(100, 250, 500 and 1000 m; Toivonen et al., 2022). For each insect
group, a landscape variable with the highest correlation with species
richness and/or abundance was used in the analyses.
Secondly, the effects of production method and the interaction
effects of production method and crop type on the insect traits were
analysed using models with crop type, production method and their
interaction as fixed factors, arable land cover as a covariate and farm
as a random factor. These models were fitted to data from five crop
types: oat, rye, faba bean, grass ley and arable pasture. For oilseeds
and cabbage, the production methods could not be compared because
only three organic oilseed fields and no organic cabbage fields were
studied. Fallows were excluded because the management of long-
term fallows is not affected by the production method.
The residuals for the models were checked using simulateResi-
duals() function of the DHARMa package (Hartig, 2020). Due to resid-
ual heteroscedasticity in the model predicting bumblebee tongue
length with crop type, ln-transformed values of tongue length were
used in that model. Type II Wald χ2 tests with the function Anova() of
the R package car (Fox & Weisberg, 2019) were used to test the sta-
tistical significance of the parameters in the models. If the landscape
variable of arable land cover had no statistically significant effect, it
was removed from the model. In case of non-significant interaction of
production method and crop type, the interaction effect was
removed, and the final model included production method and crop
type as single factors. Values of p < 0.05 were considered statistically
significant. The sequential Bonferroni correction was not used to con-
trol the Type I errors due to the multiple tests, as it substantially
increases the risk of Type II errors (Moran, 2003). Where a significant
effect of crop type or a significant interaction effect of production
method and crop type was found, post-hoc pairwise comparisons with
4 TOIVONEN ET AL.
 14619563, 0, D
ow
nloaded from
 https://resjournals.onlinelibrary.wiley.com/doi/10.1111/afe.12629 by Duodecim Medical Publications Ltd, W
iley Online Library on [13/09/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on W
iley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
Tukey p-value adjustment for multiple comparisons were conducted
using emmeans() function of the package emmeans (Lenth, 2021) to
determine which crop types or crop type production method combi-
nations differed from each other.
RESULTS
The insect data used to calculate the community trait values consisted of
559 butterflies representing 27 species (Table S1), 835 bumblebees
representing 15 species (Table S2) and 4208 carabid beetles representing
63 species (Table S3). Several functional traits of the studied insect com-
munities in the study fields were correlated with each other based on
the community-weighted mean values (Table S4): Large mean wingspan
of butterflies was associated with butterfly overwintering in advanced
development stages (adult or pupa). For bumblebees, late mean emer-
gence time was associated with long mean tongue length and small body
size. For carabid beetles, the proportion of poor flyers was positively cor-
related with the proportion of carnivores and negatively correlated with
the proportion of adult hibernators.
Effect of crop type
Crop type affected the measured species traits of the studied insect
communities except bumblebee size and the overwintering stage of
carabid beetles (Table S5). The mean wingspan of butterflies was
shortest in fallows and longest in rye and faba bean fields (Figure 1a).
Adult habitat breadth of butterflies was narrowest in fallows and pas-
tures and broadest in oilseed and cabbage fields (Figure 1b). Fallows
had the highest share of butterflies overwintering in early develop-
ment stages (Figure 1c).
The mean tongue length of bumblebees was longest in oat and
fallow fields and shortest in oilseed and rye fields (Figure 2b). The
mean emergence time of bumblebee queens was earliest in oilseed
fields and latest in fallows (Figure 2c).
The proportions of carabid beetles with poor flight ability and car-
nivorous diet were highest in leys and fallows and lowest in cabbage
fields (Figure 3a,b). Pastures did not significantly differ from leys and
fallows in these traits (Figure 3a,b).
The cover of arable land in the surrounding landscape was signifi-
cantly related to bumblebee size, the flight ability and diet of carabid
beetles (Table S5). The mean size of bumblebees increased with
increasing arable land cover in the 500 m buffer (Figure 4). For carabid
beetles, the proportion of poor flyers and carnivores decreased with
increasing arable land cover in the 250 m buffer (Figure 4).
Effects of production method and the interaction of
production method and crop type
The overwintering stage of carabid beetles differed between the pro-
duction methods (Table S6). The proportion of adult hibernators was
higher in organically than in conventionally managed fields (Figure 5).
Interaction between crop type and production method affected
the studied species traits except butterfly habitat breadth, carabid
beetle diet and overwintering stage (Table S6). For the wingspan and
overwintering stage of butterflies, post-hoc pairwise tests found no
significant difference between the crop type-production method com-
binations due to the p-value adjustment for multiple comparisons. On
average, the wingspan of butterflies was shorter, and the proportional
abundance of species overwintering as egg or larva was higher in
organically managed oat fields than in conventional oat fields
(Figure 6a,b). In the other crop types, differences between the produc-
tion methods were small or non-existent (Figure 6a,b).
The mean size of bumblebees was largest in organic rye fields and
lowest in organic ley, oat and faba bean fields (Figure 6c). The mean
tongue length of bumblebees was longest in organic oat fields
(Figure 6d). For the emergence time of bumblebee queens, post-hoc
pairwise tests found no significant difference between the crop type
production method combinations. On average, the emergence time of
bumblebee queens was latest in organically managed pastures
(Figure 6e).
The proportion of carabid beetles with poor flight ability was
highest in conventionally managed leys and lowest in oat fields, con-
ventional pastures and organic rye and faba bean fields (Figure 6f).
The cover of arable land in the surrounding landscape was signifi-
cantly related to the wingspan and adult habitat breadth of butterflies
F I G U R E 1 Wingspan (a), adult habitat breadth (b) and
overwintering stage (c) of butterflies in eight crop types.
Overwintering stages were 1 = egg, 2 = larva, 3 = pupa and
4 = adult. Statistically significant differences between the crop types
in post-hoc tests (p < 0.05) are indicated by different letters. Mean
values are indicated by black squares. The boxes display median
values and upper and lower quartiles, with whiskers extending to
minimum and maximum values within 1.5 times the interquartile
range. Outliers are shown as circles.
INSECT FUNCTIONAL TRAITS ON ARABLE LAND 5
 14619563, 0, D
ow
nloaded from
 https://resjournals.onlinelibrary.wiley.com/doi/10.1111/afe.12629 by Duodecim Medical Publications Ltd, W
iley Online Library on [13/09/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on W
iley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
and the flight ability and diet of carabid beetles (Table S6). With
increasing arable land cover in the 250 m buffer, the mean wingspan
and habitat breadth of butterflies increased, and the proportion of
carabid beetles with poor flying ability and carnivorous diet decreased
(Figure 7).
DISCUSSION
Our results showed that crop type has an important role in shaping
the functional trait composition of insect communities on arable
land. Despite a wide array of research conducted on the influence
of agricultural land use on insect traits, few earlier studies have
focused on the effects of crop types (but see Eyre et al., 2013).
One reason for this gap may lie in the emphasis of previous
research on the effects of agricultural management intensity
F I GU R E 3 The logit-scaled proportion of carabid beetles with
poor flight ability (a), carnivorous diet (b) and overwintering as adult
(c) in eight crop types. Statistically significant differences between the
crop types in post-hoc tests are indicated by different letters. Mean
values are indicated by black squares. For description of boxplot
features, see Figure 1 legend.
F I G U R E 4 Bumblebee size (a) and the proportion of carabid
beetles with poor flight ability (b) and carnivorous diet (c) in relation
to arable land cover within 500 m radius (bumblebees) or 250 m
radius (carabid beetles). Lines depict predicted values based on the
models with all crop types and arable land cover as a statistically
significant covariate (Table S5).
F I G U R E 5 The logit-scaled proportion of carabid beetles
overwintering as adult in conventionally and organically managed
fields. Statistically significant differences are indicated by different
letters. Mean values are indicated by black squares. For description of
boxplot features, see Figure 1 legend.
F I GU R E 2 Size (a), tongue length (b) and spring emergence time
(c) of bumblebees in eight crop types. Emergence time was expressed
as month number and ranged from April (4) to June (6). Statistically
significant differences between the crop types in post-hoc tests are
indicated by different letters. Mean values are indicated by black
squares. For description of boxplot features, see Figure 1 legend.
6 TOIVONEN ET AL.
 14619563, 0, D
ow
nloaded from
 https://resjournals.onlinelibrary.wiley.com/doi/10.1111/afe.12629 by Duodecim Medical Publications Ltd, W
iley Online Library on [13/09/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on W
iley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
instead of crop type, resulting in many studies reporting functional
traits to modify insects’ responses to changes in agricultural inten-
sity (Hanson et al., 2016; Rader et al., 2014; Rusch et al., 2013).
However, changes in management intensity are usually associated
with changes in crop type, and thus, the effects of the two can eas-
ily be confounded with each other.
Importance of fallows for insect functional diversity
Among the studied crop types, long-term fallows had the most dis-
tinct trait composition of insect communities. Fallows promoted poor
dispersers, habitat and diet specialist species, and species of which life
cycle timing was not typical to cultivated fields. This strengthens the
evidence that long-term fallows have an important role in protecting
not only taxonomic but also functional diversity of insects on arable
land (Alanen et al., 2011; Feng et al., 2021; Toivonen et al., 2016), sim-
ilarly to other extensive grasslands (Börschig et al., 2013). To fully
understand the contribution of fallows to insect conservation and
insect-driven ecosystem services, more studies on functional trait dif-
ferences between fallows and cultivated fields are needed. For exam-
ple, in our study fields, the species richness of bumblebees did not
differ between oilseed fields and fallows (Toivonen et al., 2022). How-
ever, oilseed fields were strongly dominated by short-tongued and
F I GU R E 6 Butterfly wingspan (a) and overwintering stage (b); bumblebee size (c), tongue length (d) and spring emergence time (e) and the
logit-scaled proportion of carabid beetles with poor flight ability (f) in five crop types under conventional and organic production. Butterfly
overwintering stages were 1 = egg, 2 = larva, 3 = pupa and 4 = adult. Bumblebee emergence time was expressed as month number and ranged
from April (4) to June (6). Statistically significant differences between the groups in post-hoc tests are indicated by different letters. Mean values
are indicated by black squares. For description of boxplot features, see Figure 1 legend.
F I GU R E 7 Wingspan (a) and adult habitat breadth (b) of
butterflies, and the proportion of carabid beetles with poor flight
ability (c) and with carnivorous diet (d) in relation to arable land cover
within 250 m radius. Lines depict predicted values based on the
models without oilseed, cabbage and fallow fields, and with arable
land cover as a statistically significant covariate (Table S6).
INSECT FUNCTIONAL TRAITS ON ARABLE LAND 7
 14619563, 0, D
ow
nloaded from
 https://resjournals.onlinelibrary.wiley.com/doi/10.1111/afe.12629 by Duodecim Medical Publications Ltd, W
iley Online Library on [13/09/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on W
iley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
early emerging species, whereas fallows harboured more long-
tongued and late-emerging species. The latter are of special conserva-
tion interest due to their documented declines in several countries
(Bommarco et al., 2012; Dupont et al., 2011; Goulson et al., 2005).
Furthermore, the loss of long-tongued species may reduce pollination
success of flowers with long corollas with negative implications for
the yields of certain crops (Bommarco et al., 2012).
The differences between fallows and the other crop types in
the functional traits of butterfly and bumblebee communities can
be explained by several reasons. Relatively high plant diversity in
fallows (Toivonen et al., 2022) supported the diversity of floral
resources and butterfly larval host plants, which, along with the
longevity of sward and the low level of disturbance, allowed spe-
cies with narrow diet or habitat specialization and poor dispersal
capacity to colonize and persist. For dispersal capacity, the differ-
ences between the crop types were found for butterflies but not
for bumblebees. Bumblebees are relatively mobile central place for-
agers, which can efficiently find rewarding floral resources in the
landscape around their nest, as far as the resources are within their
foraging range (Osborne et al., 2008). Many butterfly species,
instead, are relatively sedentary, and thus, more dependent on per-
manent and long-term habitats (Alanen et al., 2011; Toivonen
et al., 2016). The overwintering stage of butterflies was coupled
with dispersal capacity: species overwintering in advanced devel-
opment stages had, in average, larger wingspan than species over-
wintering in earlier stages. In addition, species overwintering as egg
or larvae spend a relatively long time from autumn to spring in
stages with very limited mobility and high vulnerability to habitat
disturbances (K}orösi et al., 2022). This may explain the high propor-
tion of species overwintering in early development stages in fallows
compared to most other crop types, even perennial leys.
For bumblebees, fallows probably provided a relatively good
availability of nesting sites and floral resources throughout the season.
These resources are most critical for late-emerging bumblebee species
that, compared to early species, have harder to find free nesting sites
and suffer more from the scarcity of mid- and late-season flower
resources in contemporary agricultural landscapes (Blasi et al., 2023;
Fitzpatrick et al., 2007; Persson et al., 2015). Besides, fallows in our
study were rich in Fabaceae, such as Lathyrys pratensis L., Vicia sp. and
Trifolium sp., which are important pollen and nectar sources for bum-
blebees and, in particular, for late-emerging and long-tongued species
(Goulson et al., 2005).
In our study, fallows were situated in landscapes with higher for-
est cover and lower arable land cover than the other crop types. Due
to the landscape difference, it is not possible to reliably distinguish
the effect of fallow from the effect of forested landscape. For exam-
ple, grassland butterflies with poor dispersal capacity and narrow hab-
itat breadth have been reported to benefit from forest matrix (Berg
et al., 2011; Öckinger et al., 2012; Toivonen et al., 2017). However,
our results show that long-term fallows, as they are typically located
in our region, are highly valuable for insect functional diversity on
arable land.
Differing effects of annual and perennial crops on
insect functional traits
Among actual crop fields, long-tongued bumblebees were proportion-
ally most abundant in organic oat fields. This can be explained by the
occurrence of flowering weeds with long corolla, such as Galeopsis
sp. and Vicia sp. in some of these fields in late summer (Toivonen
et al., 2022). In organic pastures and leys, the proportional abundance
of late-emerging bumblebee species tended to be high, which may be
due to the occurrence, and sometimes abundance, of clovers and
other legumes with long flowering periods in these fields. The results
corroborate the importance of maintaining weeds at tolerable levels
(Nicholls & Altieri, 2013) and growing legumes (Bommarco
et al., 2012; Dupont et al., 2011; Goulson et al., 2005) for maintaining
bumblebee diversity on arable land.
The functional traits of carabid beetle communities were more
affected by the perenniality of the sward than any specific crop type.
Besides long-term fallows, pastures and leys promoted species with
poor flight ability and carnivorous diet, whereas annual crops were
dominated by good flyers, omnivores and herbivores. Carabid beetles
have generally lower dispersal capacity than bees and butterflies.
Flightless species can move a few hundreds of meters by walking but
dispersal over longer distances requires flying (den Boer, 1990).
Therefore, it can be expected that the occurrence of carabid beetles
in such ephemeral habitats as annual crops is restricted by species-
specific flight ability. However, the results of previous studies on the
effects of management intensity, or associated crop type, on carabid
beetles with different flight abilities have been mixed. Ribera et al.
(2001) reported higher frequency of long-winged species with
increasing land use intensity, whereas Rusch et al. (2013) showed that
the mitigation of agricultural intensification increased the proportion
of species with good flight ability, and Hanson et al. (2016) found
higher proportion of good flyers in perennial grasslands than in winter
wheat and sugar beet fields. The differences between the studies may
stem from differences in landscape composition, and in particular, the
cover of grasslands. The share of carabid beetles with good flight abil-
ity increases in grasslands with grassland fragmentation, and thus, iso-
lated grasslands are dominated by good flyers (Hanson et al., 2016;
Hendrickx et al., 2009). In our study region, landscapes are relatively
rich in perennial grasslands, fallows and field margins, which probably
allowed poor dispersers to persist in grasslands.
The high proportion of carabid beetles with carnivorous diet in
the perennial crop types contrasts with some previous studies that
have reported carnivores to be proportionally more common in annual
crops than in perennial grasslands and grassy field margins (Birkhofer
et al., 2014; Hanson et al., 2016; but see Eyre et al., 2013). Further-
more, in an earlier study in Finland, perennial fallows were character-
ized by a high number of individuals of the genus Amara, which are
predominantly herbivorous (Kinnunen & Tiainen, 1999). The differ-
ences between the studies are likely to reflect variation in the avail-
ability of food resources for carabid beetles within crop types. Annual
crops can offer ample prey in the form of arthropod pests such as
8 TOIVONEN ET AL.
 14619563, 0, D
ow
nloaded from
 https://resjournals.onlinelibrary.wiley.com/doi/10.1111/afe.12629 by Duodecim Medical Publications Ltd, W
iley Online Library on [13/09/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on W
iley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
aphids, which can attract a high number of predatory carabid beetles
to the fields (Holland et al., 2004; Knapp et al., 2019). If the sampling
of carabid beetles coincides with high pest abundance (as in Birkhofer
et al., 2014), carnivores may be particularly abundant. Similarly, weeds
increase the availability of plant-derived food resources for herbivores
and omnivores (Diehl et al., 2012). In our study, 65% of the annual
crop fields were managed without chemical weed control, and 19%
were undersown with grass-clover mixtures, sometimes supplemen-
ted with other annual and/or perennial plants. The relatively high
plant diversity in the annual crop types probably increased the abun-
dance of carabid beetles with omnivorous and herbivorous diets. Die-
tary traits of carabid beetle communities have implications for pest
and weed control. From pest control perspective, supporting carni-
vores in annual crop types is important, as annual crops are typically
associated with higher pest damage than perennial crops. This could
be done by creating perennial habitats next to annual crops, for exam-
ple, by widening field margins or creating in-field perennial strips
(Kromp, 1999; Toivonen et al., 2018).
The overwintering stage of carabid beetles did not differ between
perennial and annual crop types. Based on earlier studies, adult hiber-
nators may be more resistant to intensive land use and soil distur-
bance than species overwintering in their larval form (Kinnunen &
Tiainen, 1999; Müller et al., 2022; Ribera et al., 2001). However, till-
age regime may be a more important determinant of overwintering
success than crop perenniality: species vulnerable to conventional
ploughing may tolerate reduced tillage and favour no-till systems
(Müller et al., 2022). Tillage also modifies microclimate at soil surface.
In spring and early summer, ploughed fields may provide suitable
microclimate for breeding for adult hibernating species, which are
mostly day-active and thermophilic (Hatten et al., 2007; Kinnunen &
Tiainen, 1999). In our study, organically managed fields were mostly
ploughed, whereas, in conventionally production, the main tillage
method was reduced tillage. This possibly explains why adult hiberna-
tors were relatively more abundant in organically than in convention-
ally managed fields. The high proportion of adult hibernators may
benefit pest control services in organic crop fields: adult hibernators
are active in spring and early summer, when pest numbers are low,
and thus, they can effectively prevent or delay the pest population
growth (Welch & Harwood, 2014).
Our study focused on the effects of crop type and production
method on dominant traits in insect communities. Besides dominant
traits, functional diversity of the traits influences ecosystem resilience
and ecosystem service provision (Greenop et al., 2018; Woodcock
et al., 2019) and may also differ between crop types and production
methods (Feng et al., 2021; Goded et al., 2019). On a landscape scale,
increasing crop diversity, along with increasing the area of crop types
with the most distinct trait composition, is likely to support functional
diversity across insect taxa. Further studies comparing both dominant
traits and trait diversity of insects across various crop types and pro-
duction methods would contribute to our understanding of how ara-
ble land use changes affect insect conservation and ecosystem
functioning.
Role of the surrounding landscape
Besides crop type and production method, surrounding
landscape affected the functional traits of insect communities. Land-
scapes with low arable land cover supported poor dispersers as well
as carnivorous carabid beetles and habitat specialist butterflies. This
highlights the importance of non-arable habitats in the landscape in
buffering insect communities, and especially poor-dispersing and spe-
cialist species, against the adverse effects of intensive arable farming
(Gámez-Virués et al., 2015; K}orösi et al., 2022; Tscharntke
et al., 2005). Landscapes with low arable land cover supported largely
similar traits as long-term fallows, suggesting that the lack of non-
arable habitats in intensively cultivated landscapes can be partly com-
pensated by increasing fallow area.
CONCLUSIONS
Our study shows that crop choice affects functional trait composi-
tions of butterfly, bumblebee and carabid beetle communities on
boreal arable land, which has potential implications for pollination and
pest control services. Long-term fallows harboured the most distinct
insect communities, with high relative abundance of poor dispersers,
specialist species and species with differing life cycle timing compared
to cultivated fields. This shows the high value of long-term fallows in
conserving functional diversity of insects on boreal arable land.
Among cultivated fields, annual and perennial crops differed from
each other especially in terms of functional traits of carabid beetle
communities. However, differences between the results of this and
previous studies suggest that the effects of agricultural management
on carabid beetle communities are strongly context-dependent. The
effects of production method were less clear than those of crop type.
Still, certain management practices that are influenced by, yet not
entirely tied to production method, such as ploughing, sowing
legumes in grasslands or refraining from the use of herbicides, proba-
bly affect functional traits of insect communities. To promote insect
species sensitive to agricultural disturbances and land use changes,
and their associated ecosystem services, it is recommended to
increase the area of long-term fallows and perennial crops and
enhance within-field plant diversity while maintaining landscape
heterogeneity.
AUTHOR CONTRIBUTIONS
Marjaana Toivonen: Conceptualization; data curation; formal analysis;
funding acquisition; investigation; methodology; project ad-
ministration; visualization; writing – original draft. Erja Huusela:
Conceptualization; funding acquisition; investigation; methodology;
writing – review and editing. Terho Hyvönen: Conceptualization;
funding acquisition; investigation; methodology; writing – review and
editing. Ari Järvinen: Investigation; writing – review and editing.
Mikko Kuussaari: Conceptualization; funding acquisition; methodol-
ogy; project administration; writing – review and editing.
INSECT FUNCTIONAL TRAITS ON ARABLE LAND 9
 14619563, 0, D
ow
nloaded from
 https://resjournals.onlinelibrary.wiley.com/doi/10.1111/afe.12629 by Duodecim Medical Publications Ltd, W
iley Online Library on [13/09/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on W
iley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
ACKNOWLEDGEMENTS
We would like to thank all the farmers who let us collect data on their
fields; Jaana Grahn, Anna-Elina Karimaa, Sanna Keronen, Elina Laurén,
Sami Lindgren and Anne Muotila who participated in fieldwork and
Susu Rytteri and Juha Pöyry who helped to find information on bum-
blebee species traits. Ben Woodcock and two anonymous reviewers
provided valuable comments on the manuscript. This work was sup-
ported by the Strategic Research Council of the Academy of Finland
(decision no. 352638).
CONFLICT OF INTEREST STATEMENT
The authors declare no conflicts of interest.
DATA AVAILABILITY STATEMENT
The data that support the findings of this study is available in the
Knowledge Network for Biocomplexity (KNB) Data Repository at
https://doi.org/10.5063/F11N7ZK5.
ORCID
Marjaana Toivonen https://orcid.org/0000-0002-9215-8643
REFERENCES
Alanen, E.-L., Hyvönen, T., Lindgren, S., Härmä, O. & Kuussaari, M. (2011)
Differential responses of bumblebees and diurnal Lepidoptera to
vegetation succession in long-term set-aside. Journal of Applied Ecol-
ogy, 48, 1251–1259.
Barbieri, P., Pellerin, S., Seufert, V. & Nesme, T. (2019) Changes in crop
rotations would impact food production in an organically farmed
world. Nature Sustainability, 2, 378–385.
Berg, A., Ahrne, K., Öckinger, E., Svensson, R. & Söderstrom, B. (2011) But-
terfly distribution and abundance is affected by variation in the
Swedish forest–farmland landscape. Biological Conservation, 144,
2819–2831.
Birkhofer, K., Wolters, V. & Diekötter, T. (2014) Grassy margins along
organically managed cereal fields foster trait diversity and taxonomic
distinctness of arthropod communities. Insect Conservation and Diver-
sity, 7, 274–287.
Blasi, M., Carrié, R., Fägerström, C., Svensson, E. & Persson, A.S. (2023)
Historical and citizen-reported data show shifts in bumblebee phe-
nology over the last century in Sweden. Biodiversity and Conservation,
32, 1523–1547.
Boetzl, F.A., Krauss, J., Heinze, J., Hoffmann, H., Juffa, J., König, S. et al.
(2021) A multitaxa assessment of the effectiveness of agri-
environmental schemes for biodiversity management. Proceedings of
the National Academy of Sciences of the United States of America, 118,
e2016038118.
Bommarco, R., Lundin, O., Smith, H.G. & Rundlöf, M. (2012) Drastic
historic shifts in bumble-bee community composition in Sweden. Pro-
ceedings of the Royal Society B, 279, 309–315.
Börschig, C., Klein, A.M., von Wehrden, H. & Krauss, J. (2013) Traits of
butterfly communities change from specialist to generalist character-
istics with increasing land-use intensity. Basic and Applied Ecology,
14, 547–554.
Brooks, M.E., Kristensen, K., van Benthem, K.J., Magnusson, A.,
Berg, C.W., Nielsen, A. et al. (2017) glmmTMB balances speed and
flexibility among packages for zero-inflated generalized linear mixed
modeling. The R Journal, 9, 378–400.
den Boer, P.J. (1990) Density limits and survival of local populations in
64 carabid species with different powers of dispersal. Journal of Evo-
lutionary Ecology, 3, 19–48.
Diehl, E., Wolters, V. & Birkhofer, K. (2012) Arable weeds in organically
managed wheat fields foster carabid beetles by resource- and
structure-mediated effects. Arthropod-Plant Interactions, 6, 75–82.
Dupont, Y.L., Damgaard, C. & Simonsen, V. (2011) Quantitative historical
change in bumblebee (Bombus spp.) assemblages of red clover fields.
PLoS One, 6, e25172.
Ehlers, B.K., Bataillon, T. & Damgaard, C.F. (2021) Ongoing decline in
insect-pollinated plants across Danish grasslands. Biology Letters, 17,
20210493.
Esri. (2020) ArcGIS Desktop: Release 10.8.1. Redlands, CA: Environmental
Systems Research Institute.
European Commission. (2020) Communication from the Commission to
the European Parliament, the Council, the European Economic and
Social Committee and the Committee of the Regions: a Farm to Fork
Strategy for a fair, healthy and environmentally-friendly food system.
COM(2020) 381 final. Brussels, 20.5.2020.
Eurostat. (2016) Agriculture, forestry and fishery statistics – 2016 edition.
Luxembourg: European Union. Available at: https://ec.europa.eu/
eurostat/documents/3217494/7777899/KS-FK-16-001-EN-N.pdf/
cae3c56f-53e2-404a-9e9e-fb5f57ab49e3?t=1484314012000
[Accessed 11th March 2024].
Eyre, M.D., Luff, M.L. & Leifert, C. (2013) Crop, field boundary, productiv-
ity and disturbance influences on ground beetles (Coleoptera, Carabi-
dae) in the agroecosystem. Agriculture Ecosystems & Environment,
165, 60–67.
Feng, L., Arvidsson, F., Smith, H.G. & Birkhofer, K. (2021) Fallows and per-
manent grasslands conserve the species composition and functional
diversity of carabid beetles and linyphiid spiders in agricultural land-
scapes. Insect Conservation and Diversity, 14, 825–836.
Finnish Environment Institute. (2018) CORINE land cover 2018 dataset.
Available at: https://www.avoindata.fi/data/en_GB/dataset/corine-
maanpeite-2018 [Accessed 11th March 2024].
Finnish Food Authority. (2021) Peltolohkot 2020 (Field parcels 2020).
[dataset].
Fitzpatrick, U., Murray, T.E., Paxton, R.J., Breen, J., Cotton, D.,
Santorum, V. et al. (2007) Rarity and decline in bumblebees—a test
of causes and correlates in the Irish fauna. Biological Conservation,
136, 185–194.
Fox, J. & Weisberg, S. (2019) An R companion to applied regression, 3rd edi-
tion. Thousand Oaks, CA: Sage.
Gámez-Virués, S., Perovic, D.J., Gossner, M.M., Börschig, C.,
Blüthgen, N., de Jong, H. et al. (2015) Landscape simplification fil-
ters species traits and drives biotic homogenization. Nature Commu-
nications, 6, 8568.
Geiger, F., Bengtsson, J., Berendse, F., Weisser, W.W., Emmerson, M.,
Morales, M.B. et al. (2010) Persistent negative effects of pesticides
on biodiversity and biological control potential on European farm-
land. Basic and Applied Ecology, 11, 97–105.
Goded, S., Ekroos, J., Azcárate, J.G., Guitián, J.A. & Smith, H.G. (2019)
Effects of organic farming on plant and butterfly functional diversity
in mosaic landscapes. Agriculture, Ecosystems & Environment, 284,
106600.
Gossner, M., Lewinsohn, T., Kahl, T., Grassein, F., Boch, S., Prati, D. et al.
(2016) Land-use intensification causes multitrophic homogenization
of grassland communities. Nature, 540, 266–269.
Goulson, D. & Darvill, B. (2004) Niche overlap and diet breadth in bumble-
bees; are rare species more specialized in their choice of flowers?
Apidologie, 35, 55–63.
Goulson, D., Hanley, M.E., Darvill, B., Ellis, J.S. & Knight, M.E. (2005)
Causes of rarity in bumblebees. Biological Conservation, 122, 1–8.
Goulson, D., Lye, G.C. & Darvill, B. (2008) Diet breadth, coexistence and
rarity in bumblebees. Biodiversity and Conservation, 17, 3269–3288.
Greenleaf, S.S., Williams, N.M., Winfree, R. & Kremen, C. (2007) Bee forag-
ing ranges and their relationship to body size. Oecologia, 153,
589–596.
10 TOIVONEN ET AL.
 14619563, 0, D
ow
nloaded from
 https://resjournals.onlinelibrary.wiley.com/doi/10.1111/afe.12629 by Duodecim Medical Publications Ltd, W
iley Online Library on [13/09/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on W
iley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
Greenop, A., Woodcock, B.A., Wilby, A., Cook, S.M. & Pywell, R.F. (2018)
Functional diversity positively affects prey suppression by inverte-
brate predators: a meta-analysis. Ecology, 99, 1771–1782.
Guenay-Greunke, Y., Trager, H., Bohan, D.A., Traugott, M. &
Wallinger, C. (2023) Consumer identity but not food availability
affects carabid diet in cereal crops. Journal of Pest Science In press,
97, 281–296.
Hanson, H.I., Palmu, E., Birkhofer, K., Smith, H.G. & Hedlund, K. (2016)
Agricultural land use determines the trait composition of ground bee-
tle communities. PLoS One, 11, e0146329.
Hartig, F. (2020) DHARMa: residual diagnostics for hierarchical (multi-
level/mixed) regression models. R package version 0.3.2.0.
Hatten, T.D., Bosque-Pérez, N.A., Labonte, J.R., Guy, S.O. &
Eigenbrode, S.D. (2007) Effects of tillage on the activity density and
biological diversity of carabid beetles in spring and winter crops.
Environmental Entomology, 36, 356–368.
Hendrickx, F., Maelfait, J.P., Desender, K., Aviron, S., Bailey, D.,
Diekotter, T. et al. (2009) Pervasive effects of dispersal limitation on
within- and among-community species richness in agricultural land-
scapes. Global Ecology and Biogeography, 18, 607–616.
Hendrickx, F., Maelfait, J.-P., Van Wingerden, W., Schweiger, O.,
Speelmans, M., Aviron, S. et al. (2007) How landscape structure,
land-use intensity and habitat diversity affect components of total
arthropod diversity in agricultural landscapes. Journal of Applied Ecol-
ogy, 44, 340–351.
Holland, J.M., Winder, L., Woolley, C., Alexander, C.J. & Perry, J.N. (2004)
The spatial dynamics of crop and ground active predatory arthropods
and their aphid prey in winter wheat. Bulletin of Entomological
Research, 94, 419–431.
Kinnunen, H. & Tiainen, J. (1999) Carabid distribution in a farmland
mosaic: the effect of patch type and location. Annales Zoologici Fen-
nici, 36, 149–158.
Knapp, M., Seidl, M., Knappová, J., Macek, M. & Saska, P. (2019) Temporal
changes in the spatial distribution of carabid beetles around arable
field-woodlot boundaries. Scientific Reports, 9, 8967.
Komonen, A., Grapputo, A., Kaitala, V., Kotiaho, J.S. & Päivinen, J. (2004)
The role of niche breadth, resource availability and range position on
the life history of butterflies. Oikos, 105, 41–54.
K}orösi, A´., Dolek, M., Nunner, A., Lang, A. & Theves, F. (2022) Pace of life
and mobility as key factors to survive in farmland – relationships
between functional traits of diurnal Lepidoptera and landscape struc-
ture. Agriculture, Ecosystems & Environment, 334, 107978.
Kovács-Hostyánszki, A., K}orösi, A´., Orci, K.M., Batáry, P. & Báldi, A. (2011)
Set-aside promotes insect and plant diversity in a Central European
country. Agriculture, Ecosystems & Environment, 141, 296–301.
Kromp, B. (1999) Carabid beetles in sustainable agriculture: a review on
pest control efficacy, cultivation impacts and enhancement. Agricul-
ture, Ecosystems & Environment, 74, 187–228.
Kuussaari, M., Saarinen, M., Korpela, E.L., Pöyry, J. & Hyvönen, T. (2014)
Higher mobility of butterflies than moths connected to habitat suit-
ability and body size in a release experiment. Ecology and Evolution,
4, 3800–3811.
Kuussaari, M., Toivonen, M., Heliölä, J., Pöyry, J., Mellado, J., Ekroos, J.
et al. (2021) Butterfly species’ responses to urbanization: differing
effects of human population density and built-up area. Urban Ecosys-
tem, 24, 515–527.
Lenth, R.V. (2021) emmeans: estimated marginal means, aka least-squares
means. R package version 1.5.4.
Lindroth, C.H. (1985, 1986) The Carabidae (Coleoptera) of Fennoscandia and
Denmark. Fauna Entomologica Scandinavica 15, part 1, part 2. Copen-
hagen: Scandinavian Science Press.
Martínez-Núñez, C., Kleijn, D., Ganuza, C., Heupink, D., Raemakers, I.,
Vertommen, W. et al. (2022) Temporal and spatial heterogeneity of
semi-natural habitat, but not crop diversity, is correlated with land-
scape pollinator richness. Journal of Applied Ecology, 59, 1258–1267.
Martins, K.T., Albert, C.H., Lechowicz, M.J. & Gonzalez, A. (2018) Comple-
mentary crops and landscape features sustain wild bee communities.
Ecological Applications, 28, 1093–1105.
Marttila, O., Haahtela, T., Aarnio, H. & Ojalainen, P. (1990) Suomen päivä-
perhoset [The Finnish butterflies]. Helsinki: Kirjayhtymä.
Moran, M.D. (2003) Arguments for rejecting the sequential Bonferroni in
ecological studies. Oikos, 102, 403–405.
Müller, P., Neuhoff, D., Nabel, M., Schiffers, K. & Döring, T.F. (2022) Tillage
effects on ground beetles in temperate climates: a review. Agronomy
for Sustainable Development, 42, 65.
Nicholls, C.I. & Altieri, M.A. (2013) Plant biodiversity enhances bees and
other insect pollinators in agroecosystems. A review. Agronomy for
Sustainable Development, 33, 257–274.
Öckinger, E., Bergman, K.O., Franzen, M., Kadlec, T., Krauss, J.,
Kuussaari, M. et al. (2012) The landscape matrix modifies the effect
of habitat fragmentation in grassland butterflies. Landscape Ecology,
27, 121–131.
Osborne, J.L., Martin, A.P., Carreck, N.L., Swain, J.L., Knight, M.E.,
Goulson, D. et al. (2008) Bumblebee flight distances in relation to the
forage landscape. Journal of Animal Ecology, 77, 406–415.
Persson, A.S., Rundlöf, M., Clough, Y. & Smith, H.G. (2015) Bumble bees
show trait-dependent vulnerability to landscape simplification. Biodi-
versity and Conservation, 24, 3469–3489.
Pollard, E. & Yates, T. (1993) Monitoring butterflies for ecology and conserva-
tion. London: Chapman and Hall.
Rader, R., Bartomeus, I., Tylianakis, J.M. & Laliberté, E. (2014) The winners
and losers of land use intensification: pollinator community disas-
sembly is non-random and alters functional diversity. Diversity and
Distributions, 20, 908–917.
Ranta, E. & Lundberg, H. (1980) Resource partitioning in bumblebees: the
significance of differences in proboscis length. Oikos, 35, 298–302.
Ribera, I., Dolédec, S., Downie, I.S. & Foster, G.N. (2001) Effect of land dis-
turbance and stress on species traits of ground beetle assemblages.
Ecology, 82, 1112–1129.
Rusch, A., Bommarco, R., Chiverton, P., Öberg, S., Wallin, H., Wiktelius, S.
et al. (2013) Response of ground beetle (Coleoptera, Carabidae) com-
munities to changes in agricultural policies in Sweden over two
decades. Agriculture, Ecosystems & Environment, 176, 63–69.
Scheuchl, E. & Willner, W. (2016) Taschenlexikon der Wildbienen Mitteleuro-
pas. Wiebelsheim: Quelle & Meyer.
Sekar, S. (2012) A meta-analysis of the traits affecting dispersal ability in
butterflies: can wingspan be used as a proxy? Journal of Animal Ecol-
ogy, 81, 174–184.
Söderman, G. & Leinonen, R. (2003) Suomen mesipistiäiset ja niiden uhana-
laisuus. Helsinki: Tremex Press Oy.
Stein-Bachinger, K., Gottwald, F., Haub, A. & Schmidt, E. (2021) To what
extent does organic farming promote species richness and abun-
dance in temperate climates? A review. Organic Agriculture, 11, 1–12.
Toivonen, M., Herzon, I. & Kuussaari, M. (2015) Differing effects of fallow
type and landscape structure on the occurrence of plants, pollinators
and birds on environmental fallows in Finland. Biological Conserva-
tion, 181, 36–43.
Toivonen, M., Herzon, I. & Kuussaari, M. (2016) Community composition
of butterflies and bumblebees in fallows: niche breadth and dispersal
capacity modify responses to fallow type and landscape. Journal of
Insect Conservation, 20, 23–34.
Toivonen, M., Huusela, E., Hyvönen, T., Marjamäki, P., Järvinen, A. &
Kuussaari, M. (2022) Effects of crop type and production method on
arable biodiversity in boreal farmland. Agriculture, Ecosystems & Envi-
ronment, 337, 108061.
Toivonen, M., Huusela-Veistola, E. & Herzon, I. (2018) Perennial fallow
strips support biological pest control in spring cereal in Northern
Europe. Biological Control, 121, 109–118.
Toivonen, M., Peltonen, A., Herzon, I., Heliölä, J., Leikola, N. &
Kuussaari, M. (2017) High cover of forest increases the abundance
INSECT FUNCTIONAL TRAITS ON ARABLE LAND 11
 14619563, 0, D
ow
nloaded from
 https://resjournals.onlinelibrary.wiley.com/doi/10.1111/afe.12629 by Duodecim Medical Publications Ltd, W
iley Online Library on [13/09/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on W
iley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
of most grassland butterflies in boreal farmland. Insect Conservation
and Diversity, 10, 321–330.
Tscharntke, T., Klein, A.M., Kruess, A., Steffan-Dewenter, I. & Thies, C. (2005)
Landscape perspectives on agricultural intensification and biodiversity:
ecosystem service management. Ecology Letters, 8, 857–874.
Wagner, D.L. (2020) Insect declines in the Anthropocene. Annual Review of
Entomology, 65, 457–480.
Welch, K.D. & Harwood, J.D. (2014) Temporal dynamics of natural enemy-
pest interactions in a changing environment. Biological Control, 75,
18–27.
Woodcock, B.A., Garratt, M.P.D., Powney, G.D., Shaw, R.F., Osborne, J.L.,
Soroka, J. et al. (2019) Meta-analysis reveals that pollinator func-
tional diversity and abundance enhance crop pollination and yield.
Nature Communications, 10, 1481.
Woodcock, B.A., Redhead, J., Vanbergen, A.J., Hulmes, L., Hulmes, S.,
Peyton, J. et al. (2010) Impact of habitat type and landscape struc-
ture on biomass, species richness and functional diversity of ground
beetles. Agriculture, Ecosystems & Environment, 139, 181–186.
Yvoz, S., Cordeau, S., Zuccolo, C. & Petit, S. (2020) Crop type and within-
field location as sources of intraspecific variations in the phenology
and the production of floral and fruit resources by weeds. Agriculture,
Ecosystems & Environment, 302, 107082.
SUPPORTING INFORMATION
Additional supporting information can be found online in the Support-
ing Information section at the end of this article.
Table S1. The list of butterfly species observed in 78 fields, their func-
tional traits and the number of individuals.
Table S2. The list of bumblebee species observed in 78 fields, their
functional traits and the number of individuals.
Table S3. The list of carabid beetle species sampled in 75 fields, their
functional traits and the number of individuals.
Table S4. Correlations between the functional traits of bumblebee,
butterfly and carabid beetle communities in 78 study fields. The corre-
lations were calculated using community-weighted means for the
fields. Missing values were replaced by the mean of the non-missing
values of the respective trait. In the trait names, ‘B’ refer to bumble-
bee traits, ‘L’ to butterfly traits and ‘C’ to carabid beetle traits. Statis-
tically significant correlations (p < 0.05) are bolded.
Table S5. χ2 test results, and marginal and conditional R2 for the
models with all crop types included.
Table S6. χ2 test results, and marginal and conditional R2 for the
models without oilseed, cabbage and fallow fields.
How to cite this article: Toivonen, M., Huusela, E., Hyvönen,
T., Järvinen, A. & Kuussaari, M. (2024) Crop type rather than
production method determines functional trait composition of
insect communities on arable land in boreal agricultural
landscapes. Agricultural and Forest Entomology, 1–12. Available
from: https://doi.org/10.1111/afe.12629
12 TOIVONEN ET AL.
 14619563, 0, D
ow
nloaded from
 https://resjournals.onlinelibrary.wiley.com/doi/10.1111/afe.12629 by Duodecim Medical Publications Ltd, W
iley Online Library on [13/09/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on W
iley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License