Strip cropping shows promising increases in ground beetle community diversity compared to monocultures

Insects are the largest and most diverse group of animals, comprising approximately 80% of all animal species. They inhabit nearly every place on Earth and play important and varied roles in ecosystems, from serving as a food source for other animals and as recyclers of organic matter and nutrients, to acting as crop pollinators and pest controllers.

Sadly, insect biodiversity is declining worldwide, with agriculture being a major contributor to this decline. For example, monoculture farming is a common form of farming where only one crop species is grown at a time, providing limited habitats and food resources for insects. By cultivating a diversity of crops in narrow, alternating strips, a technique known as strip cropping, farmers might make fields more suitable for insects, without reducing crop yield and productivity. However, so far, it was unknown if strip cropping can indeed increase insect biodiversity.

Croijmans et al. set out to investigate whether strip cropping can increase the biodiversity of ground beetles. Ground beetles play a key role in agricultural ecosystems, preying on common insect pests and weeds. They are sensitive to changes in farming practices and are often used as an indicator of agricultural sustainability. For this purpose, the researchers analysed four years of data from four organically managed experimental farms in the Netherlands, which included a diverse set of crops.

Croijmans et al. found that strip-cropped fields have more beetle species and more individual beetles than monocultures. As different ground beetle communities have natural preferences for specific crops, it is thought that the higher number of ground beetle species in strip-cropped fields is mostly due to the combination of two crop-related communities, rather than species unique to strip cropping. For example, if cabbage is strip-cropped with wheat, one would mostly find the ground beetle species corresponding to both cabbage and wheat, but few additional species. Interestingly, some ground beetle species preferred strip-cropped fields, while others preferred monocultures.

In conclusion, strip cropping can be a strategy to increase ground beetle biodiversity without losses to crop production. Therefore, farmers wanting to increase biodiversity might consider this approach instead of standard monocultures. Many biodiversity-increasing measures, such as flower strips or hedgerows, take up land that might otherwise be used for crop production. Strip cropping allows a more biodiverse field while keeping all land in production, making it a biodiversity measure that enables farmers to maintain the same level of crop production.

Insects account for 80% of the animal species in the world and, therefore, the recently reported unprecedented rate of insect decline is cause for alarm about the state of biodiversity on Earth (Dirzo et al., 2014; Hallmann et al., 2017; van Klink et al., 2020). An array of biodiversity metrics provides strong evidence for declines of especially terrestrial insects across all continents (van Klink et al., 2020). This includes declines in abundance of both common and rare species (Hallmann et al., 2017; Pilotto et al., 2020; Seibold et al., 2019; van Klink et al., 2024) and in total species richness and changes in assembly composition (Blowes et al., 2022; Wagner et al., 2021). Insects are essential for crop production through their role in decomposition, pollination, pest control, and sustaining food webs. Therefore, erosion of insect communities can have potentially devastating effects on ecosystem functioning, provision of ecosystem services, and ultimately on human civilization (Dirzo et al., 2014). The main drivers of insect biodiversity decline are habitat loss due to conversion to agriculture, pollution, invasive species, and climate change (Díaz et al., 2019; Müller et al., 2024; Wagner et al., 2021). Strategies for biodiversity conservation in conjunction with adequate food production require understanding of how biodiversity responds to agricultural management (Cozim-Melges et al., 2024; Mupepele et al., 2021; Saunders, 2020). Ideally, sustainable agricultural practices retain yield and enhance biodiversity, preventing the need to convert natural habitats to agricultural land to maintain food production (Tscharntke et al., 2021).

Increasing crop heterogeneity can facilitate biodiversity conservation in highly productive agricultural landscapes without compromising yield (Martin Guay et al., 2018; Sirami et al., 2019). Crop diversification can enhance niche complementarity by creating heterogeneous habitats and increasing availability and diversity of resources (Lichtenberg et al., 2017; Tamburini et al., 2020). A promising crop diversification strategy is strip cropping, where crops are grown in alternating strips, wide enough for using standard agricultural machines yet narrow enough to facilitate ecological interactions among crops (Ditzler et al., 2021). Crops that are grown in strips may benefit from increased resource use efficiency and the suppression of pests and diseases (Croijmans et al., 2024; Karssemeijer et al., 2024; Rakotomalala et al., 2023) without major yield compromises (Campanelli et al., 2023; Juventia and van Apeldoorn, 2024; van Oort et al., 2020). Growing multiple crops on a field may foster a larger diversity of organisms than monocultures through greater plant species richness that cascades into richer herbivore and predator communities (Crutsinger et al., 2006; Cuperus et al., 2023). Moreover, the expected increase in available and potentially complementary niches within the agricultural field due to higher spatial diversity in crops can result in admixture of communities related to individual crops (Hummel et al., 2012), or the creation of completely new communities by enhanced richness of agriculture-related species, and/or the occurrence of species rarely found in agricultural fields (Rischen et al., 2021). So far, it is not well understood if and how insect communities respond to strip cropping across distinct crops.

Here, we present data of 4 years of pitfall trapping of ground beetles at several moments during the growing season in 14 crops at four organic experimental farms across the Netherlands where strip cropping was compared to monocultures. Ground beetle communities are sensitive to changes in farming practices and are frequently used to examine agricultural sustainability (Holland and Luff, 2000; Makwela et al., 2023; Turin, 2022). Furthermore, ground beetle species are important for maintaining ecological functions as they comprise scavengers and predators of (weed) seeds, detritivores (e.g., collembolas and earthworms), and herbivores (e.g., aphids and caterpillars). We first examine whether strip cropping fields have greater ground beetle activity density, species richness, evenness, and diversity than monocultural fields. We also test for 12 abundant ground beetle genera whether their activity density is higher in strip cropping than in monocultures. Lastly, we evaluate whether ground beetle community changes are caused by admixture of communities, whether these assemblages promote species associated with agricultural or with natural ecosystems, and whether they contain rare species.

A total of 48,108 ground beetles belonging to 71 species were caught using pitfall traps over 4 years at four different organically managed experimental farms in The Netherlands: 40,153 at Almere; 3777 at Lelystad; 1126 at Valthermond; and 3052 at Wageningen (Supplementary file 1 and 2).

Strip cropping enhances ground beetle richness

Strip cropping fields had on average 15% higher ground beetle taxonomic richness than monoculture fields after rarefaction to the number of samples of the least-sampled crop configuration (β₀=0.151, SE = 0.044, p<0.001; Figure 1c and d). However, strip cropping fields did not harbor more species than monocultural fields with the highest ground beetle richness (β₀=–0.008, SE = 0.037, p=0.821; Figure 1d). The difference in field-level taxonomic richness could not be explained by an increase in the number of ground beetle species per crop in strip cropping compared to monocultures. At crop level, the 5% increase in ground beetle taxonomic richness in strip cropping was not statistically significant (Figure 1f). Similarly, crop-level absolute evenness, inverse Simpson index, and Shannon entropy did not differ significantly among crop configurations (Figure 1h–j, Figure 1—figure supplements 1 and 2). The effect of crop configuration on crop-level taxonomic richness was variable and was not associated with location or crop species. The effect ranged from 56% more species in potato in monoculture at Wageningen in 2022 to 136% more species in barley in strip cropping at Wageningen in 2020 (Figure 1k).

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Crop configuration alters abundance of abundant genera

We tested how crop configuration affected the abundance of twelve abundant ground beetle genera (Figure 2—source data 1). We analyzed this separately per location as some genera only occurred at specific locations. Four genera were more abundant in strip-cropped fields in at least one location (Anchomenus, Bembidion, Harpalus, and Nebria), whereas four genera were more abundant in monocultures in at least one location (Amara, Calathus, Pterostichus, and Trechus) (Figure 2). The other four common genera (Blemus, Clivina, Loricera, and Poecilus) were not significantly influenced by crop configuration. Furthermore, no ground beetle genus showed significantly contrasting responses to crop configuration among locations.

Figure 2

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Figure 2—source data 1

Abundances of the 12 most abundant ground beetle genera in monoculture and strip cropping fields in four locations.

The first value indicates the estimated mean, in brackets the confidence interval and letters indicate significant difference between monocultures and strips per location. A stripe indicates that there was no significant difference for the genus for that location. Dark cells indicate that two or fewer individuals were found at this location, and the location was excluded from the model. At Wageningen, we only found a few Anchomenus in the monoculture, whereas we found none in the strip-cropped field. As there was no variation in the strip-cropped field, this location could not be included in the model for this genus, and the mean here indicates the actual mean. The model for Pterostichus did not fit well when the data from 2020 Almere were included, as catches were much higher in this year than in the other years. Therefore, we conducted separate analyses for 2020 and 2021–2022.

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It is well established that different crop types have distinct ground beetle communities (Eyre et al., 2009; Holland and Luff, 2000) and that increasing habitat diversity by including multiple crops in a field can enhance ground beetle diversity (Cuperus et al., 2024; Puliga et al., 2023). Our study shows that strip cropping increased field-level ground beetle richness by 15%. However, ground beetle communities within the same crop in a strip or in monoculture were mostly similar. This indicates that the 15% increase in richness at the field level can be mostly attributed to the higher number of crops in strip-cropped fields that harbored crop-related ground beetle communities, and that there was only limited mixing of ground beetle species among crops in strip crops. This is in line with earlier findings that ground beetle movement is reduced by crop edges (Allema et al., 2014; Anderson et al., 2024). Further research on movement behaviors of ground beetles at crop edges might help explain how ground beetles distribute themselves within a strip cropping field, and whether they utilize the different resources provided by a more diverse cropping system.

The 30% higher ground beetle activity density in strip cropping fields compared to monoculture fields may be explained by a more stable and diverse habitat with refuges and alternative resources in strip crops (Ratnadass et al., 2012). Crop diversification enhances prey biomass for ground beetles (Lichtenberg et al., 2017) and increases weed seed richness as compared to monocultures (Ditzler et al., 2023), both of which reduce bottom-up control by increased food provision (Carbonne et al., 2022). Alternatively, the increase in activity density might be caused by higher movement of ground beetles, which in turn can be the consequence of food starvation (Wallin and Ekbom, 1994). Indeed, several papers show reduced abundances of herbivorous insects in strip- and intercropping (Alarcón‐Segura et al., 2022; Cuperus et al., 2023; Rakotomalala et al., 2023), which may be potential prey for ground beetles (Turin, 2000). Furthermore, strip cropping systems may support high herbivore reproduction in combination with a high predation pressure resulting in herbivore populations dominated by early life stages (Karssemeijer et al., 2024). Therefore, strip cropping may favor ground beetles that predate on, for instance, herbivore eggs, such as Bembidion spp. and Anchomenus dorsalis, which were more abundant in strip cropping (Finch, 1996), and disadvantaging ground beetles that feed on larger prey, such as Pterostichus melanarius, which was less abundant in strip cropping. Multi-taxa evaluation of future strip cropping studies will be a valuable approach to increase understanding of biomass flows and trophic interactions within diverse agricultural systems.

The 15% increase in ground beetle species richness through strip cropping is mostly caused by an increase in species (relatively) common to agricultural fields, rather than an increase in rare species or species otherwise common in other habitat types. A keystone species here might be P. melanarius, the most common Pterostichus species in our samples. P. melanarius can be very dominant in pitfall traps, as we regularly found hundreds of individuals per trap. This relatively large ground beetle might compete with or predate on other ground beetle species (Roubinet et al., 2018). P. melanarius is especially well-adapted to highly intensive agriculture (Turin, 2022; Turin, 2000), which explains why it was more abundant in monocultures previously (Ditzler et al., 2021). In turn, the reduced P. melanarius populations in strip cropping might allow other species to persist, such as Harpalus spp. and A. dorsalis. Therefore, strip cropping might facilitate ground beetle biodiversity by reducing the dominance of a common, competitive species.

Biodiversity gains in our study did in most cases not coincide with productivity losses (Supplementary file 5). Earlier studies on crop yield in the strip cropping fields in Lelystad and Wageningen during the years of pitfall trapping show a yield decrease when strip cropping cabbage (Carrillo-Reche et al., 2023) and wheat (Ditzler et al., 2023), whereas potato yield was unaffected by crop configuration (Ditzler et al., 2023). In Almere, bean and parsnip yields were higher in strip cropping, whereas oat and onion yields were lower (Juventia and van Apeldoorn, 2024). In Valthermond, crop productivity was similar for monocultures and strip cropping (Supplementary file 5). Therefore, biodiversity gains through strip cropping do not compromise agricultural production.

We show that changing crop configuration from monoculture to strip cropping, on average, enhances ground beetle richness by 15% and activity density by 30% within agricultural fields. These results show that strip cropping can lead to increases in biodiversity that approach those achieved by shifting from conventional to organic farming practices (+19% richness, Lichtenberg et al., 2017; +23% richness, Gong et al., 2022) and by other in-field diversification measures, like hedgerows and flower strips (+23% richness, Lichtenberg et al., 2017; +24% richness, Beillouin et al., 2021). While organic management or in-field diversification measures generally lead to lower productivity (Gong et al., 2022), strip cropping can maintain crop productivity without taking land out of production (Campanelli et al., 2023; Juventia and van Apeldoorn, 2024; van Oort et al., 2020), although occasional yield reductions have been reported (Carrillo-Reche et al., 2023; Ditzler et al., 2023). The biodiversity gain in our study was achieved without additional crop-level diversification strategies designed for biodiversity conservation, such as cover cropping or (flowering) companion plants. This increase in biodiversity stacks on top of already higher biodiversity achieved through organic management (Gong et al., 2022; Lichtenberg et al., 2017). Furthermore, the estimated richness effects of strip cropping, that is, no change at crop level and 15% increase at field level, are conservative because they only consider the pairwise comparison of biodiversity of two or three crops in a strip cropping configuration to monocultures, whereas the inclusion of more crops within strip cropping could further enhance ground beetle biodiversity. Also, the effect of strip cropping on ground beetle biodiversity might be more pronounced in conventional agriculture and at larger field sizes, as the relatively small-scale, organic fields that we studied might have already had a relatively high ground beetle biodiversity (Tscharntke et al., 2021). The ground beetle communities in our study were dominated by farmland and eurytopic species and contained only two rare species (Turin, 2022). Future research could test whether the inclusion of other in-field diversification measures within strip-cropped fields, such as the establishment of perennial semi-natural habitats (SNHs) (Rischen et al., 2021; Sirami et al., 2019) or uptake at larger spatial extents (Tscharntke et al., 2021) would allow ground beetles with other habitat preferences to establish in agricultural fields.

Data and scripts are publicly available from 4TU.ResearchData.

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Author details

1. Luuk Croijmans

   1. Laboratory of Entomology, Wageningen University and Research, Wageningen, Netherlands
   2. Farming Systems Ecology, Wageningen University and Research, Wageningen, Netherlands

   Contribution

   Conceptualization, Data curation, Formal analysis, Supervision, Investigation, Visualization, Methodology, Writing – original draft, Writing – review and editing

   Contributed equally with

   Fogelina Cuperus

   For correspondence

   luuk.croijmans@wur.nl

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0001-7690-8988

2. Fogelina Cuperus

   1. Farming Systems Ecology, Wageningen University and Research, Wageningen, Netherlands
   2. Field Crops, Wageningen University and Research, Lelystad, Netherlands

   Contribution

   Conceptualization, Data curation, Formal analysis, Supervision, Investigation, Visualization, Methodology, Writing – original draft, Writing – review and editing

   Contributed equally with

   Luuk Croijmans

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0003-0616-7920

3. Dirk F van Apeldoorn

   1. Farming Systems Ecology, Wageningen University and Research, Wageningen, Netherlands
   2. Field Crops, Wageningen University and Research, Lelystad, Netherlands

   Contribution

   Conceptualization, Supervision, Funding acquisition, Visualization, Project administration, Writing – review and editing

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0003-0636-1977

4. Felix JJA Bianchi

   Farming Systems Ecology, Wageningen University and Research, Wageningen, Netherlands

   Contribution

   Conceptualization, Writing – original draft, Writing – review and editing

   For correspondence

   felix.bianchi@wur.nl

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0001-5947-9405

5. Walter AH Rossing

   Farming Systems Ecology, Wageningen University and Research, Wageningen, Netherlands

   Contribution

   Conceptualization, Funding acquisition, Writing – original draft, Project administration, Writing – review and editing

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0003-2294-2368

6. Erik H Poelman

   Laboratory of Entomology, Wageningen University and Research, Wageningen, Netherlands

   Contribution

   Conceptualization, Supervision, Funding acquisition, Methodology, Writing – original draft, Project administration, Writing – review and editing

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0003-3285-613X

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