Cover crop mixture diversity, biomass productivity, weed suppression, and stability
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Abstract
The diversity-productivity, diversity-invasibility, and diversity-stability hypotheses propose that increasing species diversity should lead, respectively, to increased average biomass productivity, invasion resistance, and stability. We tested these three hypotheses in the context of cover crop mixtures, evaluating the effects of increasing cover crop mixture diversity on aboveground biomass, weed suppression, and biomass stability. Twenty to forty cover crop treatments were replicated three or four times at eleven sites using eighteen species representing three cover crop species each from six pre-defined functional groups: cool-season grasses, cool-season legumes, cool-season brassicas, warm-season grasses, warm-season legumes, and warm-season broadleaves. Each species was seeded as a pure stand, and the most diverse treatment contained all eighteen species. Remaining treatments included treatments representing intermediate levels of cover crop species and functional richness and a no cover crop control. Cover crop seeding dates ranged from late July to late September with both cover crop and weed aboveground biomass being sampled prior to winterkill. Stability was assessed by evaluating the variability in cover crop biomass for each treatment across plots within each site. While increasing cover crop mixture diversity was associated with increased average aboveground biomass, we assert that this was the result of the average biomass of the pure stands being drawn down by low biomass species rather than due to niche complementarity or increased resource use efficiency. At no site did the highest biomass mixture produce more than the highest biomass pure stand. Furthermore, while increases in cover crop mixture diversity were correlated with increases in weed suppression and biomass stability, we argue that this was largely the result of diversity co-varying with aboveground biomass, and that differences in aboveground biomass rather than differences in diversity drove the differences observed in weed suppression and stability.
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Numerous studies have failed to show any benefit of species diversification in cover crops to the average performance of a range of criteria, or even on the stability of that performance.
“Numerous studies have failed to show any benefit of species diversification in cover crops to the average performance of a range of criteria, or even on the stability of that performance (Florence et al., 2019;Florence and McGuire, 2020;Smith et al., 2020).”
From Performance of innovative cropping systems diversified with oilseeds and protein crops: identification and resolution of methodological issues, using the Syppre experimental network as a case study (2024) · cites this paper as Florence et al., 2019
The Conclusions report little evidence of a diversity benefit for weed suppression or biomass stability after controlling for biomass, and no advantage over a productive single species. But they also report that increasing diversity was associated with increased average cover crop biomass productivity. The claim's unqualified statement about no benefit to average performance therefore overstates what this paper found. (LLM, conf=0.88)
Experimental — accuracy not yet measuredThe most diverse cover crop mixtures (more than 5 to 10 species) were never the best performing.
“These studies showed that the most diverse mixtures (more than 5 to 10 species) were never the best performing.”
From Performance of innovative cropping systems diversified with oilseeds and protein crops: identification and resolution of methodological issues, using the Syppre experimental network as a case study (2024) · cites this paper as Florence et al., 2019
The Discussion (Diversity-stability hypothesis; page unavailable) describes cover-crop studies in which the most diverse mixture was neither the most productive nor the most stable. The paper’s Conclusions likewise find no benefit from mixing over a productive single species. Neither passage establishes that mixtures with more than 5 to 10 species were *never* the best performing across the studies invoked by the citing sentence. (LLM, conf=0.85)
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Gramig, G., Pull, Z. A., & Carr, P. M. (2023) · Farming System · doi:10.1016/j.farsys.2023.100070 · Cited by 7 · see citing works
Impacts of Smooth Pigweed (Amaranthus hybridus) on Cover Crops in Southern Ontario
VanVolkenburg, H., Guinel, F. C., & Vasseur, L. (2020) · Agronomy · doi:10.3390/agronomy10040529 · Cited by 7 · see citing works
Breeding for cold tolerance in common annual legume cover crops
Thapa, R. K., Hanson, S. J., Hua, J., & Moore, V. M. (2025) · Crop Science · doi:10.1002/csc2.70059 · Cited by 6 · see citing works
Tillage and cover crop mixtures interseeded in maize–soybean in the upper Midwest
Rabin, K. C., Johnson, G. A., Strock, J. S., Jordan, N. R., & y García, A. G. (2023) · Agronomy Journal · doi:10.1002/agj2.21322 · Cited by 6 · see citing works
Assessing the Efficacy of Living and Dead Cover Crop Mixtures for Weed Suppression in Sweet Corn
Yurchak, V. L., Leslie, A. W., & Hooks, C. R. R. (2023) · Agronomy · doi:10.3390/agronomy13030688 · Cited by 6 · see citing works
Improving semi-arid agroecosystem services with cover crop mixes
Moore, E. A., & Norton, U. (2024) · PLoS ONE · doi:10.1371/journal.pone.0306567 · Cited by 5 · see citing works
A critical analysis on multifaceted benefits of mixture of cover crops over pure stand
Zupanič, M., & Kramberger, B. (2022) · Symbiosis · doi:10.1007/s13199-022-00888-3 · Cited by 5 · see citing works
Carbon accounting per unit of food and unit of land in food production systems of Argentina
Viglizzo, E. F., & Ricard, M. F. (2023) · Universal Journal of Carbon Research · doi:10.37256/ujcr.1220232202 · Cited by 5 · see citing works
Winter cover crops and dynamics of weeds in agricultural systems of the Argentine Rolling Pampas
Buratovich, M. V., & Acciaresi, H. A. (2022) · International Journal of Pest Management · doi:10.1080/09670874.2022.2118893 · Cited by 5 · see citing works
Longis, S., Cadoux, S., Toupet de Cordoue, A.-L., Tauvel, P., Estienne, M., Onzon, P., Lescourret, F., Rouillon, C., & Aubertot, J.-N. (2024) · OCL · doi:10.1051/ocl/2023022 · Cited by 4 · see citing works
“Numerous studies have failed to show any bene fi t of species diversi fi cation in cover crops to the average performance of a range of criteria, or even on the stability of that performance (Florence et al. , 2019; Florence and McGuire, 2020; Smith et al. , 2020).”
BackgroundGagliardi, L., Tramacere, L. G., Antichi, D., Frasconi, C., Sbrana, M., Sileoni, G., Monacci, E., Pagano, L., Darra, N., Kriezi, O., et al. (2026) · Agronomy · doi:10.3390/agronomy16020220 · Cited by 4 · see citing works
Dawson, E. K., Boyhan, G. E., Coolong, T. W., Basinger, N. T., & McNeill, R. (2021) · HortTechnology · doi:10.21273/horttech04870-21 · Cited by 4 · see citing works
Clever Cover Cropping : Litter trait diversities and elemental flows
Porre, R. J. (2020) · doi:10.18174/531407 · Cited by 5 · see citing works
Zhao, X., Sun, Y., Li, Y., Jiao, Y., Xing, M., Shi, J., Yang, G., Yang, C., Tang, W., Xu, Y., et al. (2025) · Food and Energy Security · doi:10.1002/fes3.70112 · Cited by 3 · see citing works
Groß, J., Kümmerer, R., Heuermann, D., Gentsch, N., Schweneker, D., Feuerstein, U., Guggenberger, G., Nicolaus Von Wirén, A., & Bauer, B. L. (2024) · Frontiers in Agronomy · doi:10.3389/fagro.2024.1416379 · Cited by 3 · see citing works
The Ecological Base of Nonchemical Weed Control
Nosratti, I., & Chauhan, B. S. (2023) · Wiley eBooks · doi:10.1002/9781119709763.ch4 · Cited by 3 · see citing works
Mechanisms of Weed Suppression by Cover Crops, Intercrops, and Mulches
Smith, R. G., Lounsbury, N. P., & Palmer, S. A. (2023) · doi:10.1002/9781119709763.ch10 · Cited by 3 · see citing works
Kümmerer, R., Heuermann, D., Laidig, F., Piepho, H., & Bauer, B. L. (2025) · Field Crops Research · doi:10.1016/j.fcr.2025.110014 · Cited by 2 · see citing works
Can nitrogen fertilization of cover crops promote N uptake and reduce N leaching?
Porre, R. J., Elhakeem, A., van der Werf, W., Deyn, G. B. D., van Dam, J. C., Bastiaans, L., & Hoffland, E. (2025) · Nutrient Cycling in Agroecosystems · doi:10.1007/s10705-025-10457-x · Cited by 2 · see citing works
Zupanič, M., Žnidaršič, T., Podvršnik, M., Sem, V., Kristan, B., Rihter, L., & Kramberger, B. (2025) · Plants · doi:10.3390/plants14050726 · Cited by 2 · see citing works
Soil health responses to cover crop functional group and richness in semiarid Montana
Jones, C. A., Zabinski, C. A., Miller, P. R., D'Agati, K., Tallman, S. M., & Housman, M. L. (2025) · Soil Science Society of America Journal · doi:10.1002/saj2.70045 · Cited by 2 · see citing works
Citizen science reveals opportunities for improving sustainability outcomes of cover crops
Sutton, E. A., & Blesh, J. (2025) · Journal of Soil and Water Conservation · doi:10.1080/00224561.2025.2582438 · Cited by 2 · see citing works
Płaza, A., & Górski, R. (2024) · Australian Journal of Agricultural Research · doi:10.1071/cp23153 · Cited by 2 · see citing works
On productivity, resource capture, and yield stability of cover crop species mixtures
Elhakeem, A. (2021) · doi:10.18174/540613 · Cited by 2 · see citing works
Drainage water management practices affect water quality, soil properties, and crop production
Kaur, H. (2023) · doi:10.32469/10355/96988 · Cited by 2 · see citing works
Wojciechowski, A., Seassau, C., Alletto, L., & Lamichhane, J. R. (2025) · Field Crops Research · doi:10.1016/j.fcr.2025.110165 · Cited by 1 · see citing works
Do Cover Crops Reduce Downside Production Risk?
Aglasan, S., Rejesus, R. M., Bowman, M., & Goodwin, B. K. (2025) · Agricultural Economics · doi:10.1111/agec.70082 · Cited by 1 · see citing works
Summer legumes following winter wheat boost soil nitrogen availability and corn yield
Adhikari, M., Jasa, P. J., Birru, G., & Iqbal, J. (2026) · Agronomy Journal · doi:10.1002/agj2.70269 · Cited by 1 · see citing works
Zou, X., Zhang, H., Ren, J., Ou, H., Lin, Y., Shi, X., & Wang, J. (2026) · Plant and Soil · doi:10.1007/s11104-026-08845-5 · Cited by 1 · see citing works
Panyi, A. F., Maples, W. E., Tewolde, H., Hall, M. A., Shankle, M. W., Kingery, W. L., & Feng, G. G. (2026) · Agronomy Journal · doi:10.1002/agj2.70493 · Cited by 1 · see citing works
Yurchak, V. L., Leslie, A. W., & Hooks, C. R. R. (2023) · Weed Technology · doi:10.1017/wet.2023.82 · Cited by 1 · see citing works
Cover crops and fall residual herbicides for managing Italian ryegrass
de Sanctis, J. H. S., Cahoon, C. W., Everman, W. J., Gannon, T. W., Jennings, K. M., Taylor, Z. R., Dean, B. A., Forehand, J. C., & Lee, J. H. (2025) · Weed Technology · doi:10.1017/wet.2024.107 · Cited by 1 · see citing works
To Mix or Not to Mix: Performance and Management of Diverse Cover Crop Mixtures
Wolters, B. (2020) · VTechWorks (Virginia Tech) · Cited by 1 · see citing works
Ground Cover Vegetation Promotes Ecological Intensification of Pear ProductionPreprint
Ji, X., Wang, J., Dainese, M., Zhang, H., Chen, Y., Cavalieri, A., Jiang, J., & Wan, N. (2021) · Research Square · doi:10.21203/rs.3.rs-201470/v1
Górski, R., & Płaza, A. (2023) · Folia Pomeranae Universitatis Technologiae Stetinensis Agricultura Alimentaria Piscaria et Zootechnica · doi:10.21005/aapz2023.66.2.2
Kumari, A., Price, A. J., Li, S., Gamble, A. V., & Jacobson, A. L. (2025) · Frontiers in Agronomy · doi:10.3389/fagro.2024.1502864
The impacts of legume and non-legume cover crops on the performance of HLB-affected citrus trees
Brewer, M., Strauss, S. L., Kanissery, R. G., & Kadyampakeni, D. M. (2025) · Journal of Plant Nutrition · doi:10.1080/01904167.2025.2474031
Mersha, Z., Ibarra-Bautista, M. A., Birru, G., Bucciarelli, J., Githinji, L. J. M., Shiferaw, A. S., Ren, S., & Rutto, L. K. (2025) · Agronomy · doi:10.3390/agronomy15081893
Restoration Technique to Mitigate Plant Invasion
Odedra, K. N., Bhutiya, K. B., Kodiyatar, N., & Jadeja, B. A. (2025) · IGI Global eBooks · doi:10.4018/979-8-3693-8724-5.ch004
Growth and development of six cover crops in the Czech Republic
Kubíková, Z., Kintl, A., Sobotková, J., Huňády, I., Brtnický, M., Holátko, J., Kincl, D., Kabelka, D., Smutný, V., & Elbl, J. (2025) · Acta Agriculturae Scandinavica Section B - Soil & Plant Science · doi:10.1080/09064710.2025.2586838
The decomposition and nutrient release dynamics of mixed cover crops in a no‐till row crop rotation
Iamjud, K., Fultz, L. M., Bridges, K., Negrete, P. M., & Carrillo, P. A. (2025) · Agronomy Journal · doi:10.1002/agj2.70262
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Summer cover crops impact weed biomass and C and N acquisition
Larney, F. J., Catton, H. A., Geddes, C. M., Lupwayi, N. Z., Forge, T. A., Chau, H. W., Helgason, B. L., & Iheshiulo, E. M. (2026) · Agronomy Journal · doi:10.1002/agj2.70286
Integrating almond crop residues for whole orchard regenerative managementPreprint
Cho, S. L., Andrews, E. M., Gaudin, A. C. M., Brown, P. H., & Khalsa, S. D. S. (2026) · SSRN Electronic Journal · doi:10.2139/ssrn.6331700
Omokanye, A. T., Bhattarai, S., Hernandez‐Ramirez, G., Azuhnwi, B. N., Zavala, Y. J., Gyamfi, H., & Shrestha, B. M. (2026) · Agrosystems Geosciences & Environment · doi:10.1002/agg2.70325
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Species complementarity sustains productivity and multifunctionality in tropical cover crop mixtures
Baptistella, J. L. C., de Oliveira, S. M., Cherubin, M. R., & Mazzafera, P. (2026) · Rhizosphere · doi:10.1016/j.rhisph.2026.101351
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Impact of simulated drought and cover crop mixtures on subsequent spring wheat yield
Halvorson, J. J., Hendrickson, J., Archer, D. W., Clemensen, A. K., Carrlson, A. J., & Johnson, H. A. (2026) · Agrosystems Geosciences & Environment · doi:10.1002/agg2.70397
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Genotype by environment interaction and local adaptation of commercial legume cover crop genotypes
Thapa, R., Douglas, J., Englert, J., Dial, H., Robbins, K., & Moore, V. M. (2026) · Frontiers in Plant Science · doi:10.3389/fpls.2026.1903778
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Cover crop composition, not diversity, drives agroecosystem multifunctionality and wheat yield
Sone, B. M., Lynch, L., Osburn, E. D., Lucas, J. M., & Strickland, M. S. (2026) · Journal of Applied Ecology · doi:10.1111/1365-2664.70576
- Authors
- Florence, A. M., Higley, L. G., Drijber, R. A., Francis, C. A., & Lindquist, J. L.
- Affiliations
- University of Nebraska–Lincoln (US)
- Corresponding author
- Lindquist, J. L.
- Venue
- PLOS ONE
- Publisher
- Public Library of Science (PLoS)
- Published
- 2019-03-14
- Volume
- 14
- Issue
- 3
- Pages
- e0206195
- ISSN
- 1932-6203 (ISSN-L)
- PMC
- PMC6417710
- OpenAlex
- W2944746124
- Cited by
- 99
- References
- 47
- Citations by year
- 2026: 14 · 2025: 20 · 2024: 10 · 2023: 17 · 2022: 9 · 2021: 20 · 2020: 8 · 2019: 1
- FWCI
- 13.35
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Licence
- Locations
- PLoS ONE (publishedVersion, cc-by)
- bioRxiv (Cold Spring Harbor Laboratory) (acceptedVersion, cc-by)
- Europe PMC (PubMed Central) (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- Lincoln (University of Nebraska) (submittedVersion, cc-by)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
- Figshare (submittedVersion, cc-by)
- Funders
- National Science Foundation (1041000)
- North Central SARE
- Topics
- Life Sciences › Agricultural and Biological Sciences › Agronomy and Crop Science › Agronomic Practices and Intercropping Systems
- Physical Sciences › Environmental Science › Nature and Landscape Conservation › Ecology and Vegetation Dynamics Studies
- Life Sciences › Agricultural and Biological Sciences › Plant Science › Weed Control and Herbicide Applications
- Keywords
- cover crop mixtures, species diversity, weed suppression, aboveground biomass, biomass stability, functional groups, pure stands, niche complementarity, resource efficiency, winterkill
- MeSH terms
- Agriculture, Seasons, Seeds, Ecosystem, Biomass, Crops, Agricultural, Biodiversity, Weed Control
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