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

Effect of crop configuration (monoculture versus strip cropping) on ground beetle biodiversity.
(a) Location of experimental sites in the Netherlands. (b) Field set-up of the two crop configurations: monoculture and strip cropping. At Lelystad and Wageningen, strip cropping consisted of 3-m-wide crop strips of two crops (pairs), and multiple crop pairs were assessed. At Almere and Valthermond, strip cropping consisted of 6-m-wide crop strips of eight crops combined (Figure 1—figure supplements 3–14). (c) Sample-based species accumulation curves of all year series from monocultures (brown) and strip cropping (green), in Almere from 2021 and 2022. This was the only location where an equal number of samples were taken in the monocultures and in strip cropping on a similar area. Ground beetle species include Poecilus cupreus (left) and Pterostichus melanarius (right). (d, e) Overall relative change in field-level ground beetle (d) taxonomic richness and (e) activity density. Positive values indicate higher richness or activity density in strip cropping, negative values in monocultures. (f–j) Overall relative effect of crop configuration on ground beetle (f) taxonomic richness, (g) activity density, (h) absolute evenness, (i) inverse Simpson index, and (j) Shannon entropy. (k) Effect of crop configuration on ground beetle taxonomic richness for each combination of location, year, and crop. Barley mixture consists of a mixture of barley-bean (2020) or barley-pea (2021). Squares indicate estimated means, the bar indicates the 95% confidence interval, both based on generalized linear mixed models. Asterisks indicate significant differences among the crop configurations. Empty panels indicate combinations of years and locations that were not sampled. When no estimated mean and confidence interval are shown, crops were not grown or sampled in that year. Open circles indicate individual year series to visualize sample size (Supplementary file 6).
© 2011, Ortwin Bleich. Figure 1 panel c ground beetle images were reprinted from http://www.eurocarabidae.de/ with permission from Ortwin Bleich. It is not covered by the CC-BY 4.0 license and further reproduction of this panel would need permission from the copyright holder.

Effect of crop configuration on ground beetle activity density and absolute evenness.
Effect of crop configuration on activity density (a) and absolute evenness (d) per combination of location (columns), year (rows), and crop (x-axis); and the effect of crop configuration on activity density (b) and absolute evenness (c) in the case study in Wageningen, consisting of two early spring sampling rounds in 2022. Activity density here is the total number of ground beetles captured in year series, which might differ in sampling frequency (Supplementary file 6). Absolute evenness is calculated from the total number of ground beetles captured in year series. Empty panels indicate combinations of years and locations that were not sampled. Squares indicate estimated means, the bar indicates the 95% confidence interval, both based on generalized linear mixed models. When no estimated mean and confidence interval are shown, then those crops were not grown or sampled in that year. Asterisks indicate significant differences among the crop configurations (α=0.05). Open circles indicate individual year series (Supplementary file 6).

Effect of crop configuration on ground beetle inverse Simpson index and Shannon entropy.
Effect of crop configuration on inverse Simpson index (a) and Shannon entropy (d) per combination of location (columns), year (rows), and crop (x-axis); and the effect of crop configuration on inverse Simpson index (b) and Shannon entropy (c) in the case study in Wageningen, consisting of two early spring sampling rounds in 2022. Inverse Simpson index and Shannon entropy were calculated from the total number of ground beetles captured in year series. Empty panels indicate combinations of years and locations that were not sampled. Squares indicate estimated means, the bar indicates the 95% confidence interval, both based on generalized linear mixed models. When no estimated mean and confidence interval are shown, then those crops were not grown or sampled in that year. Asterisks indicate significant differences among the crop configurations (α=0.05). Open circles indicate individual year series (Supplementary file 6).

Field map of experimental lay-out at Almere in 2020.
Colors indicate distinct crops. Brighter colors with black lining around the field indicate areas in which samples included in this study were taken.

Field map of experimental lay-out at Almere in 2021.
Colors indicate distinct crops. Brighter colors with black lining around the field indicate areas in which samples included in this study were taken.

Field map of experimental lay-out at Almere in 2022.
Colors indicate distinct crops. Brighter colors with black lining around the field indicate areas in which samples included in this study were taken.

Field map of experimental lay-out at Lelystad in 2019.
Colors indicate distinct crops. Brighter colors with black lining around the field indicate areas in which samples included in this study were taken.

Field map of experimental lay-out at Lelystad in 2020.
Colors indicate distinct crops. Brighter colors with black lining around the field indicate areas in which samples included in this study were taken.

Field map of experimental lay-out at Lelystad in 2021.
Colors indicate distinct crops. Brighter colors with black lining around the field indicate areas in which samples included in this study were taken.

Field map of experimental lay-out at Valthermond in 2020.
Colors indicate distinct crops, whereas dotted patterns indicate a crop mixture. Brighter colors with black lining around the field indicate areas in which samples included in this study were taken.

Field map of experimental lay-out at Valthermond in 2021.
Colors indicate distinct crops, whereas dotted patterns indicate a crop mixture. Brighter colors with black lining around the field indicate areas in which samples included in this study were taken.

Field map of experimental lay-out at Wageningen in 2019.
Colors indicate distinct crops. Brighter colors with black lining around the field indicate areas in which samples included in this study were taken.

Field map of experimental lay-out at Wageningen in 2020.
Colors indicate distinct crops. Brighter colors with black lining around the field indicate areas in which samples included in this study were taken.

Field map of experimental lay-out at Wageningen in 2021.
Colors indicate distinct crops. Brighter colors with black lining around the field indicate areas in which samples included in this study were taken.

Field map of experimental lay-out at Wageningen in 2022.
Colors indicate distinct crops. Brighter colors with black lining around the field indicate areas in which samples included in this study were taken.

Ground beetle species associated with crop configuration (monoculture [a] versus strip cropping [b]).
Results obtained by generalized linear mixed models on the twelve most common genera of ground beetles with the four locations analyzed separately (Figure 2—source data 1). Only those genera are given for which cropping system significantly influences activity density within at least one location (α=0.05). Locations where the genus had a higher activity density in one of the cropping systems are indicated between brackets (Al = Almere; Le = Lelystad; Va = Valthermond; Wa = Wageningen). For Pterostichus, data from Almere in 2020 was analyzed separately, as models did not fit elsewise. The asterisk ‘*’ next to Pterostichus indicates that only in 2020 the difference between monoculture and strip cropping was significant.
© 2011, Ortwin Bleich. Figure 2 panel b ground beetle images were reprinted from http://www.eurocarabidae.de/ with permission from Ortwin Bleich. It is not covered by the CC-BY 4.0 license and further reproduction of this panel would need permission from the copyright holder.
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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.
- https://cdn.elifesciences.org/articles/104762/elife-104762-fig2-data1-v1.docx

The effect of crop configuration on ground beetle community composition using redundancy analysis (RDA).
Data from all datasets that included species-level data (Almere [○], Lelystad [▽], Wageningen [▲]) are used for visualizing the first RDA and PC axis. Each axis shows the percentage explained variation. Color of the dots indicates crop configuration (brown = monoculture, green = strip cropping), and the direction of each crop configuration on the RDA axis (x-axis) is mentioned. Red arrows indicate species placement and name codes are given close to the tip of the arrow, meaning of the name codes can be found in .

The effect of crop configuration on ground beetle community composition in Almere, using redundancy analysis (RDA).
The first RDA and PC axis are visualized. Each axis shows the percentage explained variation. Shape of the dots indicates year (▽=2021, ▲=2022), color of the dots indicates crop configuration (brown = monoculture, green = strip cropping), and the direction of each crop configuration on the RDA axis (x-axis) is mentioned. Red arrows indicate species placement and name codes are given close to the tip of the arrow, meaning of the name codes can be found in Supplementary file 1.

The effect of crop configuration on ground beetle community composition in Lelystad, using redundancy analysis (RDA).
The first RDA and PC axis are visualized. Each axis shows the percentage explained variation. Shape of the dots indicates year (○=2019, ●=2020, ▽=2021), color of the dots indicates crop configuration (brown = monoculture, green = strip cropping), and the direction of each crop configuration on the RDA axis (x-axis) is mentioned. Red arrows indicate species placement and name codes are given close to the tip of the arrow, meaning of the name codes can be found in Supplementary file 1.

The effect of crop configuration on ground beetle community composition in Wageningen, using redundancy analysis (RDA).
The first RDA and PC axis are visualized. Each axis shows the percentage explained variation. Shape of the dots indicates year (○=2019, ●=2020, ▽=2021, ▲=2022), color of the dots indicates crop configuration (brown = monoculture, green = strip cropping), and the direction of each crop configuration on the RDA axis (x-axis) is mentioned. Red arrows indicate species placement and name codes are given close to the tip of the arrow, meaning of the name codes can be found in Supplementary file 1.

The effect of crop configuration on ground beetle community composition, using non-metric multidimensional scaling (NMDS).
Data from all datasets that included species-level data (Almere [○], Lelystad [▽], Wageningen [▲]) are used for visualizing the two NMDS axes. Color of the dots indicates crop configuration (brown = monoculture, green = strip cropping), and the centroids of each crop configuration are indicated with a large asterisk (*).

The effect of crop configuration on ground beetle community composition in Almere, using non-metric multidimensional scaling (NMDS).
The two NMDS axes are visualized. Shape of the points indicates year (▽=2021, ▲=2022), color of the dots indicates crop configuration (brown = monoculture, green = strip cropping), and the centroids of each crop configuration are indicated with a large asterisk (*).

The effect of crop configuration on ground beetle community composition in Lelystad, using non-metric multidimensional scaling (NMDS).
The two NMDS axes are visualized. Shape of the points indicates year (○=2019, ●=2020, ▽=2021), color of the dots indicates crop configuration (brown = monoculture, green = strip cropping), and the centroids of each crop configuration are indicated with a large asterisk (*).

The effect of crop configuration on ground beetle community composition in Wageningen, using non-metric multidimensional scaling (NMDS).
The two NMDS axes are visualized. Shape of the points indicates year (○=2019, ●=2020, ▽=2021, ▲=2022), color of the dots indicates crop configuration (brown = monoculture, green = strip cropping), and the centroids of each crop configuration are indicated with a large asterisk (*).

Effect of crop configuration and crop on ground beetle community composition in Wageningen.
Here, we use data from Wageningen in 2021 (left column; a, c, e) and 2022 (right column; b, d, f) from three fields including three crop pairs being cabbage and oat (a, b), barley and pumpkin (c, d), and grass and potato (e, f). The first and second RDA axis are visualized. Each axis shows the percentage explained variation. Color of the dots indicates crop configuration (brown = monoculture, green = strip cropping), shapes indicate crop (□=barley, ■=pumpkin, ○=grass, ●=potato, △=oat, ▲=cabbage). The red lines and abbreviated names indicate how specific ground beetles species correlate with the RDA axes, species names are given in Supplementary file 1.
Tables
Effect of crop configuration on ground beetle community composition.
Results from permanova analysis using Hellinger’s transformation for data from the three locations with species-level data (see Supplementary file 3 for analyses per location). Crops were a nested variable within years, as these differed among years. Years were nested in locations as the years that were studied differed among locations. Bold indicates significant effects (α=0.05).
Predictor | Df | Sum sq | R2 | F | p |
---|---|---|---|---|---|
Crop configuration | 1 | 1.68 | 0.01 | 7.32 | 0.001 |
Location | 2 | 47.7 | 0.29 | 103.8 | 0.001 |
Location: Year | 6 | 10.7 | 0.06 | 7.72 | 0.001 |
Location: Year: Crop species | 31 | 33.3 | 0.20 | 4.67 | 0.001 |
Crop configuration: Location | 2 | 0.63 | 0.00 | 1.37 | 0.149 |
Crop configuration: Location: Year | 6 | 2.31 | 0.01 | 1.67 | 0.012 |
Crop configuration: Location: Year: Crop species | 31 | 14.8 | 0.09 | 2.08 | 0.001 |
Residual | 245 | 56.3 | 0.34 | ||
Total | 324 | 167.4 | 1.00 |
Additional files
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Supplementary file 1
All ground beetle species found among the four locations, and their species codes as used in several figure supplements.
- https://cdn.elifesciences.org/articles/104762/elife-104762-supp1-v1.docx
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Supplementary file 2
Total number of ground beetles caught per species (or genus), per location.
The number of year series is given per location in brackets. For some locations, ground beetles were identified up to genus level, these are underlined. ‘N/A’ indicates that this taxa was identified to a different taxonomic level for the specific location. Locations are indicated with abbreviations (Al=Almere; Le = Lelystad; Va = Valthermond; Wa = Wageningen). Rarity indicates the rarity of the species according to waarneming.nl (1 = common, 2 = relatively common, 3 = rare, 4 = very rare). Affinity indicates the habitat affinity group as by table A.1 of Turin et al. (2022), a * here indicates that species are more eurytopic.
- https://cdn.elifesciences.org/articles/104762/elife-104762-supp2-v1.docx
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Supplementary file 3
Effect of crop configuration on ground beetle community composition.
Results from permanova analyses using Hellinger’s transformation for data from the three locations with species-level data. ‘Crop species’ is a nested variable within years, as these differed among years. Years were nested in locations, as the years that were studied differed among locations. p-values in bold indicate significant effects (α = 0.05).
- https://cdn.elifesciences.org/articles/104762/elife-104762-supp3-v1.docx
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Supplementary file 4
Effect of crop configuration and crop species on ground beetle community composition.
Results from pairwise permanova analyses for crop pairs pumpkin-barley, cabbage-oat, and potato-grass in 2021 (a) and 2022 (b) in Wageningen. Values show F-values for the comparison between the crop configurations and crops in crossing rows and columns. Bold numbers indicate significant differences between combinations of crop configurations and crops (α = 0.05).
- https://cdn.elifesciences.org/articles/104762/elife-104762-supp4-v1.docx
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Supplementary file 5
Effect of crop configuration on crop yield.
Yield results were retrieved from published and unpublished studies on effects of strip cropping on crop yield in similar locations, years, and crops as this study. Mean crop yield is presented in ton per hectare (t/ha). When known, standard deviations of mean crop yield are given (± SD). When a crop is indicated with NC (not collected), the crop yield was not collected due to an inconsistent sampling method (potato, 2020, Almere), crop failure (broccoli, 2020, Almere; celeriac, 2021, Almere), unavailable machine-harvest data (grass, Almere, 2020, 2021, 2022), and undocumented reasons (barley/beans, 2020, Valthermond). Unavailable data include cabbage (2019) and potato (2020) in Lelystad; and pumpkin (2020, 2021, 2022), barley (2020, 2021, 2022), oat (2021, 2022), potato (2021, 2022), grass (2021, 2022), and cabbage (2022) in Wageningen.
- https://cdn.elifesciences.org/articles/104762/elife-104762-supp5-v1.docx
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Supplementary file 6
The total number of pitfall traps placed per location, year, and crop.
Rounds indicate the number of times pitfall traps were placed and were pooled within year series.
- https://cdn.elifesciences.org/articles/104762/elife-104762-supp6-v1.docx
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Supplementary file 7
Plant species composition of flower strips adjacent to the strip cropping fields in Lelystad and Valthermond.
- https://cdn.elifesciences.org/articles/104762/elife-104762-supp7-v1.docx
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MDAR checklist
- https://cdn.elifesciences.org/articles/104762/elife-104762-mdarchecklist1-v1.docx